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THE 


TRANSACTIONS 


OF 


THE LINNEAN SOCIETY OF LONDON. 


SECOND SERIES—VOLUME XVII. 
ZOOLOGY. 


THE PERCY SLADEN TRUST EXPEDITION 


TO 
THE INDIAN OCEAN IN 1905, 
UNDER THE LEADERSHIP OF 
Mr J. STANLEY GARDINER, M.A. 
Vou. VI. 


LONDON: 
PRINTED BY J. B. PEACE, M.A., AT THE CAMBRIDGE UNIVERSITY PRESS. 
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE, PICCADILLY, W. 1 
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW, E.C. 4 


1914—1921. 


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MERLE P56 HR 


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REPORTS 


OF THE 


PERCY SLADEN TRUST EXPEDITION 


TO 


THE INDIAN OCEAN IN 1905, 


UNDER THE LEADERSHIP OF 


Mr J. STANLEY GARDINER, M.A. 


VOLUME THE SIXTH 


[BEING THE SEVENTEENTH VOLUME OF THE SECOND SERIES, ZOOLOGY, OF THE 
TRANSACTIONS OF THE LINNEAN SOCIETY OF LONDON.] 


LONDON: 


“PRINTED BY J. B. PEACE, M.A., AT THE CAMBRIDGE UNIVERSITY PRESS. 
_ SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON HOUSE, PICCADILLY, W. 1 
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW, E.C. 4. 


1914—1921. 


IL. 


II. 


LINE 


VI. 


\Aut, 


CONTENTS. 


PART J.—Dercemesr, 1914. 


A Rewision of the recent colonial Astreide possessing distinct Corallites. (Based 
on material from the Indo-Pacific Ocean and the collections of Paris, Berlin, 
Vienna, Copenhagen, London and Glasgow.) By Grorce Matrruat, B.A. (Research 
Student of Emmanuel College), Zoological Laboratory, Cambridge. (Communi- 
cated by Prof. J. Sranuey Garpiner, V.A., F.R.S., FL.S.) (Plates 1-38.) 
pages 1-140 
Coleoptera: Cucwjide, Cryptophagide. Par A. Grouvetitn. (Figures 1-6 dans 
le texte.) Avec une description de la larve et de la nymphe de Prostomama Con- 
veaiuscula Grouvelle [Cucwjide] par P. pp PrynrimHorr. (Figures A-F dans 
le texte.) (Communiqué par M. le Professeur J. Stantey Garpiner, M.A, 


JE LB Soy Molly, 9 ‘ : : 5 = 9 LAL=159 
Mallophaga, Aphaniptera, and Diptera Pupipara. By Huew Scorr, M.A., 
E.LS., FES. (Text figures 1-4.) : : : ; : Bai Uoy/ 


IE ele ViR CHONG 


Meduse from the Indian Ocean. (Collected by Prof. StraNLEY GARDINER, 7 
H.M.S.“Sealark” in 1905.) By Evwarp T. Browns, M.A,, F.L.S. (Plate 39.) 
169-210 


Report on the Hexactinellid Sponges (Triaxonida) collected by H.M.S. “Sealark” 
im the Indian Ocean. By Anraur Denpy, D.Sc., F.RS., F.L.S., Professor of 
Zoology in the University of London (King’s College). (Plates 40-43.) 211-224 
Report on the Homosclerophora and Astrotetraxonida collected by H.MLS. 
“ Sealark” in the Indian Ocean. By ArtHur Denvy, D.Sc., F.R.S., FLLS., Pro- 


fessor of Zoology in the University of London (King’s College). (Plates 44-48.) 


225-27 1 


PART III.—Septemper, 1917. 


Rhynchota. Part II: Suborder Homoptera. By Wm. Lucas Distant. (Com- 
municated by J. Stantey Garpiner, M.A., F.RS., F.L.S.) (With Plates 49-51.) 
i 273-322 


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VIII. On the Pontoniine. By L. A. Borrapatnn, VW.A. (Lecturer in Zoology in the 


IX. 


XI. 


University of Cambridge, Fellow, Dean, and Lecturer of Selwyn College). (Com- 
municated by Prof. J. Stanuey Garpiner, V.A., F.R.S., F.L.S.) (Plates 52-57.) 

323-396 
On Carides from the Western Indian Ocean. By L. A. Borrapaine, M.A. 
(Lecturer in Zoology in the University of Cambridge, Fellow, Dean, and Lecturer 
of Selwyn College). (Communicated by Prof. J. Stantey GarpDINER, MV.A., 
F.RS., F.L.S.) (Plates 58, 59.) : : : : : . 397-412 


PART IV.—January, 1921. 


Les Espéces d’ Alpheide rapportées de Vocéan indien par M. J. Stantey GARDINER. 
Par M. le Professeur H. Courthre. (Communiqué par M. le Professeur 
J. STANLEY GARDINER, W.A., F.R.S., F.L.S.) (Planches 60-64.) 413-428 
On the Cephalopoda obtained by the Percy Sladen Trust Expedition to the Indian 
Ocean in 1905. By G. C. Rogson, B.A. (Published by permission of the Trustees 
of the British Museum.) (Communicated by Prof: J. Staxtey Garviner, W.A., 
F.RS., PLS. (Plates 65, 66.) (Text-figures 1-6.) .. : : . 429-449 


Index . : : ‘ ; ; p : : 5 : : . 448-455 


No. [—A REVISION OF THE RECENT COLONIAL ASTRAIDA 
POSSESSING DISTINCT CORALLITES. 


(Based on Material from the Indo-Pacific Ocean and the Collections of Paris, Berlin, 
Vienna, Copenhagen, London and Glasgow.) 


By Grorce Marruat, B.A. 
(Research Student of Emmanuel College), Zoological Laboratory, Cambridge. 


(ComMUNICATED BY Pror. J. Stantey Garpriner, M.A., F.R.S., F.L.S.*) 


(Plates 1—88 ¢.) 


Read 18th June, 1914. 


CONTENTS. 
PAGE 
I. INTRODUCION . : : : : : : : : : : : : : : 2 
II. ANATOMY OF THE POLYPS. Methods—Body-layers—Mesoglea—Calicoblastic Layer of 
Ectoderm—Dissepiments and Polypal Growth—Nematocysts (Types I, II, I11)—Nervous 
System—Endodermal Musculature—Ectodermal Musculature—Oral Disec—Column Wall— 
Edge Zone and Ceenosare—Tentacles—Stomodzum—Mesenteries —Mesenterial Filaments— 
Acontia and Cinclides—Reproductive Organs—Zooxanthelle  . : : i : ; 6 
III. CLASSIFICATION. Gemmation and _Fissiparity—Astreide with Distinct Corallites— 
Summary of Characters—Keys to Genera (Polyps and Corallum) . : : 6 ; 33 


* [Mr Matthai wishes me to add in a footnote the part I have played in the preparation of this 
Report during the three years that he has worked in my Department. We originally intended to 
examine Cyphastrea in collaboration, and indeed we worked together on this genus for over six months, 
before I retired solely owing to pressure of other work. Since then I have constantly seen Mr Matthai 
and he has demonstrated to me on his sections and specimens most of his facts; we have discussed 
these facts but the deductions therefrom are entirely Mr Matthai’s, though with most I concur. The polyp 
sections were cut by the aid of my laboratory attendants, and will remain with me or be deposited 
in some place where they will be available for future research by serious students; sections of. coral 
were also cut professionally. Mr Matthai in the course of his research re-examined all my polyp material 
of Canopsammia, Flabellum, etc. From’ time to time I have been in communication with Professor 
G. C. Bourne, Dr Wayland Vaughan, Professor 8. J. Hickson and Mr Cyril Crossland. In particular 
Professor G. C. Bourne favoured me with a series of most valuable suggestions, as a result of which 
Mr Matthai spent three months in re-examining his polyp material. A further study of the cytology 
of the polyps, or some species of coral polyp, is requisite, but it should be undertaken with fresh 
material.—J. STANLEY GARDINER. | 

+ The Royal Society has generously given a grant of £70 from their Publication Fund towards the 
cost of the plates which accompany this paper. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 1 


WiLIBRARy|O 


= 


2 PERCY SLADEN TRUST EXPEDITION 


IV. SYSTEMATIC. 


PAGE 


Group I. 1. Cyphastrea . : : : : . ‘ : 6 6 ; : : 38 

2. Echinopora . ‘ f é : : 0 : : : 6 2 , 48 

3. Galaxea . 4 : : : j : ‘ é , ; i ‘ : 58 

4. Leptastrea . $ 6 : 0 6 : : . . ¢ . 9 66 

5. Dviploastrea, nov. . : : : é : : : 6 é : : 72 

6. Doubtful Genera . : P ; : ‘ : : : : ‘ ; 74. 

Group II. 7. fava . : : : : : : ; 3 ; : 2 ; ; 17 

8. Goniastrea . i : 3 : F : , ; : 5 3 ‘ 115 

9. Doubtful Genus . . j : : : : : ; ‘ F : 122 

V. LITERATURE OF POLYP ANATOMY . ; . é : F : 3 5 j 124 
VI. EXPLANATION OF PLATES 1—38 . : F : : : : . F : . 129 


EE INTRODUCTION: 


The morphology of the soft parts of fourteen species of Astra@id@ has hitherto been — 
investigated. Professor Bourne described Mussa corymbosa and EHuphyllia glabrescens in 
1888 (12), and Dr Fowler Galaxea esperi in 1890 (48, p. 410). Dr J. E. Duerden in 
1902, in a memoir on “ West Indian Madreporarian Polyps” (82), described some eleven 
species, viz., Astrangia solitaria Lesueur, Phyllangia americana Ed. and H., Cladocora 
arbuscula (Lesueur), Orbicella annularis (Ell. and Sol.), Solenastrea hyades (Dana), 
Favia fragum (Hsper.), Dichocenia stokesi Ed. and H., Isophyllia dipsacea Dana, 
Mancina areolata (Linn.), Colpophyllia gyrosa (Ell. and Sol.), Meandrina labyrinthica 
(Ell. and Sol.). He based a new classification of the Madreporaria on a study of 
altogether sixteen species from which, as far as the Astraide are concerned, I have 
entirely to disagree for reasons which will become apparent in the course of this paper. 

At the suggestion of Professor Stanley Gardiner, I undertook, in October 1911, a 
comparative study of the soft parts of Astrzeid Corals in his collections from the Indo- 
Pacific Ocean, mainly with a view to determining the natural relationships of the genera 
and species. In April 1912, Professor Gardiner received further preserved coral polyps 
from the Red Sea and Ceylon. From these collections I have attempted to study those 
Astreid genera in which the individual polyps remain separate, the meandering forms 
being reserved for subsequent examination. Serial sections of about one hundred and fifty 
polyps from seventy-five colonies were carefully examined, resulting in the determination 
of seven genera and twenty-seven species. The structure of the soft parts of none of 
these species had hitherto been described. 

A thorough examination of over 700 Astreeid specimens (possessing distinct corallites) 
in Cambridge™ left me in considerable doubt regarding the scientific value of the so-called 
generic and specific characters, on which systematists had based their schemes of classifica- 
tion. The characters usually regarded by them as of value were as follows: (1) the form 
of growth of the colony (flat or incrusting, massive, branching or irregular with hillocks 
and valleys) ; (2) the nature of the inter-calicinal peritheca (broad or narrow, grooved or 


* T have had the advantage of discussing these specimens with Prof. Gardiner, who had studied their 
growth-forms in relation to positions on the reefs. 


MATTHAI—RECENT COLONIAL ASTRAIDA 3 


flat, rough or smooth, vesicular, cellular or dense, the extent to which the spaces between 
the thecz were filled with calcareous deposit); (3) the flat or arched character of the 
dissepiments ; (4) the presence or absence of an epitheca; (5) the character of the theca 
(visible or sunk in the peritheca, its comparative thickness) ; (6) the number of septal 
cycles and of the septa constituting each cycle; (7) the nature of the septa (oblique or 
perpendicular, narrow or broad, exsert or not, presence or absence of teeth on their 
margins, sides rough or smooth) ; (8) the presence or absence of paliform lobes* and their 
shape, size and surface characters; (9) the nature of the coste (present or absent, visible 
or sunk in the peritheca, rough or smooth, thick or thin); (10) the nature of the 
columellaj, and the presence or absence of upright rods upon it. 

None of these characters were found to have any constant value, and therefore the 
distinctions based on them would appear to be arbitrary. Not only were there gradational 
stages connecting genera like Orbicella, Pavia and Prionastrea, and connecting species of 
the same genus, but in many single large specimens the skeletal characters varied to such 
an extent that pieces cut off from different parts, if regarded separately, had to be 
relegated to different species. The descriptions of many coral species have, in this manner, 
been taken from isolated specimens, hence the enormous multiplication of synonymy in the 
Madreporaria. 

Much attention has been paid by various authors to the epitheca. Its origin is still 
obscure. Von Koch, from his study of the development of Astroides calycularis (78), 
regarded it as the continuation of the basal plate, and, as is evident from his figure (fig. 4), 
a secretion by the ectoderm of the column-wall where it passes into the base or foot. It 
would therefore be similar in origin to von Heider’s eutheca (64). Bourne (12, pp. 36—39), 
on the other hand, defined the epitheca as formed “from the free edge of the soft tissues 
on the exterior of the corallum, as they retreat farther and farther from the original surface 
of attachment,” hence similar in origin to the perithecat deposited between the corallites. 
Indeed—and in this I am inclined to agree with him—he regards the exotheca, peritheca, 
ecenenchyme and epitheca as homologous structures, differences, if there be any, depending 
on quantity and texture. Lacaze Duthiers (60, p. 225) also was doubtful about the true 
nature of the epitheca and how to determine its presence or absence. With reference to 


* True pali, arising from the basal plate and situated between the columellz and septa, have not been 
seen in any of the Astrzids I have examined. 

+ From the sections of the coralla it is not possible to determine if true columelle, viz. rods arising 
from the basal plate and formed in folds of the basal dise of the polyps, are present in Astreids. The 
so-called “dark centres” are no test as they are seen in all the columellar parts, even in some undoubted 
septal trabecule which have united with and formed part of the columelle. 

{ I use this term as Professor Gardiner defined it in his paper on Canopsammia, viz., ‘‘That part of 
the corallum of colonial madreporaria which is deposited outside and subsequently to the theca.” Bourne 
(18, p. 217) in a subsequent paper modified the strong views he had at first held on the subject of the epitheca 
and reverted to von Koch’s original contention in these words: “‘...I1 may give my adherence to von Koch’s 
definition of epitheca, that it is a more or less conspicuous offset of the basal plate, which lies on the outer side 
of the body-wall, but no longer forms a part of the surface of attachment, and in the majority of corals has the 
form of an investment of the column (die Gestalt eines Kegelmantels). As such it is readily recognisable in 
the anthoblast of /ungia, and in some cases I have been able to detect traces of an analogous structure in the 
anthocyathus (fig. 16, ep.).” 


an) 


a 


é 
4 PERCY SLADEN TRUST EXPEDITION 


Flabellum, characterised by the absence of a “‘ Randplatte” or “edge-zone,” Bourne con- 
siders the apparent theca as “ really a basal structure which has grown upwards to form a 
calyx” and hence agrees with von Koch’s view that it is really the epitheca. As far as 
the colonial Astrzids are concerned the term epitheca may, for the present, be confined to 
the thin, usually foliated lamina seen at the edges of the corallum. 

A study of the arrangement of the mesenteries in the polyps has shown that it is 
highly misleading to regard the septal cycles, as seen in the dry corallites, as representing 
the true succession. In genera characterised by the hexameral arrangement of the mesen- 
teries as in my Group I, it often happens that the secondary septa in the entocceles of the 
secondary mesenteries meet the columella, thus simulating the true primary septa; in 
other cases the secondaries may be quite as narrow as, or even narrower than, the tertiary 
septa. The true order of succession of the septa can thus be ascertained only in relation 
to the mesenterial couples. No such arrangement of the septa into orders can be recognised 
in the genera belonging to my Group I, which have lost the hexameral disposition of the 
mesenteries ; in these some of the subsidiary septa may meet the columella and some of 
the principal septa may not; if a cyclical arrangement be recognised in the dry corallites, 
it is always a later formation, the numbers of the cycles and of the septa composing them 
being subject to so great variation that they are of little value for purposes of classi- 
fication. 

The above is part of a general indefiniteness in the arrangement of the skeleton. It 
is not surprising since the corallum lies entirely outside the external ectoderm, the part 
most open to the free play of the environmental conditions and most liable therefore to be 
modified by changes in these conditions. This variation is so considerable that a system 
of classification based on the corallum alone can scarcely be a sure guide to the actual 
relationships of the animals, for a proper knowledge of which it is essential to make 
a comparative study of the morphology of the polyps. Bourne, on the other hand, in 1888 
regarded any such attempt as futile: “every fresh form that is examined convinces me 
that the expectations formed of founding a new classification of the Madreporaria on the 
anatomy of the polyp are to meet with disappointment. There is singularly little variation 
in the forms hitherto examined. Hence I believe that a re-modelled classification must 
depend on a much more intimate study of the structure of the corallum than has hitherto 
been attempted.” (12, pp. 44—45.) 

As a result of my study of a limited group of the Astreeidze, it is clear that a thorough 
re-casting, not only of the species but also of the genera concerned, is necessary. I am 
further led to suggest that only by a comparative study of both the polyps and their 
skeletons will a scientific classification be evolved. In the allied group of Actiniz the 
importance of the internal characters of the polyps in classification has been fully 
recognised since R. Hertwig’s work on the “Challenger” Actinians. 

As an instance of the errors that may be made in basing a classification of corals 
solely on the hard parts, the case of Orbicella, as later extended by Gardiner, may be 
cited. This genus has to disappear in the light of the new relationships revealed by 
a study of the polyps of the species included in it. For O. minikoiensis Gard.—which 
has been found to be identical with Heliastrea heliopora, Ed. and H.—a new genus has 


MATTHAI—RECENT COLONIAL ASTRAIDA i) 


been created; the species of Leptastrea Ed. and H., which Gardiner brought under 
Orbicella, have again been transferred to the former genus, while all the remaining species 
have been assigned to Fava, Oken. In the same manner Heliastrea forskelana Ed. 
and H., and Orbicella mammillosa Klunz. have been found to be identical with 
Echinopora gemmacea Lam. LHchinopora hirsutissima Ed. and H. and Favia favus 
(Forsk.) may be mentioned as examples of species combining a wide range of skeletal 
variation with identity in polyp-structure. 

In order to settle the synonymy of the species it became imperative to examine the 
type specimens, as it was impossible from descriptions, often without adequate figures, to 
get any clear idea of their characters. In the summer of 1913 I visited the following 
museums: Muséum d'histoire naturelle, Paris; Museum fiir Natiirkunde, Berlin; Natur- 
historische Hofmuseum, Vienna; Universitets Zoologiske Museum, Copenhagen ; British 
Museum, London, and Hunterian Museum, University of Glasgow. I therein examined 
the existing types of Lamarck, Milne Edwards and Haime, Ehrenberg, Klunzinger, 
Forskal, Quelch, and Ellis and Solander ; unfortunately many of Forskal’s and of Ellis 
and Solander’s types have been lost. In the Berlin Museum I also examined Prof. Studer’s 
“Gazelle” types and some of Dr Ortmann’s examples from Ceylon and Dar-es-Salaam. 
Altogether my work led me to examine 590 specimens in these museums. For great 
assistance rendered in connection with this part of my work I wish to thank the following 
gentlemen: Dr Charles Gravier, Dr Lamy, Profs. Dr Brauer and Dr Weltner, Hofrat 
Dr Steindachner, Prof. Dr Jungerson, Dr Harmer, Prof. Graham Kerr and Dr Agar. 
Particularly I desire to thank Dr Gravier, Dr Weltner and Dr Mortensen for sending me 
photographs of some of the type specimens in their museums to be reproduced in this 
paper. I am indebted to the managers of the F. M. Balfour Fund, Cambridge, for financial 
aid in connection with these visits. 

I also had the great pleasure and advantage of discussing species in Madreporaria 
~with Dr Emil v. Marenzeller, whose beautiful work on corals is so well known. It would 
be an impertinence were I to speak of his great power of discrimination, particularly in his 
study of Favia savigny: Ed. and H. and F. okent Ed. and H., species with which I have 
had to deal. The collections he studied were extensive and he visited practically all the 
European museums which possess types. I differ from him in considering that the 
Madreporaria must be studied in their entirety, not on the skeleton alone. 

In settling the nomenclature of a known species, either the earliest recorded name 
may be taken, as was Milne Edwards and Haime’s aim in their great work on corals, or, 
as Dr Marenzeller held, selection may be made of the particular name which is accompanied 
by the best description and figures. In the latter case due credit would be paid to the 
author who had undertaken the most complete study of the species, but in a group hke 
the Madreporaria, in which the synonymy has been so enormously complicated with 
specific descriptions of varying merit, opinions would of course differ as to which was the 
best account ; hence a determination of this kind would be largely left to the fancy of the 
particular worker. I have adopted the method of employing the earliest names wherever 
possible, even though the descriptions be of the scantiest nature. This in Madreporarian 
corals can, in my opinion, be done in the case of a species only if its first recorded type 


6 PERCY SLADEN TRUST EXPEDITION 


specimen is available. Fortunately most of the earliest types of the species discussed in 
this paper are still existent in the various museums visited. Regarding the species of the 
American authors I have referred only to those whose characters could be made out either 
from descriptions or from figures. 

In the systematic part of this work the plan followed has been to base species and 
genera on the characters of the soft as well as of the hard parts by a comparative 
examination in every possible case of polyps from different colonies and of as many hard 
specimens as were available. Thus it has been possible in many cases to study the varia- 
tion that the corallum undergoes, the principal forms being shown in the photo-plates. 
Owing to lack of polyp-material, only the probable specific rank of some species can be 
indicated by recording their skeletal characters. Notes on the specimens examined in the 
various museums are also given under each species. 

Finally I would remark that there should be little cause for complaint in the task of 
cataloguing examples of known species in a museum. In my visits to museums I found 
that any species whose limits had once been settled by the study of both polyps and hard 
parts could be recognised later from the hard parts alone. Only in the case of undeter- 
mined species should it be necessary to examine the polyps. 


Il. ANATOMY OF THE POLYPS (Plates 1—10). 


Method. The material had been in the first place fixed in a saturated solution of 
corrosive sublimate and preserved in 90°/, pure alcohol; some were killed first in formic 
aldehyde poured into seawater. From each specimen two pieces were chipped off, one to 
be decalcified for the soft parts and the other to be macerated for the hard parts; the 
decalcification was done in weak solutions of nitric acid. Polyps to be sectioned were then 
embedded in the usual way™, and sections cut to thicknesses of 12, 10pm, 8p, 6p, 4p. 
After repeated experiments with various stains, I found that Haidenhain’s iron hematoxylin 
followed by eosin gave the best results; the former stains the nuclei, muscle-fibres and 
nematocyst threads dark, and the latter colours the cytoplasm light pink and the mesoglea 
deeper pink. All the polyps whose histology I have studied were treated by this method. 

Body-layers. From an examination of sections of coral polyps I am led to think 
that the ectoderm and endoderm are not composed of definite units known as cells. The 
usual appearance presented by each layer is that of a sheet of protoplasm with nuclei 
either closely aggregated as in the ectoderm, or somewhat scattered as in the endoderm ; 
an extreme case is that of the stomodzal ectoderm, in which the nuclei are so numerous 
and so closely packed together that it is difficult to see how there could be cell-limits. 


* In connection with the Meandroid Astreids, I cut some sections in gelatin with Aschoff’s CO, freezing 
microtome, a method recently employed by Mr J. F. Gaskell (‘A Method of cutting frozen Sections by embedding 
in Gelatin,” Journ. of Pathology and Bacteriology, vol. xvu, No 1, p. 58,1912). The advantages of this method 
consist in the avoidance of absolute alcohol, xylol or paraffin, which usually shrink and distort the internal 
organs of the polyps, and in the better chance of preserving the histological condition of the tissues as the object 
has not to be subjected to a temperature above that of the room, but there is great difficulty in making serial 
sections and no method has yet been devised of dissolving out the gelatin from the sections.’ These sections in 
general confirm the account of polyp cytology given subsequently. 


MATTHAI—RECENT COLONIAL ASTRAIDA 7 


Gardiner too could hardly distinguish cell outlines in the ectoderm and endoderm in his 
sections of Cenopsammia and Flabellum; with regard to the calicoblastic layer of the 
latter genus he says (50, p. 139), ‘“‘I can only regard the layer as an enormous syncytium, 
and for the growth of a septum there would seem to be a flowing up of the protoplasm on 
either side.” The only structures with definite boundaries which may be regarded as cells 
are, aS suggested below, the nematocysts and reproductive eléments. Whether the 
syncytial appearance of the body-layers is a secondary feature due to the fusion of cells or 
is the original condition persisting in the adult cannot at present be decided. It is possible 
that the two layers may arise and continue as sheets of protoplasm with the nuclei 
multiplying by division without corresponding fission of the protoplasm. But of course no 
detinite view as to the real nature of these layers can be formed without studying their 
development and without fresh polyp-tissue. 

Structures similar to the gland cells (both mucous and granular) of Actinians as 
described by various authors are present in the ectoderm. In my sections they look like 
vacuoles, their cellular nature bemg doubtful. They often appear non-nucleated or with 
more than one nucleus in their walls. The mucous type (figs. 2 and 3) is oval or flask- 
shaped, usually with the broader end turned towards the free surface ; it occurs in varying 
numbers in all the species, in the oral-disc, edge-zone, tentacles and the convolutions of 
the mesenterial filaments, and contains some kind of mucous material which stains brown. 
The granular type (figs. 11 and 66) is either flask-shaped or elongated, the granular 
contents, whose nature and function are uncertain, staining dark; it is usually present in 
the mesenterial filaments, less frequently elsewhere in the ectoderm. 

The Hertwigs (66) supposed the granular cells to be stages in the development of the 
mucous ones, whereas H. V. Wilson (122) regarded the two types as distinct, since they 
are distinguishable from the start and have a different distribution in the early develop- 
mental stages of Mamcina areolata. 

Small oval sacs, showing finety granular contents stained light pink, are usually 
present in the ectodermal ridges of the stomodeeum as well as in the mesenterial filaments 
(figs. 23 and 45). These perhaps belong to the granular type and their contents may be 
digestive in function. 

The endoderm is often vacuolated, the vacuoles being oval or goblet-shaped ; nuclei 
are less numerous than in the ectoderm and occur between the vacuoles, or the latter may 
appear multi-nucleated. The vacuoles are usually more or less transparent, except in the 
mesenteries, where in some species, e.g. Cyphastrea serailia (Forsk.), they stain’ reddish- 
brown (fig. 4). The exact nature of these vacuoles is unknown. 

Willem (120) regarded the endoderm as mainly absorptive and excretory and the 
mesenterial filaments as capable of secreting a digestive fluid. In addition to extra- 
cellular digestion Willem also refers to intra-cellular digestion in the endoderm, whereas 
Gardiner (49, p. 375) rejected the possibility of the endoderm having any digestive 
function, regarding the layer as being homologous with the mesoderm of Triploblastica. 
Bourne (17, p. 225), however, later suggested that in Heterocyathus equicostatus the 
“olandular-looking cells” 
Gardiner. 


may secrete a digestive fluid, and thus disagrees with 


8 PERCY SLADEN TRUST EXPEDITION 


Mesoglea. This name was employed by Bourne (11) to replace the term “meso- 
derm” in denoting the supporting lamina of Coelenterates, since by usage ‘“‘ mesoderm ” 
had become identical with the mesoblast of the Triploblastica. The supporting lamina 
has been described under various names: Milne Edwards and Haime refer to it as 
the “tunique musculaire,” the Hertwigs use the terms “mesoderm,” “stiitzlamelle,” 
“stiitzsubstanz,” while among some English authors names like ‘supporting lamella” 
(H. V. Wilson) and “structureless (or basement) membrane” (Gardiner) have been 
resorted to; McMurrich (100, p. 270) compared it with “the limiting membrane which 
occurs Just below the ectoderm” in the higher Metazoa. Bourne’s new name was accepted 
by Haddon (55) and van Beneden (8), and is now in common use among most English 
and American authors (Hickson, McMurrich, Duerden, etc.). 

Lacaze Duthiers (92) and Faurot (40) have, however, questioned the necessity for 
a new name in preference to the older term “mesoderm.” The latter author, after careful 
study, regarded it as composed of lamellee (“ plissements”), which in Actinians are dis- 
posed circularly in the body-wall and longitudinally in the mesenteries and are capable 
of being separated from one another; these lamellze could be dissociated still further into 
fibrille. The granulations on the mesenterial pleats (‘“‘feuillets”) seen in transverse 
sections, which the Hertwigs (and others after them) regarded as the cut ends of the 
endodermal muscle-fibres, are regarded as sections of “barbelures,” which are part of 
the intermediate substance. In a subsequent work (42) Faurot still adhered to the term 
“mesoderm” for two reasons; “la lame mésodermique, bien que n’étant revétue qu’en 
partie par des fibrilles musculaires, est contractile dans toute son étendue. Certaines 
régions du corps, susceptibles de se contracter tres fortement, ne présentent sur les 
coupes, aucune trace de ces fibrilles. Le disque pédieux notamment, en est completement 
dépourvu, et cependant certaines Actinies telles que | Actinia mesembryanthemum peuvent 
se déplacer sans l’intervention des cloisons, & l’aide de ce disque. Les éléments ecto- 
dermiques musculaires dits “‘immergés” n’existent pas dans le mésoderme de la colonne 
et cependant cette paroi peut se contracter longitudinalement. Et d’ailleurs, un tissu 
contractile doit-il nécessairement, soit renfermer des éléments musculaires, soit en étre 
recouvert ?” (p. 367); and again, “un autre motif pour ne pas renoncer, jusqu’a présent 
chez les Actinies, 4 la désignation de ‘mésoderme’ c’est qu il n’a pas encore été formelle- 
ment démontré que cette couche intermédiaire n’est pas, en réalité, chez ces animaux, le 
premier rudiment d’un véritable mésoderme. Il me parait certain d’ailleurs, ainsi qu’on 
le verra par la suite de ce travail, quelle joue un trés grand role dans la formation du 
pharynx, des cloisons, et aussi, fait remarquable, de certaines parties (loges) de la cavité 
du corps” (p. 369). Faurot further suggested from certain observations on the changes 
in the colour-markings of the tentacles of Peachia hastata and on the irregular size of 
the “plissements” in the column-wall of Bunodes thallia that the contractility of the 
mesoderm is amceboid. In his latest paper (48, p. 347) he regards the usual view that 
the supporting substance is a secretion of the ectoderm or endoderm as vague and not 
based on accurate observation. 

At first the supporting lamella appeared to be a homogeneous substance which 
uniformly stained in eosin or picro-nigrosin and did not contain any cell inclusions. 


MATTHAI—RECENT COLONIAL ASTRAIDA 9 


Subsequent examination revealed some nuclei and fibrous appearances, whereas Bourne 
and Duerden did not observe any such structures in the coral-polyps they examined. 
But owing to the uncertainty that still prevails over the true nature and origin of this 


iG 


intermediate substance the term “ mesoglea” is used in this paper for descriptive pur- 
poses without any of the significance that Bourne attached to it. 

Calicoblastic layer of ectoderm (PI. 1, fig. 1, Pl. 3, fig. 25 and Pl. 5, fig. 53). The 
ealicoblastic layer (von Heider) is discontinuous only where the soft parts are attached 
to the corallum by means of the wedge-shaped processes described below. Its nuclei are 
quite characteristic in being large oval sacs arranged in a single layer, in each of which 
is a central darker spot, possibly the nucleolus. At the skeletal attachments of the 
mesenteries the calicoblastic layer is considerably thickened and vacuolated in some 
species (cf. Galaxea fascicularis, Linn.), its protoplasm appearing in transverse sections 
as thin columnar strands. A structureless membrane covering the external surface of 
the calicoblastic layer and separating it from the corallum, as described by Bourne, could 
be made out, although torn in places. This membrane is similar in nature and appear- 
ance to the structureless mesoglea of the polyp. ‘The calicoblastic layer is better de- 
veloped in the edge-zone, ccenosare and the peristomial region of the body-wall, while 
towards the base of the polyp it is attenuated. 

The soft parts are attached to the corallum by means of wedge-shaped processes, 
the latter being particularly conspicuous at the skeletal attachments of the mesenteries 
(fig. 43). Later observations show that these processes are formed not in distinct cells 
(Bourne's desmocytes) but by modification of the calicoblastic cytoplasm. They may be 
formed anywhere in the calicoblastic ectoderm. Stages in their development have been 
noticed, but these require further study. It is unlikely that the strize in these processes 
mark the attachments of the muscular fibres of the mesenteries to the corallum as 
Gardiner suggested for his species of Cenopsammia, since they are found even at the 
very base of the body-wall and at places other than at the skeletal attachments of the 
mesenteries, where there are no traces of muscular fibres. 

Nematocysts have been sometimes observed in the calicoblastic layer. Duerden 
(37, p. 32) also found “small oval nematocysts with a close spiral thread” scattered 
throughout the ecalicoblastic ectoderm of Stderastrea radians. 

Dissepiments and polypal growth. Bourne, in his account of the anatomy of 
Mussa corymbosa and Huphyllia glabrescens, thus described the formation of dissepi- 
ments: ‘There are probably periods of active coral secretion alternating with periods of 
reproduction in these polyps. During the latter period the thin dissepiments are formed 
by the basal tissues, whilst in the former period, the septa increase greatly in height, the 
polyp is, as it were, moved higher up upon the stem, and deserts the old dissepiments 
upon which it was resting. Then follows a new period of reproduction during which 
new dissepiments are formed.” 

It is difficult to see how such a view can be held when we consider that the polyp is 
firmly attached all along its height to the corallum by means of the wedge-shaped 
processes which would prevent any periodical moving up of the polyp. My observations 
show that there is always some tissue in the cavities of the corallites below the base of 

SECOND SERIES—ZOOLOGY, VOL. XVII. 2 


LIBRARY). 
wh OD = 


10 PERCY SLADEN TRUST EXPEDITION 


the polyp, viz. in the spaces formed by the dissepiments. This could not be the case if 
the whole of the polyp actually moved up the corallite, and often the dissepiments are not 
complete partitions stretching right across the calyx. In the face of these facts, it seems 
to me that the dissepiments are partitions periodically formed across portions of the 
corallite and of the polyp. When a full series of such horizontal partitions is formed 
between the septa, etc., the polyp would be divided into two parts, the lower part being 
cut off from that above; the lower part degenerates while the polyp soon recovers its 
normal height. Of course where a dissepiment is to be formed the body-wall has to be 
folded horizontally inwards, and between the two layers of calicoblastic ectoderm thus 
formed the calcareous material is secreted. By the successive formation of dissepiments 
in this way the corallites grow in height, with the living polyps occupying only the 
uppermost chambers. A further reason for holding the view suggested here is the 
fact that synapticulz are actually formed as horizontal rods between septa piercing 
through the intervening mesenteries and a dissepiment is nothmg more than a 
flattened-out synapticulum partitioning off an interseptal space*. Indeed, every stage 
between a rod-like synapticulum and a complete dissepiment can be seen in the dry 
specimens. 

Ogilvie (108, p. 157) referred to a similar origin of the dissepiments in the following 
words: “The aboral wall draws itself up gradually during the period of active deposit at 
the septal edge, forming during its updrawal the arched development. Finally the period 
of ‘pause’ sets in, and the body-wall between the septa remains supported on its many- 
arched floor, which is then completed and thickened.” 

Nematocysts. Boulenger (10) used the term nematocyst for “the actual stinging 
capsule” and nematoblast for “the cell in which the former is embedded, and of which 
the cnidocil and the stalk are part.” In this paper I use the term nematocyst to mean 
the whole apparatus, viz., the sac, thread and basal filament (if present), the reason for 
which will become evident from the account of their probable development. 

I am inclined to agree with Hadzi and Boulenger that nematocysts are ectodermal 
structures ; their occasional presence in the endoderm of certain polyps may be due either 
to subsequent displacement or to migration. 

Three main types of nematocysts have been met with in the polyps studied, which 
I refer to as types I, Il and III; all of these are liable to variations in the different 
species. R. Hertwig (67) recognised, after a general survey of nematocysts in Actinians, 
that their form and the nature of their thread was not the same everywhere, and 
suggested that they might some day become of systematic importance. The subsequent 
work of van Beneden (8) on Cerianthides and of Carlgren (20 and 21) on Actinians 


* Tt is doubtful if there is any real difference between true and pseudosynapticula. The former 
according to Pratz and Ogilvie are said to possess centres of calcification, whereas the latter are described as 
formed by the union of septal granulations, and hence devoid of any such centres. The Astrzeidz are said to 
have only such pseudosynapticula, but in some of my microscopic sections of their coralla distinct dark centres 
are visible. Indeed, if the hard parts are formed in the way Bourne has shown, i.e. outside the calicoblastic 
layer of ectoderm, and not by a calcification of its “cells,” the dark centres and lines will have none of the 
significance that Ogilvie attached to them. Costal synapticula are seen in sections of species of Mavia and 


Goniastrea. 


MATTHAI—RECENT COLONIAL ASTRAGID At itl 


brought forward further evidence in support of Hertwig’s suggestion. My observations 
on nematocysts in coral polyps lead to the same conclusion. 

Type I (Pl. 1, figs. 2, 3, Pl. 4, fig. 35, Pl. 6, figs. 63 and 65). ‘This is the so-called 
“tentacular type.” It occurs not merely in the tentacles but in the edge-zone, oral-disc, 
ectodermal ridges of the stomodeum and in the straight regions of the mesenterial 
filaments (rarely in their coiled regions). There is no doubt, as Gardiner has shown, that 
Bourne mistook nematocysts of this type in Euphyllia glabrescens for immature stages 
of a totally different type, the so-called ‘“ mesenterial.” A fully developed nematocyst 
consists of a long narrow sac somewhat trumpet-shaped, widest at its upper end and 
gradually narrowing towards its base; the wall of the sac appears to be an elastic 
membrane against whose inner surface lies a slender spirally-coiled filament stained dark, 
the turns of the spiral being somewhat oblique and its number variable in the different 
species from fifteen up to fifty. Even under the highest magnification (Zeiss Oc. 12 x ob. 
homog. immers. 2°5mm.), I have not been able to detect a spiral round the filament as in 
type III described below. It merges at the bottom of the sac into a little granular 
. mass. Usually from the base of the sac a slender process passes down which, in the 
unextruded nematocysts could be traced to the mesoglea, possibly neuro-muscular, for 
keeping the nematocyst in position and for the conduction of nervous stimu. Normally, 
the nematocysts lie with their upper ends touching the surface of the ectoderm, but, 
when brought into action, are extruded partly or wholly with the filament ejected but 
never seen completely uncoiled. The nematocysts attain to their full size only in the 
tentacular ectoderm, where they are closely arranged to form batteries, elsewhere being 
shorter with fewer turns of the spiral and never forming batteries. 

This type of nematocyst appears to be derived from an ectoderm nucleus. The 
nuclear membrane grows in size to form the elastic wall of the nematocyst; within it the 
chromatin-mass breaks up into minute granules (one of the granules im the middle of 
the sac is conspicuous by its large size), which, by their union, form the coiled filament. 
As the nematocyst grows in length the turns of the spiral are added to from above 
downwards. The filament after it has attained its full length terminates in the residue 
of the chromatin left at the bottom of the sac. The filamentar process passing from the 
nucleus to the mesoglea becomes the neuro-muscular process of the nematocyst. In the 
tentacular ectoderm a nucleus, as it becomes modified, migrates towards the surface of 
the layer, hence between the row of fully formed nematocysts at the periphery of 
a battery and the small round nuclei just above the mesoglea there are various 
intermediate stages in the development of this type. According to this interpretation 
every ectoderm-nucleus may be regarded as potentially capable of taking part in the 
formation of a nematocyst. 

Gardiner, from his study of Cenopsammia, has given a totally different interpretation 
of the history of his “tentacular” nematocysts. He regards them as entirely cytoplasmic 
in origin, the filament being formed by the union of granules in the protoplasm, resembling 
“both in its development and appearance the elastic fibres in the connective tissue of 
vertebrates,” and states that ‘“‘no nucleus can in any stage be distinguished within this 
(nematocyst) membrane,” but that “generally the basal end of the nematocyst lies in 


2—2 


12 PERCY SLADEN TRUST EXPEDITION 


a finely granular mass of protoplasm with a nucleus either oval or rod-shaped.” I have 
carefully examined the serial sections of his polyps of Canopsammia and have found that 
the appearances of the tentacular and oral-dise ectoderm in iron-hematoxylin are 
identical with those in my polyps. All the stages in the modification of a round nucleus 
into a nematocyst can be made out. The nucleus in Gardiner’s fig. 7 does not of course 
belong to the nematocyst, but is simply one of the many nuclei in the ectodermal 
syncytium. 

The Hertwigs (66) have given figures of nematocysts from Actinians which resemble 
type I (Pl. 2, figs. 12 and 13, Pl. 3, figs. 5 and 11). In Pl. 4, figs. 2 and 5, each of 
these nematocysts is represented as arising in the cytoplasm of an ectodermal cell with 
the nucleus lying by its side when it is fully developed. Jourdan (74) has recorded 
nematocysts of the same type from the tentacular ectoderm of the Zoantharia he studied 
(Pl. 3, figs. 10 and 11; Pl. 12, figs. 83—85). In Pl. 5, figs. 32—36 (Actinia equina) and 
in Pl. 15, fig. 108 (Balanophyllia regia) he has figured cnidocils which I have not found 
in my sections. The type of nematocysts which, according to van Pesch (111) is so 
abundant in the tentacular batteries of Antipathides, is doubtless the same as type I 
(Pl. 4, figs. 5 and 6).* 

Type If (Pl. 1, fig. 8, Pl.2, fig. 18, Pl. 3, figs. 26, 33, 34, Pl. 4, figs. 37 a—f, 38, 
Pl. 6, figs. 58a and b, 59, and 62). The nematocysts belonging to this type are large 
oval sacs, broad at one end and bluntly pointed at the other, found everywhere in the 
ectoderm but principally in the convoluted regions of the mesenterial filaments, where 
they may be closely arranged to form batteries. Each of these consists of a membranous 
wall containing somewhat granular contents. A narrow cylindrical structure or axis 
extends from the upper end of the nematocyst to about two-thirds its length. This 
consists of a dark-stained axial core, usually straight, sometimes slightly bent, always 
pointed at its outer extremity, and enclosed in a thin membranous sheath against the 
inner surface of which lies a fine close spiral, also stamed dark; the sheath leaves the 
pointed extremity of the axial core free. This type resembles in appearance the nemato- 
cysts figured by Hadzi (59) from certain Hydromeduse, and by Boulenger (10) from 
certain Craspedote Medusz. In a few cases I have noticed the pointed tip of the axis 
projecting beyond the sac, but there is no further evidence of the protrusibility of this 
nematocyst-organ in any of the polyps I have examined. The nematocyst attains its 
full size in the convolutions of the mesenterial filaments, being smaller in the ectoderm of 
the oral-dise and edge-zone. It usually takes up a position at right angles to the surface 
of the ectoderm, the pointed tip of the axis slightly pushing up the ectodermal surface. 

In the tentacle, nematocysts of this type occur in the terminal batteries interspersed 
among those of type I, but always in a modified form, being much narrower and somewhat 
longer, and the axis not extending beyond a third of the length of the sac. In some of 
these there is a faint suggestion of a coiled thread. ‘This variety, owing to its constancy 
in shape and distribution may be termed type II (PI. 3, figs. 26 and 33). 

In the genus Gomastrea these nematocysts are much narrower and longer and the 


* The second type of nematocysts yellow in colour and without any spiral which van Pesch has observed in 
the tentacular batteries of Hucirripathes rumphi may not unlikely be a variety of type IT. 


MATTHAI—RECENT COLONIAL ASTRAIDA 13 


axis shorter ; this variety may be termed type Il c (Pl. 6, figs. 59 and 62); they are usually 
arranged close together in the coils of the mesenterial filaments. In G. retzformis (Lam.) 
the axis is about three-fifths the length of the sac and somewhat bent, whereas in 
G. solida (Ed. & H.) and G. pectinata (Kd. & H.) it is about a quarter of the length of 
the nematocyst and somewhat swollen in its middle; in the last two species there is also 
a suggestion of a coiled thread in the sac. 

I have met with nematocysts of this type in the calicoblastic layer of certain well- 
preserved polyps whose corallum resembles that of Mycediwm okent Ed. & H.*. Obviously 
they cannot be of any use in this layer as they are hidden by the corallum, but their 
presence proves that the power of developing these structures is inherent in the ectoderm 

everywhere. 
In certain species, e.g. Favia doreyensis, Ed. & H. (PI. 5, fig. 45), I have seen this 
type of nematocyst in the endoderm, though not in any great abundance as in the 
ectoderm. Their presence in that layer may be explained in two ways, either that the 
nematocysts have arisen in the endoderm, or more probably, that nematoblasts have 
migrated into this layer from the ectoderm and have developed there f. 

It is highly probable that the dark-stained axis is a modified nucleus. Certain 
appearances suggest that a nucleus wanders to the surface of the ectoderm, elongates and 
becomes modified into the axis, its chromatin forming the axial core and the fine spiral 
and the nuclear membrane constituting the surrounding sheath. This process is accom- 
panied by the formation of a membranous wall in the surrounding cytoplasm, the latter 
forming the contents of the sac. A nematocyst of this type would accordingly be a 
complete cell (Pl. 4, figs. 37 a—f). 

In Gardiner’s polyps of Cenopsammia nematocysts of this type are present in the 
convolutions of the mesenterial filaments, but are somewhat narrow, and the dark-stained 
axis of each extend usually to about a third of the length of the sac, rarely to its middle. 
Around the axis (Gardiner’s “‘eversible portion”) is a continuous spiral as in my polyps, 
not a “spiral row of short hairs.” I haye not seen any distinct coiled thread as Gardiner 
has represented in his diagrammatical fig. 14, nor any definite nucleus in the nematocyst 
as in his figs. 17 and 18. In a few cases the axis has been forcibly ejected (not everted), 
as could be seen in Gardiner’s section shown in fig. 13, leaving a corresponding space 
within the sactf. 

Type IIT (Pl. 2, figs. 15, 16, 17, Pl. 4, fig. 40, Pl. 6, figs. 57, 60, and 61). The 
largest nematocysts that I have examined belong to this type. Hach of them is broader 
in the middle than at the ends, and consists of a membranous wall, enclosing protoplasmic 
contents of the same nature as in type Il. Running through the middle of the sac, along 
its whole length, is a somewhat thick protoplasmic core, stained deeper pink than the rest 
of the contents. In addition to this, there is a long thread spirally coiled along the mner 

* The description of this species is reserved for a subsequent paper. 

t For migration of nematocysts see Schneider, Hadzi and Boulenger. 

{ The “larger nematocysts resembling those of the body-wall but usually much longer and narrower,” 
which Bourne (17, p. 224) observed in the tentacular batteries of Heterocyathus equicostatus interspersed 


among the “small nematocysts of the usual form,” may, I venture to suggest, be similar to type II 6, but 
Bourne has not figured any of them. 


14 PERCY SLADEN TRUST EXPEDITION 


surface of the membranous wall, much thicker than in type I, but with far fewer turns of 
the spiral. In the unextruded condition, it is difficult to make out the exact nature of 
the thread, but, as in some of my polyps, e.g. Favia favus (Forsk.), most of these nema- 
tocysts are found with the thread partially ejected, I have been able to study its structure. 
It consists of an axial strand stained homogeneously dark, enclosed in a thin membranous 
sheath against the mner surface of which lies a fine closely-wound spiral stained dark. 
The tip of the axial strand is pointed, and projects in spike-like fashion beyond the 
sheath. In some of the nematocysts the sheath is partly torn and the spiral broken 
here and there along the thread. The central protoplasmic core is never everted; indeed 
it does not appear to have any connection with the thread at all. As in type I, the 
thread ends at the base of the nematocysts in a small granular mass. This type varies 
hke I and II in the different species, an extreme case being that of Goniastrea retiformis, 
in which the sac is much narrower (Pl. 5, fig. 49). 

There is little doubt that the so-called “ripe nematocysts” of Bourne from Huphyllia 
glabrescens, judging from his figures (Pl. 4, figs. 10 and 12) really belong to this type; 
its “axial body,” which the author believes to be eversible, being probably the central 
protoplasmic core. I have not met with any peculiar armature at the extremity of the 
thread, as in Bourne’s fig. 11, in any of my nematocysts. 

In the three species of Hchinopora examined, this type has undergone a characteristic 
modification, in that the sac is not only narrow, as in Goniastrea retiformis, but the 
longitudinal protoplasmic core is absent and the turns of the coiled thread more numerous. 
This variety may be termed type IIIb (Pl. 2, figs. 16 and 17). 

The large nematocysts with the long coiled thread which Jourdan (74) found in the 
ectoderm of the body-wall of Corynactis viridis (Pl. 8, figs. 54 and 55) and in the 
mesenterial filament of Balanophyllia reyia (Pl. 15, fig. 113), come under my type. 
Resembling the latter are also the two nematocysts which Bourne (17) has figured from 
the ectoderm of the body-wall of Heterocyathus equicostatus, but as in Gardiner’s 
representations (Cenopsammia, figs. 15 and 16), the everted thread is covered with long 


” 


barbs arranged in a spiral. The “medium-sized elongate” nematocysts which Bourne 
describes in the same paper from the ectoderm of Dendrophyllia gracilis (Pl. 4, figs. 25 
and 26 b), as well as the nematocysts from the mesenterial filaments of an anthocyathus 
of Fungia (18, fig. 28), resemble IIT b (PI. 2, figs. 16 and 17); I have little doubt that the 
thread in these cases, if carefully examined, will be found to possess a closely wound 
spiral. On the other hand, I have not seen the “large elongate oval” nematocysts 
(Pl. 4, figs. 26 c—e, which are the same as Pl. 3, fig. 18); they are similar to Gardiner’s 
fig. 14 or 19, but have a longer “eversible portion.” 

Type III is essentially similar to type II, consisting of a membranous wall, proto- 
plasmic contents and a thread which, like the axis of II, has a dark-stained central core, 
a thin membranous sheath, and a surrounding fine spiral. The different parts appear 
also to have the same origin as those of type II, the thread being probably a modified 
nucleus (the axial strand and the spiral formed perhaps of the chromatin and the sheath 
of the nuclear membrane) and the membranous wall developed independently in the 
surrounding cytoplasm. The granular mass at the base of the nematocyst into which the 


MATTHAI—RECENT COLONIAL ASTRAIDA 15 


axial strand merges may be the residue of the chromatin left after the former has grown 
to its full length. Type III, like II, would then be a complete cell. 

If the above study of the nematocysts be confirmed by living material, then the 
trumpet-shaped body (a modified nucleus) of I would be comparable only with the axis 
of IT and with the thread of III, its wall and filament * being homologous respectively 
with the sheath and spiral of the last two types, but without a chromatic central 
core. 

Of the four kinds of nematocysts described and figured by Gosse (58, Pl. 11), the 
“spiral enidee (cnidz cochleate),” which are confined to the walls of the tentacles and 
whose thread (“ecthoreeum”) is without any armature, is in all probability the same 
as type I, the “chambered cnidze (cnidze camerate)” and “the tangled cnidse (cnidee 
come near type III, the latter having no “barbed bristles” (‘ pterygia ”) 
on the spiral (‘‘strebla”) surrounding their thread, while the “globate cnide (enide 


” 


glomiferze) 


globatee)” may be glandular structures and not nematocysts. 

Previous authors who have made the study of nematocysts agree that they arise 
in interstitial cells of the epithelium. At first a vacuole is formed in the cytoplasm 
of such a cell which becomes the capsule of the nematocyst. Opinion is divided as 
to the nature and mode of development of the filament. Of the many views that have 
been put forward the two main ones are (a) that the filament is formed inside the capsule 
(Bedot, 4), (b) that it develops outside the capsule, becoming subsequently invaginated 
into the latter (Jickeli, 73; Nussbaum, 107; Murbach, 106). Iwanzoff (72), on the other 
hand, holds an intermediate position that the filament is at first partially invaginated by 
its distal end, the invagination becoming complete with the full development of the 
nematocyst. According to these authors the filament is a cytoplasmic product, the 
nucleus of the nematocyst persisting outside the capsule. 

In the last two types I have not been able to make out a neuro-muscular process as 
in J, and in none have I observed any process suggestive of a enidocil. 

After completing the above studies of the nematocysts my attention was drawn to 
a paper by Theodor Moroff (‘‘Entwicklung der Nesselzellen bei Anemonia,” Archiv 
Zellforsch. iv. p. 142, 1909), in which the author has traced the origin of the filament to 
nuclear elements. Moroff has described two kinds of nematocysts from the tentacles 
of “ Anemonia sulcata,” (a) “ spirocytes,” which are similar in appearance to type I, and 
(b) “nematocytes” which resemble type II]. As far as I have been able to understand 
his description, each of these has a double wall (in transverse sections of my types 
the walls consist, in every case, of a single membrane), the outer “sklera” being 
cytoplasmic in origin, while the inner “propria” is chromatic and continuous with the 
thread which is also formed by the union of chromidia; he regards the axis of the 
“nematocytes ” (fig. M) as continuous with and having the same origin as the thread and 
capable of being everted. The nuclei as they become modified into the nematocysts 
wander from the base of the ectoderm to its periphery. 


* TI have applied the term “filament” to the structure in type I, while the use of the term “thread ” 
is restricted to the complex structure in type III, which consists of an axial strand surrounded by a 
sheath and by a fine spiral. 


16 PERCY SLADEN TRUST EXPEDITION 


Haddon and Shackleton (57) observed nematocysts in the endoderm of Jsawrus 
asymmetricus and of Zoanthus coppingert; those figured on Pl. 64, fig. 2 bear 
a certain resemblance to type I. Van Beneden (8) also found nematocysts in the 
endoderm of some larval Cerianthide, and gave many figures; he distinguished two main 
types of nematocysts, the first type closely resembling I, while the second comes near III, 
but the thread is much shorter in Ovactis brasiliensis (Pl. 3, figs. 2, 13, 14, 15). The 
nematocysts consisting of three sorts from the “botrucnides”* and “ enidorages” 
of Hensenanthula (Pls. 12, 14, 15) bear some resemblance to II, but the pointed 
extremity of the “fil rectiligne” is directed towards the base of the nematocyst 
(cp. Pl. 12, figs. 13 and 14, with my PI. 6, fig. 58 a, b), but one of these in the exploded 
condition shows a long tapering thread. According to van Beneden each of these 
nematocysts has a semilunar nucleus attached to it, the “fil rectiligne” being pre- 
sumably protoplasmic, as may be inferred from the developmental stages (Pl. 12, 
figs. 15—18). 

The respective functions of these types of nematocysts and the different ways in 
which they are used can be ascertained only by means of actual experiments on living 
polyps. Nos. I and III in all probability have a stinging function, which may account for 
their mutually exclusive distribution, the latter restricted to the coils of the mesenterial 
filaments, the former found mainly in the tentacular batteries. On the other hand, 
type II may not have a stinging function ; in this connection it may be pointed out that 
their sacs sometimes take a brownish tinge in iron hematoxylin and eosin like the 
mucous vacuoles. Could they then have an adhesive function and could that be the 
reason why they are found interspersed among the nematocysts I in the terminal 
tentacular batteries as well as among those of type III in the coils of the filaments ? 

In Gardiner’s polyps of Cenopsammia nematocysts III + are present in the 
convolutions of the mesenterial filaments, but the protoplasmic core has not been met 
with in any of them. From Gardiner’s description and figures it would appear that he 
regarded these as the final stage in the development of type II. His “mesenterial 
nematocyst ” in fig. 14 is a reconstruction and a combination of types II and III, having 
both the axis (“‘eversible portion”) and the coiled thread. In a few cases I have seen 
structures similar to fig. 15 which presumably belong to III, with the greater part of the 
thread broken off. The appearance of a row of hairs may be due to the breaking of 
the turns of the spiral surrounding the axial strand; such a condition is presented by 
some of the nematocysts III of my polyps of Pavia favus (Forsk.) in which there is no 
doubt whatever that the spiral round the axial strand has been broken at places. It is 
also to be noted that while most of the authors cited above place the nucleus in the 
protoplasm surrounding the nematocyst-capsule, Gardiner in his figs. 17 and 18 has 
shown it inside the capsule. 


* « Botrucnides” is the name given by van Beneden to certain curious endodermal structures having the 
form of bunches of rounded grains which take the place of acontia in Cerranthula, Hensenanthula and 
Calpanthula ; the individual grains which he has termed “Cnidorages ” fall off into the gastro-vascular cavity 
on attaining their full size. 

7 Nematocysts II and III in Gardiner’s polyps do not appear to have been well fixed. 


MATTHAI—RECENT COLONIAL ASTRAIDA 17 


Gardiner has recorded the general occurrence of nematocysts in the endoderm of the 
polyps of Flabellum on each side of the upper top ends of the septa. These undoubtedly 
belong to III, which the following considerations render it probable that they may have 
been at first formed in the calicoblastic layer, their subsequent presence in the endoderm 
being perhaps due to displacement or migration : (a) nematocysts of this type are present 
in the calicoblastic layer—compare type II in the calicoblast of Mycediwm okeni Ed. 
and H.—usually towards the top ends of the septa, some of them being entire with sac, 
protoplasmic core and coiled thread and lying parallel to the septa with the calicoblastic 
layer raised over them or in an oblique position partly in the calicoblast and partly in the 
endoderm; (>) in the endoderm no entire nematocysts are present, the wall being, in 
every case, absent and the coiled thread usually broken; (c) they have no definite 
position, but lie loose in the endoderm ; (d) the nematocysts in the calicoblastic layer are 
well stained, whereas in the endoderm they have taken very little stain, probably because 
of subsequent degeneration. 

Gardiner makes no record of such nematocysts in the mesenterial filaments of 
Flabellum, but oval, pink areas with a suggestion of coiled threads can be seen in the 
convolutions of the filaments*. 

Nervous system (Pl. 1, fig. 3, Pl. 2, fig. 24, Pl. 5, figs. 47, 52, Pl. 6, fig. 66). Nerve 
cells and fibres? as figured by the Hertwigs for the Actinians have not been met with in 
my sections. The ectoderm just above the mesoglea is more granular than usual, being 
the only indication of a nervous stratum, perhaps representing their “ Nervenfaserschicht” 
and von Heider’s “Interbasalsubstanz.” This is seen in the edge-zone, oral-disc, 
tentacles, ectodermal ridges of the stomodzeum and the mesenterial filaments, and best 
of all in the tentacles, perhaps because of their bundles of nematocysts. 

Endodermal musculature. This constitutes the main musculature of the polyp; it 
is present in two regions, viz., oral-disc and mesenteries. 

_ Attached to the lower surface of the mesoglea of the oral-disc (Pl. 5, figs. 51, 52) is 
a thin layer of muscle-fibres disposed circularly, but much less developed than in Actinians 
and without any mesogleal pleats which are so prominent in many of the latter (wide 
R. Hertwig, Haddon and van Beneden). Around the outer margin of the bases of the 
outer cycle of tentacles the muscular layer is somewhat thickened to form a sphincter, the 
so-called Rotteken’s muscle ; from the appearance of the spirit specimens it may be inferred 
that its action is to reduce the opening of the peristome by drawing its lateral wall over 
the oral-disc and hence over the tentacles, which consequently are bent over the mouth. 
Faurot (23) does not regard the sphincter as having any retractile action on the tentacles 


* In Cenopsammia Gardiner’s figs. 7 and 8 are type 1; 9 and 10 are appearances in the protoplasm ; 14 is 
a combination of types II and III; 15 and’ 16 are probably type III; 1719 are type II but the nucleus is 
an optical appearance. 

+ Havet (61) employing better methods has been able to distinguish both sensory and motor nerve cells 
and fibres in the ectoderm and endoderm of an Actinian Metridium dianthus, and has given excellent figures 
illustrating their arrangement and distribution. Van Beneden could not make out any ganglion cells in his 
sections of larval Cerianthide, although a distinct “assise nerveuse” was present in both the ectoderm and 
endoderm at the base of the “assise épithélioide” and just above the “‘assise musculaire.” Faurot could not 
find such a nervous layer in the endoderm of the Actinians he studied. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 3 


18 PERCY SLADEN TRUST EXPEDITION 


of Actinians. The oral-dise muscle is continued as a much weaker layer into the outer 
wall of the edge-zone. R. Hertwig (42) laid great stress on the systematic value of the 
circular muscle for both the genera and species of Actinians. 

The mesenterial musculature consists of a longitudinal layer of muscle-fibres on either 
side of every mesentery attached to its mesoglea (PI. 1, fig. 4, Pl. 2, fig. 13, Pl. 3, fig. 27). 
As a rule, on all the mesenteries except the directives, the entoccelic layer is much better 
developed than the exoccelic, the mesenteries bearing entoccelic mesogleal pleats or 
muscle-banners (Faurot’s “feuillets”) for the insertion of the fibres. These, usually known 
as the “retractors” (Fowler, Bourne, etc.), are attached from the oral-dise to a varying 
distance below the enterostome, and usually extend from the stomodzum to the column- 
wall; they are best developed in the stomodzeal region of the mesenteries. No specialised 
parietal muscles are present as in Actinians; the entoccelic muscular layer may, however, 
be regarded as representing on that side both the parietal and the “ faisceau unilateral ” 
of Faurot. By the contraction of this entoccelic layer, a downward pull is exerted on 
the whole oral-disc, thus bringing about the shortening of the whole polyp. In the 
non-pleatal region of the primary mesentery, a varying distance from the stomodeeal 
attachment of the latter, the entoccelic muscle-fibres are obliquely directed inwards and 
lie within the mesoglea. They are obviously attached to the stomodzeum, and by their 
contraction bring about the widening of the stomodzeum which is a characteristic 
condition of the retracted polyps. The exoccelic layer has about the same vertical 
extent, but the fibres are quite perpendicular, fewer and thinner; by their contraction 
they can only aid the entoccelic fibres in shortening the polyp, and hence the term 
‘“protractors” as applied to them by Fowler (26, p. 252) and others is a misnomer~. 
The exoccelic fibres are more distinct on the inner half of the primary mesentery in the 
stomodzeal region, presumably because the entoccelic fibres in the inner half function for 
widening the stomodzeum rather than for shortening the polyp. 

Fowler (44) believed that the exoccelic muscle-fibres of the mesenteries could be 
continued into the tentacles to form their external longitudinal coat, but to this view 
there are two obvious objections: (a) the fibres have, on such a view, to pierce their way 
through the mesoglea of the oral-disc in their upward course to the tentacles; (b) the 
external tentacular muscle-fibres are said to be ectodermal, whereas the exoccelic fibres 
are endodermal ; moreover Fowler's fig. 5 is only a diagram. Exoccelic pleats are present 
only in a few species. 

The condition of musculature is reversed on the directive mesenteries. 

Hollard (71) and the Hertwigs described the muscular fibres on the faces of the 
mesenteries of Actinians opposite the longitudinal retractor muscles as running trans- 
versely, while Faurot could find neither fibres nor pleats on that side and suggested 
that those authors might have mistaken the simple folds formed by the contraction 
of the mesoglea for permanent pleats. The parietal muscle of Actinians is unrepresented 
on. the exoccelic side of the mesenteries of coral polyps. 


* Gardiner observed a more or less similar condition in Cenopsammia. On p. 366 he says “On the faces 
of the mesenteries, opposite to the great retractor muscles, there are a few isolated longitudinal muscles with a 
similar course.... There do not appear to be any definite protractor or transverse muscles.” 


MATTHAI—RECENT COLONIAL ASTRAIDA 19 


Ectodermal Musculature. This is extremely weak in all the polyps and consists of 
(a) the filamentar processes occurring in the ectoderm of the oral-dise, edge-zone and 
tentacles whose course may be traced from the nuclei to the mesoglea, perhaps being 
neuro-muscular offsets, and (b) a thin layer of fibres with a longitudinal or oblique 
disposition, which has been recognised in the larger tentacles of some of the polyps. 
The latter layer begins at the oral-disc and extends up along the entire height of the 
tentacles, but does not appear to have any connection with the mesenterial mange Tm, 
It perhaps assists in shortening the tentacles. 

In my polyps I have not found a circular endodermal or a longitudinal ectodermal 
muscular layer in the column-wall nor are there ectodermal muscular fibres arranged 
radially in the oral-dise as described by van Beneden for certain Cerianthide (8, Pl. 5, 
ime, 1) 
Oral-disc (Pl. 5, figs. 51,52). The terms “oral-disc ” (“ disque buccal,” van Beneden) 
and “ peristome”* have been loosely applied to the upper surface of the Anthozoon polyp. 
In this paper the former name, which is synonymous with the “mouth-dise” (‘‘Mundscheibe” 
of German authors) of Fowler and Gardiner, is used for the circum-oral area delimited by 
the outermost cycle of tentacles, this being conterminous with Rotteken’s muscle, while 
“»eristome” is employed to denote the circum-oral space. The mouth is situated in the 
centre of the oral-disc. 

The ectoderm of the oral-disc has a columnar facies, with oval nuclei of varying size 
arranged along its middle. In these nuclei chromatin granules are visible, usually round 
a somewhat larger darker spot, apparently the nucleolus. Above this layer of nuclei the 
protoplasm is, as a rule, opaque, containing vacuoles usually of the mucous type, rarely of 
the granular, while below the protoplasm is somewhat transparent and finely granular. 
In many of the species deeper-stained protoplasmic areas are visible, each containing 
a nucleus; these diverging from the nuclei towards the free surface of the ectoderm. 
Round nuclei are much fewer, more or less homogeneously stained, and occur in the lower 
half of the ectoderm. Type I nematocysts (see p. 11) are present in varying numbers 
in the upper half of the ectoderm, occasionally type II also. 

The endoderm varies in its thickness and in the extent of its vacuolization, the nuclei 
being smaller than in the ectoderm and more or less homogeneously stained. Algze are 
usually abundant, sometimes massed so that the endoderm is hardly distinguishable. 

When it is considered that the convolutions of the mesenterial filaments containing 
nematocysts are frequently protruded through the oral-disc, that the peristomial ectoderm 
is provided with nematocysts and mucous vacuoles, the latter secreting an adhesive fluid, 
and that the tentacles which arise from the oral-dise are charged with batteries of 
nematocysts and also with mucous vacuoles, it is evident what an eflective apparatus the 
peristome is for capturing prey. 

* Faurot (42) takes objection to both these terms and introduces a new name “ oro-tentacular disc ” (disque 
oro-tentaculaire), which according to him makes its appearance, in Hexactinians, at the same time as the first 
eight tentacles. This name has no advantage over the older “oral-disc.” Indeed, the latter is capable of wider 
application, since in the coral genus Hydnophora—which I have recently begun to study—the tentacles are 


arranged in circles round the conical eminences or monticules, without any reference to the mouth teres which 
lie in the depressions between the monticules. 


3—2 


20 PERCY SLADEN TRUST EXPEDITION 


Column Wall (PI. 8, fig. 25 and PI. 5, fig. 53). This term denotes the vertical portion 
of body wall* outside the oral-disc. The greater part of its ectoderm (ze. the part 
lying against the corallum in the fully expanded condition) is the ealicoblastic layer already 
described. The mesoglea between the calicoblastic layer and the inner endoderm is much 
attenuated except at the attachments of the mesenteries to the corallite-walls (Pl. 2, 
fig. 22). The inner endodermal layer is usually thin in the region above the enterostome 
and contains a single layer of small more or less homogeneously stained nuclei. Towards 
the base of the polyp the endoderm becomes, as a rule, highly vacuolated and consequently 
appears much swollen, reticulated and transparent; nuclei are either absent, as in my polyps 
of Galaxea, or, when present, arranged along its free margin. Algze here are rare or quite 
absent. The mesoglea is always extremely thin over the calicoblastic layer (Pl. 2, fig. 22). 

The column then bears a strong contrast to that of Actinians, in the presence of 
the calicoblastic layer of ectoderm, the thinness of the mesoglea and the absence of an 
external or internal muscular layer. 

Edge-zone and Coenosarc (PI. 1, figs. 1, 2, and Pl. 2, fig. 24). I use these terms in 
the sense in which Professor Gardiner has defined them, viz., “The ccenosare is that part 
of the polyps in a colony which lies outside but not above (7.e. in expanded state) the thecze 
of the several corallites. The ‘ Rand-platte’ of Heider and von Koch, and ‘edge-zone’ of 
Ogilvie, is then that part of the ccenosarc which lies over the free portions of the 
corallites” (49, p. 361). It is to be noted that the distinction between the two structures 
is a physiological not a morphological one. 

The edge-zone has an outer and an inner wall, the former being an extension of 
the oral-dise, and the latter of the body-wall, and enclosed between those is an extension 
of the gastro-vascular cavity of the polyp. All the mesenteries are extended into the 
edge-zone. The structure of the outer wall of the edge-zone is more or less similar to that 
of the oral-disc, and of the inner wall to that of the column-wall. The extent of the 
edge-zone depends upon the degree of exsertness of the corallite ; in the case of Galaxea 
fascicularis (Linn.), in which the corallites project conspicuously above the peritheca, 
the edge-zone is very extensive as it covers the entire free surface of the corallite, whereas 
in Gonastrea retiformis (Lam.), in which the corallites do not project above the general 
surface of the colony, the edge-zone is absent. 

The ccenosarc covers the free surface of the peritheca, being only a continuation 
of the edge-zone with which it is similar in structure. The extent of the ccenosare 
between neighbouring edge-zones depends upon the perithecal distance between the 
corallites. In species like Goniastrea retiformis (Lam.), in which the corallites are 
so closely aggregated that there is no peritheca between them, the ccenosarc is also 
absent. 

Tentacles (Pl. 1, figs. 3, 11, 12, Pl 2, fig. 19, Pl. 8, figs. 26, 28 and Pl. 5, fig. 44). 


The tentacles arise from the oral-dise as hollow vertical outpushings of the inter-mesenteric 


* Body-wall comprises the entire wall of the polyp surrounding the gastro-vascular cavity, viz. the oral- 
dise with the tentacles above, the basal-disc below, and laterally between these the column-wall. The basal-dise 
is pushed up by the columella and paliform lobes, its ectoderm being the continuation of the calicoblastic layer 
of the column-wall and its endoderm usually vacuolated as in the lower part of the column-wall. 


MATTHAI—RECENT COLONIAL ASTRAIDA 21 


chambers; following Fowler’s terminology those from the entocceles are known as “entoccelic 
tentacles” (“tentacules loculaires” of Faurot) and those from the exocceles as “ exoccelic 


” 


tentacles” (“tentacules interloculaires” of Faurot). They do not always agree in numbers 
with those of the entocceles and of the exocceles. In the retracted condition of the polyps, 
I have not been able to make out more than two cycles—an inner and an outer—the 
former being composed of the entoccelic tentacles, and the latter of the exoccelic tentacles. 
The term ‘‘cycle”* refers to those tentacles which form a circle, 7.e. which arise from 
about the same radial distance from the mouth. In some species one of the two cycles is 
absent. When the polyps are retracted, the tentacles are usually bent over the oral-disc, 
so that, in transverse section, they are cut longitudinally; they are also capable of 
complete introversion, 7.¢. of being turned outside in. A tentacle can be distinguished, 
from any mere fold of the peristomial wall, by the swellings caused by the presence 
in its ectoderm of batteries of nematocysts. 

A large terminal battery is always present which makes the tentacular tip knobbed 
or bluntly pointed; such a battery consists of a peripheral row of closely arranged type I 
nematocysts with IIb ones interspersed among them. Much smaller subterminal batteries 
are present in most of the polyps; the number constituting a longitudinal row is of 
specific value. In these batteries only type I nematocysts are present; they spread 
outwards towards the periphery, hence their arrangement in each battery appears fan- 
shaped in transverse section. Nuclei are numerous below the peripheral row of 
nematocysts; those towards the mesoglea are round and homogeneously stained, while 
those towards the nematocysts are elongated, saccular and granular. There are no 
diverging tracts of protoplasm as in the oral-disc and edge-zone, but like the latter 
filamentar processes (neuro-muscular) pass from the nuclei towards the mesoglea. 
As described on p. 17, a longitudinal muscular layer and a granular nervous layer are 
present. 

The endoderm varies considerably in thickness from being extremely thin to being so 
thick as almost to occlude the lumina of the tentacles. 

Stomodeum (Plates 7—10). Van Beneden in 1898 questioned the use of terms like 
stomach, pharynx, cesophagus to denote the tubular structure of the Anthozoa. He 
objected also to the use of “stomodeum” on the ground that the organ in question is 
not homologous with the stomodeum of the higher Metazoa. In like manner, he held 
that any general use of the word “ mouth” would lead to confusion. Both terms are 
here retained without having recourse to new names like ““Actinopharynx ” and “Actino- 
stome.” On the other hand, van Beneden’s “enterostome” is a much needed name to 
denote the lower opening of the stomodzeum into the gastro-vascular cavity or ccelenteron. 

The stomodzum (‘‘stomatodeum” of Fowler and MeMurrich) in coral polyps 
is usually flattened from side to side in common with most of the Anthozoa, rarely 
being circular in transverse section. The mouth in many polyps forms a more or less 


* Faurot (40, p. 58), uses “cycle” in practically the same sense, 7.e. to denote the rings of tentacles, 
and “order” when referring to the sequence of the circles of mesenteries. According to him, in Actinians, the 
exoceelic tentacles, which are smaller than the entoccelic ones, constitute the last cycle but are formed at 
successive periods in the exoceeles, following the appearance of the orders of mesenteries. 


22 ae PERCY SLADEN TRUST EXPEDITION 


conspicuous rim. Over the attachments of the mesenteries the ectoderm is raised up into 
ridges which differ in width, thickness and shape in the different species, these characters 
being of specific importance. By the presence of these ridges, longitudinal furrows 


? 


(“sillons actinopharyngiens” of van Beneden) are formed. In the genera of Group I 
below, characterised by the presence of two couples of directive mesenteries and both 
bilateral and radial symmetry*, two longitudinal grooves are present facing the directive 
couples of mesenteries, being usually deepened by the folding of the stomodzeal wall into 
the directive entocceles. The stomodzeal ectoderm is ciliated, but the cilia do not appear 
to be longer in either of the grooves, nor is the ectoderm lining them specially thickened. 
Although these grooves are not so pronounced in my sections as the so-called sulcus 
of Peachia or the sulcus and sulculus of Bunodes, they are not less evident in transverse 
sections than the same grooves in some examples of Hdwardsia and Zoanthus that I have 
examined (they are well seen in my sections of Hchinopora lamellosa, Leptastrea 
roissyana and Galaxea fascicularis). On the other hand, Duerden denies the existence 
of any such groove (‘“‘siphonoglyph”) in the Madreporaria (21, p. 23); Bourne also is 
inclined to take the same view. Duerden further says ‘a siphonoglyph is generally 
wanting only in the lowest actinians and in alcyonarians, and its absence in coral polyps 
would suggest their more primitive nature.’ These two grooves are here termed directive 
groovest, as they le in the directive entocceles; none of the other names hitherto 
employed to denote them in the Anthozoa are suitable for coral polyps. 

Great confusion prevails over the orientation of Actinian polyps and in particular 
over the terminology employed to denote these two grooves. Hollard in 1851 (p. 274) 
referred to them as ‘“‘deux demi-canaux” ; Gosse in 1860 (p. 4) termed them “ gonidial 
grooves (canales gonidiales)”; the Hertwigs in 1879 distinguished them as “ ventral” 
and ‘dorsal,’ following upon Kolliker’s (1872) use of “ventral” for the side of a 
Pennatulid polyp turned towards the stem and “dorsal” for the opposite side. Following 
Haddon objection may well be raised against the extension of these terms to the 
Actinie. In 1884 Andres (p. 73) substituted “gonidium” and “gonidulum” for 
Gosse’s gonidial grooves. Hickson in 1883 (p. 693) invented “siphonoglyphe” for the 
ciliated groove of Aleyonarians which is said to be homologous with the single groove of 
Peachia and the ventral grooves of other Actinians; Parker (110) in a foot-note on p. 260 
drops the final e with Hickson’s assent, as etymologically unnecessary ; both he and 
McMurrich apply “siphonoglyphs” to the two grooves in Actinians. 

Haddon in 1889 (p. 300) introduced “sulcus” for the more important groove of 
Actinians (viz. ventral groove of the Hertwigs) and “sulculus” for the opposite (dorsal) 
one. Haddon’s nomenclature, which has been accepted by Bourne, van Beneden and 
others, is as objectionable as the Hertwigs’ “ventral” and “dorsal” when applied 
to adult Actinians in which there is no recognisable difference in the size of the two 
grooves. The distinction between ventral and dorsal, sulear and sulcular, may be 


* Boveri (18) uses biradial symmetry to denote the combination of both the bilateral and radial symmetry 
in Actinians,. 

{ Perhaps Faurot’s commissural grooves (‘‘sillons commissuraux”) and commissural mesenteries may be 
preferable to “directive grooves” and “ directive mesenteries.” 


MATTHAI—RECENT COLONIAL ASTRAIDA 23 


appreciated in the case of forms like Edwardsia, where the longitudinal muscles of all the 
non-directive mesenteries are turned to one pole, but in adult coral polyps, with the 
hexameral arrangement of the mesenterial couples and with no appreciable difference 
in the relative size of the two grooves, it is impossible to distinguish one pole from 
the other. Parker (110, p. 268) pointed out the same difficulty in the “diglyphic” forms 
of Metridium. In my Group II, in which neither directive mesenteries nor directive 
grooves are present in the adult condition, the distinction breaks down. Reliable criteria, 
if there be any for determining one pole from the other, can be expected only from 
a comparative study of the developmental sequence of the mesenteries. Van Beneden, 
homologising a cerianthid larva with the segmented larva of Amphzoxus or the embryo of 
Peripatus, substituted anterior for ventral, posterior for dorsal, the oral-disc being 
regarded as dorsal or “face neurale.” Faurot, after his exhaustive embryological studies 
of Hexactinians, prefers to use ventral and dorsal instead of sulcus and sulculus. 

In my Group II, in which directive mesenteries and bilateral symmetry are wanting, 
no two grooves are distinguishable from the others, but in many polyps the grooves appear 
(in transverse section) on the whole deeper than in Group I, owing to the folding of the 
stomodzeal wall into the imter-mesenteric chambers. G. Y. and A. F. Dixon (27—80), 
Carlgren (20), McMurrich (98, 99) and Parker (110) have observed variations in the 
number of directive grooves above and below their normal number (also total absence) 
within the same species of certain genera, viz. Sagartia, Bunodes, Metridium, and have 
correlated it with a corresponding variation in the number of directive couples of 
mesenteries. Parker and McMurrich studied this variation more thoroughly in the case 
of Metridium marginatum and Sagartia spongicola, emphasising the numerical corre- 
spondence between directive couples and grooves. Perhaps a similar relation subsists 
in coral polyps also, as will be seen by a comparison of Group I with Group II. 

The stomodeeal ectoderm has a faintly striated border below the row of cilia. Its 
nuclei are round, rod and spindle-shaped, and more or less homogeneously stained dark ; 
they are massed together in each ridge below the striated border, around a deeper, some- 
what granular region, which is almost devoid of nuclei. The nature and arrangement of 
the nuclei give this ectoderm a characteristic appearance. Through the central proto- 
plasmic region, slender filaments generally pass backwards into the mesoglea. Nematocysts 
are never numerous; when present, they may belong to any of the three types. Small 
vacuoles, as described on p. 7, are usually present in the peripheral protoplasm above 
the layer of nuclei. 

The endoderm is, as a rule, extremely thin, and alge are comparatively rare. The 
mesoglea may or may not be thickened at the attachments of the mesenteries ; when 
thickened, it projects into the ridges. 

Mesenteries. In coral literature there has been a great confusion in the terminology 
employed to denote the soft radial partitions of the polyps and the calcareous partitions of 
the corallites, “septa” having been used for both the structures by some English and 
German authors and ‘‘cloisons” by French authors. The name “mesenteries” (“les 
méséentéroides” of Lacaze Duthiers) is now well established for the radial partitions of the 


* A trace of bilateral symmetry is however left in Group II in the lateral compression of the stomodzum, 


24 PERCY SLADEN TRUST EXPEDITION 


polyp, and “septa” (‘“cloisons”) is restricted to the calcareous radial partitions of the 
corallites. 

Hollard in 1851 described at length the arrangement of the mesenteries of Actinians 
in two's, with their respective longitudinal muscles facing each other; this arrangement 
has since been recognised in all Hexactinians. 

In polyps of Group I the couples of mesenteries are arranged in alternating cycles, 
each cycle consisting of six couples of almost uniform width or of a multiple of six, the 
mesenteries of every succeeding cycle being narrower than those of the preceding one. In 
the polyps examined thus far, up to four cycles of mesenteries are present, termed 
respectively primary, secondary, tertiary and quaternary. The so-called “directive” 
couples* of mesenteries belong to the primary cycle, and are situated at the ends of the 
longer diameter of the stomodzeum, each enclosing a directive groove ; their arrangement 
of the mesogleal pleats with muscular layers is reversed f. 

For such a disposition of the mesenteries in the adult polyps I prefer to use the term 
“couples,” and “pairs” when referring to their developmental sequence. The word 
“eycle” is used to include all the couples of mesenteries having more or less the same 
radial extent, irrespective of their time of development, while “order” has been employed 
by some authors (Faurot) to denote the mesenteries which have appeared at about 
the same time to form a circle, e.g. primary order, secondary order, ete. The couples of 
mesenteries composing the first cycle are usually “complete,” 1.e. they extend from the 
column-wall to the stomodzeum; when one of the mesenteries of a couple does not reach 
the stomodzeum, the couple is spoken of as “incompletet.” 

In Diploastrea heliopora (Lam.) the primary cycle consists of twelve couples of 
mesenteries, which may be due to the true secondary couples having grown and met the 
stomodzum. Most polyps of Group I possess the full number of secondary couples, 
viz. six; in a few polyps the tertiary cycle is also complete with twelve couples, while 
quaternary couples of mesenteries have only occasionally been met with, viz. in Galaxea 
fascicularis (Linn.), the cycle itself being quite imperfect. Both Lacaze Duthiers and 
Faurot have shown that in the Hexactiniz, while the primary couples are formed in pairs 
on each side of the sagittal plane, the couples of the subsequent cycles are formed in the 
exocceles. . 

; In my Group II directive couples of mesenteries are absent, nor is there a sequence of 
mesenterial cycles as in Group I. The only distinction that could be made between the 
couples is between those that meet the stomodzeum and those that do not reach it; the 

* The term “directive” mesenteries (‘‘ Richtungssepten”) was first employed by the Hertwigs, as they 
believed that these two couples might be of use in fixing the orientation of an Actinian polyp; Schneider and 
Rotteken (114) had carefully described their arrangement in 1871. Lacaze Duthiers in 1897 (p. 221) doubted 
the suitability of Hertwigs’ terminology, as the two couples in question did not indicate any direction in the 
polyp. 

+ MeMurrich (99, p. 132) suggests that the reverse arrangement of the muscles on the directive mesen- 
teries may have ‘something to do with the preservation of the lumen of the siphonoglyphe when the lips of the 
stomodzeum elsewhere are in contact.” 

{ Faurot and van Beneden use “complete” and “incomplete,” not with reference to the couples or pairs, 


but to the individual mesenteries that either meet the stomodeum or do not reach it. The adjectives “ perfect ” 
and “imperfect” have also been used in the same sense. 


MATTHAI—RECENT COLONIAL ASTR AIDA 25 


former I term ‘‘principal” couples and the latter “subsidiary” couples. In any species of 
this group, the number of principal couples varies within narrow limits, while that of the 
subsidiaries may vary considerably ; sometimes the latter may be entirely wanting between 
two adjacent principal couples. Moreover, some of the principal couples may be incom- 
plete. The subsidiary couples differ regularly in their radial and vertical extent, some 
of them being very narrow and short; two mesenteries constituting a subsidiary couple 
may also differ in their relative size. This variation is no doubt due to the differences in 
the time of formation of the mesenteries. The number of subsidiary mesenteries in any 
exoccele is also liable to great variation™. 

All the mesenteries are attached above to the oral-dise, and along their outer margins 
to the corallum, by means of the wedge-shaped mesoglwal processes, but they vary con- 
siderably in their depth, the primary and principal mesenteries reaching very nearly to the 
base of the polyp. The arrangement and structure of the mesenteries, and the numbers 
of cycles and of the couples composing them are of considerable value in the classification 
of corals. Following Fowler's terminology (44, p. 578), the part of the gastro-vascular 
cavity included within a couple is known as an ‘‘entoccle” (‘“loge” of French authors) 
and that between two neighbouring couples as an “exocele” (“interloge” of French 
authors). 

As the two directive couples and grooves are situated at the ends of the longer 
diameter of the stomodzeum and as the mesenteries are arranged on the hexameral plan, 
the polyps of Group I are divisible into two similar halves along this plane, viz. the 
sagittal (“plan commissural” of Faurot, p. 59, and ‘‘plan médian” of van Beneden, p. 12); 
they may also be divided into two similar halves along a plane at right angles to the 
sagittal, viz. the transversal (“plan médian” of Faurot). In some polyps the bilateral 
symmetry is not quite perfect owing to the incompleteness of the last cycle of mesenteries. 
Typically, however, Group I may be regarded as bilaterally symmetrical along two planes. 
Their mesenterial couples are also arranged radially with reference to the oro-aboral axis 
(‘‘axe-vertical” of Faurot, “axe oro-aboral” or “axe du corps” of van Beneden), passing 
through the centre of the mouth at equal distances from the two directive grooves. Like 
the couples of mesenteries the disposition of the tentacles may also be assumed to be 
symmetrical both bilaterally and radially, although from preserved polyps I have not been 
able to settle this point. 

In Group II, on the other hand, the bilateral symmetry is wanting owing to the 
absence of the directive couples of mesenteries and of the grooves ; a trace of the bilateral 
symmetry is, however, left in the lateral compression of the stomodzum. The radial 
symmetry is interrupted by the indefinite number and the irregular width of the 
subsidiary couples of mesenteries and by the incompleteness of some of the principal 
couples. In Gyractis, Boveri (19) regarded the absence of directive mesenteries as only 
apparent, their presence being disguised by the development of a mesentery on either side 
of each of the two Edwardsian directive couples, but McMurrich (99) believed that the 
disappearance of the directives may be due to “the mesenteries which really represent 

* Parker (110) has made similar observations on the numerical variation and on the irregular arrangement 
of the incomplete mesenteries in the “ monoglyphic” forms of Metridium marginatum. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 4 


26 PERCY SLADEN TRUST EXPEDITION 


them having developed their longitudinal muscles on adjacent faces” (p. 119). Which of 
these two views is applicable to the forms included in Group II can be determined only 
after making a comparative study of their development. 

Faurot (40, 42) applied the term “couple” to two mesenteries that appear at the 
same time, one on each side of the sagittal plane, and ‘‘ pair” to two adjacent mesenteries, 
which lie on the same side of the sagittal plane, facing each other. According to this 
nomenclature the twelve primary mesenteries are formed in couples which appear arranged 
in pairs in the adult, whereas the mesenteries of the succeeding orders are formed in pairs, 
one pair in each exoccele. In 1900, Bourne (16), on the other hand, used these words in 
exactly the opposite sense ; in a footnote on p. 39 he added that “it is convenient when 
speaking of the adult arrangement of the mesenteries to use the word ‘couple, when of 
their developmental sequence to use the word ‘pair.’” But in 1905 (17). he reverted to 
Faurot’s use of “pair.” Bourne's original use of ‘“‘ pair and couple” is, however, to be 
preferred when speaking of forms which are bilaterally symmetrical, and since the 
Anthozoan polyp is said to have a primary bilateral symmetry, “pair” is to be used to 
denote the mesenteries that appear on either side of the sagittal plane, and couple when 
referring to their adult arrangement*. The mesogleal pleats of the mesenteries are either 
simple or sub-divided and vary in their shape, size and radial distribution in the different 
species. In the stomodzal region of the polyp the mesenterial mesoglea is devoid of 
entoccelic pleats for a variable distance from its stomodzeal attachment. Below the 
stomodzeum, the entoccelic pleats usually become broader and cover nearly the whole of 
the width of the mesentery. In a few species, e.g. Favia favus (Forsk.) (Pl. 4, fig. 36) 
and EHchinopora hirsutissima (Kd. and H.) (Pl. 2, fig. 20), short exoccelic pleats are present 
to a short distance from the stomodzal attachments of the primary mesenteries, on which 
the muscle-strands are better developed; this may perhaps be because the polyps in this 
species have assumed an oblique position. In the polyps of Mycediwm okent Ed. and H. 
that I have examined the exoccelic pleats are much broader and thicker than the entoccelic 
ones and extend over the inner half or two-thirds of each principal mesentery, the latter 
being as a rule restricted to its outer quarter. This peculiarity may be due to the 
completely horizontal position which the polyps have assumed in that species. The 
mesoglea is not of uniform thickness along the entire width of the mesentery. 

Though the contraction of the polyps may affect the condition of the mesenterial 
mesoglea, I find its characters, viz., relative thickness, the shape, size and distribution of 
the pleats (both entoccelic and exoccelic), retaining a specific constancy. The Hertwigs and 
the Actinologists after them attached the same value to the mesogleal pleats of the oral- 
dise. The endoderm is usually vacuolated. The vacuoles when definite are either oval or 
goblet-shaped (Pl. 1, fig. 4) and, as I have suggested, may be capable of ingesting and 
digesting food-particles. The endoderm is not usually of even thickness along the entire 
width of the mesentery. Its relative thickness varies in the different species. Behind 
the attachment of the mesenterial filament, the endoderm is usually swollen, pad-like in 
transverse section, and is constricted off from the filament; its two wings were termed 


* H. V. Wilson (122) used “ pair” with reference to both the developmental history and the adult arrange- 
ment of the mesenteries. i 


MATTHAI—RECENT COLONIAL ASTRAIDA 27 


“ mesenterial lobes” by H. V. Wilson, which he regarded ‘“‘merely as a device to support 
the filament” (p. 225). 

A short distance below the stomodzeum, the margin of each primary mesentery is 
drawn out into long ribbon-shaped prolongations which, with the filament continued along 
their free edges, lie coiled together in the gastro-vascular spaces. The free margins of 
mesenteries which do not meet the stomodzum are also drawn out into such processes. 
When these processes are cut across they appear as bars of mesenterial tissue, usually 
curved, with the filament on either side of each bar (PI. 3, fig. 30). They are also found 
extended into the inter-mesenteric spaces of the edge-zone but, as a rule, are scarce in the 
region above the enterostome, while below it they may be massed together in the gastro- 
vascular chambers. These convolutions are often found in the peristome, protruded 

through pores (? artefacts) in its floor (this observation is based on the preserved material) ; 

they may also be ejected through the mouth, but this is less frequently the case as 
in sections I have noticed them in the stomodza of only Echinopora and Fava ananas 
(Ell. and Sol.). 

Mesenterial filaments (PI. 1, fig. 6 and Pl. 3, fig. 30). Every mesentery which 
meets the stomodeeum and most of the others bears a filament (the “ craspedum ” of Gosse 
and ‘“‘enteroide” of Faurot) on the free margin of its straight region and along the entire 
edge of the coiled ribbon-shaped processes into which the mesenterial margin is prolonged. 
The filament terminates a short distance from the insertion of the mesentery to the basal 
disc. The structure of the filament can be best made out in a primary mesentery. It 
consists of a median part and two lateral extensions (the “main body” and the two 
“ventro-lateral tracts” of H. V. Wilson), the former is the continuation of an ectodermal 
ridge of the stomodzeum, while the latter arise from the stomodzeal ectoderm on each side 
of the ridge. The lateral extensions lie like two flaps over the mesenterial endoderm to 
a short distance below the stomodzeum (PI. 1, fig. 5). The stomodzal ectoderm is also 
reflected upwards to some distance, this being continuous with the downward lateral 
extensions referred to. In the region of this upward reflection, the stomodseal endoderm 
comes to lie between two layers of ectoderm. 

The mesoglea of the stomodeeal wall, however, is continued upwards and downwards 
along with the ectoderm, separating the latter everywhere from the endoderm. ‘The 
lateral extensions are soon tucked inwards and come to lie behind the median tract. The 
mesoglea in the filament then appears T-shaped in transverse section, the handle of the T 
is the termination of the mesenterial mesoglea, while the two arms are due to the 
downward extensions of the stomodzeal mesoglea. The so-called forking of the margin of 
the mesoglea is therefore not a case of bifurcation at all. In the earlier sections the 
filament appears somewhat triangular while, lower down the straight region, it becomes 
more consolidated and appears hemispherical or almost circular in transverse section. The 
straight region of the filament is ciliated. The nuclei of the filament are of the same 
nature as in the stomodzeal ectoderm, massed together concentric to the periphery of 
the filament, incomplete at its base where the mesoglea enters the filament. In the 
median part of the filament granular vacuoles are usually present, their granules varying 
in size. 

49 


28 PERCY SLADEN TRUST EXPEDITION 


Nematocysts are restricted to the median region of the filament, usually somewhat 
scarce in its straight region, but when present may belong to any of the three types 
previously described ; in the coiled region, they occur in large numbers, usually arranged 
close together in the form of batteries (figs. 17 and 59). Type I is less frequently present 
than II and III. From histological appearances it seems probable that the straight 
region takes part mainly in digestion, while the convoluted region is for offensive or 
defensive purposes. This suggestion may account for the fact that the convolutions are 
frequently found protruded through the oral-dise and through the stomodzeum. On some of 
the mesenteries that do not reach the stomodzeum filaments are rudimentary or even absent. 

H. V. Wilson regarded the mesenterial filament of Manicina as equivalent to the 
trilobed filament of an Actinian as described by the Hertwigs, the ventro-lateral tracts 
corresponding histologically with the lateral ciliated bands (“ Flimmerstreifen”) and the 
median portion with the central glandular lobe (‘‘ Nesseldriisenstreif”), whereas according 
to Duerden an entire coral filament represents only the middle lobe of the trifid Actinian 
filament and that in the former “there is nothing which corresponds morphologically with 
the lateral lobes” (82, p. 472). 

Various views have been held on the origin of the filaments in the Anthozoa. 
H. V. Wilson in 1889 concluded that the filaments were ectodermal structures, those of 
the primary mesenteries being extensions of the stomodeeal ectoderm along the free 
mesenterial margins, while those of the incomplete mesenteries were formed by the 
reflection of the stomodeeal ectoderm towards the oral-dise and thence across its inner 
surface to the free edges of the mesenteries. He further held that the three lobes of an 
Actinian filament were also ectodermal in origin and suggested that they could be derived 
from the simpler larval filament of Mancina; in the latter the two lateral ciliated tracts 
and the median secretory tract are already differentiated ; “to produce the trifid filament 
it is only necessary for these tracts to become separated by the division of the mesoderm 
(=mesogleea) into three lobes” (122, p. 228). KE. B. Wilson in 1884 had regarded the 
‘“‘Flimmerstreifen ” of the Hertwigs to be ectodermal and homologous with the two longer 
dorsal filaments of an Aleyonarian, which were downgrowths of the stomodeeal ectoderm, 
provided with long cilia but with no gland cells and functioning as organs of circulation ; 
the same author believed the “ Nesseldriisenstreiten” to be endodermal and homologous 
with the remaining six shorter Alcyonarian filaments which were thickenings of the 
mesenterial edges containing gland cells and subserving a digestive function. MceMurrich 
in 1891 disagreed completely from H. V. Wilson’s and favoured E. B. Wilson’s view. 
According to him the median tract of a filament of Aulactinia was formed first by a 
differentiation of the endoderm at the free margin of the mesentery while the two lateral 
lobes were downgrowths of the stomodzeal ectoderm*; he further suggested that the same 
order was followed in the phylogenetic history of the filament. In a later paper, while 
still maintaining that the median region and the lateral tracts arose independently in 
ontogeny, he did not regard the ectoderm and endoderm of Coelenterates as having 
reached in phylogeny a degree of differentiation equal to that of the epiblast and hypo- 


* KH. B. Wilson’s figs. 20 and 23, transverse sections of the embryonic filaments of Awlactinia, are similar to 
those of adult coral filaments. 


MATTHAI—RECENT COLONIAL ASTRASID A 29 


blast of vertebrates, hence his use of “ectodermal” and “endodermal” was “merely for 
convenience and not as expressing a definite homology ” (100, p. 271). 

Gardiner in 1899, from a histological study of the filaments of Cenopsammia, also 
came to the conclusion that they were entirely ectodermal structures, and supported 
H. V. Wilson’s contention based on embryological data; he added further that “the 
lateral parts of the mesenterial filaments are similar in structure to the ‘ Flimmerstreifen ’ 
of the Hertwigs and have apparently the same function” (49, p. 375). Duerden, from 
his study of West Indian Madreporarian polyps, inferred that their filaments were 
entirely endodermal, the histological resemblance between them and the stomodeeal 
ectoderm being a secondary feature. His reasons for this view were that filaments were 
present on incomplete mesenteries which never reached the stomodeeum, that the reflected 
stomodeeal ectoderm could not be traced up to the oral-dise and thence to the free edges 
of the incomplete mesenteries, that in the upper regions of the latter filaments were absent 
or incipient while they were well developed below, that filaments appeared independently 
on some of the primary mesenteries before the latter reached the stomodzeum. Duerden 
regarded the “reflected ectoderm” as “the stomodmal ectoderm passing along the 
mesentery to establish structural continuity with the upwardly growing filament” 
(pee): 

The appearances in my sections favour the view of the ectodermal nature of coral 
filaments held by H. V. Wilson and Gardiner. In Cyphastrea chalcidicum (Forsk.) 
I have found traces of the upwardly reflected stomodeeal ectoderm as far as the oral-disc ; 
perhaps this ectoderm might be continued to the secondary mesenteries and finally course 
down their free margins as their filaments. Of course, questions like these could be 
settled only by a study of:the development of coral larvee™*. 

Acontia and cinclides. Conflicting accounts have been given by different authors 
with regard to the presence or absence of these organs in coral polyps. Fowler in 1884 
and Bourne in 1887 at first applied the term acontia to the prolongations of the 
mesenterial margins in Flabellum and Fungia respectively, and found them capable of 
being protruded through permanent openings (cinclides) in the oral-disc, the former author 
regarding “the contorted mesenterial filaments” of Moseley as identical with the acontia 
of Gosse; Bourne in 1888 withdrew his application of the term “acontia” to these 
prolongations. H. V. Wilson could find no free acontia in Manicina, but noticed that 
the filaments which were attached along their whole length to the mesenteries could be 
extruded through the mouth and through pores in the body-wall. With regard to 
Cenopsammia, Gardiner remarked that “the filaments are attached to the mesenteries 
for their whole length, and are without free portions (acontia) at their lower ends” 
(p. 367), whereas in the Turbinolids he examined in 1904 ‘the lower ends of the filaments 


* Referring to the probable formation of the stomodeum in Flabellum, Gardiner says, “The only logical 
method of conceiving the formation of the stomodeum of the adult to take place is to suppose that the external 
body-wall grows inwards catching up the edges of the mesenteries in its progress. It finally reaches the 
mesenterial filaments, which by fusion together, assisted by the down-growth of the body-wall form the 
stomodeum” (p. 151). My examination of Gardiner’s “young polyp” of Flabellum (figs. 3 and 4) confirms 
the facts on which his conclusions are based, viz. that it had no trace of tentacles or stomodzeum and that 
filaments were present on the margins of the mesenteries. 


30 PERCY SLADEN TRUST EXPEDITION 


commonly form coiled masses, the acontia” (52, p. 123). Duerden could see no 
“cinclides” in the living coral polyps he examined, but observed that the filaments with 
the contorted edges of the mesenteries could be protruded through the mouth or through 
temporary openings on any part of the column-wall or oral-disc, but that such extensions 
were different from the acontia of Sagartia. 

was first applied by Gosse to the threads in the Sagartide, 


oy 


The term “acontia ’ 
which he found protruded through special openings (which he named “ cinclides”), but 
gave no structural characterization of the threads. They were re-described by the 
Hertwigs, but the exact relation between their acontia and “ mesenterialfiden” (66, 
Pl. 8, figs. 10 and 12) is not evident. Faurot in 1895 remarked that an acontium 
has “a peu pres la méme structure histologique que l’enteroide,” and that it was for 
augmenting the digestive surface of the mesenteries and “non d’étre utilisés uniquement 
comme arme défensive” (40, p. 53). In 1897 van Beneden defined them as arising from 
the mesenteries between the terminations of the mesenterial filaments and the aboral ends 
of the polyps, and gave as the essential difference between the two structures that while 
the epithelia of acontia were endodermal, those of the filaments were ectodermal, 
being continuations of the stomodzeal ectoderm. According to Duerden, “ acontia (in the 
Sagartidee) are thread-like structures, which are but feebly attached to the mesenteries, 
and pass through permanent apertures (cinclides) in the column wall of the polyps, or 
through the mouth, the mesenteries in no ways following. If not wholly liberated from 
the polyp, the acontium can be indrawn. The extruded filaments of corals, on the other 
hand, still retain their normal position along the contorted edge of the mesentery, and 
a portion of the latter passes out along with them. The function of both is probably the 
same, as in each case the organs are strongly charged with nematocysts, and less so with 
gland cells” (82, p. 476). 

Is the name ‘‘acontia,’ then, to be restricted to thread-like structures, which are 
attached by one extremity to mesenteries below the terminations of the filaments and 
endodermal in structure, and is their presence always correlated with that of “ cinclides”? 
In my coral polyps I have not satisfactorily investigated if among the convolutions of the 
mesenterial filaments, which are so often protruded into the peristome, there are any 
threads comparable to the acontia of Sagartide. 

Reproductive Organs. Mature ova have been seen in many of the polyps 
examined (Pl. 1, fig. 10, Pl. 3, fig. 29, and Pl. 5, fig. 49). They are carried by all the 
mesenteries or by the primaries only, forming one to five or six longitudinal rows. The 
ripe ovum is large, varying in size in the different species, enclosed in a thin vitelline 
membrane; its cytoplasm is highly vacuolated, the vacuoles being somewhat small and 
round and the nucleus large, stained pink, excentrically situated, with a dark-stained 
nucleolus on one side. It lies in the mesoglea usually on its exocoelic side ; the mesoglea 
surrounding it is narrowed to a thin membrane, while the mesenterial endoderm over it is 
usually swollen and granular, perhaps for nutritive purposes. 

Surrounding the large ova are frequently seen some small cells which, for reasons 
given below, I take to be germ-cells. These lie in the mesoglea or in the surrounding 
mesenterial endoderm, each consisting of a large spherical nucleus—similar to those of the 


MATTHAI—RECENT COLONIAL ASTRASID AG 3 31 


ealicoblastic layer—containing usually a dark-stained spot and surrounded by a little 
irregular cytoplasm; sometimes a network may be discerned within the nuclear mem- 
brane*. 

In two polyps of Cyphastrea seraila (Forsk.) certain structures are present in the 
same position as the ova, which I am led to conclude are mature testes (Pl. 1, fig. 9). 
Each of these is circular in transverse section or laterally compressed, and contains 
innumerable minute triangular-shaped spermatozoa stained dark in iron-hematoxylinf. 
Germ-cells, similar to those surrounding the eggs, are also seen scattered around the testis. 

In some female polyps small groups or follicles—round or adpressed—of germ-cells 
are found to lie in the mesenterial mesoglea, behind, in front of, or on each side of the 
large ova; the cells being either massed together or surrounding a space in the centre of 
the follicle. In a polyp of Cyphastrea seraslia (Forsk.) (from the same colony from which 
the male polyps were taken) in which ova were found in three of the primary mesenteries 
arranged in single rows, germ-cells were present massed together in small follicles and 
also surrounding the eggs (PI. 1, fig. 10). In a polyp of Cylucia in which ova were seen 
in most of the mesenteries in one or more usually two rows, the follicles lay mostly behind 
the ova, each usually with a space in its centre. In some serial sections of Gardiner’s, 
named Prionastrea abdita, ova of varying size were to be seen in most of the mesenteries, 
arranged in rows up to six or eight, along with as many groups of germ-cells. In another 
polyp of Cylicia a number of such follicles { were present in the mesenteries, and occasion- 
ally structures resembling the mature testes in Cyphastrea serailia; in some of the 
follicles both germ-cells and spermatozoa appear to be present ; no ova could however be 
found anywhere in this polyp. 

From the above observations I am led to make the inference that both ova and 
spermaries may develop from the same kind of follicles, that an ovum is only one of the 
germ-cells in a group which has grown at the expense of the others§, in support of which 
it may be added that small ova were to be found in the midst of a few of the follicles. 
The cells that invariably surround the ova are, on this suggestion, to be regarded as the 
remnants of the original follicles (Pl. 5, fig. 49). Further, the case of Cyphastrea seraulia 
suggests that both male and female polyps may occur in the same colony at the same time. 

In the case of Flabellum, Gardiner regarded such groups as composed of “ sperma- 


togens” and a few ova which he noticed along with the follicles as due to protandry 


* J have found similar germ-cells surrounding some of the ova in Gardiner’s sections of the polyps of 
Cenopsammia and Flabellum. 

+ The appearance is similar to Ashworth’s figure of a spermary of Xenia hicksoni (2, p. 245, pl. 27, fig. 33), 
and to van Pesch’s figures of testis-follicles from the mesenteries of Stichopathes (111, pl. VII, figs. 2 and 4) ; 
the latter author describes the spermatozoa as possessing long tails, and observed them even in the tentacles, 
but he has not given figures of isolated spermatozoa showing the tails, Lacaze Duthiers figured mature 
spermatozoa from Cladopsammia roland, which have a triangular head with a long flagellum proceeding from 
the centre of its base (92, pl. 11, fig. 7). In the spermatozoa of Cyphastrea serailia, perhaps the tail has been 
~ lost or has not yet developed. 

{ Professor V. H. Blackman after examining these bodies tells me that they do not show any plant 
characters and cannot be spores of alge. 


§ Hickson (70) has also made a similar observation of ova developing in groups of cells in Alcyoniwm 
digitatum. 


32 PERCY SLADEN TRUST EXPEDITION 


in the genus. Ina later paper (51) he further suggested that coral polyps are male when 
young, becoming female as they get older, that all the members of a colony are either 
male or female at the same time. I have looked over his sections of Mlabellum and have 
found the cell-groups he has described to be identical in appearance with those in the 
polyps of Cylicia and Prionastrea abdita already referred to. If Professor Gardiner’s 
suggestion be confirmed, then a species like his P. abdita will have to be regarded as 
hermaphrodite, since ripe ova were present in the same mesenteries along with the 
testicular masses. If, on the other hand, the bodies I have termed spermatozoa really 
represent the final stage in spermatogenesis and not simply artefacts*, then the follicles in 
Flabellum would not be testes but groups of primitive sex-cells from which ova were 
ultimately to develop. Indeed, the stringed arrangement of these groups and of the ova 
are similar. Fresh ova would develop from these groups as the ripe ones were discharged. 
When all the possible ova have thus been formed, the follicles would altogether disappear, 
as happened to be the case in the two largest polyps of Flabellum which Gardiner 
examined. All the remaining polyps which had only the follicles would then be immature 
females. On such an interpretation none of Gardiner’s polyps could be regarded as 
protandrous+. However, on questions like these, no definite conclusions are possible till 
fresh polyps are actually examined. 

The similarity between the nuclei of the germ-cells described above and those of the 
calicoblastic layer of ectoderm has already been noted. Around some of the calicoblast- 
nuclei a thin coat of cytoplasm appears to be differentiated. Certain appearances further 
suggest that these cell-elements migrate into the mesenterial endoderm and ultimately 
into the mesoglea to form the above-mentioned folliclest. If so, this phenomenon, 
together with some of my observations on the possible migration of nematocysts from the 
ectoderm into the endoderm, will have considerable theoretical importance§. ; 

Zooxanthelle (Pl. 1, figs. 1, 4, and Pl. 6, fig. 51, etc.). All the polyps I have examined 
contain the so-called zooxanthellae in greater or less abundance, but they are invariably 
restricted to the endodermal layer. Each of these algze is round, the protoplasm staining 
pink, the nucleus excentrically placed and granular in appearance. In addition, there is, 
in most algze, a homogeneously dark-stained body—in all probability a pyrenoid—with a 
transparent ring round it. Algee are most abundant in the edge-zone and the peristome, 
often completely fillmg the endoderm, and to a less extent in the tentacles, 7.c. in the 
exposed regions of the soft parts. In a species like Galaxea musicalis, in which the 
stomodzeum seems to be imperfectly functional, the zooxanthelle are most numerous. This 
fact may be taken as an additional indication of the symbiotic nature of these alge. In 
the mesenteries they are more numerous in the non-pleatal than in the pleatal sides, which 
may be accounted for by the action of the strong musculature on the pleatal sides. 

* The definite shape of these bodies and their massed arrangement in the mesenterial mesoglea (in 
exactly the same position as the ova) do not warrant their being regarded as artefacts. 

+ Duerden’s account of the gonads in the West Indian coral species which he studied is practically the 
same as Gardiner’s, but he thinks that ova are developed first and spermaria later. His Pl. 20, fig. 140, is 
similar to the cases I have recorded. 


{ Boulenger (10) has traced the origin of sex-cells in craspedote medusz to interstitial cells of the ectoderm. 
§ The sex-cells in coral polyps have hitherto been regarded as arising from the endoderm. 


MATTHAI—RECENT COLONIAL ASTRAIID A 33 


Ill. CLASSIFICATION 


Gemmation and Fissiparity. ‘The use of these terms is in a state of great confusion 
in coral literature. This is due to workers on the hard parts trying to set up artificial 
distinctions between genera and species with regard to the exact mode of multiplication 
of their corallites. In this way various types of budding have been enumerated, viz., 
marginal, apical, intra-calicular, stolonal, coenenchymal, etc., all of which can be grouped 
under two heads, viz., (1) wtra-calicinal budding, i.e. budding inside the calyx; and 
(2) extra-calicinal budding, v.e. budding outside the calyx. The latter is easy to see, 
while the former can hardly be distinguished from fission. Olgivie admits that intra- 
calicinal budding appears like fission, but adds that “fissiparity is simply a hastened 
development of buds under certain conditions and at certain periods of “ vegetative 
growth” in the life of the coral (108, p. 165). 

Duerden recognises the value of the different types of intra-calicinal budding, but 
distinguishes them all from fission, by which term he understands complete stomodeal 
division into two equal or unequal parts (32, p. 513). I have searched in vain among his 
many figures for one representing a stomodzum in the actual process of division, a phe- 
nomenon which I have not found in any of my polyps*. Moreover, Duerden maintains 
that there is a sharp morphological distinction between budding and fission, that in 
gemmiferous corals the polyps which arise as buds pass through the same stages as those 
that develop directly from larvee and ultimately possess two directive couples of 
mesenteries and a cyclical hexameral arrangement of the mesenteries, whereas the 
products of fission in fissiparous genera have neither the directive couples nor the cyclical 
arrangement. Exceptions are made in the cases of Porites and Acropora (Madrepora), in 
which fission does not deprive the polyps of directives nor of the cyclical disposition of the 
mesenteries. Later the same author uses the term “ fissiparous gemmation” to denote 
occasional cases of fission in gemmiferous genera like Cladocora, Stephanocenia, and 
Solenastrea, in which he noticed that the fission products of certain large polyps, 
possessing more than six couples of mesenteries in each of the two mesenterial cycles, 
retained the two directive couples and the normal hexameral arrangement. On this 
phenomenon he remarks: “I conceive that gemmation may occasionally take place at 
almost any part of the free polypal wall, from the disk as well as from the column-wall ; 
and, if within the disk, then also around the oral aperture. In this last case the one 
mouth and stomodzum would be common to the bud and to the parent ; the mesenteries 
of the one would intermingle with those of the other ; two additional pairs of directives 
would be developed, as in all other buds; and the mesenteries as a whole would have the 
same ordinal and cyclic value as in buds arising on the column-wall. When the bud 
reaches its full size, it will tend to separate from its parent, and in so doing it will 
_ appear as if an enlarged polyp were undergoing fission into equal halves, whereas, strictly 
speaking, it is the components of the bud-polyp separating from the parent body” (35, 


* On p. 493 Duerden says that he has not actually observed the division of the mouth or stomodeum in 
any of his polyps. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 5 


34 PERCY SLADEN TRUST EXPEDITION 


pp. 152 and 153). Later he adds that in fissiparous gemmation “the division of the 
stomodeeum has not the same significance as in true fissiparity. It is rather a separation 
of two distinct stomodzea, one belonging to the parent and one to the bud; whereas, in 
true fission, it is the division of an enlarged stomodzeum into halves, and neither represents 
a distinct individual.” 

From this it is evident that Duerden’s original ideas of fission and budding did not 
fit in with certain facts that were observed later, and as a consequence he had to modify 
them to such an extent that the processes came to merge into each other as far as 
his ultimate results are concerned. Moreover, in some of my species in which extra- 
calicinal budding is the commoner method of asexual reproduction, e.g. Favia versipora 
(Lam.) and Favia wakayana (Gard.), directive mesenterial couples are absent, and hence 
it follows that even true gemmation need not always be followed by the presence of 
directives and the regular cyclical arrangement of the mesenteries. 

Whatever may be the exact morphological changes involved in the two processes, in 


b) 


this paper ‘‘gemmation” is used to denote the development of a polyp outside the 
tentacular ring of its parent, viz. from the edge-zone or ccenosarc, the new individual 
being a new growth and hence termed a “bud.” “‘ Fissiparity ” signifies the formation of 
fresh polyps inside the tentacular ring, viz. in the oral-dise area of an older polyp, the 
former being only parts of the latter separated off. Following this distinction all extra- 
calicinal buds would be true buds, while the various types of intra-calicinal buds would be 
regarded as fission-products. 

The presence of two couples of directive mesenteries is invariably associated in my 
polyps with two types of symmetry, viz. (1) bilateral symmetry, along two vertical planes, 
indicated by the lateral compression of the stomodzeum, by the presence of a directive groove 
and by a directive couple of mesenteries at each end of its longer axis; (2) hexameral 
symmetry, shown by each of the alternating mesenterial cycles consisting of six couples or 
of a multiple of six. 

Conversely the absence of the directive couples is marked by the disappearance of 
the two symmetries, the only distinction in the arrangement of the mesenterial couples 
being between those which meet the stomodzeum and those that do not reach it. 

In the fission-products of the genera I have studied the stomodzeum appears to arise 
in two ways: (1) as an invagination of the oral-disc* of another polyp, 7.e. as an 
independent structure secondarily effecting a communication with the gastro-vascular 
cavity, e.g. in Pavia abdita (Ell. and Sol.); (2) as a diverticulum from a parent stomodzum 
which grows towards the oral-dise and opens to the exterior, as is probably the case in 
Gonastrea pectinata (Ehrb.). 

In the case of either of the two processes I have suggested, some of the mesenterial 
couples of the old polyp would secondarily become attached to the stomodzeum of the new 
individual, and to the latter would come to belong also the tentacles which arose from the 
chambers between such mesenteries. 

Astreide with distinct corallites. Duerden bases his classification of the Madrepo- 
raria upon what he considers to be two distinet types of mesenterial succession (82, 


* This process is somewhat similar to the stomodzal formation in Flabellum as conceived by Gardiner. 


.MATTHAI—RECENT COLONIAL ASTRAIDA 35 


pp. 585541). According to him, “‘the young polyps of both gemmiferous and fissiparous 
genera are built upon exactly the same plan” (p. 539), viz., with the full complement of 
primary mesenteries (“ protocnemes”’) which, in both cases, are formed as “ bilateral pairs,” 
i.e. the two members of a pair arise, one on each side of the median axis, with the 
“retractor muscle” turned towards the same aspect of the polyp, but that the succeeding 
mesenteries (“metacnemes”) arise in two ways; in the first group (‘ Entocnemaria,” 
including only two genera Porites and Madrepora) they arise in “bilateral pairs,” as the 
primaries, but only “‘ within one or both of the directive entocoeles,” whereas in the 
second group (‘‘Cyclonemaria,” including the Astreeidze, Oculinidee, and Fungidee) they 
arise as “unilateral pairs,” 7.e. the two members arise in the same exoccele, their retractor 
muscles being vis-d-vis. The family Astreeidze is divided into two sub-families, Gemmantes 
and Fissiparantes. 

Such a division of the Astraidz is based upon the author’s theories regarding 
budding and fission, but from the above it is evident that the polyp symmetries are 
not necessarily dependent upon those two processes. Whatever may be the factors 
governing the symmetries, the presence or absence of two directive couples retains 
a constant morphological value in all the species and genera I have studied, and hence 
possesses an importance in classification over and above any physiological phenomena like 
budding and fission. 

I have accordingly divided my genera into two groups, characterised (I) by the 
presence of two couples of directive mesenteries and consequently possessing both the 
bilateral and hexameral symmetries, and (II) by the absence of the directive couples, and 
hence of the symmetries. 

Boveri (19) for Gyractis and Kwietniewski (88) for Thalassianthus regarded the 
absence of directive mesenteries as of ordinal value, creating the Holactiniz and 
Thalassianthee for their reception. On the other hand, McMurrich (99) did not regard 
their absence as having any phylogenetic significance, but viewed it as “‘a peculiarity which 
is essentially individual, possibly rising secondarily in some cases to the value of a specific 
or even a generic character....” (p. 121). 

From an examination of the sections of the coralla of the following genera I am led 
to infer that in Group I the septa are not directly continuous from corallite to corallite, 
the peritheca filling the inter-corallite spaces ; if any such septal continuity is seen, as in 
Diploastrea heliopora, it appears to be due to a secondary rearrangement of the calcareous 
elements of the peritheca. When two adjacent corallites are in contact, their walls are 
still distinct, the costee of one fitting in between those of the other. On the other hand, 
in Group II the septa of neighbouring corallites are usually continuous, the continuity 
being primary. . 

In all these genera the corallite-wall appears to be formed by the union of the outer 
thickened parts of the septa, 7.e. is a pseudotheca. In Galaxea a “eutheca,” as described 
by Ogilvie, is not seen in my sections*; the wall in this genus is also composed of the 
thickened peripheral parts of the septa, with the lines of suture quite distinct between 

* On the supposition of a true theca in Galaxea, Ogilvie separated this genus from the Astreidz and 
placed it provisionally in a new family, Stylinide (p. 162). 


5—2 


36 PERCY SLADEN TRUST EXPEDITION 


them; I have not made out any “ wall-lamelle” between the peripheral parts of the septa 
(see 108, p. 159). On the other hand, such lamelle, ve. interseptal tangential bridges 
with dark lines, are seen in the transverse sections of the coralla of certain species of 
Favia, e.g. F. doreyensis and abdita, while they are absent in allied species. It would 
appear from these observations that no great importance could be given to the dark-centres 
and lines. 

Summary of characters. The main characters that I have made use of in the 
determination of genera and species, so far as the polyps are concerned, are as 
follows : 


I. Oral-dise and Edge-zone. (1) The nature and relative thickness of the ectoderm 
and endoderm. 


Il. Tentacles. (2) The presence or absence of either entoccelic or exoccelic tentacles 
or of both. 
(8) The presence or absence of sub-terminal batteries; when present, the number 
of the batteries constituting a longitudinal row. 
(4) The nature of the ectoderm and endoderm. 


Ill. Stomodeum. (5) The nature of the ectodermal ridges at the attachments of 
the primary mesenteries—their relative breadth and thickness. 

IV. Mesenteries. (6) In Group I, the number of mesenterial cycles and of the 
couples constituting each cycle; in Group IJ, the approximate number of couples in the 
principal and the subsidiary cycles. . 

(7) The shape, relative size and distribution of the entoccelic mesogleal pleats, the 
presence or absence of exoccelic pleats, and the relative thickness of the mesenterial 
mesoglea. 

(8) The nature and relative thickness of the mesenterial endoderm. 

(9) The extent to which the margins of mesenteries are convoluted. 


V. Nematocysts. (10) The nature, distribution and relative abundance of the types 
present. 

If the reliability of the above characters should be doubted on the ground of their 
dependence on the degree of contraction, preservation, etc., of the polypal tissues, it is 
only necessary to refer to the case of species like Gonastrea retiformis (Lam.) and 
Leptastrea roissyana (Kd. and H.) whose specific characters have remained constant in spite 
of their wide distribution. There was abundant material of G. retifornis from the 
Maldives, Funafuti, Fiji and Ceylon, of L. roissyana from the Red Sea and Ceylon, and 
of Havia abdita from the Maldives and Ceylon, collected by different persons at different 
times and perhaps also treated differently. In either case the transverse sections of the 
polyps from these remote localities presented the same appearance, the specific characters 
being easily recognisable from any one of them. In the case of species with limited 
distribution, the constancy of the specitic characters is maintained in polyps taken from 
different colonies. 

Since the condition of the polypal tissues can be easily judged from the sections, 
the characters of well-preserved polyps may be relied on as normal. Under every species 


MATTHAI—RECENT COLONIAL ASTRAIDA 37 


described in this paper, mention is made of the histological condition of the polyps 


sectioned. 


Key to Genera based on Polyps. 


I. Two directive couples of mesenteries present; mesenterial couples with hexameral 
cyclical arrangement ... Sb uM bi fas Ae Group I. 
A. Six couples of mesenteries meeting stomodzum ; mesenterial mesoglea not 
much thickened. 
a. Stomodzeum well formed, entoccelic pleats of mesenteries undivided ; 
mesenterial endoderm usually swollen. 
i. Two cycles of mesenteries, each of six couples; nematocysts (III) rarely 
present aes a cn ae Sas ses as Cyphastrea. 
ii. Three cycles of mesenteries, the first two of six couples each and the 
last of twelve couples ; nematocysts (III) numerous but narrow, and without any central 
core as ROR see Bhd 200 ait ee Echinopora. 
b. Stomodzeum not well formed, often much compressed laterally and dis- 
torted ; entoccelic pleats of mesenteries sometimes or frequently sub-divided ; mesenterial 
endoderm thin. 


i. ‘“ Ovoid bodies” present ; numerous sub-terminal tentacular bat- 
teries mae A = oe Ae. a ee Galanxea. 

ii. ‘“‘Ovoid bodies” absent; up to five sub-terminal tentacular bat- 
teries se an ee sits sts Ane A Leptastrea. 


B. Twelve couples of mesenteries meeting stomodeum; mesenterial mesoglea 
much thickened aie as 8 ae My o08 Diploastrea. 


II. Directive couples of mesenteries absent; mesenterial couples without any 


eyelical arrangement ... on ee Sip sia fis Group LT. 
A. Number of principal couples of mesenteries indefinite; subsidiary couples less than 
thrice the number of principal couples; nematocysts II and III typical Fowia. 


B. Not more than six principal couples of mesenteries ; subsidiary couples 
more than thrice the number of principal couples ; nematocysts II and IIT narrower, in II, 
axis shorter... ah “oe ae is ted bf Goniastrea. 


Key to Genera based on Corallum. 


I. Corallites formed by extra-calicinal budding; septa with or without cyclical 
arrangement, number meeting columella more or less definite ec Group I. 
A. Corallite-walls imperforate. 
a. Perithecal spines often present. 
i. Corallites small and vertical; costal and perithecal spines when present 
not high, their bases never fused ; not more than twelve septa (v.e. no tertiaries) meeting 
columella... rarer) 4 gases ah a e 200 Cyphastrea. 


38 PERCY SLADEN TRUST EXPEDITION 


u. Corallites usually large and oblique. Costal and perithecal spines 
conspicuously high, their bases often fused to form perithecal ridges; more than twelve 
septa (7.e. a varying number of tertiaries also) meeting columella whe Echinopora. 

b. Perithecal spines absent. 


i. Peritheca highly vesicular; corallites usually projecting far above peritheca 


with markedly exsert septa ; inter-corallite furrows absent ... fe Galaxea. 
u. Peritheca dense ; corallites level or slightly projecting, with septa little 
exsert ; polygonal inter-corallite furrows present ... ae aa Leptastrea. 
B. Corallite-walls perforate —... a s53 ES Diploastrea. 


II. Corallites usually formed by fission, sometimes by budding; septa without 
cyclical arrangement, number meeting columella indefinite Bes Group IT. 


A. Corallite-walls usually separated by perithecal regions. of varying thickness ; 
septa not of equal width at calicular margins ; an alternating cycle of rudimentary septa 
absent or present ne Seis 500 an eG meh Fawia. 


B. Corallite-walls fused; septa of equal width at calicular margins; an alter- 
nating cycle of rudimentary septa present os i Abie Goniastrea. 


IW, SNGSME OMA, = GAR OWIP IL. 


Genus CypHasrrea, Klunzinger. 


1816. Astrea (pars), Lamarck, Hist. anim. sans vert., ii, p. 257. 

1834.  Haplanaria (pars), Ehrenberg, Korall. roth. Meer., p. 82. 

1834. Favia (pars), Ehrenberg, Korall. roth. Meer., p. 93. 

1848. Astrea, subgen. Orbicella (pars), Dana, Expl. exp. Zooph., p. 205. 

1848. Cyphastrea, Milne Edwards and Haime, Compt. rend. |’Acad. Sci., xxvii, p. 494. 
1848. Solenastrea, Milne Edwards and Haime, Compt. rend. |’Acad. Sci., xxvii, p. 494. 
1857. Cyphastrea (pars), Milne Edwards and Haime, Hist. nat. Corall., ii, p. 484. 
1857. Solenastreea, Milne Edwards and Haime, Hist. nat. Corall., ii, p. 495. 

1879. Cyphastrea, Klunzinger, Korall. Roth. Meer., p. 50. 

1884. Solenastrea and subgen. Cyphastrea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 107. 
1886. Cyphastrea, Quelch, Reef Corals, Challenger Reports, Zool., xvi, pt. xlvi, p. 106. 
1889. Cyphastrea, Ortmann, Steinkorall. Sud. Ceylons, Zool. Jahrb., iv, p. 529. 

1899. Cyphastrea, Gardiner, Proc. Zool. Soc. London, p. 761. 

1904. Cyphastrea, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 778. 


Corallum. Growth-form variable, mainly incrusting, following any irregularities on 
surface of attachment, often raised into hillocks, sometimes rounded off and free, usually 
bored into by various marine animals. Peritheca with dissepiments varying considerably 
in thickness, spines present or absent. Corallites circular, touching or to 4 mm. apart, 
level with general surface or to 2mm. high. Calices to 2°5 mm. in diameter. Three 
orders of septa: a primary entoccelic, a secondary entoccelic and a tertiary entoccelic, 
consisting of 6, 6 and 12 septa, sometimes fewer. Tertiaries very narrow, costz present 
or sunk in peritheca. Columella formed of trabeculz from septal margins. 

In colonies with uneven surface, corallites in valleys almost touching one another and 
hardly projecting, but septa more exsert than on hillocks. 


MATTHAI—RECENT COLONIAL ASTRAIDA 39 


Giant corallites* occasionally present, 2.55—3mm. in diameter, with from sixteen to 
twenty septa meeting columella (PI. 12, fig. 2). 

Polyps. Small, average height 2°5 mm. and width 1:25 mm., bearing distinct edge- 
zones. Mesenteries constituting two cycles, each typically of six couples, number of 
secondaries sometimes incomplete. Primaries with filaments, secondaries with or without 
rudimentary filaments. Tentacles of six entoccelic primaries, six entoccelic secondaries 
and twelve exoccelic tertiariest. Every tentacle with a terminal battery, sub-terminal 
ones present or absent. Stomodzeum oval, with two siphonoglyphes; its ridges not well 
developed, broader than thick. Entoccelic pleats of primary mesenteries variable in size 
and shape. A single ripe gonad on each primary mesentery. Genus comparatively 
homogeneous. Multiplication by budding. 

Duchassaing and Michelotti’st three species—Chypastrea oblita, Solenastrea ellisii 
and Solenastrea micans do not appear to belong to the present genus; in the figure 
(Pl. IX, fig. 10) of the last species the corallites are over 6 mm. in diameter, and more than 
twelve septa meet the columella. 

A small specimen from Mactan Island, referred by Quelch to Cyphastrea pleiades, is, 
unlike Madrepora pleiades, Ell. and Sol.,an undoubted Cyphastrea, but owing to its worn 
condition it is impossible to make out its specific characters. 

Distribution. Red Sea, Indian and Pacific Oceans. 


1. Cypuasrrea seraryié (Forskal). (PI. 7, fig. 4; Pl. 11, figs. 1—9; Pl. 13, fig. 8; 

Pl. 38, figs. 1 and 5.) 

1775.  Madrepora serailia (pars), Forskal, Deser. anim. in Itin. Orient., p. 135. 

1834. Havia microphthalma, Ehrenberg, Corall. roth. Meer., p. 93 (non Astrea microphthalma, Lamarck). 

1850. Cyphastrea ? botte (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 115. 

11850. Solenastrea sarcinula, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 122. 

1857. Cyphastrea bottai, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 486. 

11857. Solenastreea sarcinula, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 497, D 6, figs. 2a—c. 

1877. Cyphastreea capitata, Studer, Monat. Ak. Wiss. Berlin, p. 639.. 

1879.  Cyphastrea incrustans, Klunzinger Korall. Roth. Meer., iii, p. 53 (non Madrepora incrustans, Forskal). 

1904. Cyphastrea forskelana, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 778. 

1907. Cyphastrea forskaliana, Vaughan, Proc. U.S. Nat. Mus. Washington, xxxii, p. 253, pls. 19, 20, figs. 1 
and 2, 22, figs. 1—3. 

Corallum. Incrusting or raised into humps. Peritheca vesicular with a few low 
rough scattered spines, walls of vesicles thin. Corallites 1—3 mm. apart, projecting about 
‘75 mm., rarely 2 mm. Calices 1°5—2°5 mm. in diameter, interior somewhat obscured 
owing to thickness and spinous nature of septa. Septa somewhat exsert, sides with low 
pointed spines, inner margins with short teeth, first two orders sloping obliquely to meet 
columella ; secondaries generally thinner than primaries. Coste not prominent. Colu- 
mella finely trabecular with short upright rods projecting into calyx. 

Polyps. (1) Secondary entoccelic tentacles absent. (2) Sub-terminal batteries 

absent from all tentacles. (3) Entoccelic pleats extremely narrow and blunt, not 
* See Jeffrey Bell “On the variations observed in large masses of Turbinaria,” Jour. Roy. Micr. Soc., 1895, 
pp. 148—9. 


+ C. seralia is an exception as it has no secondary entoccelic tentacles. 
+ Mémoire Corall. Antill., Turin, p. 77 (1860). 


40 PERCY SLADEN TRUST EXPEDITION 


extending over more than the outer two-thirds of width of primary mesenteries, best seen 
in second quarter from skeletal attachment ; mesoglea much thicker than in non-pleatal 
region. (4) In outer two-thirds of width of primary mesenteries (in stomodzeal region of 
polyp) endoderm consisting of large goblet-shaped vacuoles massed together, in imner one- 
third a non-saccular layer. (5) Endoderm over stomodzum thin. (6) Filaments absent 
from secondary mesenteries. 

Remarks. A. Polyps. The terminal batteries cover the greater part of the 
exoccelic tentacles, but they are small and restricted to the ends of the entoccelic 
tentacles ; the nematocysts in these are practically all of type I, each with from twenty- 
five to thirty turns of the spiral; owing to the comparative fewness the nematocysts’ 
terminal batteries not swollen. Ectodermal muscle-fibres with a longitudinal disposition 
are visible in entoccelic tentacles. The stomodeeal ridges are thicker than broad. The 
convolutions of the mesenteries are massed together in the inter-mesenteric chambers and 
extending down to the base of the polyps. Brown sacs suggesting mucous contents are 
numerous in the coils of the mesenterial filaments, appearing above as clusters of round 
vacuoles, lower becoming oval and arranged around a common protoplasmic mass con- 
taining nuclei at its outer margin; each has a deeper stained spot in its bluntly-pomted 
inner end (Pl. 1, fig. 7). Only nematocysts II are found in the filaments. The endoderm 
is stained brown in hematoxylin and eosin except at the base of polyps, thickened and 
lobulated in the entoccelic tentacles owing to the presence of large vacuoles ; it is thin in 
the entoccelic tentacles and in the non-pleatal region of the primary mesenteries ; below 
the enterostome the mesenterial endoderm is a thin layer on either side of the mesoglea ; 
in the upper two-thirds of the secondary mesenteries there are endodermal vacuoles as in 
the primaries. A circular layer of endodermal muscle-fibres is distinctly seen in the oral- 
disc. Gonads were observed only in the primary mesenteries. In a male polyp each 
directive mesentery had a single large testis consisting of a mass of mesoglea with develop- 
ing spermatozoa embedded (PI. 1, fig. 9), in the other primaries the testes were smaller. 
They were also met with in the longitudinal sections of another polyp. In a female polyp, 
taken from the same colony, ripe ova were found in three of the mesenteries in 
single rows. 

Polyps examined, seven, all from a colony from Hulule, Maldives. 

B. Corallwm. In the Copenhagen Museum there are seven of Forskal’s originals of 
Madrepora serailia from Rode Hav. Of these six (the largest measuring 16 x 12°51 x 2 em.) 
constitute one species for which I retain Forskal’s specific name, my type specimens 
agreeing with them in every respect ; many of the corallites have the six primaries thicker 
than the six secondaries and twelve prominent costee, those of the former being somewhat 
more conspicuous than those of the latter (Pl. 11, figs. 4—9). 

Of the two specimens in the Paris Museum from the Red Sea referred by Milne 
Edwards and Haime to Cyphastrea bottai, the larger (24 x 23 x 19 em.) comes nearest the 
present species but the calices are somewhat larger, up to 3 mm., average about 2°5 mm. ; 
the specimen itself is somewhat rubbed, especially the corallites on its upper part. 
Solenastrea sarcinula is represented by the very small figured specimen (4°5 x 3 cm.), its 
only difference from C. serailia consisting in the higher projection of the corallites, about 


MATTHAI—RECENT COLONIAL ASTRAIDA 4] 


3mm. Two large specimens (25°5 x 14x13 cm.) from the Red Sea, named Solenastrea 
chaleidicum, are identical with my examples of C. seraalia. 

Ehrenberg’s type of Pavia nucrophthalma is a small specimen (6 x 6 cm.) with short 
hillocks, greatly resembling the present species, its septa have rough sides, the primaries 
being often thicker than the secondaries, both orders meeting the columella (Pl. 38, 
figs. 1 and 5). A specimen (13 x 13 em.) of the “Gazelle” expedition from New Hanover, 
referred by Studer to his new species Cyphastrea capitata resembles the present species 
in most respects. Similar to this is another small specimen (8 x 6 cm.), badly cleaned, from 
“Meermaidstrasse,’ N.W. Australia, which Studer referred to C. microphthalma (Lam.). 
Marenzelier has already poimted out Klunzinger’s error in regarding Madrepora incrustans 
(Forsk.) as a Cyphastrea, and has identified Forskal’s original to be a Turbinaria. 

Localities, Red Sea (3)*. Maldives, Hulule (2). Amirante, 16 fms. (1), 30 fms. (1). 
Saya de Malha, 20 fms. (1), 26 fms. (1). Also from New Hanover and N.W. Australia 
(Studer), and French Somaliland (Vaughan). 


2. OYPHASTREA CHALCIDICUM, [hiravammegsre (U2, fp wears, 1, 3 IEA, WO), ihiegs, 13} = JPL Té4, 


ino IN) 

1775.  Madrepora chaleidicum, Forskal, Descr. Anim. in Itin. Orient., p. 136. 

11848. Astrea (Orbicella) ocellina, Dana, Expl. exp. Zooph., p. 218, pl. 10, fig. 10. 

11848. Astrea (Orbicella) microphtalma, Dana, Expl. exp. Zooph., p. 217, pl. 10, fig. 11. 

1850. Solenastrea hemprichiana, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 121 (non 
C. hemprichana, Gardiner). 

11850. Solenastrea bowrnonii, Milne Edwards, and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 121. 

1850. Solenastrea bowerbankwi, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 122. 

1850. Solenastrea gibbosa, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 122. 

1857. Cyphastrea ? ocellina, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 487. 

1857. Cyphastrea ?danai, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 487. 

1857. Solenastrcea hemprichana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 495. 

1857. Solenastrwa gibbosa, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 496. 

1857. Solenastreea bowerbanki, Milne Edwards and Haime, Hist. Nat. Corall., ii. p. 498. 

1879. Oyphastreea chalcidicwm, Klunzinger, Korall. Roth. Meer., iii, p. 53, pl. 5, fig. 8; pl. 10, figs. 11 a—e. 

1886. Cyphastrea brueggemanni, Quelch, Reef Corals, Challenger Reports, Zool., vol. xvi, pt. xlvii, p. 107, 
pl. 4, figs. 4—4a. ° 

1899. Cyphastrea savignyt, Gardiner, Proc. Zool. Soc. London, p. 761, pl. 49, fig. 1 (non Cyphastrea 
savignyt, Milne Edwards and Haime). 

1901. Cpyhastrea ocellina, Studer, Zool. Jahrb., x1, p. 402, pl. 30, fig. 10. 

1904. Cyphastrea savignyi, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 779. 

11907. Cyphastrea ocellina, Vaughan, Recent Madreporaria of the Hawaiian Islands and Laysan, U.S. Nat. 
Mus., Bull. 59, p. 103, pl. 25, figs. 4, 5, 5a, pl. 26, fig. 1. 


Corallum. More or less rounded and completely covered with corallites, or raised into 
hillocks. Peritheca vesicular in parts, with scattered blunt spines, walls of vesicles some- 
what thicker than in C. serailia. Corallites about 1 mm. apart, in some places level with 
general surface, maximum projection 1 mm., walls thin. Calices usually about 2 mm. in 
diameter, deeper than in last species, clearly visible owing to comparative thinness and 
smoothness of septa. Septa more exsert than in C. serazlia, sides slightly rough or 
smooth, marginal teeth slender and spinous, in some cases approaching condition found in 
C. swvadwe ; primaries usually as thin as secondaries, both orders reaching columella. 

* The numbers in brackets refer to the numbers of specimens examined from each locality. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 6 


42 PERCY SLADEN TRUST EXPEDITION 


Costee prominent, with short spines. Columella openly trabecular, intermediate between 
C. suvadive and C. serailia, rods hardly visible. 

Polyps. (1) Secondary entoccelic tentacles present. (2) Each tentacle with three 
or four sub-terminal batteries. (3) Entoccelic pleats somewhat constricted at their bases, 
arranged close together, best developed in the outer half of width of primary mesenteries ; 
pleatal region of mesoglea thinner than in C. serailia. (4) Endoderm thickening con- 
siderably from skeletal attachments of primary mesenteries to stomodeal attachments of 
same. (5) Endoderm over stomodzeum thick. (6) Rudimentary filaments on secondary 
mesenteries. 

Remarks. A. Polyps. Both granular and mucous vacuoles are abundant in the 
ectoderm of the oral-dise and edge-zone ; sometimes the granules appear scattered in the 
ectoderm; between batteries granular vacuoles are conspicuous. Nematocysts I are 
closely packed in all the batteries; a few IIb ones are also present in the terminal 
batteries ; only nematocysts II and III have been observed in the mesenterial filaments, 
some of the latter with partly discharged threads. The stomodzal ridges are somewhat 
more thickened than in C. seraslia. The endoderm is more vacuolated than in that 
species, especially in the tentacles and primary mesenteries ; it is comparatively thin in 
the oral-disc, edge-zone and upper half of the column-wall. Below the enterostome the 
mesenterial endoderm is four or five times thicker in the inner half of the mesentery than 
in the outer half, where the protoplasm is reduced to mere strands. Ova were present in 
one polyp. 

Polyps examined, eight: three* from one specimen, three from a second specimen 
(both Red Sea), twot from a third specimen (Ceylon). 

B. Corallum. Forskal’s original of Madrepora chaleidicum is missing from the 
Copenhagen Museum. I have retained his specific name, since it is not used for any other 
species of corals and since my type specimens are similar to Klunzinger’s figured example 
(9x 6 cm.) of Cyphastrea chaleidicum. 

Milne Edwards and Haime have referred three large specimens from the Red Sea to 
Solenastrea hemprichana (two of them measuring 23 x 15x14 em. and 21x18 x13 em. 
respectively), resembling specimen no. 5 on p. 46; many of the corallites are arranged close 
together and project like cylinders, a few of them being giant ones; only in a few corallites 
do all the secondaries meet the columella. These authors have assigned seven specimens 
from the Red Sea to Solenastrea gibbosa, three of them being large (23 x 15 x 14 em., 
20x 10x10 cm. and 14x 14x 5 em.), which are identical with many of my examples of 
C. chalcidicum. Another specimen from the Red Sea named Solenastrea chalcidicum 
also comes here. Solenastrea bournoni is represented by four small specimens from 
Antilles (the largest measuring only 9 x 9 cm.), approaching the present species; a few 
giant corallites are present on them. Solenastrea bowerbanki, Kd. and H., is missing from 
the Paris museum; judging from Milne Edwards and Haime’s description it comes 

* One of these is a giant-polyp with twelve couples of primary and eight of secondary mesenteries, the 


former bearing ova. 

+ In these polyps the stomodzeum was less wide, its wall being deeply grooved at the mesenterial attach- 
ments. The entocelic pleats were somewhat thinner. These differences are due to the sudden shrinkage 
of the tissues, as doubtless the specimens were not fixed or preserved properly. 


MATTHAI—RECENT COLONIAL ASTRAID A 43 


under the present species. A large specimen from Koseir (Red Sea), named C. serailia by 
Klunzinger, belongs to the presen, :species. 

Quelch’s new species, C. brueggemanni, is represented by a single small specimen 
(4:5 x 4 em.) from Mactan Island, Philippimes, nm which the primary and secondary septa 
are similar, both orders meeting the columella; a giant corallite is present with eighteen 
septa meeting the columella. From his description it might be inferred that he had more 
than one example of the species. Another small badly cleaned specimen, from the same 
locality referred by Quelch to C. microphthalma (Lam.) also appears to belong to 
C. chalevdicum. 

Studer’s figure of C. ocellina agrees with Klunzinger’s figure of C. chalcidicum, but 
I have not been able to examine the original. The identity of Vaughan’s examples of 
C. ocellina with the present species is somewhat doubtful, since in his Pl. XXV, fig. 5 a, 
the septa appear thicker and rougher than in my specimens, in this respect approaching 
more the condition in C. serailia. 

Localities. Red Sea (5). Maldives, Goidu (4). Chagos, Salomon (2). Singapore (2). 
Ceylon (1 small). Rotuma (1). Also from Antilles (Milne Edwards and Haime), 
Philippines (Quelch), Laysan (Studer), Sandwich Islands (Dana), Hawaiian Islands 
(Vaughan). 


8. CypHasTrea micropHTHALMA (Lamarck) (Pl. 7, fig. 6; Pl. 12, figs. 4—9; Pl. 18, 
figs. 1, 2,7; Pl. 34, fig. 4). 


1775, Madrepora serailia (pars), Forskal, Descr. Anim. in Itin. Orient., p. 136. 

1797. Madrepora interstincta, Esper. Fortz. Pflanz., p. 10, pl. 34, figs. 1—3 (non Madrepora «interstincta, 
Linneus). 

1815. Astrea interstincta, Oken, Lehrb. Naturg., i, p. 66. 

1816. Astrea microphthalma, Lamarck, Hist. Amin. sans vert., ii, p. 261 —2° édit., p. 408. 

1834. Haxplanaria galaxia, Ehrenberg, Corall. roth. Meer., p. 82 (non Astrea galaxea, Lamarck). 

1850. Cyphastrea microphthalma, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, pl. 9, figs. 5, 

5a and 6, xii, p. 114. 

1850. Cyphastrea savignyi, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 116. 

1850. Solenastrea forskaliana, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 123. 

1851. Cyphastrea miilleri, Milne Edwards and Haime, Pol. foss. terr. paleoz., ete., p. 100. 

1857. Cyphastreea microphthalma, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 485. 

1857. Cyphastreea savignyi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 485. 

1857. Cyphastrea muelleri, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 486. 

1857. Solenastrea forskelana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p 497. 

1879. Cyphastrea savignyt, Klunzinger, Korall. Roth. Meer., iii, p. 51, pl. 5, fig. 7. 

1879. Cyphastrea serailia, Klunzinger, Korall. Roth. Meer., iii, p. 52, pl. 5, fig. 4 and pl. 10, figs. 12a and 6. 

1886. Cyphastreea aspera, Quelch, Reef Corals, Challenger Reports, Zool., vol. xvi, pt. xlvi, p. 107, pl. 4, 
figs. 3—3 a. 

1904. Cyphastreea microphthalma, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 779. 


Corallum. Growth-form irregular, usually raised into hillocks. Peritheca dense, 
vesicles more or less filled up with spines, more abundant than in C. chalcrdicwm and 
arranged in rows around corallites leading up to the exsert ends of septa, each spine with 
side spinules, hence appearing star-shaped from above. Corallites usually close to one 
another, not projecting, or at most up to°75 mm. Calices about 1°25 mm. in diameter, 
interior obscured owing to considerable thickness of septa. 


6—2 


44 PERCY SLADEN TRUST EXPEDITION 


Septa slightly exsert, arched above peritheca, sides with short spines rougher and 
somewhat more pointed than in C. serailia, marginal teeth slender and spinous. As 
a rule, two of the secondary septa exocoelic and appearing to belong to the third order 
(as there are usually only four secondary mesenterial couples in the polyps), and only eight 
tertiary exoccelic septa; of these ten septa meeting the columella equally thick, the 
remaining ten very narrow and alternating with them. Costz sunk in peritheca, 
represented by the rows of spines leading up to the exsert septa. Columella thicker — 
than in the last two species with higher rods projecting into calyx. Corallum rougher 
than in any other species. 

Polyps. (1) Secondary entoccelic tentacles present. (2) In some tentacles one or 
two sub-terminal batteries. (3) Entoccelic pleats not so well developed as in C. chal- 
cidicum, nor covering more than the outer half of primary mesenteries ; non-pleatal region 
of mesoglea thicker than in same species and only slightly thinner than pleatal region. 
(4) Endoderm thickening from skeletal attachments of primary mesenteries to stomodzeal 
attachments of same. (5) Endoderm over stomodzeum thick. (6) Filaments absent 
from secondary mesenteries. 

Remarks. A. Polyps. A varying number of primary mesenteries do not reach 
the stomodeeum, but at least one in every couple joins it. Three to six secondary 
couples are present, usually four. 

Polyps examined, eleven, five from one specimen, three from a second specimen, three 
from a third specimen (all Red Sea). 

B. Corallum. Milne Edwards and Haime have referred three specimens to 
Cyphastrea microphthalma, with which my examples agree completely, the largest from 
Australia measuring 18 x 12°5 x11 cm.; of the remaining two, somewhat rubbed down 
specimens from Oceania, one (5 x 4°5 x 2°5 cm.) is in Lamarck’s collection, evidently his 
type of Astrea microphthalma (Pl. 12, fig. 9). Resembling these are Milne Edwards and 
Haime’s three original examples of Cyphastrea savignyi from the Red Sea, two of which 
are large (20x 18x16 cm. and 18x 10x13°5 cm.), the third very small and broken, 
All these specimens are characterised by ten septa meeting the columella in each corallite 
and an equal number of very narrow alternating septa, dense peritheca, and prominent 
perithecal spines arranged in rows leading up to the exsert ends of the septa (PI. 18, 
fig. 1). At present there are no specimens in the Paris museum named Cyphastrea 
muellert, Ed. and H., or Solenastrea forskelana, Ed. and H., but from Milne Edwards and 
Haime’s descriptions of these two species they do not appear to be in any way different 
from CL microphthalma. A large specimen from Koseir named Solenastrea chalcidicum 
by Klunzinger belongs to C! microphthalma. 

Resembling Milne Edwards and Haime’s types above referred to are six specimens in 
the Berlin Museum, which Ehrenberg had originally referred to Haplanaria galaxia, but 
which Klunzinger later brought under Cyphastraa savignyt, one of which he has figured. 
Klunzinger’s example (13 x7 cm.) of Cyphastrea serailia is identical with the edge region 
of specimen no. 12 on p. 47, both having projecting corallites. 

Quelch’s new species Cyphastrea aspera is based upon a very small gpecimen 
(4x 3°5 cm.) from Api, New Hebrides; its calices are somewhat deeper, but in all other 


MATTHAI—RECENT COLONIAL ASTRAIDA 45 


respects it resembles C. microphthalma. Three specimens of the “ Pola” expedition which 
Marenzeller has referred to Cyphastrea savignyi, Kd. and H., are good examples of the 
present species. 

One of Forskal’s originals of Madrepora serailia, measuring 18 x 12°5 x 8 em. (Pl. 18, 
fie. 2) is identical with my Red Sea examples of C. microphthalma. Agreeing with these 
completely is Esper’s Pl. 34, fig. 3 of Madrepora interstincta. According to Milne 
Edwards and Haime, Madrepora interstincta, Linnzeus, is a fossil millepore, Helolites 
imterstincta. 

Localities. Red Sea (19). Chagos: Salomon (7); Egmont (2). Amirante 20—25 
fms. (1). Saya de Malha 29 fms. (1). Coetivy (1). Maldives: Addu 40 fms. (1); Turadu 
(1); Suvadiva 31 fms. (1). Also from Australia (Milne Edwards and Haime), ? New 
Hebrides (Quelch), loc. ? (Esper). 


4, CypHastrna suvapivm, Gardiner (PI. 7, fig. 7; 18, fig. 3; 34, fig. 6). 


1889. Cyphastrea muelleri, Ortmann, Steinkorall. Sud. Ceylons, Zool. Jahrb., iv, p. 530. 
1904. Cyphastrea suvadive, Gardiner, Fauna Geogr., Maldives and Laccadives, ii, p. 780. 
1904. Cyphastrea maldivensis, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 780. 


Corallum. Incrusting, thin, comparatively light. Peritheca vesicular, vesicles on 
surface like blisters, often with walls incomplete, but where complete granular, dissepi- 
ments very thin and wider apart than in all previous species. Corallites from 2—4 mm. 
apart, not projecting. Calices about 1°5 mm. in diameter, shallower than in other 
species. . 

Septa slightly or not at all exsert, very thin, sides smooth or slightly rough; 
primaries as thin as secondaries, both orders meeting columella and having long, pointed, 
spinous teeth extending over the axial fossa. Costee sunk in peritheca. Columella more 
openly trabecular than in C. chulcidicum, hardly any rods projecting into calyx. 

Professor Gardiner’s type specimens of C. suvadive were all very small broken pieces; 
the shallow, closed-in nature of the corallites may be due to the pieces having been broken 
off the edge of a large specimen. 

Polyps. (1) Secondary entoccelic tentacles present. (2) A single, sub-terminal 
battery on each primary entoccelic tentacle—none on others. (3) Entoccelic pleats only 
in the second quarter of width of primary mesenteries from skeletal attachment, well 
constricted at their bases, comparatively few in number (from 8—11, in last two species 
over 20), thinner and further apart than in any previous species; mesenterial mesoglea 
also thinner. (4) Endoderm a thin layer in the first quarter of width of primary 
mesenteries, thickening towards stomodeal attachment, but less than in C. mzcroph- 
thalma. (5) Endoderm over stomodzum thick. (6) Filaments absent from secondary 
mesenteries. 

Remarks. A. Polyps. Nematocysts I are closely packed in the terminal batteries, 
while type IL1b are nearly absent. Nematocysts II are of common occurrence in the 
convolutions of the mesenterial filaments, while III are less frequently found. In the 
first quarter of primary mesenteries the mesoglea is extremely thin. The endoderm 
is less vacuolated than in C. microphthalma ; in the edge-zone, oral-dise and upper half of 


PERCY SLADEN TRUST EXPEDITION 


LIST OF SPECIMENS SHOWING THE EXTENT OF VARIATION IN 


46 
No. of . 
Specimen Locality 
1 Amirante 
2 Red Sea 
3 Amirante 
4 Hulule, Maldives 
5 Red Sea 
6 Singapore 
a Red Sea 
8 Turadu, Maldives 
9 Egmont, Chagos 
10 Salomon, Chagos 


THE GENUS CYPHASTREA. 
Species * Remarks 
C. serailia Corallum (15x15 cm.) incrusting on a rounded mass 


CO. chalcidicum 


C.microphthalma 


which is covered with corallites except at its place of 
attachment. Primaries often resemble secondaries as 
in C. chalcidicum. Peritheca completely vesicular as 
in C. suwvadive and corallites 2°5 or 3 mm. apart. PI. 
11, fig. 1; Pl. 18, fig. 8. 


Corallum (15 x 12cm.) incrusting on a hillocky mass. 
Some of the corallites simulate those of C. chalcidicum— 
primaries as thin as secondaries, both with almost 
smooth sides, columella openly trabecular. Towards 
edge corallites sunk and shallow. A giant corallite 


2°) mm. in diameter with 9, 9, and 18 septa. 


A small incrusting mass (8x6 cm.). Septal teeth some- 
what long and slender at bases of calices, approaching 
condition in C. suvadive. 


Corallum incrusting (11x 8 em.). Most corallites with 
secondaries not meeting columella, resembling Ed. and 
H.’s fig. of S. sarcinula. Towards edge corallites shallow. 
Pl. 11, fig. 2. 


Corallum (18x 13x10 em.) massive with hillocks. Coral- 
lites on some hillocks agree completely with Klunzinger’s 
type of CO. chalcidicum—projecting cylinders, 1 mm. or 
more deep and 2 mm. in diameter. In valleys corallites 
level with surface and narrow (1:25 mm. wide). Pl. 12, 
fig. 1. 


Corallum (10 x 9 x 7 cm.) inerusting on an irregular sur- 
face; corallites not projecting on flat areas, on humps up 
to 1-5 mm.—completely resembles Studer’s fig. of C. 
ocellina. Two giant corallites each with 20 septa meet- 
ing columella. Pl. 12, fig. 2. 


Corallum (19 x 16 x 12cm.) incrusting but raised into 
branching hillocks, some parts simulating Gardiner’s 
examples of C. maldivensis—vesicular peritheca, coral- 
lites 2-5 mm. apart, coste sunk in peritheca, septal teeth 
long, slender, tending to be fenestrated over axial fossa. 
In valleys calices only 1 mm. in diameter. A giant 


corallite 2-5 mm. wide with 9, 9, and 18 septa, another 
in fission with diameters 3 mm. and 2 mm. and 21 septa 
meeting columella, 


Typical (10x 8 cm.). Towards edge resembles Klunzin- 
ger’s fig. of C. savignyt. 


Corallum (14 x 9 x 8 cm.) incrusting with humps. Septa 
10 and 10. As in @. seratlia primaries much thicker 
than secondaries, exsert, and sides of both rough— 
specimen resembles Klunzinger’s fig. of C. savignyt. 
Pl. 12, fig. 6. 


Corallum small (9 x 7x5 cm.). Intermediate stages in 
corallites between C. serailia and C. microphthalma— 


septa 6, 5 and 6, 4 and 10, and 10 and 10. Perithecal 
spines low, blunt, scattered. 


MATTHAI—RECENT COLONIAL ASTRAIDA 


47 


LIST OF SPECIMENS SHOWING THE EXTENT OF VARIATION IN 
THE GENUS CYPHASTREA—continued. 


No. of 
Specimen 


Locality 


Species 


Remarks 


11 


13 


14 


15 


16 


17 


18 


19 


21 


Red Sea 


Lb) 


Salomon, Chagos 


Red Sea 


Salomon, Chagos 


Red Sea 


Suvadiva, Maldives 


Hulule, Maldives 


Red Sea 


Rotuma 


C. microphthalma 


C. suvadive 


C. hemprichana 


Corallum (11 x 9x 4 cm.) incrusting. Septa thin and 
only slightly rough as in C’. chalcidicwm but no. 10, and 
10. Perithecal spines almost smooth. _ Pl. 12, fig. 7. 


Corallum (14x 8x9cm.) raised into humps, identical 
with Klunzinger’s figured specimen of C. serailia. Pl. 
12, fig. 5. 


Corallum (17 x 12 x 12cm.) incrusting, resembles Klunzin- 
ger’s fig. of C. savignyt. Corallites shallow, 2-5—3 mm. 
apart, sunk, only granules on peritheca. 


Corallum (15 x 8 x 17 cm.) large, raised into hillocks— 
resembles Klunzinger’s fig. of C. seratlia—towards edge 
similar to fig. of C. savignyi. Pl. 34, fig. 4. 


Corallum (25 x 18 x 19 cm.) raised into humps on a large 
substratum. In a flat valley corallites crowded, thin- 
walled and columella rudimentary. Three giant coral- 
lites each 2-5 mm. wide with 18 septa meeting columella. 
Pl. 12, fig. 4. 


Corallum (20 x 12 x 13 cm.) with hillocks and deep valleys. 
Towards edge corallites shallow and closed in. Two 
giant corallites, 25 mm. wide with 16 and 18 septa 
meeting columella. PI. 13, fig. 7. 


Corallum small (12 x 9x 5cm.). Similar to C. chaleidi- 
cum except in number of septa and dense columella— 
peritheca in parts vesicular where spines few, short, 
blunt, smooth, scattered, septa thin, only slightly rough. 


Corallum small (10 x 7 x 7 cm.)—great variation in number 
of septa, a few corallites with 12 and 12, many with 10 
~ and 10 or 9 and 9, some others with 8 and 8. 


Prof. Gardiner’s specimens of C. swvadive—small broken 
pieces ; all flat, thin corallites shallow with a closed in 
appearance, low thin perithecal spines, sometimes only 
11 septa meeting columella. 


Corallum large (11 x § x 5:5 em.) with even surface. Con- 
dition typical on upper surface—on lower surface hardly 
any perithecal spines, corallites not projecting, shallower, 
hence columella visible. Pl. 13, fig. 5; Pl. 34, fig. 5. 


Corallum small (6 x 6 x 4 cm.) with low humps. Calices 
somewhat shallow, hence columella visible with con- 
spicuous rods. Septa usually 6, 4 and 10, only primaries 
meeting columella. 


Prof. Gardiner’s specimen of C’. chalcidicum (16 x 10 x 7-5 
cm.)—rougher (perithecal spines and septa) than speci- 
mens 20 and 21, also corallite-wall thicker and calices 
somewhat wider (1°75 mm.), columella distinctly seen, 
well developed, with conspicuous rods, septa 6, 6 and 12 
primaries not much thicker or broader than secondaries, 
the latter in some corallites meeting columella. Pl. 13, 
fig. 4. 


48 PERCY SLADEN TRUST EXPEDITION 


body-wall it is extremely thin, while in the secondary mesenteries it is a uniformly thin 
layer on either side of the mesoglea. 

Polyps examined, seven, three from a well-preserved specimen from Suvadiva, four 
from a badly-preserved specimen from Saya de Malha. 

B. Corallum. A specimen from Ceylon in the Berlin Museum, measuring 8 x 8 cm., 
which Ortmann had referred to Cyphastrea mueller, resembles the present species in the 
-highly vesicular condition of its peritheca and in the thinness of its septa, but the septal 
teeth are not quite so long as in Cyphastrea maldivensis, Gard., and the corallites 
are slightly projecting. 

Localities. Maldives: Felidu 20—25 fms. (1); S. Nilandu 25 fms. (1); Suvadiva 
(small fragments). Saya de Malha 26—29 fms. (3). Amirante 725 fms. (1). Cargados 
30 fms. (1). Also from Ceylon (Ortmann). 


5. CYpHASTREA GARDINERI, n. sp. (PI. 13, figs. 4 and 5; 34, fig. 5.) 


1899. Cyphastrea chalcidicwm, Gardiner, Proc. Zool. Soc. London, p. 761 (non Cyphastrea chalcidicum, 
Klunzinger). 

1904. Cyphastrea hemprichana, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 779 (non Solenastrea 
hemprichana, Milne Edwards and Haime). 


Corallum. With either even surface or raised into low humps. Peritheca dense, 
dissepiments much thickened as in C. microphthalma, with conspicuous, blunt, somewhat 
smooth spines, thicker and higher than in any other previous species, and irregularly 
distributed. Corallites from almost touching to 2 mm. apart, projecting up to *75 mm. 
Calices about 1°5 mm. in diameter, deep. 

Septa exsert, sides rough, inner margins perpendicular with short blunt teeth, 
decreasing in width and thickness from primaries to tertiaries; only primaries meeting 
columella, but narrower than in other species. Costze distinct on projecting corallite- 
walls. Columella deep down in calyx, often invisible from above. 

No Polyps. 

This species differs from both Solenastrea hemprichana, Ed. and H., and Cyphastraea 
chalcidicum, Klunz., in the following respects: (1) dense peritheca, (2) perithecal spines 
numerous and prominent (in these characters approaching C! microphthalma, but the 
spines are usually blunt and somewhat smooth), (3) calices somewhat deeper, hence 
columella often invisible from above, (4) septa thicker and rougher (in this respect 
resembling C. serazlia). In the cylinder-like appearance of the corallites the species 
agrees with C. chalcidicum. Its true position in the genus can, however, be determined 
only after an examination of its polyps. 


Localities. Red Sea (1). Rotuma (1). Maldives, Hulule (1). 


Ecutnopora, LAMARCK. 


1816. Zchinopora, Lamarck, His. anim. sans. vert., ii, p. 252. 

1816. Haplanaria (pars), Lamarck, His. anim. sans. vert., ii, p. 254. 
1830. Hchinastrea, Blainville, Dict. Sci. Nat., lx, p. 327. 

1830. Tridacophyllia (pars), Blainville, Dict. Sci. Nat., 1x, p. 343. 
1834. Stephanocora, Ehrenberg, Corall. roth. Meer., p. 76. 

1834. Haxplanaria (pars), Ehrenberg, Corall. roth. Meer., p. 82. 


MATTHAI—RECENT COLONIAL ASTRASID AL A9 


1848. Hchinopora (pars), Dana, Expl. exp. Zooph., p. 277. 

1850. Astrea (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 97. 

1850. Zchinopora (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 621. 
1857. Heliastrcea (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 456. 

1857. Hchinopora (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 621. 

1879. Orbicella (pars), Klunzinger, Korall. Roth. Meer., iii, p. 47. 

1879. Hchinopora, Klunzinger, Korall. Roth. Meer., iii, p. 54. 

1884. Hchinopora, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 117. 

1889. Hchinopora, Ortmann, Steinkorall. Siid. Ceylons, Zool. Jahrb., iv, p. 530. 

1904. Hchinopora (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, p. 782. 


Corallum. Varying considerably, flat, thin and foliaceous to massive, and rising 
into branching hillocks. Peritheca vesicular or dense, the vesicles being often filled up, 
usually with upright spinulate echinulations arranged in rows connecting cost of neigh- 
bouring corallites, their bases often fused to form ridges which are specially distinct 
‘towards edges where spines are small or absent. Corallites round or oval, close or quite 
separated, level or projecting up to a height of 6 mm., usually one side more than the other, 
near the edges of corallum the sides facing same not projecting at all. Calices not deep, 
sometimes almost flat. Septa from 3—5 orders, side spinulose, exsert up to 2mm., on 
edges confluent with perithecal costze. Paliform lobes present or absent. Coste always 
present, varying in thickness, usually with upright spines. Often the last order of septa 
distinguishable only in costz. Columella spongy, formed of trabeculae from septal 
margins. 

Polyps. Circular or oval; when corallites project, edge-zones cover their entire free 
surface ; coenosarcal regions usually extensive, varying in width up to 6mm. Mesenteries 
in three cycles, 6, 6 and 12 couples, last sometimes incomplete, primaries meeting stomo- 
deeum, ali with filaments. Tentacles entoccelic and exoccelic, but number always equalling 
that of the entocceles and exocceles, each with a large knobbed terminal battery and from 
three to six smaller sub-terminal ones. Stomodeeum usually laterally compressed, with two 
directive grooves. Nematocysts IIi uniformly modified into the III b type in all polyps 
examined ; nematocysts II with the dark-stained axis extending to even more than two- 
thirds length of sac. Tentacular endoderm much thickened owing to vacuolation and 
~somewhat lobulated, the protoplasm being reduced to thin strands, appearing transparent 
owing to scarcity of alge. 

Multiplication by budding, usually from ccenosare, sometimes from edge-zone. 

Verrill rightly separated H. aspera, Ed. and H., from Hchinopora, and established a 
new genus, Trachypora, for it. Klunzinger later substituted Hchinophyllia for this genus. 
In Lamarck’s collection in the Paris Museum there are two specimens from the Indian 
Ocean named H. aspera by Milne Edwards and Haime, which obviously were Lamarck’s 
originals of Hxplanaria aspera; these resemble Mycedium okeni, Ed. and H. (Fungideze), 
of which there are two large examples in the Paris Museum. I have not been able to see 
Klunzinger’s example of Echinophyllia aspera, but from his figure it appears to be a near 
relative of H. aspera, Kd. and H., though not quite identical with it. 

Gardiner’s small figured type of H. magna has a thicker facies than EH. aspera, 
Kd. and H., resembling Milne Edwards and Haime’s larger example of Mycediwm okena. 

Distribution. Red Sea, Indian and Pacific Oceans. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 7 


50 PERCY SLADEN TRUST EXPEDITION 


1. Hcurinopors LaMELLOSA (Esper). (Pl. 8, fig. 6; 14, figs. 2—6; 15, fig. 1; 

16, fig. 6.) 

1797. Madrepora lamellosa, Esper, Forts. Pflanz., p. 65, pl. 58, figs. 1 and 2. 

1816. Lchinopora rosularia, Lamarck, Hist. Anim. sans vert., ii, p. 253—2° édit., p. 397. 

1830. Hchinastrea rosularia, Blainville, Dict. Sci. Nat. lx, p. 344, pl. 35, fig. 2—Manuel d’Actinol., p. 378, 
pl. 56, fig. 2. 

1848. Hchinopora undulata, Dana, Expl. exp. Zooph., p. 278, pl. 17, fig. 3. 

1848. Hchinopora rosularia, Dana, Expl. exp. Zooph., p. 279. 

1848. Hchinopora reflexa, Dana, Expl. exp. Zooph., p. 280, pl. 17, fig. 2. 

11848. Hchinopora horrida, Dana, Expl. exp. Zooph., p. 282, pl. 17, fig. 4. 

18 . Echinopora rosularia, Milne Edwards, Atlas grande édit. Régne anim. Cuvier, Zooph., pl. 85 ter. 

1850. Hchinopora rosularia (pars), Milne Edwards and Haime, Aun, Sci. Nat., Zool., 3° sér., xii, p. 185. 

1857. Echinopora rosularia (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 623. 

1864. Echinopora flexuosa, Vervrill, Bull. Mus. Comp. Zool. Harvard, i, p. 54. 

1877. Echinopora rosularia, Studer, Monat. Ak. Wiss. Berlin, p. 643. 

1877. LHchinopora flexuosa, Studer, Monat. Ak. Wiss. Berlin, p. 643. 

1877. Echinopora striatula, Studer, Monat. Ak. Wiss. Berlin, p. 644, fig. 10a@ and 6. 

1889. Hchinopora rosularia, Ortmann, Steinkorall. Siid. Ceylons, Zool. Jahrb., iv, p. 531. 

1904. Hchinopora rosularia, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 782, pl. 63, fig. 37. 

Corallum. Very thin, margins irregularly folded up, attached below in the centre. 
Peritheca dense, with slender spinulate echinulations, their bases usually fused to form low 
thin perithecal coste, towards edges of corallum spines disappearing but ridges remaining. 
Corallites almost circular, projecting up to 2 mm. (usually 1:25 mm.), up to 6 mm. apart 
(average 3 mm.). Calices with diameter 3°5 mm., depth 1°5 mm. ; quite shallow towards 
edges of corallum. 

Septa in three orders, tertiaries very thin, up to 4 quaternaries sometimes present, 
sides rough, perforated, edges denticulate. Primaries sometimes thicker than secondaries, 
these and up to 6 tertiaries meeting columella; septa of even thickness along their breadth 
or somewhat thickened in thecee, margins vertical, exsert to 1mm. The exsert ends of 
primaries and secondaries divided by two notches—not extending below calicular 
margins—into an inner arched lobe and two outer bluntly-pointed teeth, the last appearing 
as the first costal tooth. Coste comparatively thin, with one or two spinulate echinula- 
tions. Columella of close trabeculee, up to two-thirds width of calyx. 

Polyps. (1) Tertiary couples of mesenteries absent. (2) Directive grooves deep 
and narrow. (3) Nematocysts I less numerous in ectoderm of oral-disc and outer wall 
of edge-zone than in HL. gemmacea, but fewer in stomodeeal ridges and in straight regions 
of mesenterial filaments than in EH. hirsutissima. (4) Entoccelic pleats in stomodeeal 
region of polyp horizontal, better developed than in E. hirsutissima or E. gemmacea, 
being thick, unconstricted at their bases, extending over the outer half of primary 
mesenteries and more or less of uniform size along the entire pleatal region; mesoglea 
thickest in pleatal region ; no exoccelic pleats. (5) Mesenterial endoderm of more or less 
uniform thickness along the entire width of primary mesenteries. (6) Endoderm in outer 
wall of edge-zone as thick as the ectoderm over it, somewhat thinner in oral-disc. 
(7) Convolutions of mesenteries abundant towards bases of polyps. 

Remarks. A. Polyps. The polyps are circular in outline, relatively small, the largest 
one being 4 mm. in height and 2°5 mm. in diameter and invariably directed obliquely 


MATTHAI—RECENT COLONIAL ASTRASTID AD 51 


downwards. The convolutions of mesenteries are scarce to some distance below the 
stomodeeum, becoming more abundant towards bases of polyps. Tentacles corresponding 
in number and position with entocceles and exocceles. The stomodzeum is much compressed 
laterally, with diameters | mm. and -4 mm.; its ridges are somewhat thicker than in 
the next species, those of adjacent mesenteries of neighbouring primary couples sometimes 
fusing in the upper part of the stomodzeum as in Goniastrea retiformis. Nematocysts I 
are rare in the ridges and in the straight regions of mesenterial filaments. 

The endoderm is extremely thin in the region of the terminal batteries, non-vacuolated 
and with a row of algze, below this vacuolated and transparent with very few algze as in 
E. larsutissima but not quite so thick as the sub-terminal batteries ; stomodzal endoderm 
is thicker than in the same species owing to greater vacuolation; mesenterial endoderm 
behind filament not so thickened. 

Gonads were not present in any of the polyps sectioned. Polyps examined, four— 
two from one specimen and two from a second, both from Hulule, Maldives. 

B. Corallum.. Of the six specimens in the Paris Museum referred by Milne Edwards 
and Haime to #. rosularia, five belong to the present species; of these, two large ones 
from Seychelles (the larger measuring 26 x 19 cm.) have the same facies as many of my 
examples from Seychelles, being thin and folded with poorly developed columellie; another 
specimen from the same locality is denser, up to 1°5 cm. thick, with most of the corallites 
level, perithecal spines thicker and columella better developed, about $ width of calyx ; 
the remaining two specimens are smaller, a rubbed one from Red Sea (15 x 8 em.) 
and another (9x7 cm.) from Australia. A specimen (13°5x12cm.) from Singapore 
named H. flexuosa, Verrill, has corallites on both sides of the corallum. Four cylindrical 
examples from Fiji (the longest 9 cm.), named #. horrida, Dana, have a tree-like mode 
of growth with few perithecal spines; these are not a dendroid variety of H. hirsutissima 
as Milne Edwards and Haime had supposed, but belong to the present species. 

Of the “Gazelle” specimens in the Berlin Museum, Studer has referred three large 
ones from Salawatti to HL. rosularia, another large one from the same locality to H. flexuosa, 
Verrill, in every respect resembling the like named specimen in the Paris Museum, and 
a number of good examples to his new species H. striatula which more or less resemble 
Specimen no. 3 on p. 57; these differ from my typical condition of HL. lamellosa in that 
the perithecal spines are shorter, many of them level, corallites on both sides of the 
corallum (in this respect agreeing with ZH. flecwosa) and almost flat, columella not more 
than one-third width of calyx. A specimen from Ceylon is referred by Ortmann to 
LE. rosularia. 

There is hardly any doubt that Esper’s type of H. lamellosa, judging from the two 
figures he has given, is identical with mine. 

Localities. Maldives, Hulule (6). Chagos: Salomon (9); Coin, Peros (2). Sey- 
chelles (4). Also from Australia (Milne Edwards and Haime), ? Singapore (Verrill), Fiji 
Islands, New Holland and East Indies (Dana), Salawatti and New Britain (Studer), 
Ceylon (Ortmann), ? loc. (Esper). 

2. HcHrnopors wirsurisst4, Milne Edwards and Haime. (PI. 8, fig. 5; 9, fig. 4; 
18, figs 7 and 8 ; 15, figs 24; 17, fig. 1; 84, fig. 7.) 

7—2 


52 PERCY SLADEN TRUST EXPEDITION 


1816. Haxplanaria gemmacea, var. stellis comosis, Lamarck, Hist. Anim. sans vert., ii, p. 256; 2° édit., p. 399. 

11848. Hchinopora ringens, Dana, Expl. exp. Zooph., p. 279. 

1850.  Lchinopora rosularia (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 185. 

1850.  Hchinopora hirsutissima, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 187. 

1854.  Lchinopora hellt, Louis Rousseau, Voy. pole sud Dumont-d’Urville, Zool., v, p. 120, Zooph., pl. 27, 
fig. 3. 

1857.  Hchinopora hellr, Milne Edwards and Haime, Hist. nat. Corall., ii, p. 623. 

1857.  Hchinopora rosularia (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 623. 

1857.  Hchinopora harsutissima, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 624. 

1904. Hchinopora soladior, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 782, pl. 63, fig. 38 (non 
Echinopora solidior, Milne Kdwards and Haime). 

1904.  Hchinopora tertia, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 782, pl. 63, fig. 39. 


Corallum. Incrusting, relatively thin, much coarser than in #. lamellosa, edges 
usually somewhat folded over, free only at edges. Peritheca dense, with upright blunt 
spinulate echinulations up to 1 mm. in height, bases of spines sometimes united to form 
ridges especially seen towards edges. Corallites oval, sometimes circular, level with 
general surface to 1 mm. high, touching or up to 4 mm. apart, usually 2 or 3mm. Calices 
up to 8 mm. in diameter, average 5°5—6°5 x 4°5—5 mm., depth 2 mm. 

Septa in four orders, the fourth incomplete, comparatively broader than in 
E. lamellosa, sides spinulate, edges denticulate. Primaries thicker than secondaries ; 
both these and a varying number of tertiaries—up to 8—with their outer half or a 
little more thicker, from 5—7:5 mm., dipping down vertically till level of columella, 
then thinning out and extending horizontally inwards to meet the latter, around 
which is a ring of about 12 spinulate pali each 1°5 mm. high. Septa meeting columella 
exsert to 1°5 mm., exsert end divided over the calicular margin by a deep notch into an 
inner lobe about 1 mm. broad and an outer narrower lobe, the latter sometimes appearing 
as the first costal tooth. Septa often perforated behind pali, hence the latter appearing 
connected by slender rods to the septal margins. The tertiaries not meeting columella 
curve towards and fuse with sides of secondaries near to columella. Quaternaries very 
narrow and thin, up to 14 in number. Coste of primaries to tertiaries with 1—3 con- 
spicuous blunt upright spinulate echinulations, those of quaternaries thin and inconspicuous. 
Columella about one-third width of calyx or slightly more. 

The corallum of this species is very variable. #. tertia, Gard. (Pl. 15, fig. 3 and 
17, fig. 1) with identical polyp-structure as H. hirsutissima is only a skeletal variety, 
differing from it in the following respects :—(1) corallites projecting up to 2 mm., some- 
what further apart ; (2) costal and perithecal spines thinner and shorter, in places not 
regularly arranged; (3) septa less exsert, thinner and more perforated, the gaps often 
seen in the edges leaving slender irregular processes which tend to intercross over the 
axial fossee; (4) columella somewhat less than one-third width of calyx and formed 
of looser trabeculee. Another specimen with similar polyp-structure resembles L. Jamellosa 
(Pl. 14, fiz. 7) in skeletal characters more than #. hirsutissima—slender shorter 
perithecal spines, corallites more exsert (to 2mm.), calices circular and smaller, septa 
thinner, columella half width of calyx—but has a greater number of quaternaries, 
up to 14 as in EZ. hirsutisseoma, and a ring of about 12 pali not rising above calicular 


margin. 


MATTHAI—RECENT COLONIAL ASTRAIDA 53 


Polyps. (1) Tertiary couples of mesenteries sometimes complete with 12 couples, 
often incomplete (occasionally a quaternary couple present). (2) Directive grooves not 
deep. (3) Nematocysts I less numerous in ectoderm of oral-disc and outer wall of edge- 
zone than in £. gemmacea, but more numerous in lower halves of stomodeal ridges and 
in straight regions of mesenterial filaments than in that species. (4) Entoccelic pleats 
in stomodzal region of polyp horizontal, extremely narrow, thin, comparatively few 
and not extending to beyond the outer halves of primary mesenteries, usually restricted 
to their outer one-third ; inner one-fifth of mesenterial mesoglea much thickened, bearing 
exoceelic pleats sometimes better developed than entoccelic ones. (5) Mesenterial 
endoderm thicker on either side of inner stouter region of mesoglea. (6) Endoderm in 
outer wall of edge-zone as thick as the ectoderm over it, somewhat thinner in oral-disc. 
(7) Convolutions of mesenteries scarce towards bases of polyps. 

Remarks. A. Polyps. These are oval in outline, height about 6 mm., in retraction, 
with diameters 5mm. and 3°5mm., smaller towards edges of colonies. The primary 
mesenteries are usually attached to the sides of the septa a short distance from the 
theca, while the remaining mesenteries are attached at the angles between walls and 
septa. The convolutions of the mesenteries are scarce in the stomodzal region of the 
polyp, but are protruded into the edge-zone and through the oral-dise ; they are abundant 
below the stomodzum, becoming scarce again towards the base of the polyp. From every 
entoccele arises a tentacle, but it has not been possible to count the exact numbers of 
exoceelic tentacles ; the terminal batteries contain closely arranged nematocysts I, having 
from forty to forty-five turns of the spiral, with a few I1b interspersed among them. The 
stomodzum is oval in outline. with diameters 1:75 mm. and 1 mm.; one of the directive 
grooves is often shallow ; the ridges are thicker than broad, somewhat conical in transverse 
section, their sides sloping away, with the mesoglea not specially thickened at their bases; 
nematocysts III occasionally present, in their lower halves nematocysts I frequently found 
with fewer turns of the spiral. On the whole the stomodeum resembles in section that 
of Favia hululensis, Gardiner. 3 

In the ectoderm of oral-disc and outer wall of edge-zone mucous vacuoles are 
commonly present, nematocysts I fairly frequent, II rare. Large vacuoles are present 
in the sub-terminal batteries as in Mavia doreyensis Kd. and H. In the straight regions 
of mesenterial filaments nematocysts I are present in small numbers, occasionally those 
of II and II10; in their coils large numbers of nematocysts IIIb are closely arranged 
forming batteries, IT being less numerous. In the endoderm of the outer wall of the edge- 
zone the nuclei are arranged along its periphery, above which alge are massed together. 
In the oral-disc the endoderm is somewhat thinner and alge are less abundant; the 
circular layer of muscle-fibres is evident. The endoderm in the upper part of the 
column-wall is extremely thin, in certain parts being as thin as the calicoblast with alge 
comparatively scarce, towards the base less vacuolated, and hence less thickened than in 

most other genera. In the tentacles it is much vacuolated being as thick as or thicker than 
the sub-terminal batteries, with few alge as in E. lamellosa, sometimes blocking their 
lumina. The mesenteral endoderm thickened behind the filaments as much as the 
latter, appearing pad-like in section; beyond this it is thin along the short non-pleatal 


54 PERCY SLADEN TRUST EXPEDITION 


region. Algze not abundant in the mesenteries. Below the stomodeum the entoccelic 
pleats extend over the greater part of the width of the mesenteries, broadening out in 
the median regions, some of them secondarily cleft; the mesoglea is not thickened behind 
the filaments. . 

Polyps examined, eight, 4 from two typical specimens, 2 from specimen of tertia 
facies, 2 from specimen whose corallum has varied towards H. lamellosa, all from Hulule, 
Maldives. 

B. Corallum. Milne Edwards and Haime’s type of E. hirsutissima (Pl. 15, fig. 4) 
from the Indian Ocean, which is very likely Lamarck’s original of Haplanaria gemmacea, 
var. stellis comosis, is a large incrusting example measuring 30 x 20cm. with a hillock 
14cm. high, and has more or less the tertia facies, but its costee and costal spines are 
well developed. Resembling this is a specimen (14x11cm.) in the Paris Museum 
named £. rengens, Dana, but without locality. Louis Rousseau’s type of E. hella from 
Seychelles—later re-described by Milne Edwards and Haime—is also a large specimen 
(25 x 19x Ll em.) somewhat flat and raised into short hillocks, on the whole resembling 
specimen no. 9 on p. 57, but in places the perithecal spines are strongly developed as in 
no. 8. One of the specimens from Bourbon, measuring 13°5 x 6°5 cm., referred by Milne 
Edwards and Haime to #. rosularia, really belongs to the present species, its corallites 
projecting obliquely up to 3°5 mm. with about forty septa in each, of which about twenty 
meet the columella ; it has more or less the tertza facies. 

Localities. Maldives, Hulule (3). Chagos: Salomon (4); Coin, Peros (1). Also 
from Seychelles (Rousseau), Bourbon Island (Milne Edwards and Haime). 


3. HEcuinopora cemmacea (Lamarck). (Pl. 14, fig. 9; 15, figs. 5 and 6; 16, figs. 
5, 7, and 8; 17, figs. 2 and 3; 87, fig. 5.) 


1816. Haplanaria gemmacea, Lamarck, Hist. Anim. sans vert., ii, p. 256; 2° édit., p. 399. 

1830. Hchinastrea gemmacea, Blainville, Dict. Sci. Nat., 1x, p. 388; Manuel d’Actinol., p. 378. 

1834. Stephanocora hemprichi, forma fruticulosa and forma explanata, Ehrenberg, Corall. roth. Meer., p. 76. 

1834. Haplanaria hemprichii, Ehrenberg, Corall. roth. Meer., p. 82. 

1850. Astrea forskaliana, Milne-Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 100. 

1850. Hchinopora gemmacea, Milne-Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 186. 

1850. Hchinopora ehrenbergi, Milne-Edwards and Haime, Ann. Sci. Nat:, Zool., 3° sér., xii, p. 187. 

1850. Hcehinopora rousseaui, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 188. 

1850. Hchinopora solidior, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 189 (non Hchinopora 
solidior, Gardiner). 

1857. Heliastrea forskelana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 457, pl. D5, fig. 3. 

1857. Hchinopora gemmacea, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 622, pl. D9, fig. : 

1857. Lchinopora hemprichi, Milne Edwards and Haine, Hist. Nat. Corall., ii, p. 623. 

1857. Hchinopora ehrenbergi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 625. 

1857. Hchinopora solidior, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 626. 

1879. Orbicella forskdlana, Klunzinger, Korall. Roth. Meer., iii, p. 48. 

1879. Orbicella mammillosa, Klunzinger, Korall. Roth. Meer., iti, p. 49, pl. 5, fig. 5, pl. 10,.fig. 10 a—e. 

1879. Hchinopora fruticculosa, Klunzinger, Korall. Roth. Meer., iii, p. 55, pl. 6, fig. 4. 

1879. Hchinopora ehrenbergi, Klunzinger, Korall. Roth. Meer., iii, p. 56, pl. 6, figs. 7 and 9, pl. 10, fig. 15. 

1879. Lchinopora concamerata, Klunzinger, Korall. Roth. Meer., iii, p. 57, pl. 6, fig. 6. 

1879. Hchinopora carduus, Klunzinger, Korall. Roth. Meer., iii, p. 57, pl. 6, fig. 5, pl. 10, figs. 14@ and 6. 

1906. Orbicella forskalana, Marenzeller, Exp. “Pola,” Roth. Meer., Riffkorall., Zool. Ergeb. Wien, xxvi, 
p- 61. 


te 


MATTHAI—RECENT COLONIAL ASTR AIDA 55 


Corallum. All the variations in growth-form of the genus. Peritheca usually 
vesicular, walls of vesicles about ‘75 mm. apart, with distinct ridges connecting neighbour- 
ing coste ; spines present or absent. Corallites usually circular, projecting in different 
directions, vertical, oblique, or quite horizontal, up to 6 mm., usually about 3 mm., 
increasing in diameter from margins to bases, up to 6 mm. apart, average 2 or 2°5 mm. 
Calices with diameter up to 7 mm., depth 2 mm. 

Septa in four orders, up to 18 quaternaries, sides spinulate, edges denticulate. 
Primaries sometimes thicker than secondaries, these and up to 8 tertiaries meeting 
columella. Primaries and secondaries with their outer halves usually twice or thrice the 
thickness of the inner halves, dipping down vertically till level of columella, then thinning 
out and passing horizontally inwards to meet columella, exsert to 1 mm., the exsert end 
of each often divided by a notch over corallite-wall into a somewhat larger inner lobe and 
a smaller outer lobe, sometimes arched, not divided. Pali less than 12, about 6 or 
altogether absent. Septa perforated, the gaps in edges often leaving long, slender 
processes which in some corallites form fenestrze over the axial fossze. Those tertiaries not 
meeting columella usually curve towards and fuse with sides of secondaries, quaternaries 
also with tertiaries. Coste usually low flat ridges with spinulate, blunt, upright 
echinulations, varying in height and degree of roughness and presenting an appearance 
of large and small ridges as the costz of the last cycle are much less conspicuous, the 
latter sometimes absent. Columella formed of loosely twisted trabeculee, about one-third 
width of calyx. 

Polyps. (1) A few tertiary couples of mesenteries present, the cycle being never 
complete. (2) Directive grooves not deep. (8) Nematocysts I more numerous in 
ectoderm of oral-disc and outer wall of edge-zone than in EF. hirsutissima, but less 
abundant in lower halves of stomodeeal ridges and in straight regions of mesenterial 
filaments than in that species. (4) Entoccelic pleats in stomodzal region of polyps 
obliquely directed towards stomodzeum, narrower than in E. hirsutissima, and not 
extending beyond the outer halves of primary mesenteries, usually restricted to their 
outer one-third ; pleatal region much thicker than non-pleatal region; no exoccelic pleats. 
(5) Mesenterial endoderm thicker near stomodeal attachments than elsewhere. (6) En- 
doderm in outer wall of edge-zone two-thirds the thickness of the ectoderm over it, 
much thinner in oral-disc. (7) Convolutions of mesenteries scarce towards bases of 
polyps. 

Remarks, A. Polyps. These are circular in outline about 3°5 mm. in diameter. 
Tentacles are not present over the tertiary entocceles, and only a few exoccelic tentacles 
are present; as all the tentacles were in a retracted condition exact counting of the 
sub-terminal batteries was impossible.. The stomodeeum is 1°35 mm. long by ‘75 mm. 
broad; one directive groove is better developed than the other; the ridges are 
narrower than in H. hirsutissima. In the three polyps examined 2, 3, and 6 couples of 
tertiary mesenteries are present. 

The ectoderm of the oral-disc and of the outer wall of the edge-zone are somewhat 
thicker than in #. hirsutissima, with abundant mucous vacuoles. In most of the 
nematocysts IIIb the coiled thread is partially extruded; the dark-stained axis of II 


56 PERCY SLADEN TRUST EXPEDITION 


is usually somewhat bent. In the endoderm of the oral-dise algze are not massed 
together, hence its vacuolated condition better seen; the nuclei are not arranged along 
its periphery as in the previous two species; the circular layer of muscle-fibres is 
distinctly visible. The mesenterial endoderm appears somewhat diffuse and transparent, 
with fewer alge than in the two previous species, and is not specially thickened into 
a pad behind the filaments. Gonads were not present in any of the polyps examined 

Polyps examined, three; from a specimen from Dongonab, Red Sea. 

B. Corallum. In the Paris Museum there are three specimens referred by Lamarck 
to Hxplanaria gemmacea, later re-described by Milne Edwards and Haime as Hchinopora 
gemmacea; one (9x9 em.) of these from the Red Sea is identical with my types of 
the present species; another (10x9 cm.) from the Indian Ocean is a concavo-convex 
specimen, with a short hump and with the corallites projecting obliquely, in every respect 
resembling specimen no. 12 on p. 57; the third (8x75 cm.) from the same locality is 
a flat, somewhat worn-out example. Seven excellent specimens have been assigned by 
Milne Edwards and Haime to EL. ehrenbergi; two of these from Seychelles (30 x 21 cm. 
and 25 x 23 cm.) have typical corallites on the flat regions, but on the frequently branching 
hillocks corallites and peritheca are much coarser, as in specimen no. 10 on p. 57; a specimen 
(19°5 x 11°5 em.) from Red Sea, with humps, has obliquely projecting corallites and a few 
short perithecal spines resembling no. 12; another specimen (19 x 18 cm.) from the same 
locality has an incrusting corallum rising into branching hillocks, the flat regions 
resembling no. 13, while the hillocks are identical with Stephanocora hempricha forma 
fruticulosa of Ehrenberg; the remaining three specimens from Red Sea have the 
fruticulosa facies, but the calices are somewhat smaller, 4—5 mm. in diameter. Milne 
Edwards and Haime’s original of E. solidior (22 x 19 cm., Pl. 16, fig. 5) has an inerusting 
corallum raised into short humps, being identical with my typical specimens of £. 
gemmacea; the average diameter of its calices is about 5°5 mm.; perithecal spines are 
slender and septa thin in calices. There are no specimens at present in the Paris 
Museum named /. hemprichi, but doubtless the species comes under HL. gemmacea. 

I have carefully examined the five large specimens from Red Sea, for which Milne 
Edwards and Haime constituted a new species Heliastrea forskelana, and have no 
hesitation in referring them to H. gemmacea. 

In the Berlin Museum are three specimens referred by Klunzinger to E. ehrenbergi ; 
two of these were Ehrenberg’s originals of Stephanocora hemprichit. Klunzinger has 
figured the smaller (Pl. VI, fig. 7); the larger measures 25 x 16 cm.; the remaining one is 
a small edge specimen, 9 x 6 em, which also has been figured (PI. VI, fig. 9). Ehrenberg’s 
type of Stephanocora hemprichi forma fruticulosa is a large specimen with a dendroid 
mode of growth, inconspicuous perithecal and costal spines, costee continuous from 
corallite to corallite over the perithecal regions, and calices somewhat larger. Klunzinger 
separated it into a new species H. fruticulosa, but it is only a skeletal variety of the 
present species, as is evident from some of Milne Edwards and Haime’s examples 
of E. ehrenbergi, on which fruticulosa hillocks are seen on typical coralla (see Pl. 16, 
fig. 5). Klunzinger’s type (20°5 x 12°5 cm.) of EH. concamerata—tormerly Ehrenberg’s 
example of Haplanaria hemprichi—has some of the characters of H. lamellosa, Esper, 


MATTH AI—RECENT COLONIAL ASTRAIDA 57 


LIST OF SPECIMENS SHOWING THE EXTENT OF VARIATION IN 
THE GENUS FLCHINOPORA. 


No. of 
Specimen 


Locality 


Species 


Remarks 


bo 


10 


11 


12 


13 


14 


Hulule, Maldives 


¢ 
Coin, Peros, Chagos 


Seychelles 


Salomon, Chagos 


Hulule, Maldives 


”? 


Coin, Peros, Chagos 
Salomon, Chagos 


Dongonab, Red Sea 


” 


E. lamellosa 


E. hirsutissima 


EL. gemmacea 


” 


Corallum 14x 9x7 em. A ring of about 12 paliform 
lobes in corallites. Columella one-third width of calyx. 
Perithecal spines shorter than type. Pl. 14, fig. 3. 


Specimen large, 28 x 14x 2cm. Corallites projecting up 
to 4:6 mm. increasing in diameter from free margins 
(up to 3 mm.) to bases (up to 5 mm.). Primaries and 
secondaries similar, thinner than in type specimens. 
Pl. 14, fig. 4. 


A small edge-piece (10 x 7 cm.). Corallites level with 
surface. About 24 thin septa, 12 meeting columella, all 
slightly exsert. Columella one-quarter to one-third width 
of calyx. Largest calyx 2°5 mm. in diameter, resem- 
bling &. striatula, Stud. 


Specimen 13x11 cm. Columella indistinct. A ring of 
12 paliform lobes present. PI. 14, fig. 5. 


Specimen 13x9cm. Corallites projecting up to 4 mm., 
primaries much thicker than secondaries. Columella 
rudimentary one-fifth to one-quarter width of calyx. Pl. 
14, fig. 6. 


Corallum (14 x 10x 7 em.). with the tertia facies. 
Specimen small, simulates #7. damellosa. Pl. 14, fig. 7. 


Specimen flat (17 x 14 em.). Calices larger than in type 
specimens up to 8 mm. in diameter. Septa up to 44, 
up to 22 meeting columella; primaries usually much 
thickened in their outer halves. Pl. 15, fig. 2. 


Corallum 13 x 8 x 4 cm. with the tertia facies but calices 
larger, up to 7:25x5 mm. Septa up to 44, up to 19 
meeting columella—resembling Milne-Edwards and 
Haime’s type of #. hell. PI. 15, fig. 3. 


Corallum incrusting on a large irregular mass (33 x 30 x 23 
em.) and rising into a great many branching hillocks. 
Corallites and peritheca very coarse. Corallites on hil- 
locks projecting in all directions vertical, horizontal and 
oblique, in valleys level. Pl. 16, fig. 8. 


Corallum 17 x 13x 10¢m. On one side costae thin and 
perithecal spines slender, few and only slightly rough ; 
on another side typical. Pl. 87, fig. 5. 


Corallum (11 x8 cm.) incrusting on a concavo-convex 
surface. Corallites projecting obliquely towards edges, 
sometimes almost horizontal, height up to 9 mm. Peri- 
thecal spines very short or absent. Pl. 15, fig. .6. 


Corallum (13 x 14 em.) broken off, with a thick stem and 
short dividing branches on it. Calices shallow, some- 
times almost flat. Up to 20 septa making columella, pali 
inconspicuous. Costae thin, spines short on costae, absent 
on peritheca. Pl. 16, fig. 7; Pl. 17, fig. 3. 


Specimen (15 x 12 cm.) resembles Klunzinger’s fig. of EB. 
concamerata. Pl. 17, fig. 2. 


SECOND SERIES—ZOOLOGY, VOL. XVII. ; 8 


58 PERCY SLADEN TRUST EXPEDITION 


viz. thin fan-shaped corallum, small corallites each with not more than thirty septa, 
usually twenty-four ; but it has the rougher facies of #. gemmacea, the resemblance to 
E. lamellosa being due to its having been an edge piece. ‘Two specimens are referred 
by Klunzinger to his new species #. carduus, but this is only a variety of the present 
species, having long, rough costal spines. Klunzinger’s figured type (14 x 9°5 x7 em.) 
of Orbicella mammullosa is in no way different from Milne Edwards and Haime’s examples 
of Heliastrea forskelana, and like them undoubtedly belongs to H. gemmacea. A large 
specimen from Dar-es-salaam named £. hemprichi, Ed. and H., by Ortmann also belongs 
to the present species. 

Of the “‘ Pola” specimens in the Hofmuseum, Vienna, which come under the present 
species, Marenzeller referred three to Orbicella forskalana, two to E. ehrenberqr, and 
seven excellent examples to #. fruticulosa, the largest measuring 34 x 30 x 30 em.; these 
latter have incrusting coralla with the typical gemmacea facies, but are raised in places 
into branching hillocks with the fruticulosa facies, thus affording conclusive proof that 
Ei. fruticulosa is only a skeletal variation of H. gemmacea. 

Localities. Red Sea (13) and broken pieces of fruticulosa facies. Also from the 
Indian Ocean and Seychelles (Milne Edwards and Haime), Dar-es-salaam (Ortmann). 
A common species in the Red Sea and Indian Ocean. 


GALAXEA (OKEN). 


1815. Galawea (pars), Oken, Lehrb. Naturg., i, p. 72. 

1816. Sarcinula (pars), Lamarck, Hist. Anim, sans vert., ii, p. 222. 

1816. Caryophyllia (pars), Lamarck, Hist. Anim. sans vert., ii, p. 224. 

1820. Anthophyllwm (pars), Schweigger, Handb. Naturg., p. 417. 

1834. Amnthophyllum, Ehrenberg, Corall. roth. Meer., p. 89. 

1848. Sarcinula, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 310. 
1851. Galaxea, Milne Edwards and Haime, Pol. foss. terr. paleoz., etc., p. 70. 
1857. Galaxea, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 223. 

1879. Galaxea, Klunzinger, Korall. Roth. Meer., ii, p. 77. 

1884. Galaxea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 118. 

1899. Galaxea, Gardiner, Proc. Zool. Soc. London, Zool., p. 762. 

1904. Galaxea, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 782. 


Corallum. Distinct peritheca between the corallites which pass through it and 
stand up above its surface, so as to appear quite separate. Peritheca formed of arched 
vesicles varying from °75x°5 mm. to 2X1 mm., arranged in alternating tiers and 
appearing like blisters on the surface. Colonies rather massive, but generally much 
bored into below, this tending to break them up. Corallites from 2 to 8°5 mm. across 
somewhat conical. Septa generally markedly exsert, sloping obliquely towards a columella 
formed by their fused ends, almost flat above and dropping somewhat vertically to 
the coste, which are only conspicuous near the margins. Orders of septa from 3 to 5. 

Polyps. Varying in size. Hdge-zones extending over the free surfaces of corallites 
and continuing as the ccenosare over peritheca. Mesenteries forming two to four cycles, 
first two always complete, each of six couples; primaries meeting stomodzeum, third cycle 
sometimes incomplete, the fourth when present incomplete ; all with filaments. Tentacles 
corresponding in number and position with entocceles and exocceles, each with a large 


MATTHAI—-RECENT COLONIAL ASTRAIDA 39 


terminal battery and sub-terminal batteries if visible of varying number. Stomodzeum short, 
much compressed laterally ; directive grooves deep and narrow. Ovoid bodies* present 
everywhere in ectoderm excepting in calicoblast; they appear to be degenerate nematocysts 
of nos. III or I. Entoccelic pleats broad, constricted at their bases and often sub- 
divided, below stomodzeum extending over two-thirds breadth of mesenteries, decreasing 
in size from skeletal attachments. Entoccelic muscle-bands large. Endoderm of body- 
wall above enterostome thin, below distended by vacuolation with gradual loss of nuclei, 
so that it forms at base of polyp a highly reticulated non-nucleated layer, filling up 
the greater part of the gastro-vascular spaces. Ova in a single longitudinal row in 
every mesentery. Multiplication by budding usually from ccenosare, sometimes from 
edge-zone. 

Remarks. In the retracted condition the oral-dise is unevenly raised up by the 
highly exsert septa and deeply cleft radially by the ingrowth of the latter, leaving only 
a narrow undivided portion around the slit-like mouth. 

Owing to insufficiency of polyp material, I have not been able to determine 
satisfactorily the relationships of the many recorded species of this genus. 

G. bougainviller (Blain.), G'. pauciradiata (Blain.), G. hexagonalis, Ed. and H., and 
G. laperouseana, Ed. and H., are missing from the Paris Museum, and Anthophyllum 
musicale (Khrb.) from the Berlin Museum. Without polyps the relationships of Milne 
Edwards and Haime’s four examples of G. quoyi, Ed. and H., to the species of Galaxea 
described in this paper cannot be determined. ‘Their corallites are smaller than those 
of G. fascicularis and the corallum is lighter ; quinary septa are also wanting. 

I have only given the principal characters of Prof. Gardiner’s examples of G. hexa- 
gonalis and G. laperouseana, since without polyps it is impossible to determine their 
respective places in the genus; Milne Edwards and Haime’s type specimens of these two 
Species are missing from the Paris Museum. 

Distribution.—Red Sea, Indian and Pacific Oceans. 


1. Gataxea rascrcutaris (LINN&US). (PI. 8, fig. 4; 16, fig. 4; 88, fig. 6; 34, fig. 3; 
38, fig. 6.) 


1767. Madrepora fascicularis, Linneus, Syst. Nat., edit. 12, p. 1278. 

1786. Madrepora fascicularis, Ellis and Solander, Nat. Hist. Zooph., p. 151, pl. 30. 

1791. Madrepora cuspidata, Esper, Forts. Pflanz., i, p. 155, pl. 28, figs. 1 and 2. 

11791. Madrepora fascicularis, Esper, Forts. Pflanz., i, p. 157, pl. 29, figs. 1 and 2. 

1815. Galaxea fascicularis, Oken, Lehrb. Naturg., i, p. 73. 

1815. Galaxea cuspidata, Oken, Lehrb. Naturg., 1, p. 73. 

1816. Caryophyllia fasciculata, Lamarck, Hist. Anim. sans vert., ii, p. 226,—2° édit., p. 349. 
1818, Caryophyllia fasciculata, Blainville, Dict. Sci. Nat., vii, p. 194. 

1834. Anthophyllum fasiculare, Ehrenberg, Corall. roth. Meer., p. 89. 

1846. Anthophyllum cuspidatum, Dana, Expl. exp. Zooph., p. 401. 

1846. Anthophyllum hystrix, Dana, Expl. exp. Zooph., p. 401, pl. 28, fig. 2. 

1848. Sarcinula fascicularis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 313. 


* Prof. Bourne (15, p. 531, fig. 30) has used the term “ovoid bodies” to somewhat similar structures in the 

calicoblastic ectoderm of Caryophyllia smithii, which he regarded as degenerate nematocysts. If these bodies 

and the ones recorded above are homologous, it is noteworthy that the latter should be absent everywhere from 
the calicoblast of the two species of Galaxea, whose polyps I have studied. 


8 —2 


60 PERCY SLADEN TRUST EXPEDITION 


1848. Sarcinula ellissii, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 315. 

11848. Sarcinula irregularis, Milne Kdwards and Haime, Amn. Sci. Nat., Zool., 3° sér., x, p. 316. 

1848. Sarcinula hystrix, Milne Edwards and Haime, Ann. Sci. Nat., 3° sér., x, p. 318. 

1851. Galaxea fascicularis ellisi and irregularis, Milne Edwards and Haime, Pol. foss. terr. paleoz., etc., p. 71. 
1857. Galamwea fascicularis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 227. 

1857. Galawxea ellisi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 228. 

11857. Galaxea irregularis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 229, pl. D2, fig. 2a and 6. 
1877. Galaxea fascicularis, Studer, Monat. Ak. Wiss. Berlin, p. 637. 

1877. Galaxea cespitosa, Studer, Monat. Ak. Wiss. Berlin, p. 637. 

1879. Galaxea fascicularis, Klunzinger, Korall. Roth. Meer., ii, p. 78. 

21879. Galaxea irregularis, Klunzinger, Korall. Roth. Meer., ii, p. 78, pl. 7, fig. 11. 

1886. Galawea aspera, Quelch, Reef Corals, Challenger Reports, Zool., vol. xvi, pt xlvi, p. 72, pl. 4, figs. 5—5d. 
1886. Galaxea ellisii, Quelch, Reef Corals, Challenger Reports, Zool.. vol. xvi, pt xlvi, p. 72. 

1899. Galaea fascicularis, Gardiner, Proc. Zool. Soc., p. 763. 

1904. Galaxea fascicularis, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 783. 

1906. Galaxea irreqularis, Marenzeller, Exp. “Pola,” Rot. Meer., Riffkorall., Zool. Ergeb. Wien, xxvi, p. 54. 


Corallum. Perithecal vesicles small, °75x‘d mm. in size, usually open on surface. 
Corallites circular or oval in outline, projecting above peritheca to 15 mm. or possibly 
more, decreasing in diameter from calicular opening (8°5 mm.) to peritheca (5 mm.), 
1—2°5 mm. apart, diverging upwards from peritheca; walls °75 mm. thick towards bases, 
thinning to the free margins. Largest calices when round 8 mm. in diameter, when oval 
9x6mm.; depth from calicular opening to columella 3 mm. 

Septa in five orders, the first four complete, the fifth incomplete, very narrow, 
maximum number ten, hence most of the quaternaries exoccelic ; septa exsert vertically, 
sides smooth, edges entire, sharp; primaries thicker than secondaries, both exsert for 
2—3 mm., not swollen in thecz, and meeting columella. Tertiaries with edges bent 
towards and united with sides of secondaries, sometimes extending near to columella ; 
quaternaries oceasionally fused with sides of tertiaries, both exsert to 1 mm. Coste 
smooth, seen as low sharp ridges in upper third of corallite-wall, below as striz. Columella 
poorly developed, primary and secondary septa almost meeting in centre (as in Baryastrea 
transversa). Budding near the edge of corallum from peritheca. 

The smallest corallite measured 5°5 mm. in height and 3 mm. in diameter, with three 
complete orders of septa, the primaries and secondaries meeting columella. 

Polyps. (1) Size corresponding with corallites. (2) Entoccelic tentacles as well 
developed as exoccelic; terminal batteries with thickened ectoderm and closely packed 
nematocysts; sub-terminal batteries well defined, up to thirty or thirty-five. (3) Exoccelic 
tentacles similar. (4) Quaternary couples of mesenteries present, cycle incomplete. 
(5) Ectoderm of outer wall of edge-zone and oral-dise about the same thickness. 
(6) “Ovoid bodies” numerous. (7) Algee not very abundant. 

Remarks. A. Polyps. Quaternary couples of mesenteries in the two large polyps 
sectioned numbered 0, 0, 0, 3, 0, 2 and 0, 3, 2, 1, 2,3. The convolutions of mesenteries are 
massed together in the inter-mesenteric chambers for some distance below the stomodzeum, 
but scarce towards the base of the polyp. The stomodzeum with lumen distinct, diameters 
2x°75 mm. In some of the “ovoid bodies” a thick central core with a faint spiral 
around it is visible, suggestive of degenerate nematocysts III. The calicoblast is 
thickened near the skeletal attachments of the mesenteries about the thickness of the 


MATTHAI—RECENT COLONIAL ASTRAIDA 61 


ectoderm of the outer wall of the edge-zone. The ectoderm of the oral-dise is thick with 
nuclei massed together in its lower half, ovoid bodies rare, nematocysts I and IIb frequently 
present. In the upper half of the ectoderm of the outer wall of the edge-zone nuclei are 
arranged in layers with numerous mucous vacuoles and nematocysts I and II. In the 
terminal batteries, nematocysts IIb are more numerous than I, the latter with up to 
30 turns of the spiral. Ovoid bodies absent, but present in the sub-terminal batteries. 
Mucous vacuoles stain orange with hematoxylin and eosin as in Leptastrea rowssyana, 
Ed. and H., in contrast to species where the colour is brown. The stomodzeal ridges are 
as broad as or broader than thick, nuclei massed together towards the lumen and ovoid 
bodies towards the mesoglea; nematocysts absent. Filaments are well developed on 
primary and secondary mesenteries, rudimentary on rest; nematocysts are scarce in their 
straight regions, but I and II occur in small numbers in the coils. The mesenterial 
mesoglea thickens near the stomodzeum, on this part exoccelic fibres being better 
developed. The endoderm in the oral-disc and in the body-wall above the enterostome is 
thin, and contains a few algze; it is much thickened in the outer wall of the edge-zone, 
thin in its inner wall; in the tentacles it is as thick as the ectoderm of the sub-terminal 
batteries, and algze are more abundant in the entoccelic than in the exoccelic tentacles ; 
over the mesenteries the endoderm is comparatively thin, except near the stomodzal 
attachments of the primaries. 

Gonads were not present in the polyps examined. 

Polyps examined, three ; from two colonies from the Red Sea. One of these was a 
young polyp 4mm. in height and 2mm. in diameter, in which only four tertiary couples 
had appeared. 

B. Corallum. The corallum of this species is apparently very variable. Most of the 
examples of G. wregularis, Ed. and H., that I have examined differ from my specimen, 
whose polyps were sectioned in having larger corallites, which are laterally compressed or 
distorted. The appearance suggests that the large size and the distortion might be due to 
a somewhat retarded fission, but, of course, this can be ascertained only by an examination 
of the polyps. 

The specimens referred by Prof. Gardiner to G. fascicularis form a somewhat hetero- 
geneous lot, one of them approaching G'. musicalis in appearance. They differ from the 
specimen on Pl. 16, fig 4, in the following respects: (1) primary and secondary septa of almost 
equal thickness and more highly exsert, up to 4 mm. ; (2) primaries swollen in thece but 
thinner in calices; (3) costze more prominent; (4) columella better developed ; 
(5) corallites often irregularly compressed, tending to be sub-triangular in outline and 
with thicker walls. In other words, they have more or less the wregularis facies. 

Milne Edwards and Haime have referred nine specimens in the Paris Museum to 
G. fascicularis, of which four are in Lamarck’s collection, and four others to G. wrregularis ; 
an example of each of these species resembles my figured specimen. The same authors 
_ have assigned three specimens to G. ellisi; the same name is written on four specimens of 
Rousseau’s and one of Agassiz’s. 

Of the four examples in the Berlin Museum named Anthophyllum fasciculare by 
Ehrenberg, No. 624 (12 x7 cm., Pl. 88, fig. 6) is identical with my figured example ; the 


62 PERCY SLADEN TRUST EXPEDITION 


remaining three have much larger corallites, usually compressed or distorted, which 
Klunzinger later referred to G. wrregularis, Ed. and H. Resembling these latter are two 
of the “Gazelle” specimens referred by Studer to G. fascicularis. The large examples 
Studer referred to G. cespitosa, Dana, have corallites of the same shape and size as in my 
figured type, but the septa are rougher. 

Marenzeller has named twenty-one specimens of the ‘“ Pola” expedition G. wrregu- 
lavis, of which eight are numbered. No. 15898 resembles my type in every respect; 
in the remaining ones the corallites are larger and distorted. In one of the unnumbered 
specimens from Massawah some of the corallites are exactly as in my figure, while others 
have the irregulars facies. 

In the Copenhagen Museum is a single rubbed specimen, doubtfully regarded as 
Forskal’s original of Madrepora diergens; its characters are beyond recognition. 
Ellis and Solander’s figured type of Madrepora fascicularis is missing from the Glasgow 
University Museum, but their figure resembles my specimens from the Red Sea. 

Quelch’s type of G. aspera from Amboina measures 9 x 6 x 9 em. ; a fifth septal cycle 
is wanting in the corallites, while the fourth is incomplete, but these differences are to be 
attributed to the smallness of the specimen. His example of G. ellis, Ed. and H., from 
Mactan Island, Philippines, has somewhat the ivregularis facies, many of the corallites 
being laterally compressed, with 15—17 septa meeting the columella. The place of his 
type (9x 8x5cm.) of G. fragilis from Amboina is more doubtful; its corallites are 
smaller and thinner (both walls and septa) than in his type of G. aspera, but are wider 
apart, perithecal vesicles larger, costee prominent and sides of septa rough. 

A specimen (10 7X7 cm.) from Ponape, named Galaxea tenella, Briigg., sent from 
the Hamburg Museum, and now in the British Museum, is perhaps only a variety of 
G. fascicularis ; the corallites are close together, the septa in five cycles (fifth incomplete), 
up to eighteen meeting the columella. 

Localities. Red Sea (8 large, some with the wregularis facies and a few small ones). 
Aldabra, Lagoon Reef (6, mostly with the regularis facies). Saya de Malha (1, with the 
irregularis facies from 26 fms.). Also from Seychelles (Milne Edwards and Haime), 
Fiji Islands (Dana), Amboina and Philippines (Quelch), Ponape (Briiggemann), loc. ? (Ellis 
and Solander), China Seas (Esper). 


2. G'aLaxeA musrcatis (Linneus). (Pl. 16, figs. 2 and 3.) 
11767. Madrepora musicalis, Linneeus, Syst. Nat., edit. 12, p. 1278. 
1791. Madrepora musicalis (pars), Esper, Forts. Pflanz., p. 160, pl. 30, fig. 1. 
1815. Galaxea musicalis, Oken, Lehrb. Naturg., i, p. 73. 
1816. Caryophyllia musicalis, Lamarck, Hist. Anim. sans vert., ii, p. 227; 2° édit., p. 350. 
1817. Caryophyllia musicalis, Blainville, Dict. Sci. Nat., vii, p. 195. 
1830. Sarcinula musicalis, Blainville, Dict. Sci. Nat., 1x, p. 314; Manuel d’Actinol., p. 348. 
1848. Anthophyllum musicale, Dana, Expl. exp. Zooph., p. 399. 
1848. Sarcinula musicalis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 312. 
1848. Sarcinula erecta, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 317. 
1851. Galaxea musicalis and clavus, Milne Edwards and Haime, Pol. foss. terr. palzoz., etce., pp. 70 and 71. 
1857. Galaxea musicalis, Milne Edwards and Haime, Hist. Nat. Corall., ti, p. 225. 
1857. Galaxea clavws, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 232. 
1889. Galaxea heterocyathus, Ortmann, Steinkorall, Stid. Ceylons, Zool. Jahrb., iv, p. 534, pl. 16, fig. 12. 
1904. Galaxea musicalis, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 783. 


MATTHAI—RECENT COLONIAL ASTRAIID A 63 


Corallum. Perithecal vesicles slightly larger than in G. fascicularis ; usually closed 
on surface. Corallites oval or circular, much smaller than in last species, projecting above 
peritheca up to 3°5 mm., of even diameter or somewhat increasing from calicular opening 
to peritheca, up to 5 mm. apart, usually 3 mm., not so diverging as in last. Walls of about 
same thickness. Calices with diameters 4 x 3 mm., depth 1°75 mm. 

Septa in four orders, the first three complete, the fourth incomplete, maximum 
number 13, hence most of the tertiaries exoccelic ; septa exsert vertically, sides rough, 
edges entire sharp, primaries usually thicker than secondaries, both exsert to 1°5 mm., 
swollen in thecee and meeting columella. ‘Tertiaries with edges free and exsert to 1°5 mm. 
Costee smooth, seen as sharp ridges along almost the whole projecting part of corallite- 
wall. Columella distinct, circular or oval in outline, about -75 mm. broad, sometimes 
laterally compressed and septa meeting above it. Budding from any part of peritheca. 

Polyps. (1) Size small, corresponding with corallites. (2) Entocclic tentacles 
much less differentiated than exoccelic; terminal batteries with loosely arranged nemato- 
eysts, not swollen; sub-terminal batteries not defined, with numerous nematocysts. 
(3) Exoccelic tentacle with up to about twelve sub-terminal batteries. (4) Quaternary 
couples of mesenteries absent. (5) Hctoderm of outer wall of edge-zone not more than 
half the thickness of oral-dise ectoderm. (6) “ Ovoid bodies” fewer than in Gt. fasci- 
cularis. (7) Algee much more abundant than in G. fascicularis. 

Remarks. A. Polyps. The tertiary cycle of mesenteries was incomplete, the 
couples being 6, 3 and 1 in the three polyps examined. One of the polyps was abnormal 
in possessing a third couple of directives. The exoccelic tentacles are somewhat bluntly 
pointed. The primary septa press against the stomodeum near the enterostome, 
greatly distorting its wall and almost occluding its lumen; if this means an imperfect 
functional capacity, as I am disposed to suggest, it is correlated with a great abundance 
of alge. The ectoderm of the oral-disc is thick, with numerous nematocysts I and a few 
of If. Ovoid bodies are scarce in both the oral-dise and edge-zone; some of them are 
narrower than in G. fascicularis, with no central core but with traces of a spiral 
suggesting degenerate nematocysts I. In the terminal batteries of exoccelic tentacles, 
nematocysts II b are less numerous than in the last species. Ovoid bodies are present in 
the sub-terminal batteries. The stomodzeal ridges are not so thick as in the last species. 
In the coils of the mesenterial filaments nematocysts I are more numerous. Entoccelic 

pleats are broader and thicker than in G. fascicularis, and tend to arise in groups of 

three or four, the pleats of each group often fusing at their bases to form thick compound 
ridges. The endoderm in the oral-disc is thicker than in the last species, and is massed 
with algze ; it is much thinner in the outer wall of the edge-zone, considerably thickened 
in the tentacles owing to vacuolation, and filled with alge, occluding the lumen in the 
region of the terminal batteries of the exoccelic tentacles. The mesenterial endoderm 
is somewhat thicker on the exoccelic side where alge are massed together. 

The polyps examined were ripe females. The eggs are carried in a single row on the 
exoccelic side of every mesentery, in transverse section each egg almost fitting the 
exoceele. The appearance of a double row of eggs is sometimes presented, owing to 
either the overlapping of neighbouring eggs or to the eggs of adjacent mesenteries being 


64 PERCY SLADEN TRUST EXPEDITION 


accommodated in the exoccele, one in front of the other. Small immature ova are present 
in the endoderm surrounding the eggs. 

Polyps examined, six, from a dredged specimen from Maldives. 

B. Corallum. Milne Edwards and Haime’s type of G. musicalis, which was perhaps 
Lamarck’s type of Caryophyllia musicalis, is a small rubbed specimen. Of their three 
examples of G. clavus, one is equally damaged, but the two others resemble my examples 
of the present species. 

Ortmann’s figured type of G. heterocyathus is a long, somewhat cylindrical specimen, 
measuring 23 X16 cm. 

Quelch’s recorded example (11 x9 cm.) of G. musicalis from Somerset, Cape York, 
has corallites with broken margins. A specimen (7x7 x4 cm.) from Kandavu and 
another (4°5 x 4°5 em.) from Amboina, which he referred to G. fascicularis and G. tenella, 
Briigg., respectively, come under the present species in size of corallites, number and 
roughness of septa, thickness of columella, etc. 

Localities. Maldives (3). Coin, Peros (1). Also from Fiji (Milne Edwards and Haime 
and Quelch) ; Ceylon (Ortmann), Torres Strait and Amboina (Quelch). 


3. Ganaxea LAmARCKI, Milne Edwards and Haime. (Pl. 18, fig. 6; 16, fig. 1; 
34, fig. 2.) 
11766. Madrepora organum, Pallas, Elench. Zooph., p. 317. 
11815. Galaxea organum, Oken, Lehrb. Naturg., i, p. 73. 
1816. Sarcinula organwm, Lamarck, Hist. Anim. sans vert., ii, p. 223; 2° édit., p. 241 (non Wadrepora 
organum, Linneus). 
1816. Caryophyllia astreata, Lamarck, Hist. Anim. sans vert., li, p. 227; 2° édit., p. 250. 
1830. Sarcinula astreata, Blainville, Dict. Sci. Nat., lx, p. 314; Manuel d’Actinol., p. 348. 
1834, Anthophyllum spherula, Ehrenberg, Corall. roth. Meer., p. 89. 
1848. Sarcinula organum, Milne Edwards, Atlas grande édit. Reégne Anim. Cuvier, Zooph., pl. 85, fig. 1 
1848. Sarcinula organwm, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 311. 
1848. Sarcinula astreata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, p. 317. 
1851. Galaxea lamarcki and astreata, Milne Edwards and Haime, Pol. foss. terr. paleeoz., etc., pp. 70 and 71. 
1857. Galaxea lamarcki, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 225. 
1857. Galaxea astreata, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 230. 
1879. Galaxea lamarcki, Klunzinger, Korall. Roth. Meer., ii, p. 77. 
1886. Galaxea explanata, Quelch, Reef Corals, Challenger Reports, Zool., vol. xvi, pt. xlvi, p. 71, pl. 4, 
figs. 6—6d. 
1904. Galaxea lamarcki, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 783. 
1904. Galazea, sp.? Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 783. 


Corallum. Perithecal vesicles larger than in any previous species, up to 2X 1 mm.— 
hence corallum very ight—closed on surface. Corallites always round, smaller than in all 
other species, less projecting, maximum 2mm.,2—4 mm. apart. Walls up to ‘5 mm. thick. 
Calices usually 2 mm. in diameter, maximum 3 mm. Septa in three orders, sides rough, 
edges entire; usually primaries only meeting columella, exsert up to 1 mm., secondaries 
thinner, sometimes varying in number, rarely all the six reaching columella. Coste incon- 
spicuous. Columella circular in outline, distinct. Budding from any part of peritheca. 

Polyps useless for any detailed study. Two were sectioned from specimen from 
S. Nilandu (36 fms.) and one from another specimen from Peros (16 fms.). 

Remarks. There is no doubt that this is a distinct species. The small piece from 


MATTHAI—RECENT COLONIAL ASTRAIDA 65 


Felidu, about whose identity Prof. Gardiner was uncertain, is only a skeletal variety of 
G. lamarch, differing from the latter in having less projecting corallites and indistinct 
columella. 

The five small specimens referred by Milne Edwards and Haime to G. lamarchi, 
perhaps originally Lamarck’s examples of Sarcimula organum, are all much defaced. Of 
their three examples of G. astreata—all of which are in good condition—one is in Lamarck’s 
collection, being probably his original of Caryophyllia astreata. 

Ehrenberg’s type of Anthophyllum spherula is a small round specimen in bad 
condition, but of its place in the present species there is no doubt. 

Quelch’s large type (21 x 10 cm.) of G. explanata resembles my examples of G. lamarchi 
in possessing only three septal cycles, the primaries and most of the secondaries meeting 
columella, and large perithecal vessels, but the corallites project much higher and are further 
apart ; the primaries are thicker than the secondaries and have more prominent costee. 

Localities. Maldives: Nilandu, 36 fms. (1); Felidu, 25 fms. (1). Chagos: Diamont, 
Peros, 16 fms. (1); Coin, Peros (1). Saya de Malha, 29 fms. Also from Red Sea 
(Ehrenberg and ? Lamarck), Fiji (Quelch). 


4. GaLdéxea HEXAGONALIS (Kd. and H.) (?). 


Galaxea hexagonalis, Gardiner, Fauna Geogr. Maldives and Laccadives, 783 (1904). 


Perithecal vesicles small. Corallites smaller than in G. fascicularis, irregularly 
compressed, becoming sub-triangular in outline, projecting to 9 mm., 1—2 mm. apart. Walls 
thin. Largest calyx 8x5 mm. Septa in four orders, the fourth incomplete—up to 19— 
sides with short spines, edges denticulate ; about 15 septa meeting columella; secondaries 
as thick as primaries ; tertiaries exsert obliquely outwards. Costee prominent, raised into 
short spines at wide intervals. Columella distinct. Budding towards edge of corallum 
from peritheca, sometimes towards bases of corallites. Sometimes two corallites fuse, in 
appearance simulating fission. 

The polyps are too badly preserved to make out their specific characters. 

Locality. Minikoi. Milne Edwards and Haime’s type specimens are missing from 
the Paris Museum. 


5. G'aLAXEA LAPEROUSEANEA (Hd. and H.) (2). 
Galanea laperouseanea, Gardiner, Proc. Zool. Soc., 1899, 762. 

Perithecal vesicles small. Corallites oval or sub-triangular, projecting to about 7 mm., 
1—2 mm. apart. Walls up to 2 mm. thick. Largest calyx 10x5 mm. Septa in four 
orders, up to 14 quaternaries, sides slightly rough, edges entire, about 17 septa meeting 
columella; secondaries usually as thick as primaries, both swollen in thecee and exsert 
vertically; tertiaries exsert obliquely outwards, hence their coste appearing more 
prominent than those of primaries and secondaries. Columella distinct. Budding near 
edge of corallum from peritheca. 

The coralla are denser than those of other species of Galaxea, with corallites wider, 
walls and septa thicker. 

No polyps. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 9 


66 PERCY SLADEN TRUST EXPEDITION 


Locality. Rotuma (2). Milne Edwards and Haime’s type specimens of this species 
are missing from the Paris Museum. 


LEPTASTREA (Milne Edwards and Haime). 


1848. Leptastrea, Milne Edwards and Haime, Compt. rend. Acad. Sci., xxvii, p. 494. 
1848. Baryastrea, Milne Edwards and Haime, Compt. rend. l’Acad. Sci., xxvii, p. 495. 
1857. Leptastrea, Milne Edwards and Haime, Hist. Nat. Corall., i, p. 493. 

1857. Baryastrea, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 512. 

1879. Leptastreea (pars), Klunzinger, Korall. Roth. Meer., ii, p. 43. 

1884. Leptastrea, Duncan, Journ. Linn. Soc., London, Zool., xviii, p. 119. 

1884. Baryastrea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 119. 

1886. Leptastrea, Quelch, Reef Corals, Challenger Reports, Zool., vol. xvi, pt. xlvi, p. 108. 
1899. Orbicella (pars), Gardiner, Proc. Zool. Soc. London, p. 751. 

1901. Leptastrea, Studer, Zool. Jahrb., xl, p. 402. 

1904. Orbicella (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 774. 


Corallum. Incrusting, convex or round, sometimes raised into hillocks, dense and 
usually heavy. Corallites polygonal, oval or round, not projecting or at most up to 
1°75 mm., one side usually more than the other, arranged close together with little or no 
peritheca, as a rule separated on the surface by narrow furrows, which are always 
polygonal around corallites. Walls fused, dense, varying from *5 to 3 mm. in thickness. 
Calices oval, round or polygonal, sometimes laterally compressed and distorted, varying 
considerably in diameter in the same specimen from 2 to 5°5 mm. and in depth from almost 
flat up to 3°6 mm. Septa in four orders, the first three always complete, upper margins 
usually flat, primaries meeting columella; the fourth order complete or incomplete, some- 
times a few quinaries, last order exoccelic. Costze seen as low flat ridges when corallites 
project. Columella varying from very thin to a third or half the width ‘of calices. 
Calicular dissepiments horizontal or somewhat oblique, about *75 or 1 mm. apart. Giant 
corallites present. 

Polyps. Oval or circular, upper margins somewhat polygonal, varying in size. 
Edge-zones absent when corallites do not project, and little or no intervening ccenosarcal 
regions owing to close approximation of corallites. Mesenteries in three or four cycles, 
the first two of six couples each, the third when complete of twelve couples but often 
incomplete, quaternaries when present few in number; all with filaments. Tentacles 
corresponding in number and position with entocceles and exoceeles. Stomodzeum short, 
usually compressed laterally, with narrow directive grooves. Hntoccelic pleats similar 
in general shape to those of Galaxea but less broad and only a few sub-divided. Multipli- 
cation by budding. 

Remarks. This genus comes nearer Gialaxea than any other described genera, as 
may be seen from the following comparison of L. roissyana and G. musicalis. (1) The 
terminal batteries of entoccelic tentacles are feebly developed in both species, with neither 
thickened ectoderm nor closely-packed nematocysts, whereas those of the exoccelic 
tentacles are well defined with nearly the same shape and relative size. (2) The endoderm 
of exoceelic tentacles is so much distended as to fill their lumina in the regions of the 
terminal batteries. (3) The stomodzeum is much compressed laterally and its wall 
distorted by the ingrowth of septa. (4) Tertiary couples of mesenteries present. (5) The 


MATTHAI—RECENT COLONIAL ASTRAIDA 67 


entoccelic pleats have the same general shape and distribution in both species, broad but 
constricted at the base, hence appearing knobbed in transverse section, sub-divided usually 
in G. musicalis, exceptionally in L. roissyana. (6) The mesenterial endoderm is com- 
paratively thin. (7) The great abundance of algze. 

There are, however, some important differences between the two species, viz. : 
(1) The general appearance of the colonies is strikingly different in the two cases; in 
L. roissyana the corallites are close together and almost level with the general surface, in 
G. musicalis they are wider apart and project above the peritheca. (2) Peritheca dense 
in L. rowssyana, highly vesicular in G. musicals, the vesicles with thin walls. (3) ‘‘Ovoid 
bodies” are always met with in the ectoderm of G. musicalis, their presence being quite 
diagnostic of Galaxea ; they are absent from L. rowsyana. 

The “Challenger” examples of Leptastrea available for examination were all small 
spirit specimens. Owing to the presence of soft tissues on them, it was impossible to 
compare them with the continental types, but there is little doubt that most of Quelch’s 
determinations were wrong. 


Distribution. Red Sea, Indian and Pacific Oceans. 


1. Leprasrrea rorssy4n4, Milne Kdwards and Haime. (PI. 8, figs. 1—3; 17, fie. 4; 
18, fig. 1; 19, figs. 1 and 2; 87, fig. 4.) 

1850. Leptastrea roissyana, Milne Edwards and Haime, Ann. Sci. Nat., 3° sér., x, pl. 9, figs. 6 and 6a, and 
xii, p. 120. 

1857. Leptastrea roissyana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 494. 

1879. Leptastrea transversa, Klunzinger, Korall. Roth. Meer., iii, p. 46, pl. 6, fig. 2. 

1904, Orbicella ehrenbergana (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, p. 776. 

Corallum. Incrusting or massive and convex, rarely rising into humps. Inter- 
corallite furrows usually distinct, the exsert ends of septa ending at their margins, rarely 
absent, leaving a sharp ridge over which septa become continuous. Corallites not pro- 
jecting. Walls about °35 mm. thick, sometimes up to 1 mm. towards edge of corallum. 
Calices oval or polygonal, widest at their margins, with diameters about 4 and 3°6 mm., 
depth to columella 2-6—3°6 mm. 

Septa swollen in thecze, thin in calices, with sides smooth, edges entire, in not more 
than four orders, the first three always complete; primaries slightly broader than 
secondaries, both meeting columella, with somewhat oblique margins and exsert to ‘5 mm. 
The fourth order often incomplete, usually of about 10 very narrow septa, hence some of 
the tertiaries exoccelic; tertiaries often curve towards and meet sides of secondaries near 
to columella. Columella much compressed laterally, mdistinct, primary and secondary 
septa almost meeting in the centre and forming with the two directive septa a transverse 
partition across calyx. 

Young corallites formed by budding are seen intercalated among the larger ones. 

Polyps. About 4 mm. in height, with margins somewhat polygonal. Mesenteries in 
three cycles ; primaries of six couples all meeting the stomodzeum; secondaries of an equal 
number from half to two-thirds the width of the former; tertiaries from 2—8 couples ; all 
with filaments in whose coils nematocysts II are common and III rare. Convolutions 
of mesenteries massed together in inter-mesenteric chambers to some distance below 


9—2 


68 PERCY SLADEN TRUST EXPEDITION 


stomodzeum, not abundant at the bases of polyps. The terminal battery of each entoccelic 
tentacle swollen but without much thickening of ectoderm, and nematocysts I and IIb 
mostly restricted to its tip; the exoccelic terminal battery well formed with thickened 
ectoderm and nematocysts closely packed; 3—5 very small sub-terminal batteries ; 
mucous vacuoles* found close together in all batteries, between which lie rod and spindle- 
shaped nuclei homogeneously stained dark ; granular vacuoles also present. Stomodeeum 
much compressed laterally ; its ridges narrow with no nematocysts. 

Entoccelic pleats present in outer half of primary mesenteries in the stomodzeal region 
of polyps, best developed in outer one-third, broad and constricted at their bases, hence 
appearing knobbed in transverse section, occasionally sub-divided, below stomodzeum 
extending over two-thirds width of mesentery. Mesenterial mesoglea slightly thickened 
near stomodeal attachment, elsewhere thin. 

Ectoderm of oral-dise with columnar facies resembling that of C. chalcidicum. Endo- 
derm of body-wall thin above enterostome, with few algee and many transparent vacuoles, 
below becoming reticulated and consequently distended; tentacular endoderm hardly 
visible owing to aggregation of algee, sometimes blocking lumen of exoccelic tentacle in the 
region of its terminal battery ; endoderm of primary mesenteries somewhat thickened near 
their stomodeeal attachments, behind filaments appearing pad-like in transverse section, 
elsewhere thin ; aloze massed together in exoccelic side. Stomodzal endoderm extremely 
thin. 

Polyps examined, ten, 4 from one and 3 from a second specimen from Red Sea, 
1 from a specimen from Salomon{ and 2 from a specimen from Ceylon. 

Remarks. Corallum. Milne Edwards and Haime’s type of ZL. roissyana (locality 
unknown) is a large specimen measuring 25x 20x17 cm. It is peculiar in that the 
corallum has incrusted over projecting calcareous tubes of some polycheete worms ; conse- 
quently the corallites have undergone some modification in shape—most of them project 
slightly, some almost circular, a few drawn out ; on the general substratum, the corallites 
have the usual polygonal shape. As in my examples the septa are never in more than 
four orders, but in many corallites more than twelve septa—up to thirteen—meet the 
columella. 

In the Berlin Museum are two specimens, referred by Klunzinger to L. transversa, 
one large and flat, measuring 24x12 cm., the other small, both entirely resembling my 
type specimens of the present species. Identical with these is another small specimen 
referred by the same author to LZ. ummersa. 

Localities. Red Sea (16 specimens). Chagos, Salomon (7). Maldives: Hulule (3) ; 
Goidu (1); Addu (1). Minikoi (3). Ceylon (small specimens). Not known from the 
Pacific. 


2. LmprastRreaA EHRENBERGANA, Milne Edwards and Haime. (PI. 17, figs. 5—7; 18, 
figs. 2 and 7; 19, figs. 3 and 4; 34, fig. 8.) 
1850. Leptastrea ehrenbergana, Milne Edwards and Haime, Ann. Sci. Nat., 3° sér., x11, p. 120. 


* Mucous vacuoles stain orange with hematoxylin and eosin as in Galaxea. 


} The polyps of this specimen were in a bad state of preservation ; in the one sectioned a few disconnected 
ova were found. 


MATTHAI—RECENT COLONIAL ASTRAIDA 69 


1857. Leptastrea ehrenbergana, Milne Kdwards and Haime, Hist. Nat. Corall., ii, p. 494, pl. D7, fig. 4. 
1879. Leptastrea ehrenbergana, Klunzinger, Korall. Roth. Meer., ii, p. 46, pl. 6, fig. 3. 

1899. Orbicella klunzingeri, Gardiner, Proc. Zool. Soc. London, p. 755. 

1904. Orbicella ehrenbergana (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 776. 

Corallum. Incrusting or rounded and completely covered with corallites, sometimes 
rising into humps. Inter-corallite furrows shallower and less distinct than in L. roissyana, 
sometimes absent, septa then meeting over walls in ridge-form. Corallites not projecting, 
the larger ones usually laterally compressed and distorted, widest at their margins, 
showing considerable variation in size in different parts of the same specimen, often much 
larger in certain regions (usually eminences) than in others (valleys). Walls much thicker 
than in the last species, up to 1°5 mm., usually about ‘75 mm. Calices oval or polygonal, 
from 2—5°6 x 5 mm. in diameter, average 4°5 x 4 mm., depth up to 3°6 mm. 

Septa appearing crowded in calices, not swollen in thecse, somewhat thicker in calices 
than in L. rovssyana, usually less exsert, sides granular, edges denticulate, in four or 
sometimes five orders. Primaries broader than secondaries above columella, both with 
almost vertical margins; these and often a varying number of tertiaries (up to 10), usually 
4 or 5, meeting columella, remaining tertiaries as a rule curving towards and their edges 
fusing with sides of secondaries near to columella; up to 6 quinaries present, all the 
_quaternaries being then entoccelic. Columella dense up to 2x 1°6 mm. thick, often with 

conspicuous upright rods. 

A thicker and rougher species than L. rowssyana. Occasionally cases of fusion of two 
corallites, giving an appearance of fission, 

No polyps. 

Remarks. Milne Edwards and Haime’s original of L. ehrenbergana from Red Sea, 
in the Paris Museum, is a massive specimen, measuring 15 x 5x 12x11 cm. with typical 
corallites, being identical with specimen no. 9 on p. 71. Resembling these in the Berlin 
Museum are Klunzinger’s small figured example (7 x 6 cm.) of the same species and a 
large specimen from Dar-es-Salaam, referred by Ortmann to LZ. immersa, Klunz. 

Localities. Red Sea (5). Maldives: Hulule (1); Goidu (1) ; Turadu (1). Minikoi 
(3 and broken pieces). Chagos: Salomon (4); Egmont (1). Rotuma (1). Funafuti (1). 
Also from Dar-es-Salaam (Ortmann). 


3. Leprastrea soca (Milne Edwards and Haime). (PI. 17, figs 8 and 9; 18, 
figs 3—6 and 8; 19, figs 5 and 6.) 
1850. Cyphastrea? botte (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 115. 
1850. Baryastrea solida, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 144. 
1857. Cyphastrea bottai (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 486, pl. D7, fig. 1 (not 
“orandeur naturelle” but an enlarged view). 
1857. Baryastreea solide, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 512, pl. D8, fig. 2a and 6. 
11860. Chypastrea oblita, Duchassaing and Michellotti, Mémoire Corall. Antill. Turin, p. 77. 
1879. Leptastreea bottai, Klunzinger, Korall. Roth. Meer., iii., p. 44, pl. 5, fig. 9, pl. 10, fig. 13 @ and 6. 
1879. Leptastrea inequalis, Klunzinger, Korall. Roth. Meer., iii, p. 45, pl. 5, fig. 6. 
1904. Orbicella bottat, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 777, pl. 63, fig. 36. 


Corallum. Variations in growth-form as in B. ehrenbergana. Inter-corallite 
furrows distinct, rarely absent, in which case wall flat with septa continuous over same. 
Corallites usually round, sometimes polygonal or oval, projecting to about 1 mm., maximum 


70 PERCY SLADEN TRUST EXPEDITION 


2mm., usually one side higher than the other. Wall-75 mm. thick. Calices on the whole 
smaller than in the two previous species but of uniform width from margins to bases, 
average diameter 3 mm., depth 2 mm. 

Septa usually thicker in calices than in B. ehrenbergana, exsert to ‘75 or 1mm. 
Sides with short spines, in three complete orders, quaternaries absent or up to 6. 
Primaries ustially thicker than secondaries, both—often also 1 to 3 tertiaries—meeting 
columella ; septal edges vertical, each usually with a paliform lobe near the columella either 
upright or directed obliquely inwards with sometimes an additional blunt tooth or two 
above it. Costze as low flat ridges. Columella thicker than in B. ehrenbergana, from 
a third to half the width of calices, with sometimes short rods. 

Giant corallites 4—4°6 mm. in, diameter with from 16—22 septa meeting columella, 
frequently found among the normal ones (Pl. 18, figs. 5 and 6). Occasionally two 
corallites fuse, presenting an appearance of fission (Pl. 18, fig. 4). 

No polyps. 

Remarks. Corallum. Milne Edwards and Haime’s smaller figured example of 
Cyphastrea bottai is massive, measuring 11°5x10x8cem. (Pl. 18, fig. 8). Its calices 
range up to 3mm. in diameter (average 2—2'3mm.); the six primary septa in each 
corallite are thicker than the secondaries, the former and a varying number of the latter 
meeting the columella from which short rods project; on the whole it resembles 
specimen no. 15 on p. 71. But, as the second larger example referred by Milne Edwards 
and Haime to the same species is a true Cyphastrea (most probably C. serailia), the 
specific name botta: cannot be given to the present species. Unfortunately Milne 
Edwards and Haime’s type of Baryastrea solida is missing from the Paris Museum, 
but since the two figures agree completely with my type specimens I here applied 
the name solida to the present species. 

A specimen (13 x 8 x 9 cm.) from St Thomas in the Paris Museum named Chypastrea 
oblita by Duchassaing comes near the present species. Its chief characters are as follows: 
peritheca vesicular, with spines; calices circular up to 3mm. in diameter, average 
2°5mm.; septa in three orders of 6, 6 and 12, primaries and secondaries in every 
corallite meeting columella, each dropping vertically for the greater part of its height, 
then passing obliquely or horizontally inwards to meet columella, the latter region 
of septum with 3—5 conspicuous blunt teeth; primaries somewhat thicker than second- 
aries ; columella with short rods. 

Klunzinger’s figured example (12 x 12 x 7cm.) of ZL. bottaa has no paliform lobes ; 
in each corallite both primary and secondary septa meet the columella with which 
twelve tertiaries alternate; a few giant-corallites are present. Resembling this are 
his figured type (8 x 7 cm.) of ZL. inequalis and another small specimen from Koseir in 
the Paris Museum, to which Klunzinger has given the same name. 

Localities. Red Sea (6). Maldives: Hulule (3); Goidu (2). Minikoi (8). Chagos, 
Salomon (12). ? Also from St Thomas (Duchassaing and Michellotti). 


4. Leprasrrna? mmerss, Klunzinger. 


1879. Leptastrea immersa, Klunzinger, Korall. Roth. Meer., iii, p. 47, pl. 6, fig. 1. 
1904. Orbicella immersa, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 776. 


MATTHAI—RECENT COLONIAL ASTRAIDA 71 


LIST OF SPECIMENS SHOWING THE EXTENT OF VARIATION IN 
THE GENUS LEPTASTREA. 


No. of 
Specimen 


Locality 


Species 


Remarks 


15 


Dongonab, Red Sea 


” 


Salomon 


Rotuma 


Funafuti 


Dongonab, Red Sea 


Hulule, Maldives 


2) 


Salomon 


Hulule, Maldives 


> 


Dongonab, Red Sea 


L. roissyana 


” 


L. ehrenbergana 


L. solida 


2) 


LL. roissyana 


Corallum (11 x 11x 9 cm.) incrusting. Most corallites 
simulate B. ehrenbergana in possessing distinct columella 
with short rods and rough septa. PI. 17, fig. 4. 

Corallum (16x 11 x8 cm.) approaches LZ. ehrenbergana— 
columella half to two-thirds width of calyx with short 
rods, septa somewhat thicker and rough. PI. 19, fig. 2. 

Corallum (13 x 13 x 9 cm.) has varied towards L. solida— 
some corallites round, short paliform lobes from some 
septa meeting columella. Giant corallites with about 
19 septa meeting columella. 

Corallum (10 x 9 x 7 cm.) with a few giant corallites, 19— 
22 septa meeting columella. 

A small mass (9°5x 7x5 cm.) with humps completely 
covered with corallites, perhaps dropped off from a 
large specimen. Oorallites smaller and shallower, and 
quinaries have not appeared. Pl. 19, fig. 4. 

Corallum (15 x 7 x 7 cm.) incrusting with two humps on 
one of which corallites large with diameters 5°6 x 5 mm., 
sometimes much compressed laterally, 6 x 3 mm., about 
15 septa meeting columella, up to 6 quinaries. In the 
valley corallites small about 2 mm. in diameter with only 
12 septa meeting columella, quaternaries about 10; among 
these a giant-corallite 4 x 3°6 mm. with 20 septa meet- 
ing columella. PI. 19, fig. 3. 

Corallum (15°5 x 105 x5 em.) flat with even surface— 
shows same extent of variation as in no. 6: in one half 
corallites large with about 6 quinaries, in the other 
small with up to 10 quaternaries. 

Corallum incrusting on a large mass 27 x 21 x 13 cm. with 
humps, showing considerable variation. Corallites round, 
oval, polygonal, much compressed laterally or distorted. 
Walls up to 1-5 mm. in thickness, inter-corallite furrows 
present or absent. Calices from 2mm. wide in depressions 
to 8 x 3:5 mm. on eminences, depth up to 3-5 mm. Septa 
3—5 orders, quinaries up to 13, 12—17 meeting colu- 
mella, the latter well developed with rods or indistinct. 
In one region near edge corallites shallow, small, wide 
apart and without furrows. PI. 18, figs. 2 and 7. 

Corallum (14x 11 x8 em.) incrusting with humps—corallites 
usually round or oval slightly projecting to 1 mm., in 
valleys crowded together and smaller. PI. 17, fig. 6; Pl. 
34, fig. 8. 

Specimen small (9 x 6 x 3em.) approaching L. roissyana— 
corallites polygonal, large, 4:6 x36 mm. PI. 17, fig. 9. 

Corallum 11 x 8 x 5 cm.; septa meeting columella equally 
thin ; pali and columella not well developed. A case of 
fusion of two corallites—one giant-corallite with 21 
septa meeting columella. PI. 18, fig. 3. 

Corallum 11 x 4:5 x 4:5 em.—corallites approaching JZ. 
ehrenbergana polygonal, projecting less ‘5 mm., pali not 
distinct but columellar rods. Pl. 18, fig. 6. 

Corallum 8:5 x6x2cm. Pl. 17, fig. 8. 

Corallum incrusting, corallites level or projecting about 
‘5 mm., primary septa much thicker than secondaries, 
usually only the former meeting the columella. Short 
rods on columella. A giant-corallite 4°5 x 3 mm. with 
44 septa, of which 15 meeting columella. Pl. 18, fig. 5. 

Corallum 12 x10x9cm. Corallites in one edge almost 
circular and 1 mm. apart, in another edge circular and 
projecting to 1 mm. as in ZL. solida. Similar to Klun- 
zinger’s figured examples of L. bottai and L. inaequalis. 
Pl. 19, fig. 5. 


72 PERCY SLADEN TRUST EXPEDITION 


Klunzinger’s figured type of this species measures 16 x 12 x 8cem. Its distinguishing 
characters are as follows: (1) septa in four orders but the fourth incomplete, the maxi- 
mum numbers of quaternaries counted being ten; (2) six to ten septa meeting the 
columella; (3) the remaining septa very narrow; (4) columella quite rudimentary. 
In general appearance the specimen resembles L. roissyana, but its real position in 
the genus cannot be settled till its polyps are examined. 

The single specimen from Hulule which Gardiner has referred to L. immersa is 
identical with Klunzinger’s example. 


GENUS DIPLOASTREA, NDU.GEN. 


1816. Astrea (pars), Lamarck, Hist. Anim. sans vert., 11, p. 257. 

1850. Asérea (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 97. 
1857. Heliastrea (pars), Milne Edwards and Haime, Hist. Nat. Corall., p. 456. 

1904. Orbicella (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 774. 


This genus has been created for Orbicella minikovensis, Gardiner. Its characters 
are as follows: 

Corallum. Incrusting or massive. Corallites circular not projecting. Walls fused 
and perforate, hence peritheca almost absent. Calices shallow. Septa in not less than 
two orders, the first two entoccelic, each consisting of twelve septa, exsert, much thickened 
towards their outer ends. Columella formed of twisted trabecule from septal margins. 
Calicular dissepiments oblique. 

Polyps. Close together with narrow edge-zones, no ccenosare. Mesenteries in not 
less than two cycles, each of twelve couples, usually directly continuous from polyp to 
polyp, primaries meeting stomodzeum ; all with filaments. Mesoglea thick. Tentacles 
corresponding in number and position with entocceles and exocceles. Stomodzeum short, 
laterally compressed with two directive grooves. Multiplication by budding. 

Remarks. Owing to the presence of two directive couples of mesenteries O. mint- 
koiensis has to be separated from the old genus Orbicella (Group IL) and placed in Group I. 
But it differs from all other described genera of this sub-family in possessing, in its adult 
polyps, double the usual number of couples of primary and secondary mesenteries, viz. 
12 in each cycle, and a highly thickened mesenterial mesoglea, the mesenteries in 
consequence presenting an unique appearance in transverse section. The perforate 
nature of the corallite-walls indicates a probable variation of Diploastrea to the Perforata, 
but the perforations are larger and fewer than in the latter group and are irregular. 

Distribution. Red Sea, Indian and Pacific Oceans. 


1. Dretoastrea HELIOPoRA (Lamarck) (Pl. 20, figs. 7 and 8 ; 34, fig. 9). 


1816. Astrea heliopora, Lamarck, Hist. Anim. sans vert., ii, p. 265; 2° édit., p. 415. 
1824. Astrea heliopora, Lamouroux, Encycl., Zooph., p. 128. 

1848. Astrea heliopora, Milne Edwards, Grande édit. Régne Anim. Cuvier, Zooph., pl. 84 ter, figs. 1, la and 0. 
1850. Astrea heliopora, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 99. 

1857. Heliastrea heliopora, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 459. 

1904. Orbicella minikotensis, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 774, pl. 63, fig. 35. 

1907. Orbicella minikoiensis, Vaughan, Proc. U.S. Nat. Mus., xxxii, p. 252. 


~ 


MATTHAI—RECENT COLONIAL ASTRAIDA 73 


Corallum. Incrusting, tending to be massive, somewhat convex, not rising into 
hillocks. Corallites more or less circular in outline, not projecting. Walls up to 3mm. 
thick, somewhat perforate, neighbouring ones fused with a shallow groove in the middle. 
Calices from 8—10 mm. in diameter, shallow, varying from flat to 2 mm. in depth. 

Septa in two orders, both entoccelic, each consisting of 12—sometimes a few exoccelic 
tertiaries up to 7 (in one corallite 14)—highly exsert about 2mm. ; sides tuberculated, often 
perforated ; edges with 4—8 bluntly pointed teeth, the longest representing the most 
exsert part of septum; outer half or two-thirds of septa much thicker (up to 1 mm.) 
swollen in thece, thinning out within calices ; primaries usually thicker and somewhat 
more exsert than secondaries, both meeting columella; tertiaries when present thinner, 
of varying width, 1—3 sometimes reaching columella, edges usually curving towards 
and fusing with sides of adjacent secondaries. Hxsert ends of septa arched, with bluntly 
pointed teeth. Costze, also with blunt teeth, either continuous from corallite to corallite 
or alternating, when continuous with a distinct notch over inter-corallite furrows; in 
overhanging edges of corallum, thinner than exsert ends of septa, sometimes very thin ; 
exoccelic costee present alternating with the main ones. Columella circular in outline, 

thick, from a third to half the diameter of calices, from upper surface appearing fasciculated, 
- formed of twisted trabeculee from septal margins, in some corallites terminating in upright 
-rods. Calicular dissepiments sloping into columella at an angle of 35°, about 1 mm. apart. 

The perforate walls, shallow calices, stout twisted columella and thick exsert toothed 
septa give the corallum an unique appearance. Smaller corallites formed by budding are 
intercalated between the larger ones. Corallites at the margin of the corallum have 
thinner septa which pass over its edge as coste and loosely trabecular columelle. 

In the sections of the corallum the septa appear to be continuous from corallite 
to corallite, but this continuity is really secondary, as in the young corallites the costal 
surfaces are demarcated off. The perforate wall seems to be formed by the perithecal 
deposit uniting here and there the peripheral thickened septal parts. The peritheca 
is without any dark centres and hence appears distinct from the septal ‘‘ trabeculz.” 

Polyps. Large, almost circular in outline, about 8 mm. in height, closely aggregated 
with narrow edge-zones and no ccenosare. Oral-disc much narrowed, about 2mm. wide 
in the retracted condition. Mesenteries constituting two complete cycles, each of 
12 couples. Outer skeletal attachments of some of the mesenteries of neighbouring 
polyps tending to unite, hence appearing to be continuous from polyp to polyp. The 
mesenteries with filaments containing nematocysts II and III in their coils, primaries 
meeting stomodzeum*. Tentacles as in genus; well developed terminal batteries present 
with closely packed nematocysts I and II } interspersed; also sub-terminal batteries 
(number doubtful). Stomod:eum laterally compressed. 

Remarks. A. Polyps. The histology of the polyps could only be imperfectly 
studied owing to partial maceration of their tissues. The ectoderm of oral-disc is thick 
with narrow diverging tracts containing nuclei in the centre ; granular and mucous vacuoles 


* It is possible that the presence of twelve couples of primary mesenteries in the adult polyps may be due 
to six couples of secondaries having grown and met the stomodeum; if so the twelve secondary couples in that 
Stage would be tertiaries, but this can be settled only by examining young polyps which were not available. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 10 


74 PERCY SLADEN TRUST EXPEDITION 


present but nematocysts rare. The calicoblast is thickened near skeletal attachments of 
mesenteries, in transverse section resembling ectoderm of oral-disc. In some of the 
terminal batteries numerous long narrow bodies, homogeneously stained pink in eosin, 
are seen (as in Mavia doreyensis, Kd. and H.) which may probably be immature stages in the 
development of nematocysts IIb. The mesoglea is thick. The entoccelic pleats extend 
over the outer halves of primary mesenteries in the stomodeal region of polyps, being 
narrow, thick and unconstricted at their bases, below stomodzeum becoming broader and 
extending over the greater part of the width of mesenteries, some of them being sub- 
divided. Mesenterial mesoglea is stouter in pleatal region, thickening again towards 
stomodzeum, the connecting sheet being narrow and thin. Groups of small bodies stained 
dark in iron hematoxylin are present in the outer mesenterial mesoglea, probably spores 
of alge. The endoderm of oral-dise is not thicker than the ectoderm over it and contains 
numerous algze ; the mesenterial endoderm is thickened on either side of the inner stouter 
region of mesoglea. 

Polyps examined, three from a specimen from Minikoi. 

B. Corallum. In Lamarck’s collection in the Paris Museum are two small specimens 
(8x8x3'5 em. and 8°5x5°5x2°5 cm.) from “mers Australes” named Heliastrea heliopora 
by Milne Edwards and Haime which were doubtless Lamarck’s originals of Astrea 
heliopora (Pl. 20, fig. 8). These are similar to Gardiner’s examples of Orbicella mim- 
kovensis, but the calices are somewhat smaller with an average diameter of about 7 mm., 
and a cycle of thin coste are present regularly alternating with the main ones, those 
of neighbouring corallites usually meeting in notches, their corresponding septa being 
quite rudimentary or absent. 

Resembling Lamarck’s examples are two undescribed specimens in the Berlin 
Museum, which Dr Weltner tells me were collected by Dr Dahl in 1897 from Ralum (New 
Pommern). One of these is large measuring 30x21 cm., the other a hump, 12x11 em. 
in size; the secondary septa in their corallites are much thinner than the primaries, 
some of them not reaching the columella; an alternating cycle of thin coste are present 
and the columella is about one-third the width of the calices. 

Vaughan records a single specimen from French Somaliland, East Africa, which he 
says is identical with Orbicella minikoiensis, Gard. 

Localities. Minikoi (4). Also from Australia (Lamarck), French Somaliland 
(Vaughan), New Britain (Dahl). Not known from the Red Sea. 


ADDENDA. SOME TYPE SPECIMENS. 


The following species are not represented in my collections, but I have examined 
examples. From a study of their septal arrangement there is hardly any doubt that they 
possess both the bilateral and hexameral symmetries, but it is not possible to determine 
the genera to which they belong without examining their polyps. 

1. Stephanocenia intersepta (Esper); Ed. and H., Ann. Sci. Nat., 3° sér., x., Pl. 7, fig. 1, 

1a and 6, and Corall. 11., p. 265. 


In Milne Edwards and Haime’s three examples of this species the corallites are small 


MATTHAI—RECENT COLONIAL ASTRAIDA 75 


and polygonal, the calices being 2—2°5 mm. in diameter. In every corallite about 12 septa 
meet the columella, each usually with a conspicuous paliform lobe; six (secondary septa) 
of these septa are somewhat thinner than and alternate with the others (primary 
septa); an alternating cycle of twelve narrower septa (tertiaries) is present. The 
columellze are unique in beg quite solid rods—it would be interesting to see if the 
columella arises as such from the basal plate—which are either circular in outline or 
somewhat laterally compressed and rough at their upper ends. New corallites appear to 
arise by true budding. 

There are two more examples of this species in the Paris Museum, one of which is 
from St Thomas (Duchassaing). The types of Milne Edwards and Haime’s two remaining 
recent species of Stephanocenia, viz. S. michelini and S. dendroidea, are missing. 

Esper’s figures (p. 99, Pl. 79) of Madrepora intersepta are not adequate for satisfactory 
comparison with Milne Edwards and Haime’s types. 

When polyps are examined Stephanocenia may, in all probability, prove to be a good 
genus. Its nearest ally appears to be Leptastrea. 


2. Helastrea cavernosa (Esper); Ed. and H., Corall. i, p. 463. 
Heliastrea gigas, Ed. and H., Corall. i1., p. 458. 


In the Paris Museum are four large specimens and two small ones from “ mers 
d’Amerique,” referred by Milne Edwards and Haime to Heliastrea cavernosa. In these 
the corallites are circular, projecting more and further apart than in Heliastrea conferata, 
Kd. and H. (hence the costze are better seen), their walls increasing in thickness from the 
corallite-margins towards the peritheca. The calices are 7 or 8 mm. in diameter ; usually 
24 septa meet the columella in every corallite; alternating with these is a cycle of 
narrower septa. The columelle are well developed, being 4—4 the width of the calices 
and formed of septal trabeculz. The coste are alternately large and small, corresponding 
to the two septal cycles, those of the neighbouring corallites usually meeting in notches. 
The peritheca is highly vesicular and tends to break down; thus the corallum approaches 
the conditions in Diploastrea heliopora; when the peritheca is entirely removed, the 
corallites appear as long cylinders. 

There are three more specimens with the same name, but they are quite small and 
much rubbed. 

There is no doubt that Milne Edwards and Haime’s examples are identical with 
Esper’s figure of Madrepora cavernosa (p. 18, Pl. 37). They have also a certain 
resemblance to one of my specimens (no. 13) of Hchinopora gemmacea (Lam.). 

The only example of Heliastrea gigas in the Paris Museum has only two sectioned 
corallites, resembling those of Heliastrwa, cavernosa. 

3. Eaplanaria argus, Ehrb., Corall. ii, p. 83 [non Astrea argus, Lam., which is 
Heliastrea cavernosa (Esper) |. 
Heliastrea conferata, Ed. and H., Corall. ii., p. 460. 
Hlehiastrea lamarckana, Kd. and H., Corall. ii., Eons 


Of Milne Edwards and Haime’s examples of Heliastrea conferata (locality unknown) 
10—2 


76 PERCY SLADEN TRUST EXPEDITION 


one is small and much worn, while the other is larger (14x 10x7 cm.) and in better 
condition. In this the corallites are slightly projecting and are separated from one 
another by polygonal furrows; about 24 septa (edges toothed but no paliform lobes) 
meet the columella in the corallites, which are all equally thin in the calices, but 
somewhat thickened in the walls; alternating with these is a cycle of narrower septa. 
The columelle are about one-third the width of the calices and formed of twisted septal 
trabeculze rising into short upright rods. The costze of all the septa are similar to one 
another, those of the neighbouring corallites usually meeting in the inter-corallite 
furrows. 

The two examples of Heliastrea lamarckana in the Paris Museum are small and of 
little use. 

Ehrenberg’s two examples of Haplanaria argus ave small and defaced; they may 
probably constitute a single species along with Milne Edwards and Haime’s types 
mentioned above. 

This species differs from Heliastr@a cavernosa in the corallites being smaller, closer 
together, and somewhat less projecting. 

In the Paris Museum there are three more specimens named Heliastrea conferata, 
which appear to belong to a different species: a large one from Guadeloupe with smaller 
corallites than in Heliastrea conferata and fewer septa meeting the columelle, and two 
small ones from St Thomas (Duchassaing) in which the columelle are much thicker. 


4. Madrepora annularis, Ell. and Sol., Zooph., p. 169, Pl. 53, figs. 1 and 2. 


Astrea annularis, Lam., Hist. Anim., p. 259 (non var. 2 which is Favia acropora). 

Explanaria annularis, Ehrb., Corall. p. 84. 

? Orbicella annularis, Dana, Zooph., p. 214, Pl. 10, fig. 6. 

Hehastrea annularis, Ed. and H., Corall. ii., p. 473. 

Madrepora stellulata, Ell. and Sol., Zooph., p. 165, Pl. 53, figs. 3 and 4.. 

¢ Orbicella stellulata, Dana, Zooph., p. 215, Pl. 10, fig. 7. 

Heliastrea stellulata, Ed. and H., Corall. ii, p. 473. 

2 Plesiastrea urviller, Ed. and H., Ann. Sci. Nat., 3° sér., x., Pl. 9, fig. 2, and Corall. i1., 
p- 490. 


In Lamarck’s type of Astrea annularis (later Milne Edwards and Haime’s type of 
Heliastrea annularis) the corallites are circular and slightly projecting, the calices being 
2—2°5 mm. in diameter and shallow. In every corallite about 12 precisely similar 
septa meet the columella; alternating with these is a cycle of 12 narrower septa. 
A pali-crown is absent. The columella is spongy. The costee are thick with transversely 
extending granulations. 

In addition there are two much larger specimens in Duchassaing’s collection (1870), 
one from Antilles and the other from St Thomas, and a third from Antilles (collected by 
Schramm, 1869). The remaining three examples in the Paris Museum from “mers 
d’'Indes,” “mers d’Amerique,” and “ Australie” respectively are small and rubbed. 

Ellis and Solander’s type of Madrepora annularis in the Glasgow Museum is much 
smoothed down, but is no doubt identical with the Paris specimens mentioned above. 


MATTHAI—RECENT COLONIAL ASTRAIDA 77 


In Milne Edwards and Haime’s type (5 5x5 em.) of Heliastrea stellulata from 
West Indies, the corallites are slightly projecting and are separated by polygonal furrows, 
the calices oval or circular and about 2 mm. in diameter ; the septa are similar in nature 
and number to those of Helvastrea annularis; the corallum also shows clear signs of 


budding. ° 
Milne Edwards and Haime’s examples (5 small ones) of Plesiastrea urviller from 
Australia come near the above specimens, but it is doubtful if they belong to the same 


species. 
According to Duerden (82, p. 563) two directive couples of mesenteries are present in 


the polyps of Orbicella annularis. 


5. %¢Madrepora plecades, Ell. and Sol., Zooph., p. 169, Pl. 53, figs. 7 and 8. 


I have examined a somewhat defaced specimen in the Glasgow Museum, which, 
though not identical with Ellis and Solander’s fig. 7, closely resembles it. Its corallites 
_ are small and circular in outline, the calices being 2—2°5 mm. in diameter. Twelve septa 
meet the columella in each corallite ; alternating with these is a cycle of narrower septa 
which curve towards and fuse with the sides of the former in pairs. 

Without polyps it is not possible to determine the genus to which this specimen 
‘belongs. It appears to be more related to Cyphastrea than to any other existing 
genus. 


GROUP IL. 


GENUS FAVIA (OKEN). 


1801. Astrea, sec. 1, Lamarck, Syst. Anim. sans vert., p. 371. 

1815. Favia, Oken, Lehrb. Naturg., i, p. 67. 

1816. Astrea (pars), Lamarck, Syst. Anim. sans vert., ii, p. 257. 

1830. ubastrea (pars), Blainville, Dic. Sci. Nat., 1x, p. 334. 

1830. Dipsastrea (pars), Blainville, Dic. Sci. Nat., 1x, p. 338. 

1834, Favia (pars), Ehrenberg, Corall. roth. Meer., p. 93. 

1834. Astrea (pars), Ehrenberg, Corall. roth. Meer., p. 95. 

1846. Orbicella, subgenus I of Astrea (pars), Dana, Expl. exp. Zooph., p. 206. 

1846. Fissicella, subgenus III of Astrea (pars), Dana, Expl. exp. Zooph., p. 220. 

1848. Astrea (pars), Milne Edwards and Haime, Compt. rend. l’Acad. Sci., xxvii, p. 494. 
1848. Plesiastrea (pars), Milne Edwards and Haime, Compt. rend. l’Acad. Sci., xxvii, p. 494. 
1848. Phymastrea, Milne Edwards and Haime, Compt. rend. |’ Acad. Sci., xxvii, p. 494. 
1848. Parastrea, Milne Edwards and Haime, Compt. rend. l’Acad. Sci., xxvii, p. 495. 
1848. Qulastrea, Milne Edwards and Haine, Compt. rend. l’Acad. Sci., xxvii, p. 495. 
1848. Acanthastrea, Milne Edwards and Haime, Compt. rend. ’Acad. Sci., xxvii, p. 495. 
1848. Prionastrea, Milne Edwards and Haine, Compt. rend. |’ Acad. Sci., xxvii, p. 495. 
1857. Favia, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 426. 

1857. Goniastrea (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 444. 

1857. Heliastreea (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 456. 

1857. Ulastreea, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 488. 

1857. Plesiastrea (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 489. 

1857. Phymastrea, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 499. 

1857. Acanthastrea. Milne Edwards and Haime, Hist. Nat. Corall., li,-p. 501. 

1857. Prionastrea (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 513. 


78 PERCY SLADEN TRUST EXPEDITION 


1879. Favia, Klunzinger, Korall. Roth. Meer., iii, p. 25. 

1879. Goniastrea (pars), Klunzinger, Korall. Roth. Meer., iii, p. 32. 

1879. Prionastrea, Klunzinger, Korall. Roth. Meer., iii, p. 36. 

1879. Acanthastrea, Klunzinger, Korall. Roth. Meer., iii, p. 42. 

1879. Orbicella (pars), Klunzinger, Korall. Roth. Meer., iii, p. 47. 

1884. Favia, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 100. 

1884. Heliastrea (including subgenus Ulastrea), Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 104.' 
1884. Phymastrea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 106. 

1884. Plesiastrea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 107. 

1884. Acanthastrea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 119. 

1884. Prionastrea, Duncan, Journ. Linn. Soc. London, Zool., xviii, p. 123. 

1899. Astrea, Gardiner, Proc. Zool. Soc. London, p. 747. 

1899. Orbicella (pars), Gardiner, Proc. Zool. Soc. London, p. 751. 

1899. Prionastrea, Gardiner, Proc. Zool. Soc. London, p. 757. 

1900. avia, Vaughan, Monogr. U.S. Geol. Surv., xxxix, p. 154. 

1904. Pavia, Gardiner, Fauna Geogr. Maldives and Laceadives, p. 766. 

1904. Orbicella (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, p. 774. 

1904.  Stephanocenia, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 784. 

1904. <Acanthastrea, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 784. 

1904. Prionastrea (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, p. 785. 

1906. avia, Marenzeller, Exp. ‘‘ Pola,” Rot. Meer., Riffkorall., Zool. Ergeb. Wien, xxvi, p. 56. 
1906. Orbicella (pars), Marenzeller, Exp. “Pola,” Rot. Meer., Riffkorall., Zool. Ergeb. Wien, xxvi, p. 60. 


Corallum. Massive or incrusting, varying considerably in form of growth. Corallites 
polygonal, oval, circular or distorted, usually separated by perithecal regions of varying 
thickness, level or projecting with distinct rims, sometimes with walls completely fused as 
in Goniastrea. Calices varying much in size, from 2 to 20 mm. or more in diameter. 
Septa without any cyclical arrangement, varying considerably in number and nature in 
the different species. Costze present when walls project, those of neighbouring corallites 
usually meeting along the middle of the intervening perithecal regions. Columella formed 
of septal trabeculze, a rudimentary to well developed compact structure. 

Polyps. Varying in shape and size as the corallites. Mesenteries without directive 
couples, and without regular alternation, principal couples 6—12, varying only slightly 
in number in each species. Subsidiary couples from same number as principal couples 
to more than twice as many, varying much in number in the same species, also in relative 
height and width, this variation due to continued addition of new mesenteries in the 
course of polyp-growth, some of the later ones being so short as not to extend below the 
stomodzal regions of polyps. Stomodzum laterally compressed, no directive grooves, 
its ridges varying considerably in width and thickness. Every tentacle with a large 
terminal battery and from three to eight much smaller sub-terminal ones. Entoccelic 
pleats varying much in size, shape and distribution. All types of nematocysts excepting 
Ile and IIIb present. Stomodzeal endoderm very thin. Ova occurring in both principal 
and subsidiary mesenteries. 

Multiplication as a rule by fission, sometimes aided by budding. 

Remarks. I agree with Vaughan that the name Favia as originally employed 
by Oken should be retained for section I of Lamarck’s Astrea, typified by Madrepora 
rotulosa, Ell. and Sol. 


This genus has a wide range of variation—as is seen from a comparison of two 


MATTHAI—RECENT COLONIAL ASTRALID At 79 


extreme species like /. acropora and F. hirsuta. Indeed, it is possible that a comparative 
study of the polyps of the three known species with the largest corallites, viz. F. hirsuta, 
F. vasta, and F. favosa, may afford sufficient data for grouping them under a separate 
genus. Till then, I consider the present arrangement as the most satisfactory. 

A study of polyp-morphology has revealed how unnatural the genera Heliastraa 
and Orbicella are as constituted respectively by Milne Edwards and Haime and Gardiner. 
Of the species of Heliastraa, Ed. and H., not represented in my collection, H. conferata, 
H. cavernosa, H. lamarckana, and HH. stellulata, would, in all probability, prove to have 
bilateral and radial symmetries. H. radiata and H. quadrangularis are represented, 
in the Paris Museum, each by a small completely worn-out specimen useless for specific 
determination and H. gigas by an example consisting of only two sectioned corallites. 
Milne Edwards and Haime constituted a new genus and species,—Ulastrea crispata, 
for two small bits of coralla from the Indian Ocean, one incrusting on a Gasteropod 
shell and the other on a piece of stone; there is little doubt that these two examples 
belong to Favia but they are too small for determination. These authors also created the 
genus Phymastrea on the strength of a single skeletal character, viz. the presence of 
definite inter-corallite grooves ; a character shared by many species of Favia. 

Distribution. Indian and Pacific Oceans. Jurassic to Recent (Hastman after 
_ Zittel). 


1. Favra ravos (Forskal). (Pl. 9, fig. 2; 20, figs. 1—6; 21, figs. 1—8; 22, figs. 
1—5; 32, fig. 1; 35, figs. 1 and 2.) 


1775. Madrepora favus (pars), Forskal, Deser. Anim. in Itin. Orient., p. 132. 

1775. Madrepora cavernosa, Forska&l, Descr. Anim. in Itin. Orient., p. 132 (uon Madrepora cavernosa, Esper 
or Klunzinger). 

1786. Madrepora denticulata, Ellis and Solander, Nat. Hist. Zooph., p. 166, pl. 49, fig. 1 (non Favia denticu- 
lata, Gardiner). 

1816. Astrea denticulata, Lamarck, Hist. Anim. sans vert., ii, p. 263; 2° édit., p. 413. 

1830. Dipsastrea denticulata, Blainville, Dict. Sci. Nat., lx, p. 338; Manuel d’Actin. p. 373 

1833. Astrea ananas, Quoy and Gaimard, Voy. |’Astrol., Zooph., p. 207, pl. 16, figs. 6 and 7 (non Astrea 
ananas, Lamarck). : 

1834. Fava versipora, Ehrenberg, Corall. roth. Meer., p. 93 (non Astrea versipora, Lamarck). 

1834. avia denticulata, Ehrenberg, Corall. roth. Meer., p. 94. 

11834. Astrea deformis, Ehrenberg, Corall. roth. Meer., p. 96 (non Astrea deformis, Lamarck). 

1850. Prionastrea rowsseawi (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 131. 

11850. Prionastrea seychellensis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sév., xii, p. 132. 

1850. Parastrea denticulata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 167. 

1850. Parastrea afinis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 167 (non Fava affinis, 
Gardiner). 

1850. Parastrea deformata, Milne, Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 168. 

1850. Parastrea rousseaui, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 168. 

1850. Parastrea amicorum, Milne Edwards and Haime,:Ann. Sci. Nat., Zool., 3° sér., x, pl. 9, fig. 9, and xii, 
p. 171. 

1850. Parastrea savignyt, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 173. 

1857. Favia denticulata, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 428. 

11857. Favia aginis, Milne Edwards and Haime, Hist. Nat. Corall., i, 429. 

1857. Favia rousseawi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 429. 

1857. Favia amicorum, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 431. 

1857. Favia Jacquinoti, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 433 = Parastrea jacquinoti, Valen- 
ciennes, Mss., Catal. Mus. Paris. 


80 PERCY SLADEN TRUST EXPEDITION 


1857. Favia geoffroyi, Milne Edwards and Haime, Hist. Nat. Corall., u, p. 433 = Parastrea geoffroyi, Valen- 
ciennes, Mss., Catal. Mus. Paris. 

1857. Favia deformata, Milne Edwards and Haime, Hist. Nat. Corall., 11, p. 434. 

1857. Favia savignyi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 437. 

1857. Favia aspera, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 438 = Parastrea aspera, Valenciennes, 
Mss., Catal. Mus. Paris. 

21857. Prionastrea seychellensis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 517. 

1857. Prionastrea halicora (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 517. 

1879. Favia denticulata, Klunzinger, Korall. Roth. Meer., ii, p. Diy 

1879. Favia clouet, Klunzinger, Korall. Roth. Meer., iii, p. 29 (non Pavia clowei, Milne Kdwards and Haime). 

1879. Favia ehrenbergi, Klunzinger, Korall. Roth. Meer., iti, p. 29, pl. 3, fig. 5 (var. laticollis), ‘fig. 7 (gewohn- 
lichste Form), fig. 8 (var. sulcata), and pl. 9, figs. 1, and 1,. 

11879. Goniastrea seychellensis, Klunzinger, Korall. Roth. Meer., iii, p. 33, pl. 4, fig. 3. 

1899. Astrea puteolina, Gardiner, Proc. Zool. Soc. London, p. 749. 

21904. Havin versipora, Gardiner, Fauna Geogr. Maldive and Laccadive Archipel., 11, p. 766. 

1904. Prionastrea spinosa, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 785. 

1904. Prionastrea magnistellata, Gardiner, Fauna Geogr, Maldives and Laccadives, p. 788, pl. 64, figs. 40 
and 41. 

1904. Prionastrea suvadive, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 789. 

1906. Favia savignyt, Marenzeller, Exp. “ Pola,” Rot. Meer, Riffkorall., Zool. Ergeb. Wien, xxvi, p. 56, pl. 25, 
figs. 84—89, and pl. 24, fig. 79 (= Madrepora cavernosa, Forskal). 

1907. Favia savignyi, Vaughan, Proc. U.S. Nat. Mus. Washington, xxxii, p. 256. 


Corallum. Massive, often rounded, sometimes incrusting. Corallites polygonal, 
often distorted, sometimes oval, slightly projecting, with distinct rims, up to 5 mm. 
apart, the intervening perithecal regions being flat and vesicular. Calices varying 
much in size, up to 20x12mm., average 12x10mm., somewhat decreasing in width 
from calicular margins to bases, by depth up to 14mm, average 8 mm. 

Septa usually thin, sometimes thickened, vertical or slightly slopmg with toothed 
edges, rough sides, 60—70 in number in the largest corallites, 30-—45 in the average 
sized ones; up to 25 septa meeting columella (usually about 18), about 1 or 1:25 mm. 
exsert, with up to 14 teeth, usually becoming elongated from calicular margin to 
columella. A definite pali-crown absent. Some of the broader subsidiary septa usually 
curving towards and fusing with sides of principals. Exsert ends of septa arched or flat, 
finely toothed, meeting in notches or not actually meeting, giving an appearance of furrows 
over the perithecal regions, sometimes directly continuous from corallite to corallite. 
Columella varying in degree of development, from loosely inter-crossing septal teeth 
to twisted trabeculze. 

Multiplication by fission, more often sub-equal than unequal. 

The skeleton is highly variable in this species but two main types are to be recog- 
nized: (1) characterised by hght corallum, somewhat narrow perithecal regions, large 
and deep calices, numerous thin septa, the principals with their lower halves or one-thirds 
somewhat oblique, columella imperfectly formed (e.g. Prionastrea magnistellata, Gard., 
Favia jacqunot, Ed. and H., Favia geoffroyr, Ed. and H.); (2) a thicker and rougher 
type with heavy corallum, broader perithecal regions, smaller and shallower calices, 
fewer septa but broader at the calicular margins, thicker and rougher, the principals 
dipping vertically to the columella or their lower parts usually entire, broader, ending 


MATTHAI—RECENT COLONIAL ASTRAIDA 81 


in a simulated pali-crown. Columella formed of twisted trabecule (eg. Favia ehren- 
bergt var. laticollis, Klunz.). 

In some specimens the corallite-rims are distinct, oval or circular as in the following 
species, [’. doreyensis. 

The specimen referred by Gardiner to a new species P. suvadive is only an abnormal 
skeletal variation of the present species. The much greater thickness of the perithecal 
regions and the somewhat filled in appearance of the calices are doubtless due to 
slackened growth under unfavourable conditions ; as Gardiner suggests, ‘it was probably 
growing on the edge of a lagoon shoal and fell off into deeper water, the muddy bottom 
of which was slowly killing it.” It is remarkable that other Suvadiva specimens, cp. 
Cyphastrea maldivensis, should also show the same type of modification. 

Polyps. (1) Both entoccelic and exoccelic tentacles present. (2) Stomodeeal ridges 
as thick as or thicker than broad, with sides either parallel or constricted at their bases 
and inner surfaces somewhat flattened. (3) About ten principal couples of mesenteries. 
(4) In the stomodzeal region of polyp subsidiary couples of mesenteries about twice the 
number of principal couples. (5) In same region of polyp, entoccelic pleats narrow, 
unconstricted, undivided and not extending to middle of width of principal mesenteries ; 
inner one-sixth of mesoglea thickened with a few narrow exoccelic pleats, remaining 
part thin. (6) Mesenterial endoderm uniformly thin, except on each side of the inner 
thickened region of mesoglea where it is swollen. (7) Convolutions of mesenteries massed 
together im inter-mesenteric chambers below stomodzeum in the stomodzal region of polyp, 
a characteristic appearance in transverse section is due to convolutions of subsidiary 
mesenteries lying massed together as far inwards as the latter extend. 

Remarks. A. Polyps. Two of the typical polyps have each ten principal couples 
of mesenteries, two of which are incomplete; the remaining polyps have from nine to 
twelve couples. The number of subsidiary couples varies from twelve to eighteen. The 
exoccelic tentacles appear longer than the entoccelic ones. The terminal tentacular 
battery is large and contains in its peripheral region closely packed nematocysts I, each 
with from forty to fifty turns of the spiral ; among these are a few of type II b (mostly 
thrust out of the batteries in the sections); below the nematocysts saccular nuclei of 
varying size are crowded together. Sub-terminal batteries number from four to eight. 
A thin longitudinal layer of muscle-fibres is clearly visible on the outer surface of the 
tentacular mesoglea. 

The stomodzeum is oval in transverse section, with diameters measuring about 2 mm. 
and 1°5 mm.; its ridges are closely approximated to one another, but form deep and narrow 
grooves between. Digestive vacuoles are frequently found in the peripheral part of each 
ridge ; in this region nuclei, which have assumed the saccular appearance of those of the 
tentacular and oral-dise ectoderm, are thickly crowded together, some extending even into 
the central region of the ridge. Numerous nematocysts I are present in the lower half of 
each ridge; the development of these in such large numbers may account for the 
vesicular condition of the nuclei; nematocysts III are rarely met with. The mesoglea 
is considerably thickened in the ridges. The ectoderm of the oral-dise is typical; along 
its middle saccular nuclei (oval and elongated) are aggregated in diverging tracts of 

SECOND SERIES—ZOOLOGY, VOL. XVII. 11 


82 PERCY SLADEN TRUST EXPEDITION 


protoplasm between which mucous vacuoles are frequently present; in the lower half 
of the layer the protoplasm is clearer with scattered round nuclei. 

Filaments are present on all the mesenteries. Nematocysts I are quite common in 
the straight regions of the filaments of the principal mesenteries, less frequent in those 
of the subsidiary mesenteries ; in the coils numerous nematocysts IT and III are present, 
the former with the dark-stained axis slightly bent and the latter with the thread partly 
extruded; type I is scarce. The endoderm has a slaty-blue colour, conspicuous in 
the mesenteries; in the oral-disc it is thinner than the overlying ectoderm ; the circular 
layer of endodermal muscle-fibres 1s well seen. The tentacular endoderm is less than half 
the thickness of the sub-terminal batteries, with comparatively few alge. The mesenterial 
endoderm is only slightly swollen behind the filaments; incomplete nematocysts II are 
occasionally met with in it. Algz are comparatively rare, occurring mostly on 
the entoccelic side in the outer pleatal region and on the exocelic side in the inner 
pleatal region. To the inner exoccelic pleats thick muscle-bands are attached which 
make up for the somewhat weak musculature on the entoccelic pleats. Below the 
stomodzeal region of the polyp, the exoccelic pleats do not extend, but the entoccelic pleats 
cover almost the entire width of the mesenteries. One, two or three longitudinal rows of 
somewhat small ova were present in all the polyps, in some of the principal and subsidiary 
mesenteries. 

Number of polyps examined, fourteen, taken from ten different specimens of varying 
skeletal facies, all from Dongonab, Red Sea. 

B. Corallum. The exact position of the specimen from Minikoi, measuring 
12°5x115x6 em., which Gardiner referred to Fava versipora (Ehrb.), is doubtful 
(Pl. 21, fig. 4). As pointed out by him it comes nearest to the typical form of 
Klunzinger’s Pavia ehrenberg: (P1. III, fig. 7). Its corallites are polygonal, often elongated 
for fission, the single calices beg quite small, about 4 mm. wide and as much deep (the 
diameters of the longest calyx with signs of fusion into three being 10 and 3°6 mm.). The 
inter-calicinal walls are flat above, about 2 mm. thick. The septa are few in number, about 
20, of which 10—12 meet the columella; they are thick, vertical, with rough sides and 
4 or 5 bluntly-pointed horizontal teeth on their edges. The septa of adjacent corallites 
are usually continuous over the walls, being about ‘5 mm. exsert; the exsert ends are 
sometimes connected along the middle of the walls by thin longitudinal ridges. The 
columellee are formed of dense trabecule, often quite compact and 4—+ the width of the 
calices. Fission is sub-equal. The specimen also approaches PI. 21, fig. 3, in appearance, 
but the latter has larger corallites and thicker walls. 

Marenzeller has already pointed out Klunzinger’s error in regarding Madrepora 
favus, Forsk. as a species of Goniastrea. Five of Forskal’s examples of this species are 
at present in the Copenhagen Museum, one of which (21 x 17 x 15 cm.) comes near Milne 
Edwards and Haime’s type of Favia bertholleti. The remaining four (Pl. 22, figs. 1—4) 
are skeletal varieties of the present species; one of these (11 x 10°59 cm., fig. 4) has 
thick perithecal regions, distorted corallites, approaching the condition in Klunzinger's 
type of Fava ehrenbergi, var. laticollis. Madrepora cavernosa, Forsk. is represented by 
a single specimen (17x17 x11°5 em., Pl. 22, fig. 5), which retains the normal charac- 


MATTHAI—RECENT COLONIAL ASTRAIDA 83 


teristics of the present species but is different from Klunzinger’s example of Favia 
cavernosa. 

Ellis and Solander’s type of Madrepora denticulata is missing from the University 
Museum, Glasgow, but judging from their figure it certainly belongs here. 

In the Paris Museum there are six specimens from the Red Sea which had been 
originally referred by Milne Edwards and Haime to Pavia denticulata; of these four 
had later been assigned to Kawa savgnyi, Ed. and H., one of which (measuring 
16x 108 cm.) had been marked by Milne Edwards and Haime as their type of Favia 
denticulata; a fifth specimen (round, 11 x 11 x 8 em., with distorted corallites) was referred 
to Faia geoffroyi, Ed. and H., while the last (also round, 9x 8x5 cm.) had been 
erroneously referred by Klunzinger to his species, Kavia cavernosa. Favia savignyi, 
Ed. and H., is represented by ten excellent examples (excluding the denticulata specimens 
above referred to) from Red Sea, one of which is in Lamarck’s collection, having obviously 
been his type of Astrea denticulata (PI. 21, fig. 8); these show many of the skeletal 
variations exemplified in my collections. The single specimen (12 x 8x9 em.) of Favia 
amaicorum, Ed. and H., from Tongatabou also belongs here, but its corallum is much 
lighter, with thin perithecal regions, and fewer septa, these differences being doubtless 
due to conditions of growth. Two specimens (15 x 9°54 em. and 9x9xX6 cm.) with 
large corallites from unknown locality on which Milne Edwards and Haime had formed 
a new species, Pavia jacquwnot: (Pl. 21, fig. 7), are identical with Gardiner’s Minikoi 
example of Prionastrea magnistellata (fig. 40). These authors had also assigned a 
specimen each to Maa geoffroy: and Fawa deformata, measuring respectively 
10°5 x 10 x7 cm. (Red Sea) and 10 x 10x 8 cm. (unknown locality). Favia aspera, Ed. 
and H., is represented by five specimens from the Red Sea, the largest measuring 
20x 20x12cm. Identical with the latter specimen are a large example (23 x 23 x 14 cm.) 
from Seychelles which Milne Edwards and Haime had made the types of Favia rousseaw 
(Pl. 21, fig. 5), and five others referred by them to Prionastreaa halicora. The corallites 
of their type (18x 16 x 11 cm.) of Favia affinis have almost the same facies as that of the 
present species but the corallum is lighter (Pl. 21, fig. 6). 

In the Berlin Museum are five specimens assigned by Klunzinger to Favia ehren- 
bergi, Klunz.; two of these, which had been Ehrenberg’s originals of Favia versipora 
(=F. ehrenbergi, var. laticollis, Klunz., fig. 8), resemble one of Forskal’s examples of 
Madrepora favus. Klunzinger erroneously regarded Favia clower, Ed. and H., as practi- 
cally identical with his F. ehrenbergi var. sulcata (fig. 8); the last specimen (fig. 7) has 
smaller corallites and fewer septa, its place in the present species being doubtful. The 
appearance of pali-crowns in Klunzinger’s type (8°5 x 75 x 3°5 cm.) of Goniastrea seychel- 
lensis is due to a ring of upturned septal teeth four or five rows above the columella. 
Somewhat resembling this specimen is Studer’s large “Gazelle” example of Prionastrea 
seychellensis from New Ireland. Ortmann’s example (11°5 x 8°5 cm.) of Favia ehrenbergr 
from Dar-es-Salaam also belongs to F. favus. Ehrenberg’s type specimen of Astrea 
deformis (Pl. 35, fig. 1) perhaps belongs here, but it has also some resemblance to Favia 
berthollett. 

Marenzeller has made an extensive study of Fava savignyt, Ed. and H., to which he 

112 


84 PERCY SLADEN TRUST EXPEDITION 


) 


has referred 41 specimens of the “Pola” expedition, 26 from Idda, 10 from Kamaran, 
2 from Manuret and Hamidije, 1 each from Massawa, Bernice and Dahab. The 
Idda, Bernice and Dahab specimens are heavier, thicker and rougher, belonging to 
var. 2 of F’. favus, characterised by heavy coralla, many of them having the facies of 
Klunzinger’s type of F. ehrenbergi var. laticollis, while the rest are lighter, thinner and 
less coarse, belonging to var. 1. No. 15954 from Kamaran resembles Milne Edwards and 
Haime’s type of Favia amicorum and No. 15943 from Idda has towards its edge the 
facies of Ehrenberg’s type of Pavia complanata. 

Localities. Red Sea (104). Chagos, Salomon (6). Seychelles (4). Maldives: 
Suvadiva (1 from 25 fms.); Hulule (1 from reef). Minikoi (2 from reef). Singapore 
(1 between reef and shore). Ceylon, Weligama (1). Funafuti, Ellice Islands (2 doubtful). 
A species of wide distribution, particularly common in the Red Sea. Also from Tonga- 
tabou (Milne Edwards and Haime) and Dar-es-Salaam (Ortmann). 


2. Favta porrvensis, Milne Kdwards and Haime. (Pl. 9, figs. 1 and 3; 22, figs. 8 
and 9; 32, figs. 2—4.) 


1801. Astrea rotulosa, Lamarck, Syst. Anim. sans vert., p. 371 (non Madrepora rotulosa, Ellis and Solander). 

1816. Astrea rotulosa, Lamarck, Hist. Anim. sans vert., ii, p. 259; 2° édit., p. 405. 

1816. Astrea ananas, Lamarck, Hist. Anim. sans vert., ii, p. 260; 2° édit., p. 406 (non Madrepora ananas, 
Hillis and Solander). 

1850. Parastrea doreyensis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 168. 

1850. Parastrea urvilliana, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 169. 

1850. Purastrea rotulosa, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 170. 

1850. Parastrea ananas, Milne Edwards and Haime, Ann. Sci. Nat., Zool., Be ie, saul, 7/2, 

1850. Parastrea amplior, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 172. 

1857. Favia rotulosa, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 429. 

1857. Havia wrvilleana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 432. 

1857. avia doreyensis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 432. 

1857. Favia ananas, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 435. 

1857. Favia amplior, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 436. 

11889. Goniastrea serrata, Ortmann, Steinkorall, Siid. Ceylons, Zool. Jahrb., iv, p. 526, pl. 15, fig. 10. 

1899. Astrea okent, Gardiner, Proc. Zool. Soc. London, p, 749, pl. 47, fig. 2. 

11904. Fawia cavernosa (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, p. 767, pl. 61, fig. 13 (non 
Madrepora cavernosa, Forskal and Favia cavernosa, Klunzinger). 

1904. Havia denticulata, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 768, pl. 61 (non Astrea denticu- 
lata, Lamarck or Gardiner, Yavia denticulata, Khrenberg, Milne Edwards and Haime, or Klunzinger). 

1904. Faria laccadivica, Gardiner, Fauna Geogr. Maldives and Laccadives, p, 769. 

1904. Orbicella laxa, Gardiner, Fauna Geogr. Maldives and Laccadives, p, 775, pl. 63, fig. 33 (non Orbicella 
laxa, Klunzinger). 

1904. Orbicella borradailei, Gardiner, Fauna Geogr. Maldives and Laccadives, p, 775, pl. 63, fig. 34. 


Corallum. Massive, growth-form usually rounded. Peritheca vesicular, dissepiments 
more or less horizontal, 1 or 1°25 mm. apart. Corallites oval, distorted or sometimes 
circular, projecting to about 1mm. Walls thin or thick, 2 or 2°5mm.apart. Calices with 
diameters up to 13 x 9 mm.; average 8 x 6 mm.; of equal width from calicular margins to 
bases, depth 2—7 mm., usually 4°6 mm. 

Septa thin or thick, usually vertical, with rough sides, 20—40 in number, average 
24—34. Of these 14—20 meeting columella, each with its upper half or two-thirds 
(edge with 2—6 teeth) narrower than its lower half or one-third and as a rule with 


MATTHAI—RECENT COLONIAL ASTRAIDE 85 


a paliform lobe, exsert to about 1 mm., exsert ends usually arched sometimes flat. Costee 
thin and sharp or thick and flat, serrate or with transversely extending granulations, 
those of neighbouring corallites usually meeting in notches along the middle of the 
perithecal regions, sometimes alternating ; an alternating cycle of smaller costz (usually 
without corresponding septa) sometimes present. Columella formed of twisted septal 
trabecule, sometimes quite dense, from 4 to 4 width of calyx. 

Multiplication by unequal or sub-equal fission. 

Towards edge of specimens the corallites often project obliquely, the sides facing the 
edge projecting more—up to 3 mm.—than those facing the centre of corallum; the 
corallites are further apart (up to 4 mm.), with walls thicker, calices shallower, septa 
thicker and rougher, costze flat and thick. 

Gardiner’s examples of Orbicella laxa, Klunz. and Orbicella borradaile:, Gard. form 
a dense variety of the present species, characterised by obliquely projecting corallites with 
thick walls, well-developed paliform lobes, dense columella and a conspicuous alternating 
eycle of smaller costeze without corresponding septa. Some of the corallites appear to have 
been formed by budding. In certain regions the corallites have remained in the normal 
condition (see Pl. 32, fig. 3). The examples of Pavia laccadivica, Gard., show a somewhat 
crowded appearance of septa within the corallites; the cycle of smaller costz has rudi- 
_ mentary septa, and the exsert ends of the septa are either directly continuous between 
corallites or meet in shallow notches. One of Gardiner’s specimens of Favia cavernosa 
(Forsk.) has the corallites wide apart up to 7 or 10 mm. (average 3°6 or 4 mm.), not 
projecting ; the septa and costz are very thin, the latter directly continuous across the 
intervening perithecal regions and tending to be united by transverse ridges. 

Polyps. (1) Both entoccelic and exoccelic tentacles present. (2) Stomodezeal ridges 
broader than thick and somewhat constricted at their bases. (3) About eleven principal 
couples of mesenteries present. (4) In the stomodzal region of polyp subsidiary couples 
of mesenteries about twice the number of principal couples. (5) In the stomodzeal region 
of polyp, entoccelic pleats narrow except in middle of pleatal region, where they are some- 
times sub-divided, usually constricted at their bases and extending over outer halves of 
principal mesenteries. (6) Mesenterial endoderm vacuolated to form closely-arranged 
goblet-shaped spaces, swollen where entoccelic pleats are broader. (7) Convolutions of 
mesenteries not so abundant as to be massed together in inter-mesenteric chambers below 
stomodzeum, scarce or even absent towards bases of polyps. 

Remarks. A. Polyps. Of the four polyps examined three have each 11 principal 
couples of mesenteries (4 incomplete), while the remaining polyp has only 10 couples 
(6 incomplete). Subsidiary couples number 10—24, some not extending below the entero- 
stome ; the members of the same couple are frequently unequal. The convolutions of 
the mesenteries protrude into the edge-zone and through the oral-disc. The tentacles are 
often less in number than the entocceles and exocceles. The large terminal batteries 
consist of closely-arranged nematocysts I, each with from 35 to 40 turns of the spiral, and 
a few ILb; long narrow structures, homogeneously stained pink, are seen in large number 
in a few of the terminal batteries as in Galaxea fascicularis. Four to six sub-terminal 
batteries are present, in which large clear oval vacuoles are conspicuous. 


86 PERCY SLADEN TRUST EXPEDITION 


The stomodzeum is much compressed laterally, its long diameter being about 2 mm, 
Owing to the breadth and thickness of the stomodzal ridges grooves are formed between 
them. Rod and spindle-shaped nuclei are massed together in the ridges around a some- 
what granular protoplasmic regionyinto which slender filaments pass from the mesoglea ; 
digestive vacuoles and a few nematocysts IIT and rarely I also occur. The mesoglea is 
thickened, appearing concavo-convex in transverse section. Near the skeletal attach- 
ments of the mesenteries, the calicoblast is thickened and vacuolated, containing ‘‘ desmo- 
cytes” at various stages of development. In the oral-dise and in the outer wall of the 
edge-zone the ectoderm is thick, with a columnar facies containing a row of transparent 
oval vacuoles ; nuclei of varying shape occur in the middle of the layer, with nematocysts 
I and II above. Filaments are present on all the mesenteries, being rudimentary on 
the narrower subsidiaries ; nematocysts are comparatively few in them, in their straight 
regions occasionally types I, II and III, and in the coils a few III. In the mesenteries 
the mesogleea is thickened in the region of the wider pleats, thin in the remaining parts. 
Below the stomodzeal region of the polyp, the entoccelic pleats extend over the greater 
part of the width of the mesentery. 

The endoderm of the oral-disec is not so thick as the ectoderm over it, but has 
abundant algze; that of the body-wall is thin above the enterostome but towards the base of 
the polyp becomes highly reticulated, swollen and transparent with the nuclei arranged 
along its periphery. The endoderm of the tentacle is thinner than the batteries, with 
conspicuous vacuoles giving it a somewhat lobulated appearance, and algze are scarce. The 
mesenterial endoderm is only slightly thickened bebind the filament ; algee are compara- 
tively few, being more numerous in the pleatal region on the exoccelic side and in the non- 
pleatal region on the entoccelic side. In one of the polyps scattered nematocysts II occur 
in the endoderm, some of them incompletely formed. Ripe ova are present in three 
of the polyps in longitudinal rows of from 1 to 5 in every mesentery; immature ova 
occur in the endoderm surrounding the eggs. 

Polyps examined, four, 1 from a specimen from Hulule whose corallum is identical 
with Gardiner’s example of F. denticulata, 2 from a specimen from Hulule whose corallum 
resembles Gardiner’s example of Astrea okent and Ed. and H.’s #. amplior, 1 from an 
edge specimen from Minikoi. 

B. Corallum. In the Paris Museum Favia doreyensis, Ed. and H., and Favia 
urvilleana, Ed. and H., are each represented by a single type-specimen, measuring 
75x6x5em. (Pl. 22, fic. 8) and 8x5x45cm., both from Port Dorey; in the latter 
paliform lobes are inconspicuous. The single specimen (10x9x6 cm.) from Mers 
dAmerique, the type of Fava rotulosa, Ed. and H., is identical with some of my 
specimens and was doubtless Lamarck’s type of Astrea rotulosa (Pl. 32, fig. 4). In 
Lamarck’s collection is a specimen 7 x 6 X 2°5 cm. from Mers d’A merique originally named 
Astrea ananas by Lamarck, which Milne Edwards and Haime have later made the type of 
Favia ananas. Its corallites are often irregularly compressed, one side projecting more 
than the other, columella and paliform lobes well developed, a cycle of smaller costz 
alternating with the main ones, sometimes represented by rudimentary septa within 
corallites, in general appearance resembling Gardiner’s examples of Orbicella laxa and 


MATTHAI—RECENT COLONIAL ASTRAIDA 87 


borradaile: but calices slightly smaller and walls thinner. Similar to this example but 
with somewhat larger calices are two small specimens (from unknown locality) in the same 
collection, having the facies of Gardiner’s example of Astrea oken from Rotuma; they 
were undoubtedly Lamarck’s types of Astrea ananas var. stellis amplioribus; on these 
Milne Edwards and Haime have founded their species Favia amplior. A large specimen 
(70 x 60 x 35 cm.) from Seychelles in Rousseau’s collection named Prionastrea quoyi is 
identical with my types of F. doreyensis. 

In the Berlin Museum are two specimens from Ceylon named Goniastrea serrata, 

Ortm., the larger measuring 23 x 18x17 cm., presumably Ortmann’s type; the second 
specimen much smaller and not properly cleaned. These do not belong to’ Goniastrea 
but comes near my examples of Hava doreyensis, their main difference consisting in the 
absence of paliform lobes. 
In the Challenger collection is a small specimen from Fiji, which Quelch has referred 
to Astrea dane (Kd. and H.), in every respect identical with my types of Favia dorey- 
ensis. A broken off fragment from Mactan Island, Philippines, which the same author 
has assigned to Astrea ordinata, Verrill, resembles Gardiner’s example of Astrea okeni 
(Ed. and H.). 

Localities. Maldives: Hulule (9); Goidu (6). Minikoi (11). Seychelles (8). 
' Ceetivy (1). Chagos : Salomon (4); Coin, Peros (1); Rotuma (1). Also from American 
Seas (Lamarck), Port Dorey (Milne Edwards and Haime), Ceylon (Ortmann), Fiji and 
Mactan Island, Philippines (Quelch). Not recorded from the Red Sea. 


3. Hayi4 HuLULENSIS, Gardiner. (Pl. 9, fig. 6 ; 22, fig. 6; 35, fig. 1.) 

1834. Favia rotulosa, Ehrenberg, Corall. roth. Meer., p. 95 (non Madrepora rotulosa, Ellis and Solander, or 
Astrea rotulosa, Lamarck). i 

1899. Astrea fragilis, Gardiner, Proc. Zool. Soc. London, p. 748. 

1904. Favia hululensis, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 769, pl. 41, figs. 19—21. 

Corallum. Massive, rounded, sometimes incrusting. Peritheca vesicular as in 
previous species. Corallites circular or oval, projecting from ‘5—1 mm., closer together 
than in F. doreyensis. Walls thin. Calices somewhat smaller than in previous species, 
with average diameters 7 x 5°5 mm. and of equal width from calicular margins to bases, 
depth 3—6 mm. 

Septa thin, vertical, with toothed edges, rough sides, 20—30 in number, average 
22—25. 8—12 septa meeting columella, usually 9 or 10, each with its upper half or 
two-thirds narrower, as a rule without paliform lobes (sometimes the lowest septal tooth 
tending to be paliform), exsert to about 1 mm., exsert ends arched, entire or serrate. 
 Costze usually thin, those of neighbouring corallites either meeting in notches or alter- 
nating. Columella rudimentary. 

Multiplication by sub- equal or unequal fission. 

Towards edge of specimens the corallites project obliquely as in F. doreyensis, up 
to 2mm., and are further apart, up to 4mm. Gardiner’s specimen no. 4 (p. 770, fig. 21) 
is peculiar in that, crowded between the exsert septal ends of each corallite, are about 
20 similar ridges without traces of septa. 

In general appearance this species is like a smaller edition of Favia dore, yensis. 


88 PERCY SLADEN TRUST EXPEDITION 


Polyps. (1) Exoccelic tentacles absent. (2) Stomodzal ridges thicker than broad, 
their sides sloping away from apices to bases. (8) About seven principal couples of 
mesenteries present. (4) In the stomodzal region of polyp subsidiary couples of 
mesenteries about twice the number of principal couples. (5) In the same region, 
entoccelic pleats narrow, thick, undivided, unconstricted at their bases, directed obliquely 
towards stomodeeum, not extending beyond outer two-thirds of width of principal 
mesenteries, better seen in their outer halves. (6) Mesenterial endoderm thin in pleatal 
region, but gradually swelling towards stomodeum. (7) Convolutions of mesenteries 
abundant below stomodzum to base of polyp, massed together in intermesenteric 
chambers. 

Remarks. A. Polyps. These are smaller than in F. doreyensis but more 
regular in form—either circular’ or oval. Owing to close approximation of corallites 
there is little or no ccenosare. An appearance of binary fission frequently results 
from the presence of two oral openings placed at equal distances from the tentacular 
ring. In the two polyps examined 7 principal couples of mesenteries are present, 
2 of them being incomplete, and 13 subsidiary couples. The convolutions of the 
mesenteries are scarce in the stomodzeal region. The entoccelic tentacles equal the 
entocceles in number. In the large bluntly pointed terminal batteries long narrow 
pink bodies are found as in the previous species ; certain appearances suggest that these 
may be immature stages in the development of nematocysts IIb. About six sub-terminal 
batteries are present. 

The stomodzeum is almost circular in transverse section, its diameter being 1°5 mm., 
its ridges are much less broad than in F. doreyensis, appearing more or less conical, with 
no deep grooves between, their mesoglea only slightly thickened. Both ectoderm and 
endoderm are vacuolated as in the previous species, but the tentacular endoderm is 
thinner. Nematocysts I] are present in the coils of the filaments in addition to those 
of types I and III. In the mesenteries the mesogleea is usually thinner in the non-pleatal 
than in the pleatal region, but is thickened to a short distance from the stomodeal 
attachment; on this thickened region narrow entoccelic pleats are seen. Below the 
stomodeal region of the polyp the entoccelic pleats extend over the entire width of 
the principal mesenteries. 

Both the polyps sectioned had eggs in every mesentery, in one or two longitudinal 
rows. Immature germ cells, surrounding the eggs, are better seen than in the polyps of 
the previous species. The proliferation of the calicoblastic ectoderm is conspicuous even 
towards the bases of the polyps containing cell elements similar to those of the immature 
ova. This appearance suggests that these polyps were in a state of reproductive activity. 
When all the eggs have been formed, as in the polyps of F. doreyensis, the calicoblast 
loses all signs of its proliferating activity and the follicle-cells become fewer. 

In the Berlin Museum is a specimen (No. 739) measuring 14 x 12 x 7 cm. referred 
to Favia rotulosa by Ehrenberg (Pl. 35, fig. 1), resembling Gardiner’s example no. 1 
(p. 770, fig. 19). 

Number of polyps examined, two, from a specimen from Hulule, Maldives. 


Localities. Maldives: Hulule (5); Goidu (1). Ceylon (2). Rotuma (1). Funafuti 


MATTHAI—RECENT COLONIAL ASTRAIDA 89 


(a doubtful specimen). Ccetivy (1). Also one of Ehrenberg’s specimens from the 
Red Sea. 


4, Havra crover (Valenciennes). (Pl. 10, fig. 6; 23, figs. 1, 2 and 5; 25, fig. 3; 
34, fig. 1.) 
1797. Madrepora radiata, Esper, Forts. Pflanz., p. 74, pl. 61 (non Madrepora radiata, Ellis and Solander). 
1815. Favia radiata, Oken, Lehrb. Naturg., i, p. 68. 
1834. Faria wa, Ehrenberg, Corall. roth. Meer., p. 94 (non Madrepora wva, Esper, which is a Dichoceenia). 
11846. Astrea ( fissicella) speciosa, Dana, Expl. exp. Zooph., p. 220, pl. 11, fig. 1. 

Parastrea clowei, Valenciennes, Mss., Catal. Mus. Paris. 

1849. | Parastrea radiata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 169. 
1857. Favia okeni, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 430. 
1857. Favia clowei, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 436 (non Favia clowei, Klunzinger). 
1879. Favia cavernosa, Klunzinger, Korall. Roth. Meer., ii, p. 26, pl. 3, fig. 4 (non Madrepora cavernosa, Forskal). 
1879. Favia tubulifera, Klunzinger, Korall. Roth. Meer., ii, p. 28, pl. 3, fig. 6, and pl. x, fig. 2. 
1899. Astrea affinis, Gardiner, Proc. Zool. Soc. London, p. 750 (non Pavia affinis, Milne Edwards and Haime). 
1904. Favia cavernosa, (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 767, pl. 61, fig. 13. 
1904. Favia affinis, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 767, pl. 62, figs. 22 and 23. 


Corallum. Massive, becoming convex. Peritheca highly vesicular, vesicles on 
surface often like blisters, easily broken, dissepiments thin, 1—1°5 mm. apart, corallum 
consequently light. Corallites oval, irregularly compressed or circular, with usually 
distinct margins, level or projecting to 1°5 mm.; interior often appearing blistery; owing to 
vesicular deposition of calcareous matter; up to 7 or 10 mm. apart, average 2—4 mm. 
Calices 7—10 mm. in diameter, average depth 4 or 5 mm. 

Septa thin in calices, often swollen in thece (swollen parts hollow), usually vertical, 
with toothed edges, spinulose sides, exsert to 1 or 1°5 mm., about 35 in number ; of these 
up to 17 or 18 meeting columella, average 13—15, with the upper two-thirds or half 
much narrower than the lower one-third or half, the former with short teeth or entire 
edges, while the latter with 3 or 4 prominent teeth, the first two of which often vertical, 
simulating a pali-crown; sometimes an alternating cycle of rudimentary septa present. 
Columella formed of thin twisted septal trabeculee, 1—14 width of calyx, sometimes rudi- 
mentary. Cost prominent, thin, toothed, continued over peritheca as thin ridges usually 
meeting those of neighbouring corallites, the perithecal coste often tending to be united 
transversely by a thin heavy ridge along the middle of the perithecal regions. 

Multiplication by unequal, sometimes equal, fission. 

The species is characterised by the light corallum, perithecal costze, open calices, and 
thin septa. 

Polyps. (1) Both entoccelic and exoccelic tentacles present (the latter few in 
number). (2) Stomodzeal ridges thicker than broad, with convex inner surfaces and 
sides almost perpendicular or only slightly sloping. (8) About eight principal couples of 
mesenteries. (4) In the stomodeeal region of polyp subsidiary couples of mesenteries, 
about twice the number of principal couples. (5) In the same region of polyp, entoccelic 
pleats, somewhat narrow, unconstricted, some of them sub-divided, arising usually at right 
angles to the mesoglea, sometimes sloping towards stomodseum and extending to not 
more than middle of principal mesenteries, usually over about one-third their width ; 

SECOND SERIES—ZOOLOGY, VOL. XVII. . 12 


90 PERCY SLADEN TRUST EXPEDITION 


non-pleatal region quite thin. (6) Mesenterial endoderm swollen, more in pleatal than in 
non-pleatal region. (7) Convolutions of mesenteries abundant below stomodzeum to base 
of polyps, massed in inter-mesenteric chambers. 

Remarks. A. Polyps. Three polyps have 8 principal couples of mesenteries to each, 
while the number in another is only 7; of these 1—5 are incomplete. 17—20 secondary 
couples of mesenteries are present in the tentacular regions of the polyps, of which only 
10—15 extend down to the lower end of the stomodzea. Each of the tentacles has 
a swollen terminal battery in which incompletely-formed nematocysts II b are interspersed 
among the closely-arranged nematocysts I, the latter with between 40 and 50 turns of the 
spiral; 5 or 6 sub-terminal batteries. The tentacular endoderm is vacuolated and lobu- 
lated, about the same thickness as the overlying ectoderm, with algze massed together in 
the region of the terminal battery. 

The stomodzeum is laterally compressed, its diameters 1°5—-2 mm. and »5—1 mm. 
Nematocysts I are of common occurrence in the stomodzal ridges. The ectoderm of the 
oral-disc has numerous mucous vacuoles, the endoderm being as thick or thicker, and with 
closely-packed algze. The mesenterial endoderm is vacuolated and almost transparent, the 
vacuoles being somewhat oval, and contains few alge. Filaments are present on all 
the mesenteries; nematocysts I are present in large numbers in their straight regions, 
while in their coils type III are abundant (unfortunately not properly fixed); those 
of type II are few. The mesenterial endoderm is considerably swollen behind the 
filaments, up to three times the size of the latter. Below the stomodzeum the endoderm 
of the column-wall is reticulated and quite transparent. Gonads were not present in 
any of the polyps examined. 

The polyps of this species appear to be more nearly related to F. hululensis than to 
any other species of Havia. i 

Number of polyps examined, eight, 4, 2 and 2, all from Ceylon, also two additional 
polyps from a doubtful specimen from Minikoi. 

B. Corallum. Inthe Paris Museum there are three large specimens (38 x 38 x 12 cm., 
37 X29 x 15 em., 22 x 21 x 12 em.) from Seychelles, referred by Milne Edwards and Haime 
to Favia clouer, in which the corallites are oval or irregularly compressed laterally, slightly 
projecting, coralla light, the perithecal dissepiments about 1:5 mm. apart and the 
columellee poorly developed (PI. 25, fig. 2), on the whole resembling my Ceylon examples. 
The same authors have referred three specimens from the Red Sea to Parastrea radiata 
and later to Favia oken, the largest of which measures 16 x 12 x 8 cm. ; their only differ- 
ence from Havia clouei is in the larger size of their corallites (Pl. 25, fig. 1). 

Ehrenberg’s type of Favia uva is a large example which Klunzinger later made the 
type of Favia cavernosa. Marenzeller discovered Klunzinger’s mistake in naming this 
specimen after Forskal’s Madrepora cavernosa and correctly pointed out its similarity 
with Milne Edwards and Haime’s types of Favia okent. Klunzinger’s two examples of 
Favia tubulifera are essentially similar to Ehrenberg’s type, the only difference being that 
in the former the corallites are not so far apart as in the latter; my Ceylon specimens are 
identical with these two species of Klunzinger’s. In the Berlin Museum is also a small 
convex specimen from Ceylon (no. 3729), referred by Ortmann to Fava amplior, 


MATTHAI—RECENT COLONIAL ASTRAIDA 91 


Ed. and H., which resembles Klunzinger’s Favia cavernosa and belongs to the present 
species. 
The three small specimens from Banda, referred by Quelch to Astrea pandanus, 
Dana, Astrea doreyensis (Ed. and H.), and Astrea speciosa, Dana, are identical with 
Klunzinger’s types of Hava tubulifera and hence come under the present species. 

Localities. Maldives, Hulule (1). Minikoi (3). Chagos: Salomon (3); Egmont 
(1). Ceylon: Delft and Nainitavoe (small specimens). Also from Seychelles (Milne 
Edwards and Haime) and from the Red Sea (Milne Edwards and Haime and Klunzinger) 
4 Hast Indies (Dana), ? loc. (Esper). 


5. avira appira (Hllis and Solander). (Pl. 9, fig. 5; 29, figs. 14; 86, fig. 2.) 


1786. Madrepora abdita, Ellis and Solander, Nat. Hist. Zooph., p. 162, pl. 50, fig. 2. 

1797. Madrepora favosa (pars), Esper, Forts. Pflanz., p. 34, pl. 45a, fig. 2 (copy of Ellis). 

1816. Astrea abdita, Lamarck, Hist. Anim. sans vert., ii, p. 265; 2° édit., p. 415. 

1830. Favastrea magnifica, Blainville, Dict. Sci. Nat., 1x, p. 340; Manuel d’actin., p. 374, pl. 54, fig. 3 (1834). 

1833. Astrea abdita, Quoy and Gaimard, Voy. l’Astrol., Zooph., p. 205, pl. 16, figs. 4—5. 

1833. Astrea fusco-viridis, Quoy and Gaimard, Voy. l’Astrol., Zool., iv, pl. 17, figs. 8 and 9. 

1834. Astrea pentagona, Ehrenberg, Corall. roth. Meer., p. 96. 

1834, Astrea abdita, Ehrenberg, Corall. roth. Meer., p. 97. 

1846. Astrea fusco-viridis, Dana, Expl. exp. Zooph., p. 228, pl. 11, fig. 7. 

11846. Astrea ( fissicella) magnifica, Dana, Expl. exp. Zooph., p. 231, pl. 12, fig. 3. 

1850. Prionastrea abdita, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 128. 

1850. Prionasirea magnifica, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 129. 

1850. Prionastrea sulfurea, Milne Edwards and Haime, Ann, Sci. Nat., Zool., 3° sér., xii, p. 130 = Astrea sulfurea, 
Valenciennes, Mss., Catal. Mus. Paris. 

11850. Prionastrea quoyt, Milne Edwards and Haime, Ann. Sci. Nat., Zool , 3° sér., xii, p. 130. 

1850. Prionastrea obtusata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p: 180=Astrea obtusata, 
Lamarck, Mss. 

1850. Prionastrea crassior, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 131 (non Prionastrea 
crassior, Gardiner). 

1857. Prionastrea abdita, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 514. 

1857. Prionastrea crassior, Milne Kdwards and Haime, Hist. Nat. Corall., ii, p. 515. 

1857. Prionastrea magnifica, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 515. 

1857. Prionastreea obtusata, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 518. 

1857. Prionastreea sulfurea, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 518. 

11857. Prionastrea quoyi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 519. 

1877. Prionastrea sulfurea, Studer, Monat. Ak. Wiss. Berlin, p. 639. 

1877. Prionastrea robusta, Studer, Monat. Ak. Wiss. Berlin, p. 640. 

1879. Prionastrea gibbosa, Klunzinger, Korall. Roth. Meer., iii, p. 40, pl. 4, fig. 10. 

1899. Prionastrea abdita, Gardiner, Proc. Zool. Soc. London, p. 758, pl. 47, fig. 4. 

1899. Prionastrea fusco-viridis, Gardiner, Proc. Zool. Soc. London, p. 759, pl. 47, fig. 5. 

1899. Prionastrea purpurea, Gardiner, Proc. Zool. Soc. London, p. 760. 

1899. Prionastrea echinata, Gardiner, Proc. Zool. Soc. London, p. 760, pl. 47, fig. 6. 

1904. Prionastrea fusco-viridis, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 787, pl. 64, figs. 43 
and 44. : 


i Corallum. Incrusting, following irregularities on surface of attachment, frequently 
rising into irregular hillocks, sometimes with a corallite on top of each or more often 
ending in a ridge, with corallites on either side. Corallites polygonal, varying greatly in 
shape, often one side much higher than the other, hence with oblique calicular openings. 
Corallite-walls fused, ridged, the inter-calicinal partitions thus formed varying from being 
very thin up to 6 mm. thick, solid in section. Calices varying considerably in diameter, 
12—2 


92 PERCY SLADEN TRUST EXPEDITION 


widest at calicular margins, up to 20x12 mm., average about 11 or 12 mm., decreasing 
towards bases, depth up to 11 mm., usually about 8 mm., but much shallower in valleys, 
and near edges sometimes quite flat. 

Septa varying in thickness, sloping, with toothed edges, sides smooth to rough, from 
35 to 70 in number. Of these from 14 to 20 septa meeting columella, not more than 5 mm. 
exsert, their edges with 8 to 18 closely-arranged teeth, usually becoming elongated towards 
columella (up to 2 mm. high), round the latter tending to become upright. The broader 
subsidiary septa usually curving towards and fusing with sides of principals. An 
alternating cycle of rudimentary septa with entire margins present, these in adjacent 
corallites usually meeting the main septa at angles over the walls. Columella formed of 
twisted trabeculee, 2 or 2°5 mm. in diameter. 

Multiplication by very unequal fission towards the calicular margins. In the marginal 
region of a large corallite, one to four small corallites are frequently seen ; when definite 
walls are formed round these, they simulate buds. 

The peculiar growth-form—incrusting corallum rising into sharp-ridged irregular 
hillocks—fusion of corallite-walls, and the presence of small corallites near the calicular 
margins of the normal individuals are striking features of this species, giving the speci- 
mens an unique appearance. 

Two principal types, connected by gradational forms, are to be distinguished, one in 
which the intercalicinal walls and septa are very thin and columella openly spongy as in 
most of the abdita specimens (Pl. 29, fig. 1; Pl. 35, fig. 2), the other with thicker walls 
and septa and columella formed of closely-twisted trabeculee as in Gardiner’s examples of 
Prionastrea fusco-viridis (Pl. 29, figs. 3 and 4). 

Polyps. (1) Exoccelic tentacles absent. (2) Stomodeeal ridges usually broader 
than thick, constricted at their bases and with convex inner surfaces, in transverse section 
more or less resembling those of F. pentagona. (3) About nine principal couples of 
mesenteries. (4) In the stomodzeal region of polyp subsidiary couples of mesenteries, 
about thrice the number of principal couples. (5) In the stomodeeal region of polyp, 
entoccelic pleats very narrow, thick, unconstricted at their bases (in some mesenteries 
hardly recognisable) and restricted to outer quarter of width of principal mesenteries ; 
mesenterial mesoglea stouter than in #. favus; pleatal region thicker than non-pleatal 
region. (6) Mesenterial endoderm vacuolated and somewhat swollen in pleatal region 
and near stomodeeal attachment, the vacuoles being goblet-shaped, elsewhere quite thin. 
(7) Convolutions of mesenteries massed together in inter-mesenteric chambers below 
stomodzeum to base of polyp. 

Remarks. A. Polyps. The polyps follow the very irregular shapes of the corallites. 
Owing to their approximation the ccenosare is absent. About twenty-three subsidiary 
couples of mesenteries were counted ; some of the newly-formed couples are very narrow, 
not extending below the stomodeeal region of the polyps and without tentacles over their 
entocceles. The tentacular batteries resemble in shape and structure those of F. favus. 
The stomodzeum is oval in transverse section with the diameters measuring about 1°25 and 
‘75 mm. Nematocysts I and III are occasionally present in the ridges, but the mesoglea 
is not thickened in them. Filaments are present on nearly all the mesenteries, being 


MATTHAI—RECENT COLONIAL ASTRAIDA 93 


definitely hemispherical in transverse section ; in their straight regions nematocysts I are 
less numerous than in F, favus, while in their coils types II and III occur, the latter 
not in such large numbers as in /. favus. The endoderm stains brown when treated with 
iron—hzematoxylin and eosin, specially noticeably in the mesenteries. The goblet-shaped 
vacuoles in the mesenterial endoderm are massed together as in Cyphastrea serailia. 
Behind the filaments the endoderm is swollen to their thickness, the filaments and their 
endodermal pads giving a characteristic appearance to the polyp in transverse section. 
Below the stomodzeal region of the polyp, the entoccelic pleats are better seen and extend 
over the greater part of the width of the mesenteries. 

In two of the polyps sectioned large ova are present in all the mesenteries in one to 
four longitudinal rows, with germ-cells in the endoderm surrounding them. In another 
polyp a group of minute dark-stained bodies of unknown nature is present in the mesoglea 
of nearly every mesentery in a cavity near its skeletal attachment. 

Number of polyps examined, four, 2 from a mature ?, Hulule, Maldives, 1 from a 
specimen dredged off Salomon, 1 from a specimen from Ceylon. 

B. Corallum. In the Paris Museum Prionastrea abdita, Ed. and H., is represented 
by four large specimens (three thin, the fourth somewhat thickened), probably from the 
Indian Ocean. A large specimen from Batavia, Indian Ocean referred by these authors 
_to Prionastrea magnifica (Blain.) is similar to P. abdita, Ed. and H. Prionastrea 
erassior, Ed. and H., is represented by two type specimens, the larger measuring 
14x 12cm., and Prionastrea obtusata, (Lam.) by a small convex one from Tongatabou, 
all three being of the thick type. Two small examples from Vanikoro referred by Milne 
Edwards and Haime to Prionastrea sulfwrea (Valen.) and another smaller one from 
New Ireland to Prionastrea quoyi, Ed. and H., are perhaps also to be placed in this 
species. 
Ehrenberg’s example of Astrwa abdita has dendroid hillocks with thin corallites at 
the top of the hillocks and thicker ones on the main stems (Pl. 35, fig. 2). Klunzinger’s 
original of Prionastrea gibbosa is only a small edition of the present species. ‘Two worn- 
out specimens referred by Ehrenberg to Astrea pentagona, Ehrb. are quite unlike 
Klunzinger’s example of Prionastrea pentagona ; they resemble the present species more 
than any other. 

Ellis and Solander’s figured type of Madrepora abdita (15x15 x9 cm.) is in the 
University Museum, Glasgow ; though much damaged, its resemblance to the abdita 
‘specimens in Paris and Berlin is obvious. 

Quelch has referred two small specimens from Kandavu, Fiji, to Prionastrea flexuosa, 
(Dana) and Prionastrea obtusata, Ed. and H., and two others from Amboina to Prion- 
astrea quoyi, Ed. and H.; these four resemble the thick variety of F. abdita. A fairly 
large specimen assigned to Prionastrea robusta (Dana) agrees with the thin type. 
A small fragment from Banda, referred by the same author to Gonastrea favistella 
(Dana), is identical with Klunzinger’s type of Prionastrea gibbosa. 

Localities. Maldives: Hulule (12); Addu (1 from edge of W. reef); Goidu (5). 
Minikoi (6). Rotuma (11). Chagos: Salomon (2); Egmont, (1). Singapore (2). Ceylon 
(small specimens). Also from Tongatabou, ? Vanikoro and ? New Ireland (Milne Edwards 


94 PERCY SLADEN TRUST EXPEDITION 


and Haime), Fiji (Dana and Quelch), Amboina and Banda (Quelch), Salawatti (Studer), 
2 Rast Indies (Dana), and Red Sea (Ehrenberg and Klunzinger). 


6. ayra BERTHOLLETI (Valenciennes). (PI. 7, fig. 2; 22, fig. 7; 23, figs. 4 and 6; 

24, fig. 1). 

Parastrea bertholleti, Valenciennes, Mss., Catal. Mus. Paris. 
1850. Phymastrea valenciennesti, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, pl. 9, fig. 3, and xii, 
. 124. 

1850. ee rousseaut (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 131. 

1851. Prionastrea halicora (pars), Milne Edwards and Haime, Pol. foss. terr. palzoz., p. 102 (non Astrea 
halicora, Ehrenberg). 

1857. Favia bertholleti, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 431. 

1857. Phymastrea valenciennesi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 500. 

1857. Prionastrea halicora (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 517. 

21857. Prionastreea australensis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 520. 

1886. Phymastrea aspera, Quelch, Challenger Reports, Reef Corals, Zool., vol. xvi, part xlvi, p. 105, pl. 4, 
figs. 1—16. 

Corallum. Massive, convex; corallites longer than broad, usually hexagonal in 
shape, close together. Calices with diameters about 12 or 13mm. and 7 or 8mm. 

Septa usually sloping towards columella, with irregularly toothed edges (about 
8 teeth), exsert to about ‘5mm. 33—40 in number (in one corallite 46 septa), 11—18 
meeting columella ; the broader subsidiary septa curving towards and fusing with sides of 
principal septa near to where the latter meet columella. In some corallites an alternating 
cycle of rudimentary septa present. Columella usually formed of closely twisted septal 
trabeculee, about 4 width of calyx. 

Multiplication by unequal fission. 

Two principal types of corallum may be recognised in this species : (1) in which the 
adjacent corallite-walls are fused, the inter-calicinal walls thus formed being not more 
than 1 mm. in thickness, often thinner; over these the septa are continuous in arches, 
the septa being thin; (2) thicker-looking in which the corallite-walls are distinct, separated 
on the surface by inter-corallite grooves at the margins of which the exsert ends of the 
septa stop ; the septa are thicker and rougher. 

Polyps. (1) Both entoccelic and exoccelic tentacles present. (2) Stomodeeal ridges 
thicker than broad, with convex inner surfaces. (3) Nine or ten principal couples of 
mesenteries. (4) In the stomodzal region of polyp subsidiary couples of mesenteries about 
twice the number of principal couples. (5) In the same region of polyp entoccelic pleats 
extending over almost the entire width of principal mesenteries, better developed in their 
outer half or one-third where the pleats are broader than in any other species of Fava, 
thin, greatly subdivided, individual ones appearing club-shaped in transverse section ; 
in inner one-third pleats narrower, thicker, but subdivided to a less extent; between 
these two regions pleats often quite narrow or absent and the mesogleea slightly constricted. 
(6) Mesenterial endoderm usually thicker in inner pleatal region, and somewhat constricted 
between the two regions. (7) Convolutions of mesenteries abundant to some distance below 
stomodzeum but no blocking inter-mesenteric chambers, absent at base of polyps. é 

Remarks. A. Polyps. In three large polyps principal couples of mesenteries 
numbered nine, ten and eleven; a smaller one had only seven such couples. About 


MATTHAI—RECENT COLONIAL ASTRAIDA 95 


twenty-one subsidiary couples were present in the tentacular region, sixteen or seventeen 
extended down into the stomodzeal region. Numerous mucous vacuoles are present in the 
ectoderm of the oral-disc. The endoderm of the latter is as thick as the ectoderm, with 
closely packed alge. The tentacles have swollen terminal batteries with the usual 
structure and four to six sub-terminal batteries; nematocysts I have about forty turns 
of the spiral. The tentacular endoderm is lobulated with numerous alge but is only about 
half the thickness of the overlying ectoderm. The stomodzeum is laterally compressed ; in 
the lower halves of its ridges nematocysts III and I] are frequently present, the former 
more numerous than in any other species examined. Filaments are present on all the 
mesenteries ; in their straight regions nematocysts I are of common occurrence; in their 
coils are numerous nematocysts II and III. The mesenterial endoderm has comparatively 
few algze; it is swollen behind the filaments to the size of the latter or even larger ; into 
this the pleats do not extend. In transverse sections the entoccelic pleats have an unique 
appearance owing to their great breadth and branching nature. No gonads were present 
in any of the polyps. 

Number of polyps examined, four, 2 from a colony from Ceylon of var. 1, and 2 from 

another colony from Ceylon of var. 2. 


B. Corallum. Milne Edwards and Haime’s type of Fawa bertholleti is a large 
“specimen (24 x 17 x 17 em.) from Seychelles in which inter-corallite grooves are usually 
present (Pl. 23, fig. 4); identical with this are some of my Ceylon specimens. Of the 
eight examples referred by these authors to Prionastrea halicora, three come under the 
present species. Their type of Prionastrea australensis has a meandering tendency 
but the separate corallites resemble those of Mavia bertholle. The single small type 
of Phymastrea valencienness (an edge of a colony, measuring 5 x 4°5 x 2cm.) perhaps 
belongs to the present species; it has deep inter-corallite grooves and coarse septal sides 
and may therefore be only an extreme case of var. 2, described above, but the principal 
septa have long teeth near their union with the columella. 

One of Forskal’s types of Madrepora favus (21x17x15cm., Pl. 22, fig. 7) has the 
same facies as var. 1 of the present species. Quelch’s type of Phymastrea aspera is 
a small fragment, with deep inter-corallite grooves, which in all probability belongs here. 

Localities. Seychelles (2). Aldabra (2). Ceylon (5); also broken ones from Point 
Pedro, Nainitavoe and Delft. Also from the Red Sea (Milne Edwards and Haime and 
?Forskal), ? Australia (Milne Edwards and Haime), and Banda (Quelch). 


7. avia pEnrsAGonaé (Esper). (Pl. 10, fig. 5; 24, figs. 2—4; 36, fig. 4.) 


1797. Madrepora pentagona, Esper, Forts. Pflanz., p. 23, pl. 39, figs. 1 and 2. 

1834, Astrea melicerum, Ehrenberg, Corall. roth. Meer., p. 96. 

1850. Prionastrea gibbosissima, Milne Edwards‘and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 133. 

1850. Goniastrea rudis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 161. 

1857. Goniastrea rudis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 445, pl. D5, fig. 5. 

1857. Prionastrea melicerwm, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 521. 

1877. Plesiastreea haeckeli, Briiggemann, Abh. Ver. Bremen, v, pp. 395—400. 

1879. Prionastrea pentagona, Klunzinger, Korall. Roth. Meer., iii, p. 41, pl. 4, fig. 11 (non Astrea pentagona, 
Ehrenberg). : 

1886. Goniastrea laxa, Quelch, Reef Corals, Challenger Reports, vol. xvi, part xlvi, p. 102, pl. 3, figs. 4— 4d. 


96 PEROY SLADEN TRUST EXPEDITION 


1904. %Goniastrea favus, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 773 (non Goniastrea favus, 
Klunzinger). 

1904. Stephanocenia maldivensis, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 784. 

1904. Prionastrea pentagona, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 785. 

21907. Favia hawariensis, Vaughan, Recent Madreporaria of the Hawaiian Islands and Laysau, U.S. Nat. 
Mus., Bull. 59, p. 105, pl. 26, figs. 3 and 3a. 


Corallum. Incrusting or massive, often with low humps. Corallites polygonal. 
Walls fused, the inter-calicinal partitions thus formed up to 3°6 mm. thick, usually about 
‘75 or 1mm., ridged, solid in section. Calices up to 6 mm. in diameter, average 4—5 mm., 
depth 3—4 mm. 

Septa usually thin, sometimes thickened, vertical, with denticulate or entire edges, 
smooth or slightly rough sides, 20—30 in number, average 22—24, alternating in adjacent 
corallites or meeting at angles over the walls. 9—14 septa meeting columella, very 
slightly exsert, each with a bluntly pointed conspicuous paliform lobe. Columella spongy, 
4—4} width of calyx. 

Multiplication by unequal fission ; when definite walls are formed round the daughter 
corallites, they appear like buds intercalated among the larger corallites. 

Towards edges inter-calicinal walls become thicker (calices then decreasing in width 
from margins to bases), some of the subsidiary septa so narrow as to be hardly recognisable, 
columella rudimentary or even absent. 

Polyps. (1) Both entoccelic and exoccelic tentacles present. (2) Stomodeeal ridges 
broader than thick (some of them twice as broad as thick), constricted at their bases, and 
with convex inner surfaces. In contrast to the small size of the polyps their ridges 
broader than in all the previous species. (3) Seven to nine principal couples of mesen- 
teries. (4) In the stomodzeal region of polyps subsidiary couples of mesenteries about 
twice the number of principal couples. (5) In the stomodzeal region of polyp, entoccelic 
pleats narrow and thick—except towards middle of mesentery where they are much 
broader, thinner and occasionally sub-divided—and not extending beyond outer two-thirds 
of width of principal mesenteries ; mesenterial mesoglea thickened only in region of 
broader pleats, and near stomodzal attachment, elsewhere quite thin. (6) Mesenterial 
endoderm extremely thin, except on either side of the broader pleats and near the 
stomodeum. (7) Convolutions of mesenteries never abundant, absent towards base 
of polyp. 

Remarks. A. Polyps. These are small, circular or polygonal in outline, with little 
or no coenosare. In one polyp there are 9 principal couples of mesenteries, of which 
4 are incomplete; in 2 others 7 couples, of which 2 are incomplete. Subsidiary couples 
number about 20, of which 5 or 6 reach up to half or two-thirds the width of the 
principal mesenteries, the remaining couples narrow and hardly extending below the 
stomodeum. very tentacle has a swollen terminal battery similar in structure to those 
of F’. favus, its nematocysts I having each a spiral of about 33 turns. The stomodeum 
is oval in outline, its longer diameter measuring about 1°25mm. Nuclei are not thickly 
crowded in its ridges as is the case in F. favus; occasionally nematocysts I and III are 
present. The mesoglea is thickened in each ridge, the fibres passing from the former into 
the latter, lying close together in the middle of the central non-nucleated granular region. 


5 MATTHAI— RECENT COLONIAL ASTRAIDA 97 


In the ectoderm of the oral-disc nematocysts I are more frequently found than in the 
previous species, each with a spiral of about 20 turns; type II are rarely present. 

Filaments are attached to the principal mesenteries and to the wider subsidiary ones ; 
they are almost circular in section. A few nematocysts I are present in their straight 
regions, while IIb are frequently found in their coils. Below the stomodzal region of 
polyp the entoccelic pleats extend over almost the entire width of the mesentery, becoming 
broader towards the skeletal attachments. The endoderm of the oral-disc is never thicker 
than the overlying ectoderm, in parts only half the thickness of the latter; it has a 
columnar facies as the nuclei are aggregated along its margin where algze never occur ; 
the thin circular layer of muscle-fibres is well seen. Both tentacular and stomodeal 
endoderm are thin with few alge. Only in some of the mesenteries is the endoderm 
swollen behind the filaments, where occasionally nematocysts II are seen. The com- 
parative thinness of the endoderm and the scarcity of algze are distinctive of this species. 
Ova were present in some of the principal and subsidiary mesenteries in one or two 
longitudinal rows. _ 

Number of polyps examined, three, from a colony from Hulule, Maldives. 

-B. Corallum. In the Salomon specimens, as also in Klunzinger’s example of 
Prionastrea pentagona, up to 5 or 6 of the principal septa tend to become thicker, broader 

“and more exsert than the others; this is particularly evident in some elongated corallites 
and perhaps shows where division is ultimately to take place. This process is carried 
further in Gardiner’s type of Favia adduensis, in which many of the corallites appear 
either elongated or distorted, in all probability due to delayed fission, the diameters 
of the calices in such cases being about 6—7 and 4 mm. with 12—14 septa meeting the 
columella. The latter is a well-formed structure, either closely spongy or quite compact. 
The corallites towards the edges of this specimen are not elongated and closely resemble 
those of the Maldive examples of the present species. 

Ehrenberg’s type of Astr@a melicerum consists of three incrusting bits on a gastropod 
shell (Pl. 36, fig. 4); though somewhat defaced their specific characters may still be 
recognised. Similar to this is Klunzinger’s small figured example of Prionastrea 
‘pentagona. 

In the Paris Museum there are two large specimens named Prionastrea gibbosissima, 
Ed. and H., measuring 22 x 16 x 14 em. (loc.?) and 38 x 25 x 30 cm. (Red Sea), the former 
probably the type. Milne Edwards and Haime have also referred two specimens to 
Prionastrea melicerum, one small from Red Sea, the other large (20x19 x19 cm.) 
and with low humps from Seychelles (both re-named Prionastrea pentagona by 
Klunzinger); in the latter the calices are about 5 mm. in depth, columella distinct in 
most corallites, towards edges calices larger up to 6 mm. in diameter, columella rudimentary 
and paliform lobes conspicuous. Identical with these is another example from Seychelles 
named Prionastrea abdita, later referred by Klunzinger to Prionastrea pentagona and 
also Milne Edwards and Haime’s large figured type of Goniastrea rudis. A small piece 
named Astrangia asiatica, Mich., in Michelin’s collection from East Indies, also belongs to 
F. pentagona. 

From Quelch’s description of his new species Goniastrea laxa from Api, New 


SECOND SERIES—ZOOLOGY, VOL. XVII. 13 


98 PERCY SLADEN TRUST EXPEDITION 


Hebrides, it appears he was dealing with two specimens; of these only one (incrusting 
corallum, measuring 5°5 x 5 em.), presumably the smaller, was found in the British Museum. 
It is not in any case a Goniastrea, but comes nearest the present species. 

Of Plesiastrea haeckeli, Briigg., 1 have seen only two or three corallites, kept in the 
Berlin Museum; they are similar to those of Prionastraa melicerum, Ed. and H. 

Localities. Maldives: Hulule (6); Addu (5 small and 1 from 25 fms.) ; Suvadiva 
(1 small, 25 fms.). Minikoi (2). Seychelles (4). Saya de Malha (1 small, 29 fms.). 
Cargados (1 small, 30 fms.). Chagos, Salomon (7). ?Amirante (1, 16 fms.). Providence 
(1,29 fms.). Also from Red Sea (Ehrenberg, Milne Edwards and Haime, and Klunzinger), 
2New Hebrides (Quelch), ? East Indies (Michelin), ? (Esper). 


8. Fava ananas, Ellis and Solander. (Pl. 10, figs. 2 and 4; 28, fig. 7.) 


1786. Madrepora ananas, Ellis and Solander, Nat. Hist. Zooph., p. 168, pl. 47, fig. 6 (non Astrea ananas, 
Lamarck). f 

1857. Plesiastrea peront, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 492, pl. D7, fig. 3a and b. 

21866. Plestastrea indwrata, Verrill, Proc. Essex Instit., vol. v, part 3, p. 35, pl. 2, fig. 7. 


Corallum. Massive. Corallites circular or oval, with distinct rims. Walls about 
1—1°5 mm. thick, meeting in polygonal inter-corallite furrows. Calices 4°5—5 mm. in 
diameter, about 4 mm. in depth. 

Septa thickened towards walls, vertical, with entire or serrate edges, slightly rough 
sides, exsert to 1 or 1:25 mm., 21—24 in number; of these 8—11 meeting columella, each 
usually with a blunt somewhat thickened paliform lobe. An alternating cycle of 
rudimentary septa present. Costee of main septa conspicuous, with transversely extending 
granulations, those of rudimentary septa smaller ; the former meeting latter or the costee 
of the main septa of adjacent corallites in inter-corallite grooves. Columella low, loosely 
trabecular, +—4 width of calyx, sometimes quite rudimentary. Multiplication by fission. 

Polyps. (1) Both entoccelic (only over principal entocceles) and exoccelic tentacles 
present. (2) Stomodeeal ridges thicker than broad, with convex inner surfaces. (3) Seven 
principal couples of mesenteries. (4) In the stomodzal region of polyps about as 
many subsidiary couples of mesenteries as there are principal couples. (5) In the same 
region of polyp entoccelic pleats thick, usually unconstricted at their bases, undivided, 
extending over the greater part of width of principal mesenteries, broader and thicker in 
outer half of width; mesenterial mesoglea comparatively thick, usually stouter in its ~ 
outer half. (6) Mesenterial endoderm vacuolated, considerably swollen, of more or less 
uniform thickness along entire width of mesenteries. (7) Convolutions of mesenteries 
abundant from stomodeum to base of polyp massed in inter-mesenteric chambers, 
almost blocking the whole gastro-vascular cavity. 

Remarks. A. Polyps. These are circular or oval in shape and small in size. Four 
have each 7 principal couples of mesenteries, of which 3 are incomplete ; a smaller polyp 
has only 5 principal couples; about 10 subsidiary couples are present, 3 or 4 of which 
do not extend below the stomodzum. The tentacles are short, present over all the 
exocceles but only over the principal entocceles, the latter larger than the former; every 
tentacle has a swollen terminal battery containing a fair number of nematocysts I1b 


MATTHAI—RECENT COLONIAL ASTRAIDA 99 


in addition to closely packed nematocysts I, the larger ones having 40 to 50 turns of the 
spiral; sub-terminal batteries number 3 or 4. The stomodeeum is laterally compressed 
with its diameters in the retracted condition measuring 1°25 and °75 mm. ; owing to the 
radial contraction of the mesenteries some of the ridges are pulled outwards, appearing 
somewhat concave in transverse section. Small nematocysts I, similar to those in the 
oral-disc, are present in the stomodzal ridges, in which the mesoglea is not thickened. In 
the ectoderm of the oral-disc numerous mucous vacuoles are present ; small nematocysts I 
are fairly frequent, but II are scarce. 

Filaments are present on all the mesenteries ; nematocysts I occur in their straight 
regions as in the stomodzeal ridges, but have fewer coils ; where numerous, nematocysts III 
are arranged in rows, many with threads partly discharged; some of type II, with the 
axes usually bent a little, are also found. Convolutions of the mesenteries are pro- 
truded into the stomodeum. The endoderm is, as a rule, much vacuolated; in the 
oral-disc it is as thick as the ectoderm over it; in the tentacles it fills their lumina. 
The gastro-vascular cavity is blocked below the stomodeum owing to the swollen condition 
of the mesenterial and column-wall endoderm and to the abundance of the mesenterial 
convolutions. Algz are massed together in the endoderm of the oral-disc and tentacles, 
scarce in the mesenterial endoderm. Gonads were not present in any of the polyps. 
- Number of polyps examined, five, all from one colony from Dongonab, Red Sea. 

B. Corallum. Ellis and Solander’s figured type of Madrepora ananas is missing 
from the University Museum, Glasgow, but judging from the figure there is little doubt 1 
am dealing with a specimen similar to theirs. Since Madrepora ananas of Linnezeus is, 
according to Milne Edwards and Haime, a Cyathophyllid = Acervularia ananas (Corall. iti, 
p- 412), Ellis and Solander's specific name may be retained for the present species. 
Milne Edwards and Haime’s original of Plesiastrea peroni is missing, but their figure 
resembles my specimen, the only difference being that in the former the corallites appear 
to project more. 

Verrill’s figure of his species, Plesiastraa indurata, greatly resembles my type 
specimen, but his specimens have to be examined before the relationship of his species to 
the present can be finally settled. 

Localities. Red Sea (1 small). Also known from ? Australia (Milne Edwards and 
Haime), ? Loo Choo Islands (Verrill), ? loc. (Ellis and Solander). 


9. Favr4 taxa (Klunzinger). (PI. 24, figs. 5 and 6; 37, fig. 2.) 


1879. Orbicella laxa, Klunzinger, Korall. Roth. Meer., iii, p. 49, pl. 5, fig. 3, pl. 10, figs. 9a and 96 (non 
Orbicella laxa, Gardiner). 


Corallum. Massive, tending to be rounded off, sometimes incrusting. Peritheca 
vesicular, the dissepiments about °75 mm. apart. Corallites circular or laterally com- 
pressed, projecting about 1 mm., sometimes almost level, with distinct rims, usually about 
1—2 mm. apart, always separated by inter-corallite furrows. Calices 4—6 mm. in diameter 
depth 2—3 mm. 

Septa thickening towards walls, usually vertical, with toothed edges, crowded blunt 
spinules on sides, exsert to °5 or ‘75 mm., up to 37 in number, average 32; of these 


13—2 


100 PERCY SLADEN TRUST EXPEDITION 


12—17 meeting columella, each with a blunt paliform lobe and 3 or 4 blunt teeth above 
it, usually perforated behind paliform lobes. Subsidiary septa alternating with the 
principals, the broader ones curving towards and fusing with sides of principals. Exsert 
ends of septa arched, serrate. Costze with transversely extending granulations, usually 
meeting in inter-corallite furrows, tending to be united by transverse ridges where they 
meet. Columella well developed, trabecular, about 4 width of calyx. 

Multiplication by equal or sub-equal fission. 

The corallites towards the edges project up to 3°5 mm. and increase in diameter from 
calicular margins (6 or 7mm.) to bases (9 mm.). They are wider apart, up to 6mm., 
with calices shallower, septa sloping, thicker and rougher, and with usually an alternating 
cycle of smaller costze, hardly represented by septa within. 

The skeletal facies of this species has a general resemblance to that of Hchinopora 
gemmacea. 

Polyps. (1) Both entoccelic and exoccelic tentacles present. (2) Stomodzal ridges 
narrow, thicker than broad, as in Favia hululensis, inner surface slightly convex or flat. 
(3) About nine principal couples of mesenteries. (4) In the stomodeeal region of polyp 
about as many subsidiary couples of mesenteries as there are principal couples. (5) In 
the stomodeeal region of polyp, entoccelic pleats thick, undivided, slightly constricted at 
their bases, narrow on outer half of pleatal region, directed obliquely towards the stomo- 
dzeum and extending over outer halves of principal mesenteries ; pleatal region of mesoglea 
thicker than non-pleatal region, inner half of former stouter than its outer half and with 
broader pleats. (6) Mesenterial endoderm thicker in pleatal region. (7) Convolutions 
of mesenteries not abundant, but extending to base of polyp. 

Remarks. A. Polyps. The polyps of this species were not in a good condition for 
detailed study. 9 principal couples of mesenteries were present in each of 3 polyps, 7 and 
8 couples in two others; of these, 1 to 5 couples were incomplete; subsidiary couples 
numbered 8 to 10. The tentacles were short, with well-developed terminal batteries 
of the usual structure and with not more than 4 or 5 sub-terminal ones. The stomodeeum 
is oval in transverse section, its diameters in the retracted condition being 1°5 and 1 mm. ; 
owing to the radial contraction of the mesenteries some of the ridges appear concave 

“in transverse section ; the mesoglea is not thickened in the ridges. The endoderm in the 
tentacles and stomodzeal wall is quite thin. Gonads were not present in any of the polyps. 

Number of polyps examined, five, 3 from one colony and 2 from a second, both 
from Dongonab, Red Sea. 

B. Corallum. Klunzinger’s type of Orbicella laxa is a small specimen measuring 


d 


10x9x4cem.- In the Hofmuseum, Vienna, are three specimens of the “ Pola” expedition, 
referred by Marenzeller to Orbicella laxa, Klunz. Two of these are large examples, 
one from Sherm Abban (No. 15916) measuring 19x 19x15 cm., the other from Koseir 
(No. 2276) measuring 30 x 16 x 8 cm. 


Locality. Red Sea (5). Not recorded from elsewhere. 


10. Favra uirsura (Milne Edwards and Haime). (PI. 24, figs. 7 and 8.) 


1816. Astrea dipsacea, Lamarck, Hist. Anim. sans vert., ii, p. 262. 


MATTHAI—RECENT COLONIAL ASTRAIIDA 101 


1833. Astrea dipsacea, Quoy and Gaimard, Voy. !’Astrol., Zooph., p. 210, pl. 17, figs. 1—2. 

1834. Astrea dipsacea, Ehrenberg, Corall. roth. Meer., p. 97. 

1850. Acanthastrea hirsuta, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 145. 

1850. Acanthastrea spinosa, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 145. 

1850. Acanthastrea brevis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 146. 
1850. Acanthastrea grandis, Milne Kdwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 146. 

1857. Acanthastreea hirsuta, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 502, pl. D5, fig. 4. 

1857. <Acanthastrea spinosa, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 502. 

1857. Acanthastrea brevis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 503. 

1857. Acanthastrea grandis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 504. 

1879. Acanthastrea hirsuta, Klunzinger, Korall. Roth. Meer., ii, p. 42, pl. 5, figs. 1 and 2. 

1886. Acanthastrea irregularis, Quelch, Reef Corals, Challenger Reports, Zool., vol. xvi, part xlvi, p. 102, 

pl. 4, figs. 2—2a. 

1899. Prionastrea hirsuta, Gardiner, Proc. Zool. Soc. London, p. 760. 

1904. Acanthastrea hirsuta, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 784, pl. 59, fig. 6. 

Corallum. Partly incrusting, partly massive. Corallites polygonal. Walls fused, 
the inter-calicinal partitions thus formed up to 9 mm. in thickness, average about 4 mm., 
and dense in section. Calices varying much in size, average diameters 18 x 13 mm. (the 
largest calyx measuring 25x11 mm.), depth up to 15 mm., usually about 11 mm., 
decreasing in width from calicular margins to bases. 

Septa thick, swollen in thecz, sloping, with strongly-toothed edges, almost smooth 
-sides, exsert to 2—3mm., 35—50 in number of these 13—18 meeting columella, with 7—12 
bluntly-pointed more or less triangular hollow teeth, the first upright, swollen at its 
base, forming the most exsert part of each septum ; teeth about 2 mm. apart at the upper 
part, 1 mm. towards columella, the lowest tooth usually swollen. Exsert ends of septa of 
adjacent corallites meeting in notches or alternating, hence inter-calicinal walls appearing 
furrowed above. The broader subsidiary septa curving towards and fusing with sides of 
principals. Columella formed of intercrossing septal teeth, mostly upright. Calicular 
dissepiments passing from wall to columella at an angle of 45°, about 2 mm. apart. 

Multiplication by unequal fission towards calicular margins. 

Polyps. A small specimen in spirit which, on the authority of Prof. Gardiner, was 
taken from the figured example of var. megalostoma, had to be entirely decalcified, as it 
possessed only two polyps. Owing to the lack of sufficient material and as the sectioning 
of the two polyps was not quite successful, only a short description of the anatomy is given 
below: Both entoccelic and exoccelic tentacles present, each with a swollen terminal 
battery having the usual structure (most of the I1b nematocysts are in their earlier 
stages of development, the axes being very short and their nuclear origin quite apparent) 
and up to eight sub-terminal batteries with vacuoles in their sides ; nematocysts I with 
about 40 turns of the spiral. Tentacular endoderm vacuolated with algze scarce, hence 
almost transparent. In vertical sections the tentacles greatly resembling those of Fama 
doreyensis, Ed. and H. Stomodeal ridges thicker than broad, with convex imner surfaces. 
In the larger polyp 13 principal couples of mesenteries and 23 subsidiary couples are 
present, in the smaller polyp 12 and 5. In the stomodzal region of the polyp the ento- 
celie pleats thick, unconstricted at their bases, sometimes sub-divided, always close 
together and directed obliquely towards the stomodeum, and extending to not more than 
the middle of a principal mesentery, better developed in its outer one-third ; very narrow 


102 PERCY SLADEN TRUST EXPEDITION 


exoccelic pleats for a short distance from the stomodzal attachment. Mesenterial endo- 
derm quite thin and of more or less uniform thickness along the entire width of the 
mesentery. Convolutions of the mesenteries abundant below the stomodzeum to the 
base of the polyp, being massed in the inter-mesenteric chambers. Filaments are found 
on all the mesenteries. In the larger polyp ova present, in up to five rows in every 
mesentery, with the endoderm granular and swollen on either side of them. 

Number of polyps examined, two, from a specimen from Hulule, Maldives. 

Remarks. Milne Edwards and Haime placed Astrea dipsacea, Audouin (Expl. 
Savignyi, Deser. Egypte, xii. p. 57, pl. 5, fig. 3, 1809), under Fava savignyi, Ed. and 
H. I have not been able to see this example. If Klunzinger is correct in identifying it 
with Acanthastrea hirsuta, then the present species would be named F. dipsacea. 

Of the five specimens now in the Berlin Museum referred by Ehrenberg to Astrea 
dipsacea, only one (later re-named Acanthastrea hirsuta var. megalostoma by Klunzinger) 
comes under the present species. Klunzinger’s figured example of var. megalostoma is not 
in the Berlin Museum. 

Milne Edwards and Haime named 7 specimens (8 large, one with the microstoma 
facies, and 4 small, all from Red Sea) Acanthastraa hirsuta, Ed. and H., 4 speci- 
mens (2 large and 2 small, from unknown locality) Acanthastrea brevis, Ed. and H., 
and 4 large specimens (3 from Red Sea and 1 from unknown locality) Acanthastrea 
grandis, Ed. and H.; I could not find any differences separating these three species. 
They also named two very small specimens from Tongatabou Acanthastrea spinosa, Ed. 
and H., these doubtless also belonging to the present species. Lamarck’s original of 
Acanthastrea dipsacea is similar to the megalostoma examples. A specimen from Red 
Sea named Celoria botte, Ed. and H., also comes under the present species. Quelch’s 
type of his new species, Acanthastrea wrregularis, is a small, thin, convex specimen from 
Kandavu, Fiji, measuring 7 x 7°5 cm., in no way different from the hirsuta examples. 

Localities. Maldives, Hulule (1). Red Sea (1). ?Hllice Islands, Funafuti (a frag- 
ment). Also from Tongatabou (Milne Edwards and Haime), Fiji (Quelch). 


11. Favia acroporé (Linneeus). (Pl. 25, figs. 1 and 3; 26, fig. 4; 33, fig. 1.) 
1767. Madrepora acropora, Linnaeus, Syst. Nat., edit. 12, p. 1276.! 
1788. Madrepora acropora, Gmelin, Linn. Syst. Nat., edit. 13, p. 3767. 
1797. Madrepora acropora, Esper, Forts. Pflanz., p. 21, pl. 38, figs. 1 and 2. 
1815. Favia acropora, Oken, Lehrb. Naturg., i, p. 68. 
1816. Astrea pleiades, Lamarck, Hist. Anim. sans vert., ii, p. 261; 2° édit., p. 408 (non Madrepora pleiades, 
Ellis and Solander). 
1850. Astrea pleiades, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 105. 
1850. Parastrea lobata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 171. 
1857. Favia lobata, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 434, pl. D8, fig. 3. 
1857. Heliastrea acropora, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 477. 
1879. avia lobata, Klunzinger, Korall. Roth. Meer., ii, p. 31, pl. 3, fig. 9, pl. 10, fig. 8. 
1899. Astrea lobata, Gardiner, Proc. Zool. Soc. London, p. 749. 
1899. Orbicella acropora (pars), Gardiner, Proc. Zool. Soc. London, p. 752. 
1899. Orbicella orion, Gardiner, Proc. Zool. Soc. London, p. 752. 
1906. Orbicella lobata, Marenzeller. Exp. “ Pola.” Roth. Meer., Riffkorall., Zool. Ergeb. Wien xxvi, p. 61. 


Corallum. Massive, rising into hillocks, which usually decrease in thickness from 
their summits. Peritheca vesicular or compact. Corallites circular, dividing ones 


MATTHAI—RECENT COLONIAL ASTRAIID A 103 


elongated, slightly projecting, with distinct rims, usually ‘5—1 mm. apart (towards edges 
of corallum up to 2°5 mm.), separated by somewhat polygonal furrows. Calices varying 
little in size, 2 or 2°5 mm. in diameter, shallow, not more than 1 mm, 

Septa thickening towards walls, with vertical ragged edges, spinulose or coarsely 
rough sides, ‘5—°75 mm. exsert, 14—24 in number, 7—13 meeting columella, narrow,— 
some of these sometimes thicker than the others—mostly with rough, blunt paliform lobes, 
perforated behind. Exsert ends of septa arched. Costze conspicuous, spinulose, usually 
stopping at inter-corallite grooves, sometimes those of adjacent corallites meeting in 
notches. A cycle of very narrow septa alternating with the principals and subsidiaries 
usually without costz. Columella varying in texture, being compact or spongy, and $—4 
width of calyx, in some corallites quite rudimentary. 

Multiplication by unequal or equal fission. 

The hillocky mode of growth and the shape of the hillocks are quite characteristic of 
this species. It is also comparatively compact, varying only within narrow limits. 

The polyps were all in a badly-preserved condition; two were sectioned from a 
specimen from Rotuma, in one of them 6 principal couples of mesenteries and 9 subsidiary 
couples were counted. Tentacles were present over all the entocceles and exocceles. 


Remarks. Corallum. In the Paris Museum there are 2 specimens from the Indian 
Ocean, referred by Milne Edwards and Haime to Heliastraa acropora, one a broken off 
hump (15 x 11 em.), the other a large hillocky specimen (23 x 19 x 16 em., Pl. 26, fig. 4), 
and 4 examples from Red Sea (all humps, the largest 10x 7°5 em.) to Kavia lobata. In 
the Berlin Museum are small examples referred by Klunzinger to Pavia lobata, 2 of 
which he has figured. Marenzeller has assigned 3 specimens of the ‘“ Pola” expedition to 
Orbicella lobata, one of which, measuring 48 x 37 x 26 cm., is an excellent example with 
erowded hillocks, the latter up to 10 cm. high. 


Localities. Chagos: Salomon (13); Egmont (2). Amirante (1 from over 25 fms.). 
Seychelles (1). Aldabra (1). Funafuti (3). Rotuma (1). Also from Red Sea (Milne 
Edwards and Haime, Klunzinger and Marenzeller), ? loc. (Esper, Linneeus and Oken). 


12. Favra verstpora (Lamarck). (Pl. 28, fig. 3; 25, figs. 5, 6 and 9; 87, fig. 3.) 


1816. Astrea annularis, var. 2, Lamarck, Hist. Anim. sans vert., ii, p. 259; 2° édit., p. 405 (non Madrepora 
annularis, Ellis and Solander). 

1816. Astrea versipora, Lamarck, Hist. Anim. sans vert., ii, p. 264; 2° édit., p. 414. 

1830. Dzupsastrea versipora, Blainville, Dict. Sci. Nat., lx, p. 338; Man. d’actinol., p. 373. 

1833. Astrea annularis, Quoy and Gaimard, Voy. |’Astrol., Zooph., p. 210, pl. 17, figs. 1718. 

1850. Astrea laperousiana, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 101. 

1850. Astrea annuligera, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 103. 

1850. Plesiastrea versipora, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 118 (non Orbicella 
versipora, Gardiner). 

11850. Plesiastrea quatrefagiana, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 119. 

1857. Heliastrea laperouseana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 460. 

1857. Heliastreea annuligera, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 471. 

1857. Plesiastrea versipora, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 490, pl. D7, fig. 5. 

11857. Plesiastrea quatrefaugesana, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 491. 

1899. Orbicella acropora (pars), Gardiner, Proc. Zool Soc, p. 752. 

1899. Orbicella annuligera, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 774, pl. 63, fi 


104 PERCY SLADEN TRUST EXPEDITION 


Corallum. Incrusting. Peritheca compact. Corallites circular, unless dividing 
when elongated, projecting up to 1°5 mm., usually to ‘75 or 1 mm., up to 1°5 mm. apart, 
usually closer but separated by inter-calicinal grooves. Calices 3—4 mm. in diameter, 
1—2 mm. in depth. 

Septa comparatively thin, vertical (edges somewhat concave in the shallower calices), 
with denticulate edges, spinulose sides, exsert to ‘5—’75 mm., 16—25 in number; 
12—18 septa meeting columella, most of these with rough blunt paliform lobes (usually 
a ring of 12), which in shallower calices reach to the calicular margins ; subsidiary septa 
often curving towards and fusing with sides of principals. Exsert ends of septa arched, 
denticulate. Generally a cycle of very narrow septa alternating with the principals and 
subsidiaries, with or without corresponding costz. Costz conspicuous with transversely 
extending granulations, stopping at inter-corallite grooves or meeting those of adjacent 
corallites in notches. Columella spongy or dense, 1—4 width of calyx. 

Multiplication by budding (small corallites intercalated between larger ones) and 
by fission. 

The polyps were all badly preserved. Though budding undoubtedly takes place 
among them, neither of the polyps sectioned from a specimen from Rotuma had any 
directive couples of mesenteries ; one of them had 6 principal couples of mesenteries and 
19 subsidiary couples. In its stomodzeal region the entoccelic pleats were narrow and did 
not extend beyond the outer one-third of each principal mesentery. Tentacles were 
present over all the entocceles and exocceles. 


Remarks. Corallum. In the Paris Museum is a small incrusting specimen 
(6°5 x 6 cm.) from Australia, Milne Edwards and Haime’s Heliastrea annuligera (Pl. 37, 
fig. 3), with which Gardiner’s figured examples of Orbicella annuligera are identical. ‘Two 
other specimens are named by them Plesvastrea versipora, Kd. and H., one from Indian 
Ocean measuring 10x 5x 3°5 cm. (PI. 23, fig. 3), the other smaller from an unknown 
locality ; in the latter and on one side of the former the corallites are closer and with 
polygonal inter-corallite furrows. Plesiastrea quatrefagesana, Ed. and H., is represented 
by a small specimen with rudimentary paliform lobes from an unknown locality. In all 
these specimens multiplication is mainly by budding. Milne Edwards and Haime’s types 
of Heliastrea laperouseana are two small incrusting specimens from Vanikoro, which 
perhaps belong here (Pl. 25, fig. 9). 

A badly-cleaned fragment in the “ Challenger’ 
from Bermuda, referred by Quelch to Astrea coarctata (Duchassaing and Michelotti), 
comes nearest the present species. 

Localities. Maldives, Goidu (2). Minikoi (3). Seychelles (3). Chagos, Salomon 
(9). Rotuma (1). Also from Australia and Vanikoro (Milne Edwards and Haime) ; 
? Bermuda (Quelch). 


’ 


collection, with five or six corallites 


13. avis waxayana (Gardiner). (Pl. 25, fig. 4.) 


1899. Orbicella wakayana, Gardiner, Proc. Zool. Soc. London, p. 753, pl. 49, fig. 2. 
1899. Orbicella versipora, Gardiner, Proc. Zool. Soc. London, p. 753 (non Plesiasirea versipora, Milne Edwards ~ 
and Haime). 


MATTHAI—RECENT COLONIAL ASTRAID A 105 


1899. Orbicella heliopora, Gardiner, Proc. Zool. Soc. London, p. 756, pl. 49, fig. 4 (non Heliastreea heliopora, 
Milne Edwards and Haime). 
1899. Orbicella solidior, Gardiner, Proc. Zool. Soc. London, p. 756 (non Heliastrea solidior, Milne Edwards 


and Haime). 
1899. Orbicella funafutensis, Gardiner, Proc. Zool. Soc. London, p. 756, pl. 49, fig. 5. 

Corallum. Usually incrusting, sometimes massive. Corallites circular, oval or 
sometimes laterally compressed, projecting up to 3°5 mm., usually 1°5 or 2 mm., in some 
specimens with one side higher than the other as in F. doreyensis. Corallites usually 
closely arranged but always with inter-corallite grooves, towards edges up to 3 mm. apart. 
Calices about 4—5 mm. in diameter, when laterally compressed up to 8 x 5 mm., depth 
3—o mm. 

Septa thickening towards theca, vertical or slightly sloping, with toothed edges, 
spinulose or coarsely rough sides, slightly exsert, 17—25 in number, 10—18 meeting 
columella, each usually with a blunt paliform lobe and perforated behind it; 3—7 of the 
principal septa, as a rule, somewhat broader, thicker and rougher than the others (the 
degree of thickness varying in different examples), with conspicuous paliform lobes, and 
exsert up to 1 or 15mm. The subsidiary septa often curving towards and fusing with 
‘sides of principals. Exsert ends of septa arched, rarely flat, serrate. An alternating 
cycle of very narrow septa present, with smaller coste. Costze conspicuous, toothed, 
usually meeting in inter-corallite grooves. Columella formed of septal trabeculee, varying 
in degree of development, from 4 to 4 width of calyx and from being loosely spongy to 
compact. 

Multiplication by budding—the small corallites formed in this manner are found 
intercalated between the ordinary ones—rarely also fission. 

Towards edges corallites are wider apart, up to 3 mm., and shallower. 

The skeletal facies of the present species has a general resemblance to that of 
F. lasa. 

The polyps of this species were in a badly-preserved condition. In three polyps 
sectioned (taken from a specimen from Rotuma resembling Gardiner’s example of 
Orbicella wakayana). tentacles were present over all the entocceles and exocceles ; 8—10 
couples of principal mesenteries and 11—14 subsidiaries were counted. Entoccelic pleats 
were broader than in F. versipora and extended to about the middle of each principal 
mesentery. 

It is likely that the present species may have been previously recorded by Dana, but 
this point cannot be settled till Dana’s Astreid types are examined. A fairly large 
Specimen from Kandavu, Fiji, in the “Challenger” collection, which Quelch has referred 
to Plesiastrea indurata, Verrill, is identical. with Gardiner’s examples of Orbicella 
wakayana. 

Localities. Funafuti (8). Fiji, Wakaya (3). Rotuma (1). Known only from the 
Pacific Ocean. 


14. Faria sotrpror (Milne Edwards and Haime). (PI. 25, fig. 8; 28, fig. 1.) 


1850. Astrea solidior, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 102. 
1857. Heliastreea solidior, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 466. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 14 


106 PERCY SLADEN TRUST EXPEDITION 


1899. Orbicella curta, Gardiner, Proc. Zool. Soc. London, p. 754. 

1899. Orbicella coronata, Gardiner, Proc. Zool. Soc. London, p. 754. 

1899. Orbicella rotumana, Gardiner, Proc. Zool. Soc. London, p. 755, pl. 49, fig. 3. 

21904. Orbicella curta, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 776. a“ 


Corallum. Incrusting. Corallites circular or oval, projecting more than in 
F, wakayana, average 2°5 or 3 mm., with conspicuous rims and separated by deep grooves. 
Calices from 5—8 mm. in diameter, depth about 5 mm. 

Septa thickening towards theca, almost vertical, with toothed edges, spinulose or 
granular sides, exsert to about °75 or 1 mm., 20—28 in number; 9—15 septa meeting 
columella, average 10—13, each usually with a palitorm lobe behind which it is perforated, 
6—8 of the principal septa broader and thicker than in F. wakayana (this being a 
marked feature of the species) and exsert up to 2mm. An alternating cycle of very 
narrow septa present, with smaller costee. Costze conspicuous, more or less flat ridges, 
varying in thickness in accordance with their septa, with transversely extending 
granulations usually meeting in inter-corallite grooves. Columella poorly developed, 
spongy, +—4 width of calyx. 

Multiplication as in F. wakayana by budding, rarely aided by fission. 

This species is a larger and thicker edition of /. wakayana. 

Milne Edwards and Haime’s type of Heliastrea solidior is a small specimen 
(7x45 x5 cm., Pl. 25, fie. 8) from Tongatabou, similar to Gardiner’s example of Orbicella 
coronata. 

Localities. Funafuti (2). Rotuna (1). ?Maldives, Mahlos madulu (a fragment). 
Also from Tongatabou (Milne Edwards and Haime). 


15. avira waticora (Khrenberg). (PI. 26, figs. 3 and 5—7.) 


1834. Astrea halicora, Khrenberg, Corall. roth. Meer., p. 96 (non Prionastrea halicora, Milne Edwards and 
Haime). 

1879. Goniastrea halicora var. acuta, Klunzinger, Korall. Roth. Meer., iii, p. 33, pl. 4, fig. 1. 

889. Favia ehrenbergi, Ortmann, Steinkorall. siid. Ceylons, Zool. Jahrb., iv, p. 526 (non Favia ehrenbergt, 
Klunzinger). 

1904. Favia halicora, forma obtwsa, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 771. 

1904. Prionastrea crassior, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 786, pl. 64, figs. 45 
and 46 (non Prionastrea crassior, Milne Edwards and Haime). 


Corallum. Incrusting or massive. Corallites polygonal, sometimes oval or circular, 
level or projecting to 1—2 mm., usually separated by polygonal furrows. Walls ‘75—2 mm. 
in thickness. Calices oval or circular, sometimes polygonal, with diameters 9—11 mm. 
and 7—8 mm. (when circular 6—8 mm.), depth 3—6 mm.; much shallower towards 
edges of corallum. 

Septa thin in calices, thickened or swollen in theca, up to 1 mm., with toothed edges, 
slightly rough to spinulose sides, exsert up to 1 mm., 25—40 in number, average about 30; 
of these up to 20 meeting columella, usually 15—17, with upper two-thirds or half narrow 
and usually vertical; on each septum 6—10 teeth, longer and closer together towards 
columella ; lower one-third or half of principals much broader, these or the ring of last 
septal teeth forming a pali-crown. Septa oblique in some of the calices—particularly in 
the shallower calices, Subsidiary septa usually curving towards and fusing with sides of 


SS 


MATTHAT—RECENT COLONIAL ASTRAIDA 107 


principals, a group of 1—4 meeting a principal septum. Coste: conspicuous when inter- 
eorallite grooves are present, with transversely extending granulations ; alternating with 
these a cycle of smaller coste represented by very narrow septa in the thin-walled 
eorallites; costee of neighbouring corallites meeting in inter-corallite grooves, directly 
continuous over walls when grooves are absent. Columella formed of closely-twisted 
trabeculee, up to 4 width of calyx, usually +—1 width. 

Multiplication by very unequal fission towards calicular margins. When walls are 
formed round the small fission-products, they simulate buds. 

This species has a variable skeletal facies. Between the thin-walled polygonal level 
eorallites without surrounding vrooves and the oval or circular thick-walled corallites with 
distinct furrows there are many intermediate stages. (Compare PI. 26, figs. 5—7.) 

Gardiner’s example (2) of Favia halicora, forma obtusa, is missing. 

Some of my examples from Salomon show a certain resemblance to Havia abdita in 
having an alternating cycle of very narrow septa in every corallite and in fission being 
marginal. The dredged specimen from Peros, Diamant, has further the hillocky mode of 
growth so characteristic of Pavia abdita. 

The two large specimens from Aldabra are ‘dantitea! with Khrenberg’s type of Pavia 
halicora (= Gomastrea halicora var. acuta, Klunz.). 

In the two small specimens referred by Gardiner to forma acuta, Klunz., the inter- 
ealicinal walls are thinner (about 1 mm.), the calices smaller (about 6 mm. in diameter), and 
the total number of septa is less (22—26) than in the large specimen which he assigned to 
obtusa. Nevertheless, they belong to the present species. 

Ehrenberg’s original of Astrea halicora, which Klunzinger re-described and figured 
as the type of Goniastrea halicora var. acuta, is a large example (Mus. No. 733) 
measuring 25 x 21 x 16 cm., in most parts without inter-corallite grooves. Identical with 
this are my two large examples from Aldabra. The specimen from Ceylon which Ortmann 
referred to Favia ehrenbergi measures 23 x 16 x 10 cm. 

Localities. Maldives: Hulule (1); Goidu (1); Turadu (1). Minikoi (3). Chagos: 
Salomon (9); Egmont? (1); Peros, Diamant (1 from 15 fms.). Aldabra (2. from lagoon 
reefs). Ccetivy? (1). Singapore (1). Also from the Red Sea (Ehrenberg) and Ceylon 
(Ortmann). 


16. Favra Hompront (Rousseau). (Pl. 26, figs. 1 and 2; 38, fig. 2.) 


1854. Parastreea hombronii, Louis Rousseau, Voy. au pole Sud de Dumont-d’Urville, Zool., v, p. 122, Zooph. 
pl. 28, fig. 3. 

1857. Favia hombroni, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 435. 

1866. Astrea (Favia) hombroni, Verrill, Proc. Essex Inst., v, p. 33. 

1904. Favia hombroni, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 771, pl. 62, fig. 27. 


Corallum. Massive, often becoming rounded off. Peritheca more or less compact. 
Corallites polygonal, close together, with margins very thin when distinct, otherwise 
inter-calicinal walls flat. Calices with diameters about 3—4 mm. and 2°5—3°5 mm., 
depth 2—2°5 mm. Calicular dissepiments about 1°25 mm. apart. 

Septa of even thinness along their entire breadth, with vertical, entire or finely- 

142 


108 PERCY SLADEN TRUST EXPEDITION 


serrate edges, smooth or slightly rough sides, 17—24 in number, average about 20; 
5—10 septa meeting columella, usually 6—8, exsert about -75 mm., most of them with 
slender paliform lobes. Most of the subsidiary septa narrow. Usually an alternating 
cycle of rudimentary septa present, recognisable only with a lens. Septa continuous from 
corallite to corallite, their exsert ends being flat or slightly arched, giving a characteristic 


appearance to the corallum. Columella usually rudimentary, formed of a few septal 
trabeculz. 


Fission equal or sub-equal. 

A species showing comparatively little variation of the corallum. 

Milne Edwards and Haimes’ type of Favia hombroni from “Oceanie?” is a large 
specimen measuring 23 x 16 x 11 cm., in every respect resembling my examples. 

Localities. Maldives, Hulule (1). Minikoi (1). Chagos: Salomon (8). Also known 
from Oceania (Milne Edwards and Haime), ? Sandwich Islands (Verrill). 


17. #avra vasra (Klunzinger). (Pl. 27, figs. 3, 5 and 6.) 


1879. Goniastrea halicora var. obtusa, Klunzinger, Korall. Roth. Meer., iii, p. 33, pl. 4, fig. 2, and pl. x, 
figs. 3a and 6. 


1879. Prionastrea vasta, Klunzinger, Korall. Roth. Meer., iii, p. 38, pl. 4, figs. 8 and 12, pl. 10, figs. 4a and 6. 


Corallum. Massive, more or less flat. Corallites polygonal, usually hexa- or 
pentagonal, with walls of neighbouring ones fused, the inter-calicinal partitions thus 
formed 1°5—2°5 mm. in thickness. Calices widest at the calicular openings, with diameters 
up to 22 and 15 mm., depth up to 13 mm. 

Septa sloping or with upper two-thirds or halves vertical and narrower, those of 
adjacent corallites continuous over inter-calicinal walls, being exsert to 1 or 1°5 mm., with 
edges well toothed, sides rough or sometimes smooth, up to 55 in number (average about 
40); of these up to 24 meeting columella (average about 20), each with 8—10 teeth 
becoming longer towards columella, the lower somewhat broader parts of the septa with 
the longer teeth sometimes simulating a pali-crown. The subsidiary septa often curving 
towards and fusing with sides of principal septa, up to 3 or 4 meeting one of the latter. 
An alternating cycle of very narrow septa invariably present, those of neighbouring 
corallites being continuous over inter-calicinal walls; hence from surface a characteristic 


alternating appearance of high and low septal arches. Columella formed of closely-twisted 
trabecule, often rising into points. 


Multiplication by marginal fission. 

Klunzinger has described two varieties of Prionastrea vasta, In one (Pl. 4, fig. 12, 
type measuring 8x7 x4'5 cm.) the corallites are deep, with their diameters not very 
unequal, the septa somewhat thin and the columell well developed. Agreeing with this 
are two of my specimens from Salomon (the larger measuring 13 x 8 x 4m.) with calices 
about 20x15 cm. wide and 10—12 mm. deep (Pl. 27, fig. 5). These resemble Pavia 
abdita (Ell. and Sol.) in (1) marginal fission, (2) the subsidiary septa usually fusing with 
the sides of the principal septa in groups of up to 3 or 4, (3) an almost equal number 
of septal teeth of similar appearance to F. abdita, (4) the invariable presence of an 
alternating cycle of very narrow septa, and (5) the columellz formed of closely-twisted 


MATTHAI—RECENT COLONIAL ASTRAIDA 109 


trabecule ; but the corallites are much larger than in F. abdita, the septa thicker and 
further apart, and the alternating septa meet their fellows over the walls and not the 
main septa of the adjacent corallites. 

Klunzinger’s var. superficoalis (Pl. 4, fig. 8, type measuring 9x 5°5 x5 em.) has 
shallow corallites about twice as long as broad, average diameters 20 and 10 mm., inter- 
ealicinal walls and septa thicker, and the columelle either rudimentary or absent. This 
variety approaches Fama hirsuta (Ed. and H.) in the following respects: (1) heavy 
corallum, presenting a similar appearance in transverse section, (2) the inter-calicinal 
walls, septa and septal teeth thickened but not to the same extent (septal sides almost 
smooth), (3) the corallites with similar shape, being widest at the calicular openings and 
gradually narrowing towards the bases. Resembling Klunzinger’s type is one of my 
specimens from Egmont, Chagos, measuring 23 x 13x 11lem. (Pl. 27, fig. 3), but in the 
latter the columelle are well formed, of twisted septal trabecule. 

Klunzinger’s var. obtusa of Gonmastrea halicora (type measuring 18 x 10 x 4 cm.) 
has the same facies as his Prionastrea vasta (Pl. Iv, fig. 12), but has a somewhat thinner 
build and smaller corallites. Resembling this type are two of my specimens, with 
inerusting coralla: one from Aldabra (11°5 x 10°5 cm., Pl. 27, fig. 6), which has thin 
_sharp-ridged inter-calicinal partitions (not more than | mm. in thickness), calices about 
. 15x 10mm. wide by 8 or 9mm. deep, with more or less distinct pali-crowns; the other, 
from Salomon, smaller, but with thicker walls and septa. 

In the Paris Museum are two large specimens of Prionastrea vasta, Klunz., from 
Koseir, representing Klunzinger’s two varieties, and also a good example of Goniastrea 
halicora var. obtusa from the same locality. 

Localities. Chagos: Salomon (3); Egmont (1). Aldabra (1). Previously recorded 
only from the Red Sea. 


18. Favzra compianara, Ehrenberg. (Pl. 80, figs. 1—3.) 


1834. Havia complanata, Ehrenberg, Corall. roth. Meer., p. 93. 

1834. Astrea tesserifera, Ehrenberg, Corall. roth. Meer., p. 97. 

11846. Astrea tesserifera, Dana, Expl. exp. Zooph., p. 248, pl. 13, fig. 9. 

1850, Prionastrea michelini, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 132. 
1851. Prionastrea tesserifera, Milne Edwards and Haime, Pol. foss. terr. palzoz., etc., p. 102. 
1857. Prionastrea tesserifera, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 517. 

1879. Prionastea tesserifera, Klunzinger, Korall. Roth. Meer., iii, p. 37, pl, 4, fig. 9. 


Corallum. Incrusting, following irregularities on surface of attachment, often rising 
into hillocks, which usually end in ridges with corallites on either side of each such ridge 
from light to heavy, and with a thorny appearance, generally distorted. Corallites polygonal, 
on the hillocks, with one side much higher than the other, in the narrow valleys with 
a tendency to be drawn out, more regular in shape on the level regions. Inter-calicinal 
walls varying from very thin and sharp to 5 or 6mm. broad, Calices with diameters up 
to 18 by 12 mm., depth up to 10 mm. 

 Septa varying considerably in thickness, sloping or with their upper two-thirds 
vertical and narrower, those of adjacent corallites usually continuous over the inter- 
calicinal partitions, exsert to ‘5 or 1mm., septal edges and exsert ends well toothed, sides 


110 PERCY SLADEN TRUST EXPEDITION 


spinulose, up to 55 in number, average about 40; of these 12—18 meeting columella, 
usually 15—17, each with up to 10 spinulose solid teeth, usually becoming longer towards 
columella. The subsidiary septa often curving towards and fusing with sides of principals 
near to columella, up to 3 or 4 on one side of a principal septum. An alternating cycle of 
very narrow septa present, continuous over inter-calicinal walls as in F. vasta (better seen 
over the walls than within the corallites) and alternating with exsert ends of the main 
septa. When the inter-calicinal walls are broad, the continuous exsert ends of both cycles 
are sometimes connected by an incomplete narrow ridge or appear grooved along the 
middle of the walls. Columella formed of septal trabeculze, loosely spongy or closely 
twisted, rising into points, about 1—4 width of calyx. Multiplication by marginal 
fission. 


On level regions and especially towards edges the corallites are more regular in shape, 
hexa- or pentagonal, calices oval or circular but smaller in size, about 10 mm. in diameter 
and shallower (4 or 5mm. in depth). The inter-calicinal walls are thickened to about 
2°5 or 3mm. and appear furrowed; the septa are also thickened, especially towards the 
walls, with the teeth reduced to 3 or 4 in number but swollen at their bases, the lowest 
ones becoming upright and simulating a pali-crown ; the columellz are formed of inter- 
lacing septal teeth. These regions of the corallum vary towards the hirsuta facies, 
but the corallites are much smaller and the corallum much less massive. Ehrenberg’s 
type of Favia complanata, which is an edge-piece, has exactly the same appearance. 

This species is variable to even a greater extent than F’. abdita, all the stages from 
a light corallum with thin walls and septa to a heavy corallum with thick walls and septa 
being present. It resembles 7. abdita in (1) having the same type of hillocky growth, 
with distorted corallites, (2) fission being marginal and very unequal, (3) up to 3 or 4 
subsidiary septa fusing with one side of a principal septum ; but differs from it in that the 
septa are further apart and less in number, the corallum more spiny and on the whole 
having a thicker facies. 

The present species also approaches /’. vasta (Klunz.) in that the two cycles of septa 
are continuous from corallite to corallite over the intervening walls, the consequent 
alternating appearance of large and small exsert ends being a characteristic feature of 
both ; but in F. vasta the corallites are larger and the corallum is without the hillocky 
mode of growth. ; 

Most of my specimens have the same facies as Ehrenberg’s type (19 x 8 x 9 cm.) of 
Astrea tesserifera. In the Paris Museum are eleven good examples (the largest 
30 x 23 em.) from the Red Sea, referred by Milne Edwards and Haime to Prionastrea 
tesserifera, showing much of the variation possible in this species. 

Locality. Red Sea (12). Not recorded from any other locality. 


19. Favia HEuPRIcui (Ehrenberg). (Pl. 27, figs. 1, 2 and 4; 36, fig. 3.) 
1834. Astrea hemprichivi, Ehrenberg, Corall. roth. Meer., p. 96. 
11834. Astrea dipsacea (pars), Ehrenberg, Corall. roth. Meer., p. 97. 
1851. Prionastrea hemprichi, Milne Edwards and Haime, Pol. foss. terr. palzoz., p. 102. 
1857. Prionastrea hemprichi, Hist. Nat. Corall., 11, p. 521. 
1879. Prionastrea spinosa, Klunzinger, Korall. Roth. Meer., iii, p. 39, pl. 4, fig. 7 and pl. 10, fig. 5. 
11879. Acanthastrea hirsuta var. microstoma, Klunzinger, Korall. Roth. Meer., iii, p. 42, pl. 5 2 


MATTHAI—RECENT COLONIAL ASTRAIDA 111 


1904. Acanthastrea hirsuta var. aperontean, Gardiner, Fauna and Geogr. Maldives and Laccadives, ii, p. 784, 
pl. lix, fig. 6. 

Corallum. Usually incrusting, flat or convex, tending to become massive. Corallites 
hexa- or pentagonal, sometimes meandering (two or three corallites in a row being without 
intervening partitions). Inter-calicinal walls 1—5 mm. in thickness, average about 
2° or 3mm. Calices with average diameters 12 and 8 mm. (the longest meandering 
calyx being 25 mm.), depth 7 or 8mm. 

Septa usually sloping, swollen towards walls, those of adjacent corallites continuous 
over inter-calicinal walls, with conspicuously toothed edges, sides usually smooth, some- 
times rough. Septa up to 45 in number, average about 35; of these 12—18 meeting 
columella, each with 6—8 smooth pointed teeth (hollow or solid), towards calicular 
opening usually becoming longer, obliquely directed and swollen at their bases, the first 
tooth (situated over the inter-calicinal wall) being longest (up to 3mm.) and vertical and 
forming the most exsert part of each septum; between two such rows of teeth the inter- 
calicinal partition appearing furrowed. Subsidiary septa often curving towards and 
fusing with sides of principal septa in groups of up to 3 or 4 as in F. complanata. 
Sometimes a few alternating rudimentary septa but the cycle never complete. Columella 
formed of interlacing septal teeth, often with points above, 1—+ width of calyx. 

Multiplication by unequal or equal fission, rarely by marginal fission. 

I apply the name Pavia hemprichiz to this species with some hesitation, as Ehren- 
berg’s type of Astrea hemprichi is a small incrusting specimen measuring 9°5 x 8°5 cm. 
(Pl. 36, fig. 3). Judging only from the corallum, the grouping of my specimens 
under Ehrenberg’s species appears to be the most satisfactory. Perhaps they resemble 
Klunzinger’s type (8°5 x 7 x 4x4 em.) of Prionastrea spinosa somewhat more than Ehren- 
bere’s example, but the former is in no essential respect different from the latter ; it has 
polygonal corallites, well developed columellee, conspicuous septal teeth which are longer 
and erect on the exsert septal ends, and its main septa are continuous over the walls. 

This species is closely related to F. complanata, having more or less the same facies 
as the edge-regions of the latter, but differs from it in (1) the corallum being flat or convex, 
without the hillocky mode of growth so characteristic of 1. complanata, (2) the corallites 
more uniform in shape and size, (3) fission sub-equal or equal, rarely marginal, (4) the 
general absence of smaller exsert ends alternating with those of the main septa, (3) the 
teeth being smooth and pointed and on the whole more conspicuous, increasing in length 
towards the calicular opening, hence the corallum appearing more thorny, The actual 
relationship of these two species can be settled only after a comparative study of their 
polyps. 

Owing to the swollen nature of the septa in the theca and of the teeth, the species 
also resembles #. hirsuta. However the specimen, which Gardiner referred to Acantha- 
strea hirsuta var. macrostoma (Klunz.) (Pl. 27, fig. 4) appears to be more related to the 
present species than to var. megalostoma, its calices being much smaller than in the latter 
and of almost uniform size (diameters about 10 and 8°5mm., depth 7 or 8mm.), the 
inter-calicinal walls thinner (about 2°5 mm.) and the septa less in number (30—35). The 
specimen serves to show the degree of thickness possible in this species. The specimens 


112 PERCY SLADEN TRUST EXPEDITION 


from Aldabra and Seychelles (Pl. 27, figs. 1 and 2) constitute an intermediate series 
between Gardiner’s example from Minikoi and the thin forms from Salomon. The latter 
specimens show some resemblance to F’. berthollet: owing to the thinness of their walls and 
septa and the comparative shortness of their septal teeth ; hence their place in F. hem- 
prichw is doubtful. 

Of the five specimens in the Berlin Museum named Astrea dipsacea, Ehrb., four 
have been later referred to Acanthastrea hirsuta var. nicrostoma by Klunzinger—one, 
measuring 19 x 14x 10cm. is figured—and the fifth to var. megalostoma. The former, 
though possessing the thickened character of the latter, have smaller corallites and fewer 
septa ; I am therefore led to think that they, like the similarly named example of Gardiner’s, 
form an extreme case of skeletal variation in the present species. Klunzinger’s figured 
type of var. megalostoma is not in the Berlin Museum. 

Milne Edwards and Haime’s type of Prionastrea hemprichi is a small incrusting 
specimen (11 x 9 em.) from the Red Sea; it undoubtedly is identical with both Ehrenberg’s 
and Klunzinger’s types. The spines on the exsert septal ends are even longer than in 
Klunzinger’s example, the corallite-walls are ridged, 2—4 of the principal septa are 
thicker and broader, but to a less extent than in Gardiner’s type of Pavia adduensis, 
and the corallites have a tendency to meander as in my examples. In the Paris Museum 
there is also a large specimen from Koseir named Prionastrea spinosa. 

Localities. Seychelles(5). Minikoi (1). Chagos, Salomon (5, thinner than the rest). 
Previously known only from the Red Sea. 


20. Mavra ravosa (Ellis and Solander). (PI. 28, fig. 2.) 


1786. Madrepora favosa, Ellis and Solander, Nat. Hist. Zooph., p. 167, pl. 50, fig. 1. 

1797. Madrepora favosa (pars), Esper, Forts. Pflanz., p. 34, pl. 45a, fig. 1 (copy of Ellis and Solander’s figure). 
1830. Dipsastrea favosa, Blainville, Dict. Sci. Nat., 1x, p. 338; Manuel d’actinol., p. 373. 

1850. Prionastrea magnistellata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xi, p. 129. 

1850. Prionastrea favosa, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 132. 

1857. Acanthastrea bowerbanki, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 503, pl. D6, fig. 1. 

1857. Prionastrea magnistellata, Milne Kdwards and Haime, Hist. Nat. Corall., ii, p. 516. 

1857. Prionastrea favosa, Milne Edwards and Haime, Hist. Nat. Corall., 11, p. 520. 

I refer to this species a single massive convex specimen from Red Sea measuring 
19x17 x 145 em. which closely resembles Ellis and Solander’s figure of Madrepora favosa 
(the type itself is missing from the Glasgow Museum). It has polygonal corallites, the 
ealices of more or less uniform width along their entire height with diameters up to 
20 and 14mm. (average 18 and 13mm.) and depth up to 11 or 12mm. The inter- 
ealicinal walls are flat, being about 3mm. The total number of septa vary up to 50 
(average about 42); of these 16—18 septa meet the columella, their upper two-thirds 
being narrow and vertical while their lower thirds are broader and simulate a pali-crown. 
The septa have rough sides, short teeth (about 10) on their edges, and are exsert for 
1 or 1‘5mm. The exsert ends are usually arched, those of neighbouring corallites either 
alternating or meeting, and are usually connected along the middle of the walls by a 
thin ridge. Owing to the arched condition of the exsert ends, the walls appear grooved 
above. The columella is formed by loosely interlacing septal trabeculee and is {—+4 the 
width of the calyx. New corallites are formed by unequal fission. 


MATTHAI—RECENT COLONIAL ASTRAIDA 113 


This specimen as well as Ellis and Solander’s figure of Madrepora favosa have much 
the same facies as some of my examples of Hawa favus, but possess, on the whole, larger 
corallites. Polyps are, however, necessary to determine if the specimen represents a good 
species or is only a large form of F. favus. 

In the Paris Museum Proonastrea favosa and magnistellata ave each represented by 
a large specimen and Acanthastrea bowerbanki by a very small one. I fail to see any 
differences separating these three species. They have, on the whole, larger corallites 
than my specimen, with the calices widest at the calicular openings and gradually narrow- 
ing towards their bases; moreover the inter-calicinal walls are ridged, the septa slope 
down towards the columell, those of neighbouring corallites meeting over the walls. 

In the Glasgow Museum I have examined a specimen (16:5 x 14 x 9°5 em.) which 
resembles Milne Edwards and Haime’s P. magnistellata or P. favosa but it is doubtful 
if it is Ellis and Solander’s figured type. Prof. Graham Kerr does not include it in his 
list* of specimens described by these authors. 

According to Milne Edwards and Haime, Madrepora favosa (pars), Linnzus, is a 
rugose species Staurza astreiformis (Cor. ili, p, 325). 


21. Favia rracum (Esper). 


A single convex specimen (7 x 5°5 x 4:5 em.) from St Vincent, collected by Mr C. 
Crossland. I hope soon to give a full account of this interesting species in the scientific 
Report of the British Antarctic Expedition based on material collected off South Trinidad. 
I have also seen examples of it in the Paris Museum. 


22. ?Mayia parvimuratTa, Gardiner. 

1904. Favia parvimurata, Gardiner, Fauna Geogr. Maldive and Laccadives Archipel., ii, p. 771, pl. 62, 
fig, 25. 

The only existing representative of this species is Gardiner’s type (11°5 x 10 x 4cm.) 
from Hulule. The principal characters of this specimen are the penta- or hexagonal shape 
of the corallites, the thinness of the inter-calicinal walls, the last septal teeth being 
upright, high (1°5 mm.), broad and swollen at their bases, with their outer edges sloping 
away from their pointed tips and inner margins vertical. There is a conspicuous pali- 
crown; but towards the edge of the specimen the teeth are thin and short and hence 
the pali-crown is inconspicuous, the corallites approaching those of Fava berthollet in 
appearance. 

Somewhat resembling this specimen is Milne Edwards and Haime’s type (small and 
flat) of Favia bowerbanki from Australia and a specimen (no. 3999) in the Berlin Museum 
from Ralum. A larger series of specimens is required in order to settle whether this is a 
true species. . 


93. Mavra sp. ? 


1899. Astrea denticulata, Gardiner, Proc. Zool. Soc. London, p. 748, pl. 47, fig. 1. 
1899. Astrea pallida, Gardiner, Proc. Zool. Soc. London, p. 748. 


* “ Remarks upon the Zoological Collection of the University of Glasgow,” The Glasgow Naturalist, vol. iu, 
no. 4, p. 111, 1910. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 15 


114 PERCY SLADEN TRUST EXPEDITION 


1899. Astrea rotumana, Gardiner, Proc. Zool. Soc. London, p. 750, pl. 47, fig. 3. 
1899. Astrea affinis, Gardiner, Proe. Zool. Soc. London, p. 750. 


The two large specimens (the larger measuring 19 x 15 x 9 em.) from Funafuti, which 
Gardiner referred to Astrea denticulata (Hl. and Sol.), the two small convex ones (the 
larger measuring 9 x 7 x 6 cm.) from Rotuma, which he assigned to Astrea pallida (Dana), 
and the fragment from Wakaya named Astrea affinis (Eid. and H.) resemble one another 
so closely that there is hardly any doubt they belong to the same species. They do not 
agree with any of the type specimens I have examined. It is, however, likely that this 
species may have been previously recorded by Dana, but no satisfactory determination 
is possible from his descriptions or figures. 

In all these specimens the corallum is massive and convex, the corallites are polygonal, 
often irregularly compressed, close together but usually separated by narrow grooves. 
Sometimes the walls are fused. The average diameters of the calices are 8—11 mm. and 
6—8 mm. and the depth 5 or 6 mm. The septa are vertical, with rough sides and either 
toothed or entire edges, their total number varying from 25—35 ; of these 9—15 (average 
11—13) meet the columella, their upper two-thirds being narrower than their lower one- 
third. At the calicular margins the principal septa are much broader (about 2 mm.) and 
thicker (‘5 mm.) than the subsidiary septa and are more highly exsert (1‘5—2 mm.). (This 
feature, as has been pointed out, is characteristic of many of Gardiner’s Pacific specimens.) 
The exsert ends of the septa of neighbouring corallites either meet in notches or alternate. 
The columella is spongy, being formed of thin septal trabeculee and is from 4 to 4 the 
width of the calices, sometimes quite rudimentary. New corallites are formed by equal 
or sub-equal fission, which is usually effected by two of the thickened principal septa 
meeting across the calices. 

The small specimen (10 x 6 x 4°5 em.) from Rotuma, which Gardiner has made the 
type of a new species, Astr@a rotwmana, is closely related to the above-mentioned 
specimens but its corallum is lighter, the corallites somewhat further apart and the septa 
fewer. These differences are perhaps to be accounted for by the fact that the specimen 
was an edge-piece. It has also a general resemblance to my Salomon examples of Favia 
clouet. 


24. Favta sp. ? 
1899. Prionastrea tenella, Gardiner, Proc. Zool. Soc. London, p. 761. 


A small convex specimen from Rotuma with an incrusting corallum 4°8 cm. long and 
4-2 em. broad, which Gardiner referred to Prionastrea tenella, Dana, appears to be a good 
species of Favia, being quite different from any other specimen I have examined. Its 
corallites are polygonal, the diameters of the calices measuring on an average 5 mm. and 
4mm. and depth 3 mm.; the inter-corallite walls are ridged, being about 1 mm. thick. 
The septa are sloping with uneven edges and rough sides, their total number varying from 
20—26 ; alternating with these is a cycle of very narrow septa. 12—15 septa meet the 
columella, of which 6—8 are broader than the others, these giving the corallites a star- 
shaped appearance. All the septa are very slightly exsert, those of adjacent corallites 
meeting over the inter-corallite walls. The columella is thin and spongy. Fission is 


MATTHAI—RECENT COLONIAL ASTRAIDA 115 


equal or sub-equal. In the specimen there are seven double corallites of which two are 
Celoria-like, being without partitions, while in the others the partitions have grown only 
up to the middle of the calices. 


25. Havia rorutosa (Ellis and Solander). 

1786. Madrepora rotulosa, Ellis and Solander, Nat. Hist. Zooph., p. 166, pl. 55, figs. 1—3 (non Astrea rotulosa, 
Lamarck, and’ favia rotulosa, Khrb. and Kd. and H.). 

There is no example of this species in my collections. 

In Ellis and Solander’s type (12x10x9 cm. and identical with their fig. 1) of 
Madrepora rotulosa, now in the Glasgow Museum, the corallites are round or oval, 
slightly projecting, becoming somewhat oblique towards the edges; the calices are 
4—5 mm. in diameter. The total number of septa is up to 16 or 20, of which 6—8 meet 
the columella in every corallite, each with a conspicuous bluntly-pointed paliform lobe ; an 
alternating cycle of rudimentary septa can be made out in some of the corallites. 

Similar to this and of about the same size is another specimen in the Glasgow 
Museum, but as it is not one of Ellis and Solander’s figured types its history cannot be 
determined. 

In the Paris Museum are two large specimens named Heliastrea annularis, the 
larger (26 x 25x16 cm.) from “mers d’Amerique” and the other from West Indies, 
- which are identical with Ellis and Solander’s type. 


GONIASTREA (Milne Edwards and Haime). 


1816. Astrea (pars), Lamarck, Hist. Anim. sans vert., ii, p. 257. 

1830. Dipsastrea (pars), Blainville, Dict. Sci. Nat., Ix, p. 338. 

1834, Astrea (pars), Ehrenberg, Corall. roth. Meer., p. 95. 

1848. issicelli subgenus iii of Astrea (pars), Dana, Expl. exp. Zooph., p. 220. 
1849. Goniastrea (pars), Milne Edwards and Haime, Compt. rend. de l’Acad. des Sci., xxvii, p. 495. 
1857. Goniastrea (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 444. 
1879. Goniastrea (pars), Klunzinger, Korall. Roth. Meer., iii, p. 32. 

1884. Goniastreea, Duncan, Jour. Linn. Soc. London, Zool., xviii, p. 102. 

1899. Goniastrea, Gardiner, Proc. Zool. Soc. London,-p. 746. 

1904. Goniastrea, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 772. 

1904. Prionastrea (pars), Gardiner, Fauna Geogr. Maldives and Laccadives, p. 785. 


Corallum. Massive or incrusting. Corallites polygonal, often with a tendency to 
meander, walls completely fused, usually thin and ridged. Septa narrow, slightly exsert, 
all of more or less equal width at the calicular margin; when dipping vertically presenting 
an appearance of remarkably even-sized plates and making the walls appear thicker than 
they really are. The principal septa with paliform lobes; no true pali. An alternating 
eycle of rudimentary septa present. The main septa in neighbouring corallites either 
continuous over the walls or meeting the rudimentary ones, in the latter case appearing to 
alternate when seen with the naked eye. Columella little developed, sometimes quite 
absent.. 

Polyps. The following characters distinguish the polyps from those of Fava: 
(1) the presence of not more than six principal couples of mesenteries ; (2) the number of 
subsidiary couples of mesenteries proportionately more than in Favia, viz. more than 
thrice (usually four or five times) the number of principal couples; (3) nematocysts II 

15—2 


116 PERCY SLADEN TRUST EXPEDITION 


modified into the Ile type; usually arranged in rows in the coils of the mesenterial 
filaments ; (4) nematocysts III, when present, narrower and longer than in Fava. 

The polyps are polygonal, with no edge-zone. Both entoccelic and exoccelic tentacles 
are present, their number not always equalling that of the entocceles and exocceles, each 
with a swollen terminal battery ; sub-terminal batteries present but number doubtful. 
The stomodzeum is comparatively narrower than in Favia. The subsidiary couples of 
mesenteries vary considerably in their respective widths; their number in a primary 
exoccele also varies up to 5. The coils of the filaments are usually found protruded 
through the oral-dise. Fission of polyps equal or sub-equal. 

The genus resembles Favia in many respects but the species studied form a homo- 
geneous group. Until conclusive evidence is available for merging these genera, it is 
advisable to keep them separate. 

Remarks. In the Berlin Museum there are two large specimens with irregular lobes, 
referred to Scaphophyllia lobata, Stud., their corallites showing a pronounced meandering 
facies. There is little doubt that these specimens belong to G'onzastrea, the corallites on 
the humps having thin septa and resembling those of G. pectinata (Ehrb.), while those 
towards the edges are shallower with conspicuous paliform lobes simulating the condition 
in G. planulata, Kd. and H. The irregular mode of growth of the corallum is, however, 
a striking difference from both G. pectinata and G. planulata. I did not examine the 
type-specimen of Scapophyllia cylindrica, Ed. and H., in the Paris Museum, not realising 
at the time the probable relationship of that genus to Goniastrea. If, on examination of 
polyps, the identity of the two genera should be established, Scapophyllia would, in 
accordance with the claims of priority, have to replace Goniastrea. The three specimens 
on which Quelch instituted his new species G. multilobata (figured specimen 13 cm. long 
and 12 em. high) resemble in every respect the Berlin examples of Scaphophyllia lobata, 
Stud. ; the hillocks in these specimens sometimes end in ridges as in Favia abdita (EIlL. 
and Sol.). . 

The following is a list of specimens originally assigned to G'oniastrea by various 
authors but which really belong to Fawa: 


pees Size of specimens Locality Sp neon By whom pe a 
1 large, convex Unknown | G. rudis, Kd. and H. Milne Edwards | /. pentagona (Esp.) 
and Haime 

1 18 x 10 x 4 cm. Red Sea | G. halicora (Ehrb.) var. | Klunzinger F. bertholleti, Ed. & 
obtusa H. 

1 25 x 21 x 16 cm. ba G. halicora (Ehrb.) 5 F. halicora (Erhb.) 
var. acuta (= Astrea 
halicora Ehrb.) 

1 85x75 x 3:5 em. 3 G. seychellensis, Klunz. t 1 Ff. fawus (Forsk.) 

(= Astrea deformis, Ehrb.) 
1 Corallum incrusting | Api, New | G@. Jaxa, Quelch Quelch F. pentagona (Esp.) 
5-5 x 5 em. Hebrides 

1 A fragment Banda G. favistella (Dana) “ F. abdita (Ell. & Sol.) 

1 23 x 18 x 17 cm. Ceylon G. serrata, Ortm. Ortmann F, doreyensis (Oken) 

1 6 x 4:5 x 2°5 cm. Suvadiva |? G. favus (Forsk.) Gardiner F. pentagona (Esp.) 


MATTHAI—RECENT COLONIAL ASTRAID Ai 117 


In the Hofmuseum, Vienna, are eight large specimens (‘‘ Pola” expedition), referred 
by Marenzeller to G. halicora (Ehrb.). Without polyps it is difficult to decide if they 
belong to Goniastrea or Favia. No. 15961 (20x 19x16 cm.) resembles Klunzinger’s 
example of G. seychellensis ; the corallites on its side are circular or oval, shallow and 
wider apart with peritheca between. Nos. 15928 and 15933 are somewhat similar to 
Milne Edwards and Haime’s example of Prionastrea profundicella (perhaps a Goniastrea), 
while No. 15929 agrees with these authors’ type-specimens of Favia denticulata [= F. favus 
(Forsk.)]. No. 15934 is the largest specimen, measuring 35 x 35 x 16 cm., with also the 
largest corallites. 

Distribution. Indian and Pacific Oceans. Cretaceous to recent (Hastman after 
Zittel). 


1. Gonrastrna sotipa (Milne Edwards and Haime). (Pl. 10, fig. 1; 28, figs. 3 
and 4; 31, fig. 1; 33, fig. 4; 38, fig. 3.) 
1830. Dipsastrea solida, Blainville, Dict. Sci. Nat., 1x, p. 338 ; Manuel d’Actinol., p. 373. 
1850. Goniastrea solida (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 160 (non 
Madrepora solida, Forsk.). 
1857. Goniastrea solida (pars), Milne Kdwards and Haime, Hist. Nat. Corall., ii, p. 444. 


1879. Goniastrea favus (pars), Klunzinger, Korall. Roth. Meer., iii, p. 35 (non Madrepora favus, Forsk.) 
1904. Goniastrea solida, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 772, pl. 62, fig. 28. 


Corallum. Massive, convex. Corallites never meandering. Inter-calicinal walls 
1 or 1°5 mm. thick, sometimes rounded. Calices about 4 mm. in diameter, up to 4 mm. 
deep, usually 3 or 3°5 mm., shallower towards the edges. 

Septa slightly sloping or vertical, edges serrate, with sides spinulose, up to 32 in 
number, average 25—30, usually meeting the rudimentary septa of neighbouring corallites. 
8—11 septa meeting in the centre; sometimes the pali-crown represented by the lower 
thickened parts of their inner edges. Columella more or less compact, short, 4—+ width 
of calyx. ; 

Polyps. (1) In the stomodzeal region of polyp, subsidiary couples of mesenteries, 
about four or five times the number of principal couples. (2) In same region of polyp, 
-entoccelic pleats usually absent from principal mesenteries, when present very narrow 
and not extending to outer one-third of width of mesentery ; pleatal region thickened. 
(3) Nematocysts IIb somewhat more numerous in the terminal tentacular batteries than 
type I. (4) Nematocysts Ile with the dark-stained axis about + length of sac. 
Remarks. A. Polyps. In the polyps examined, 5 or 6 principal couples of 
mesenteries are present, of which 1 couple is incomplete, and 21—26 subsidiary couples, 
the latter varying greatly in their width. Entoccelic pleats make their appearance below 
the stomodzal region of polyp but are short, thick and blunt. The convolutions of 
mesenteries are abundant below the stomodzeum, also protruded through the oral-disc. 
The stomodum is oval in transverse section, its diameters being about 1°25 mm. and 
‘75 mm.; its ridges are thicker than broad, with crowded nuclei, and with some nemato- 
cysts I occurring in their lower halves. The ectoderm of the oral-disc is thin, resembling 
that of Cyphastrea. The nematocysts I in the terminal batteries of the tentacles have 


118 PERCY SLADEN TRUST EXPEDITION 


25—30 turns of the spiral in each. Filaments are present on all the principal 
mesenteries and the wider subsidiaries ; nematocysts I occur in small numbers along their 
whole lengths ; II ¢ nematocysts are numerous, as closely arranged in the convolutions as 
IIL} in Echinopora. Type III is absent. The endoderm in the oral-disc is as thin as the 
ectoderm over it, in the tentacles vacuolated and lobed with numerous alge, in the 
mesenteries somewhat swollen on each side of the thickened region of the mesoglea, 
beyond which it is thin. Algze are scarce in the mesenterial endoderm. Gonads were not 
present in any of the polyps. The polyps of this species bear greater resemblance to those 
of G. pectinata than to those of G. retiformis, although the condition is reversed with 
regard to the corallum. 
Number of polyps examined three, from a specimen from Amirante, 16 fms. 


B. Corallum. Klunzinger’s suggestion that Madrepora solida, Forsk. was a 
Porites has later been confirmed by Marenzeller’s examination of Forskal’s type-specimens 
in the Copenhagen Museum. Marenzeller has also made the discovery that Madrepora 
favus, Forsk., is not a Gonastrea. Of the two specific names I have retained the former 
since it comes first in Forskal’s paper. I could not examine Klunzinger’s type-specimen of 
G. favus, as it was not in the Berlin Museum, but judging from his figure it appears to 
belong to the present species. Of the five specimens referred by Milne Edwards and 
Haime to G. solida (2 from Red Sea and 3 from Seychelles) two (one from each locality) — 
belong to G. retiformas, the figured specimen (Ann. Sci. Nat., 3° sér, x, pl. 9, fig. 7) being 
one of them ; all these specimens were later re-named G’. favus by Klunzinger. 

In the Hofmuseum, Vienna, are 19 specimens (“ Pola” expedition) referred to 
G. favus by Marenzeller, of which 12 are numbered; of these nos. 15919, 15921, 15922 
and all the unnumbered specimens are undoubtedly G. retiformes. The remaining ones 
may be assigned to G. solida; no. 15918 has corallites showing a meandering tendency. 

Localities. Seychelles (1). Maldives, Hulule (1). Amirante (1 small, 16 fms.). 
Known also from the Red Sea (Milne Edwards and Haime, and Marenzeller). 


2. GoONIASTREA RETIFORMIS (Lamarck). (PI. 10, fig. 3; 31, figs. 1—5; 33, fig. 3; 
38, figs. 2 and 4.) 


1816. Astrea retiformis, Lamarck, Hist. Anim. sans vert., ii, p. 265; 2° édit., p. 415. 

1834. Astrea spongia, Ehrenberg, Corall. roth. Meer., p. 96. 

1850. Goniastrea solida (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° séer., xii, p. 160. 

1850. Goniastrea retiformis, Milne Edwards and Haime, Ann, Sci. Nat., Zool., 3° sér., xii, p. 161. 

1850. Goniastrea bournoni, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 162. 

1857. Goniastrea solida (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 444. 

1857. Goniastrea retiformis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 446. 

1857. Goniastrea bournoni, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 446. 

1879. Goniastrea retyformis, Klunzinger, Korall. Roth. Meer., iii, p. 36, pl. 4, fig. 5. 

1889. Goniastrea retiformis, Ortmann, Zool. Jahrb., iv, p. 527. 

11879. Goniastrea favus (pars), Klunzinger, Korall. Roth. Meer., iii, p. 35, pl. 4, fig. 4 and pl. 10, fig. 7 (non 
Madrepora favus, Forsk.). 

1892. Goniastrea retiformis, Ortmann, Zool. Jahrb., vi, p. 661. 

1899. Goniastrea eximia, Gardiner, Proc. Zool. Soc. London, p. 747. 

1899. Goniastrea solida, Gardiner, Proc. Zool. Soc. London, p. 747. 

1904. Goniastrea retiformis, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 772. 


MATTHAI—RECENT COLONIAL ASTRAIDA 119 


Corallum. Massive, convex. Corallites penta- or hexagonal, often drawn out for 
fission. Inter-calicinal walls thinner than in G. solida, never more than ‘75 mm. Calices 
about 4 mm. in diameter, depth 2—3 mm. 

Septa sloping gradually to a short distance, then dipping vertically ; edges serrate, 
sides rough ; total number 18—22, usually meeting the rudimentary septa in neighbouring 
corallites; 7—11 septa meeting columella. Clefts behind paliform lobes sometimes 
extending below level of columella. Columella spongy or dense, +—% width of calyx, 
sometimes quite rudimentary. 

Polyps. (1) In the stomodzal region of polyp, subsidiary couples of mesenteries 
about three or four times the number of principal couples. (2) In the same region, 
entoccelic pleats narrow, slightly constricted at their bases, occasionally sub-divided and 
restricted to outer one-third of width of principal mesentery ; pleatal region thicker than 
in G. solida. (3) Nematocysts I1b much fewer in the terminal tentacular batteries than 
in G. solida. (4) Nematocysts Ile with the dark-stained axis in each about three-fifths 
length of the sac. 

Polyps examined 14, 3 and 2 from two specimens from Hulule, Maldives, 2 from 

a Ceylon colony, and 7 from five Rotuma colonies. 
Remarks. A. Polyps. 5 or 6 principal couples of mesenteries are present, of 
which 1—3 are incomplete, and as many subsidiary couples as in G. solida. The pleatal 
region of the mesenterial mesogleea, when compared with the very thin non-pleatal region 
is much thicker than in any species studied ; below the stomodzeal region of polyp, the 
entoceelic pleats become broader and sub divided and extend over the outer half or two- 
thirds of the width of the mesentery. ‘The convolutions of the mesenteries are scarce, 
hence the inter-mesenteric chambers appear more or less empty. 5 or 6 sub-terminal 
batteries are present on the tentacles. The stomodzeum is narrower than in G‘. solida, its 
diameters being ‘75 mm. and 65 mm. Its ridges are thicker than broad, and tend to 
unite in the upper half of the stomodeeum, two belonging to mesenteries of adjacent 
couples forming each compound ridge. Nematocysts I and III rarely occur in the ridges. 
Nematocysts I and IIc are found occasionally in the straight regions of the mesenterial 
filaments, while in their convolutions types I1¢ and III occur in large numbers, the axis 
of the former being somewhat bent into a drawn out 8. The calicoblastic layer of ecto- 
derm is vacuolated and deeply columnar near the skeletal attachments of the mesenteries. 
The endoderm is vacuolated everywhere ; in the oral-disc it is thicker than the overlying 
ectoderm, resembling that of Favia pentagona in that the algee are not closely arranged 
and the nuclei gathered along its peripheral region ; on the mesenteries the endoderm 
is very thin on either side of the non-pleatal region, swollen in the pleatal region where 
the vacuoles are arranged in the form of a row of oval sacs on either side, conspicuously 
swollen (pad-like in transverse section) behind the filaments. Gonads were not present in 
any of the polyps. 

B. Corallum. In Lamarck’s collection in the Paris Museum is a small specimen 
named Astrea retiformis, Lam., on which apparently Lamarck founded his species. 
A small specimen from the Red Sea, which Milne Edwards and Haime assigned to 
Heliastrea acropora, and which was later referred to G. favus by Klunzinger, resembles 


120 PERCY SLADEN TRUST EXPEDITION 


G. retiformis more than any other species, but has somewhat larger calices. Milne 
Edwards and Haime’s examples of G. bowrnoni and G. retiformis number respectively five 
(locality unknown) and two (large, from Seychelles). Ehrenberg’s type-specimen of 
Astrea spongia (later referred by Klunzinger to G. retiformis) is a large example 
measuring 20x15 x11cm. In the Hofmuseum, Vienna, are two fine specimens (‘“ Pola” 
expedition), referred to G. retiformis by Marenzeller. Quelch has referred a large speci- 
men from Kandavu, Fiji, to G. ceriwm (Dana) ; it is identical with G. retiformis. 

Localities. Maldives: Hulule (8); Goidu (2). Minikoi (4). Seychelles (13). Red 
Sea (5). Aldabra (2). Rotuma (4). Ceylon (small specimens). Singapore (1). Also 
from Fiji (Quelch) and Dar-es-Salaam (Ortmann). A common species in the Indo-Pacifie 
Ocean. 

3. Goni4stREA PecTINATA (Khrenberg). (Pl. 28, fig. 6; 87, fig. 1.) 

1834. Astrea pectinata, Ehrenberg, Corall. roth. Meer., p. 96. e 

1850. Goniastrea solida (pars), Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 160. 

1850. Goniastrea quoyi, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 162. 

1850. Goniastrea grayi, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 163. 

1857. Goniastrea solida (pars), Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 444. 

1857. Goniastrea quoyt, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 447. 

1857. Goniastrea grayi, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 447. 

1879. Goniastrea pectinata, Klunzinger, Korall. Roth. Meer., ui, p. 34, pl. 4, fig. 6 (non Goniastrea pectinata, 
Gardiner). 

1886. Goniastrea coronalis, Quelch, Reet Corals, Challenger Reports, vol. xvi, part xlvi, p. 101, pl. 3, figs 33a, 

1907. Goniastrea pectinata, Vaughan, Proc. U.S. Nat. Mus. Washington, xxxii, p. 257. 

Corallum. Incrusting, raised into rounded humps. Corallites irregular with a 
tendency to meander. Inter-calicinal walls thin. Single calices larger than in the 
previous two species, average size 7 x 6 mm.—the longest meandering calyx without any 
sign of fission 16 mm.—depth about 7 mm. 

Senta vertical, of equal width at calicular margin (I—1°25 mm.), here appearing as 
one series, edges serrate, sides with blunt spinules, up to 30, average about 25; septa 
continuous in arches over walls of neighbouring corallites, perhaps meeting rudimentary — 
septa; 10—15 septa (average 12) approaching centre of corallite, each with a blunt 
paliform lobe. Columella quite rudimentary or absent. 

Towards edges calices are shallower, septa and inter-calicinal walls thicker, the latter 
up to 3 mm. 

Polyps. (1) In the stomodeeal region of polyp, subsidiary couples of mesenteries about 
four or five times the number of principal couples. (2) In the same region, entoccelic 
pleats when present very narrow and blunt, and not extending beyond outer half of 
principal mesenteries, usually over outer one-third, sometimes absent; pleatal region 
thickened. (3) Nematocysts I1b few in the terminal tentacular batteries as in G. reti- 
formis. (4) Nematocysts II ¢ resembling those of G. solida, each with its dark-stained 
axis not more than one-third length of sac. 

Three polyps sectioned from a specimen from Donganab, Red Sea. 

Remarks. A. Polyps. As in G. solida the convolutions of the mesenteries are 
abundant below the stomodzeum and also protruded through the oral-disc. The stomo-~ 
deeum is somewhat wider than in G. solida, its diameters being 1°5 and 1 mm.; its 


MATTHAI—RECENT COLONIAL ASTRAIDA 121 


ridges are thicker than broad with nematocysts I present in their lower halves, these 
occurring also in the mesenterial filaments. Nematocysts Ilc¢ are arranged close together 
asin G@. solida. Nematocysts III are absent. The endoderm is somewhat swollen on either 
side of the thickened region of the mesenterial mesoglea, with few algee. 

In one of the polyps sectioned two stomodzea are present with six couples of 
mesenteries attached to each. The appearance suggests that one has been formed as an 
upward diverticulum from the base of the other, since the former opens into the latter and 
some of the original principal mesenteries are still found passing from one to the other. 
' This polyp recalls some of the figures which Duerden has given of transverse sections 
of polyps of West Indian Acropora (Madrepora; 33, figs. 4, 5 and 7), but he interprets 
those appearances on the hypothesis of stomodeeal fission. 

B. Corallum. Ehrenberg’s specimen of Astrea pectinata (=G. pectinata, Klunz.) 
in the Berlin Museum measures 8°5 x 6 x 5°5 em., but has somewhat narrower corallites 
than my examples. In the Paris Museum G. quoy, Ed. and H., is represented by a 
specimen from Tongatabou, measuring 11 x11x7 cm., and G. grayi, Ed. and H., by a 
smaller example from Australasia. Klunzinger has rightly referred two of Milne Edwards 
and Haime’s specimens to G. pectinata, one from Red Sea originally assigned to G. solida 
and another without locality to Prionastrea seychellensis, Ed. and H. In the Hof- 
museum, Vienna, are two fine specimens collected by the “Pola” exhibition, one from 
Idda and the other (larger, 17 x 11 x 14 em.), from Bernice, referred by Marenzeller to 
G. pectinata. 

Quelch’s new species G. coronalis is founded on a small, thin, somewhat convex edge- 
piece from Banda, measuring 8 x 8 em. ; it is, in all probability, the same as G. pectinata, 
the only differences being that the columelle are somewhat better developed and the 
paliform lobes broader. The two specimens (one, large, from Somerset, Cape York, the 
other, small, from Banda), referred by Quelch to G. quoyi, differ from Milne Edwards and 
Haime’s examples of the same species in that (1) the columellee are composed of closely- 
twisted trabecule, and (2) the septa are somewhat sloping, very narrow at the calicular 
margins, about 18 meeting columella. The two small specimens from Mactan Island, 
Philippines, which the same author has referred to G. grayi, resemble his specimens of 
G. quoyr, but have slightly larger and deeper calices. 

Localities. Red Sea (4). Chagos, Salomon (1). Also from Tongatabou and 
Australasia (Milne Edwards and Haime), ? Banda (Quelch), French Somaliland (Vaughan). 


4. (Gonrastrea pLANuLATA, Milne Edwards and Haime. (PI. 28, fig. 5; 381, figs. 7 
and 8.) 


1850. Goniastrea planulata, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., xii, p. 162. 
1857. Goniastrea planulata, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 447. 
1904. Goniastreea pectinata, Gardiner, Fauna Geogr. Maldives and Laccadives, p. 773. 


Corallum. Incrusting, becoming massive. Corallites with tendency to meander as 
in G. pectinata. Inter-calicinal walls thicker, up to 3 mm., usually about 1°5 mm., ridged. 
Single calices somewhat smaller, 5—6 x 4 mm. (the longest meandering calyx without any 
mark of fission 11 mm.), depth about 5 mm. 

Septa vertical, of equal width at calicular margins, but thicker and arranged closer 

SECOND SERIES— ZOOLOGY, VOL. XVII. 1€ 


122 PERCY SLADEN TRUST EXPEDITION 


together than in G. pectinata, edges serrate, sides with crowded pointed spinules, total 
number 22—29, usually continuous from corallite to corallite; 9—11 septa meeting 
columella, with conspicuous paliform lobes (75—1 mm. broad). Columella somewhat 
better developed than in G. pectinata. 

Towards edges calices shallower, but septa and inter-calicinal walls thinner. 

A thicker and rougher species than G’. pectinata, but polyps necessary for determining 
whether it is different from the latter or not*. Milne Edwards and Haime’s type of 
G. planulata (locality unknown) is 13 em. long and 10 em. broad, with incrusting corallum, 
but somewhat shallower calices than in my specimens (PI. 81, fig. 7). 

Localities. Maldives: (1 dredged); Goidu (1); Turadu (1). Chagos: Salomon (38) ; 
Egmont (2). Red Sea (1). Not recorded from the Pacific Ocean. 


ADDENDA. SOME TYPE SPECIMENS. 


Astrea deformis, Ehrb., Corall., p. 96 (non Astrea deformis, Lam.). 

Prionastrea seychellensis, Hd. and H., Corall. ii, p. 517. 

I have examined Ehrenberg’s large type of Astrea deformis and Milne Edwards and 
Haime’s two large examples of Prionastrea seychellensis}, and have no doubt that they 
belong to the same species. There is also a large series of this species from different 
localities in my collections showing a wide and interesting range of skeletal variation. 
I hope to receive polyps of this important species in a short time, when a full account 
of its hard and soft parts will be given. At present it cannot be settled whether the 
species is a Mavic or Goniastrea—probably the latter. 


tAPHRASTREA (Milne Edwards and Haime). 


¢ ApHRASTR#A DEFORMIS (Lamarck). 
1816. Astrea deformis, Lamarck, Hist. Anim. sans vert., ii, p. 264; 2° édit., p. 414. 
1830. Dipsastrea deformis, Blainville, Dict. Sci. Nat., lx, p. 338; Manuel d’Actinol., p. 273. 
1849. Aphrastrea deformis, Milne Edwards and Haime, Ann. Sci. Nat., Zool., 3° sér., x, pl. 9, fig. 11, and 12, p. 165. 
1857. <Aphrastrea deformis, Milne Edwards and Haime, Hist. Nat. Corall., ii, p. 452. 
1904. Aphrastrea deformis, Gardiner, Fauna Geogr. Maldives and Laccadives, ii, p. 773, pl. 63, fig. 31. 

The four specimens from the Maldives, referred by Gardiner to Aphrastrea deformis 
(Lam.), are identical with Milne Edwards and Haime’s figure of the species, the type being 
missing from the Paris Museum. No polyps of this species were available for examina- 
tion, but from the appearance of the corallites it may well be presumed that the species 
does not possess the bilateral symmetry characteristic of Group 1. The question whether 
it constitutes a separate genus or is to be placed in the genus Favia can be settled only 
after a study of its polyps. 

The principal characters of Gardiner’s specimens are as follows : Corallum more or less 
flat. Corallites penta- or hexagonal, more or less uniform in size. Inter-calicinal walls, 
much thickened, up to 5 or 6 mm., average 3 mm.; this thickness due to secondary 
deposition of calcareous material in the form of vesicles within the corallites} filling up 


* Three sectioned polyps of this species from a specimen from Maldives were useless for histological purposes. 

+ The intra-corallite deposition of calcareous matter is very evident in transverse sections of the corallum 
and is much more than the inter-corallite deposit (2.e. peritheca). The significance of this phenomenon can be 
determined only after examining the polyps. 


MATTHAI—RECENT COLONIAL ASTAIRIDA 123 


the spaces between the septal synapticulz and the wall (Milne Edwards and Haime’s 
“endotheca”); thin low ridges present along middle of inter-calicinal walls, presumably 
marking the position of the original thecz and therefore the true boundaries of the 
corallites. Calices usually circular, 4—5 mm. in diameter, depth 2 or 2°5 mm., with 
tendency to get filled in. 

Septa thin, appearing narrow within calices (owing to their outer regions being buried 
in the intra-corallite calcareous deposit), slightly exsert (not more than °5 mm.), with 
slightly rough sides, toothed vertical edges; septa 30—36 in number, about 11—13 
meeting columella, each with a conspicuous cylindrical blunt paliform lobe 1°5 mm. high 
(behind which septum perforated) and about 4 teeth. Only the exsert ends of about half 
the number of subsidiary septa visible. Septa of adjacent corallites either meeting over 
walls or alternating, connected along middle of walls by thin ridges. Columella 
closely spongy, 4—+ width of calyx, formed of septal trabeculee, sometimes rising into 
rough points. 

Multiplication by fission. 

Vaughan records a specimen of this species from French Somaliland, but gives no 


figure. 


16—2 


19. 
20. 


PERCY SLADEN TRUST EXPEDITION 


V. LITERATURE RELATING TO POLYP ANATOMY. 


AsHworta, J. H. “The Stomodeum, Mesenterial Filaments and Endoderm of Xenia.” Proc. Roy. 
Soc. London, |xiui, p. 443, 1898. 


Ip. “The Structure of Xenia hicksoni, noy. sp., with some Observations on Heteroxenia 
elizabethew, Kollicker.” Quart. Jour. Micr. Sci. xl, p. 245, 1899. 


AWERINZEW, 8. “Studien iiber parasitische Protozoen.” Trav. Soc. Naturlistes St Pétersbourg, 
Bd. 38, Lief 2, S. 1—139, T. 1—3, 1908. 


Bepot, M. “Recherches sur les cellules utricanthes, 1 Vélellides, Physalides.” Rec. Zool. Suisse, 
vol. 4, p. 51, 1886. 


Ip. “Note sur les Cellules utricanthes.” Rev. Suisse Zool., vol. 3, p. 588, 1896. 


BENEDEN, EpouarD vaN. “Une Larve voisine de la Larve de Semper.” Archiv. de Biologie, x, 
p. 485, 1890. 


. Ip. “Recherches sur le développement des Arachnactis, Contributions & la Morphologie de 


Cerianthides.” Archiv. de Biologie, xi, p. 115, 1891; and Bull. Acad. r. s. Belgique, 3° sér., xxi, 
p. 179, 1891. 


Ip. “Les Anthozoaires de la Plankton-Expedition.” Resultats de la Plankton-Expedition der 
Humboldt-Stiftung, u, k. e., 1897. 


BoULENGER, CHARLES L. “On Moerisia lyonsi: a new Hydromedusan from Lake Qurun.” Quart. 
Jour. Mier. Sei, lu, p. 357, 1908. 


Ip. “On the Origin and Migration of the stinging cells in Craspedote Medusz.” Quart. Journ. 
Mier. Sci., vol. lv, p. 763, 1910. 


Bourne, G. C. “The Anatomy of the Madreporarian Coral Fungia.” Quart. Jour. Micr. Sci, 
XXvll, p. 293, 1887. 


Ip. “The Anatomy of Mussa and Luphyllia and the Morphology of the Madreporarian Skeleton.” 
Quart. Jour. Micr. Sci., xxviil., p. 21, 1888. 


Ip. “On the Postembryonic Development of Fungia.” ‘Trans. Roy. Dublin Soce., v (series ii), iv, 
p: 205, 1893. 


Ip. “A criticism of the Cell-Theory; being an Answer to Mr Sedgwick’s Article on the 
Inadequacy of the Cellular Theory of Development.’ Quart. Jour. Mier. Sci., xxxvili, p. 187, 
1895. 


Ip. “Studies on the Structure and Formation of the Caleareous Skeleton of the Anthozoa.” 
Quart. Jour. Micr. Sci., xli, p. 499, 1899. 


Ip. “The Anthozoa.” A Treatise on Zoology, edited by E. Ray Lankester, Part m1, chap. v1, 1900. 


Ib. “On the Solitary Corals.” Ceylon Pearl Oyster Fisheries, Suppl. Report, No. xxx, p. 187, 
1905. 


Boveri, TH. von. “Ueber Entwicklung und Verwandtschaftsbezeichungen der Aktinien.” 
Zeits. Wiss. Zool., xlix, p. 461, 1889. 


Ip. “Das Genus Gyractis.” Zool. Jahrb., vii, p. 241, 1893. 


CaRLGREN, Oskar. “Studien tiber Nordische Actinien.” Kongl. Svenska Vetenskaps-Aka- 
demiens Handlingar, Stockholm, Bd. 25, No. 10, 1893. 


21 


22. 
23. 
24. 
25. 


26. 
27. 


28. 
29, 
20. 
81. 


32. 


31. 
38. 
39, 
40. 
41. 
42. 


43. 


MATTHAI—RECENT COLONIAL ASTRAIDA 125 


CaRLGREN, OsKAR. “Ostafrikanische Actinien. Gesammelt von Herrn Dr F. Stuhlmann 1888 und 
1889.” Mitt. Nat. Mus. Hamburg, xvii, 1900. 


Ip. “Beobachtungen iiber die Mesenterienstellung der Zoantherien nebst Bemerkungen tiber die 
bilaterale symmetrie der Anthozoen.” Festskrift for Lilljeborg, Upsala, p. 149, 1896. 


Ip. “Zur Mesenterienentwickelung der Actinien.” Ofv. k. vet. Akad. Féhr., Stockholm, vol. 54, 
p. 159, 1897. 


CERFONTAINE, Paut. “Notes préliminaires sur l’organisation et la développement de differentes 
formes d’Anthozoaires.’ Bull. Acad. r. s. Belgique, 3° sér., xxi, p. 29. 1891. 


Ip. “Notes préliminaires sur organisation et le développement de différentes formes de Antho- 
zoaires.” Bull. Acad. r. s. Belgique, 3° sér., xxii, p. 128, 1891. 


Deace, Y. ET E. HkrouarD. “Les Celentérés.” Traité de Zoologie Concrete, 11, 2, 1901. 


Dixon, F. “On the Arrangement of the Mesenteries in the Genus Sagartia, Gosse.” Sci. Proc. 
Roy. Dublin Soe., N.S., vi, p. 136, 1888. 


Dixon, G. Y. “Remarks on Sagartia venusta and Sagartia mivea.” Sci. Proc. Roy. Dublin Soe., 
N.S., vi, p. 111, 1888. 


Dixon, G. Y. and A. F. “Notes on Bunodes thallia, Bunodes verrucosa and Tealia crassicornis.” 
Sci. Proc. Roy. Dublin Soc., N.S., vi. p. 310, 1889. 


Ip. “Report on the Marine Invertebrate Fauna near Dublin.” Proc. Roy. Irish Acad., 3rd ser., 
u, p. 19, 1891. 

DusERDEN, J. E. “On the Relations of Certain Stichodactyline to the Madreporaria.” Jour. 
Linn. Soc. Zool., xxvi, p. 635, 1898. 

Ip. “West Indian Madreporarian Polyps.” Nat. Acad. Sci. U.S.A, viii, p. 408, 1902. 

Ip. “The Morphology of the Madreporaria.” 


33. u. “Increase of Mesenteries in Madreporaria beyond the Protocnemic stage.” Johns 
Hopkins University Circulars, xxi, No. 157, p. 59, 1902; and Ann. Mag. Nat. 
Hist., ser. 7, x, p. 96, 1902. 


34. iii. “The significance of Budding and Fission.” Ann. Mag. Nat. Hist., ser. 7, x, p. 382, 
1902. 


35. iv. “Fissiparous Gemmation.” Ann. Mag. Nat. Hist., ser. 7, xi, p. 141, 1903. 
36. v. “Septal Sequence.” Biol. Bull. vii, No. 2, p. 79, 1904. 


Ip. “The Coral Siderastrea radians and its Postlarval Development.” Carnegie Institution, 
Washington, 1904. 


Duncan, P. Martin. “On some Thallophytes parasitic within recent Madreporaria.” Proc. Roy. 
Soe., xxv, p. 238, 1876—1877. 


ErpMANN, Aucust. “Ueber einige neue Zoantheen.” Jenaische Zeitsch. Naturwiss., xix, p. 489, 
1886. 


Fauror, L. “Etudes sur |’Anatomie, lHistologie et le Développement des Actinies.” Archiv. de 
Zool. Exp. et Gen., 3° sér., i, p. 48, 1895. 


Ip. “Scyphozoaires—L’Actinie.” Zoologie Descriptive, edited by Louis Boutan, vol. i. chap. VII, 
1900. 


Ip. “Développement du Pharynx des Couples et de Paires de Cloisons chez les Hexactinies.” 
Archiv. de Zool. Exp. et Gen., 4° sér., i, p. 359, 1903. 


Ip. “Nouvelles Recherches sur le Développement du Pharynx et des Cloisons chez les Hexactinies.” 
Archiv. de Zool. Exp. et Gen., 4° sér., vi., p. 333, 1907. 


126 


49. 


50. 


51. 


52. 


53. 


54. 
55. 


56. 


57. 


58. 


59. 


60. 
61. 


62. 


63. 
64. 


65. 
66. 


67. 
68. 


PERCY SLADEN TRUST EXPEDITION 


Fowter, G. H. “The Anatomy of the Madreporaria.” 


44. 1. “Flabellum, Rhodopsammia.” Quart. Jour. Micr. Sci., xxv, p. 577, 1884; and Stud. 
Owens Coll., 1., p. 243, 1886. 


45. u. “ Madrepora.” Quart. Jour. Micr. Sci., xxvii, p. 1, 1887; and Stud. Owens Coll., ii, 
p. 1, 1890. 


46. i. “Turbinaria, Lophohelia, Seriatopora, Pocillopora.” Quart. Jour. Mier. Sci., xxviii, 
p. 1, 1888; and Stud. Owens Coll., 11. p. 17, 1890. 

47. iv. “Madracis, Amphihelia, Stephanophyllia.” Quart. Jour. Micr. Sci, xxviii, p. 413, 
1888. 


48. yv. “ Duncania, Madrepora, Galaxea, Heteropsammia, Bathyactis.” Quart. Jour. Micr. Sci, 
xxx, p. 405, 1890. 


GARDINER, J. STANLEY. “On the Anatomy of a supposed new species of Cenopsammia from Lifu.” 
Willey Zool. Results, Part Iv, p. 357, 1899. 


Ip. “South African Corals of the Genus Flabellwm, with an account of their anatomy and 
development.” Marine Invest. South Africa, 11, p. 117, 1902. 


Ip. “Some Notes on Variation and Protandry in Flabellwm rubrum, and Senescence in the same 
and other corals.” Proce. Camb. Phil. Soe., xi, Pt. Iv, p. 468, 1902. 


Ip. “The Turbinolid Corals of South Africa, with notes on their Anatomy and Variation.” Marine 
Invest. South Africa, 11, p. 95, 1904. 


Gossz, P. H. “Actinologia Britannica: A History of the British Sea Anemones and Corals,” 
London, 1860. 


Haacke, W. “Zur Blastologie der Korallen.” Jenaische Zeitschrift, xii, p. 269, 1879. 


Happon, A. C. “A Revision of the British Actiniz, Part 1.” Trans. Roy. Dublin Soc., iv 
(Series It), v, p. 297, 1889. 


Ip. “A Newly-Hatched Larva of Huphyllia” Proc. Roy. Dublin Soc, vu, Part mW, XIV, 
p. 127, 1890. 


Happon, A. C., and SHACKLETON, ALIcE M. “A Revision of the British Actiniz, Part 1.” ‘Trans. 
Roy. Dublin Soc., iv (Series 11). x1, p. 609, 1891. 

Ip. “Reports on the Zoological Collections made in Torres Straits.” Trans. Roy. Dublin Soce., 
iv (Series 11), xii, p. 673, 1891. 

Hap, J. “Uber die Nesselzellwanderung bei den Hydroidpolypen.” Arb. Zool. Inst. Wien, — 
xvul, 1907. 

Ip. “Die Entstehung der Knospe bei Hydra.” Arb. Zool. Inst. Wien, xvii, 1910. 


Havet, J. “Contribution a l’étude du Systéme nerveux des Actinies.” La Cellule, xviii, fase. 2, 
p. 387, 1901. 


Herper, A. R. von. “Sagartia troglodytes—Ein Beitrag zur Anatomie der Actinien.” Sitzb. 
kais. Acad. Wien, lxxv, Abth. 1, p. 367, 1877. 


Ip. “Die Gattung Cladocora Ehrb.” Sitzb. Akad. Wissensch. Wien, lxxxiv, p. 634, 1881. 


Ip. “Korallenstudien: Astroides calycularis u. Dendrophyllia ramea.” Arbeit. Zool. Inst. Graz. 1, 
No. 3, p. 153, 1886; and Zeitsch. Wiss. Zool., xliv, p. 507, 1886. 


Ip. “Korallenstudien. Madracis pharensis.” Zeitsch. Wiss. Zool., li, p. 677, 1891. 


Hertwic, Oscar and RicHarp. “Die Actinien Anatomisch und Histologisch mit Besonderer 
Beriicksichtigung des Nervenmuskelsystems Untersucht.” Studien zur Blattertheorie, Heft 1, 
Jena, 1879. 


Hertwic, R. “The Actiniaria.” Challenger Reports, Zool., vi, Part xv, 1882. 
Ip. “ Actiniaria—Supplement.” Challenger Reports, Zool., xxvi, Part LXxtil, 1888. 


69. 


70. 
71. 
72. 


73. 


1. 


15. 
76. 


77. 


78. 


79. 
80. 
81. 


82. 
83. 
84. 


85. 
86. 
87. 
88. 
89. 


90. 


91. 


92. 


93. 
94. 


95. 


MATTHAI—RECENT COLONIAL ASTRAIDA 127 


Hickson, Sypney J. “On the Ciliated Groove (Siphonoglyphe) in the stomodeum of the 
Aleyonarians.” Phil. Trans., elxxiv, p. 698, 1883. 


Ip. “The Anatomy of Alcyonium digitatum.” Quart. Jour. Micr. Sci., xxxvu, p. 348, 1895. 
HoiarD, H. “Monographie du Genre Actinia.” Ann. Sci. Nat., 3° sér., xv, p. 257, 1851. 


Iwanzorr, N. “Ueber den Bau, Die Wirkungsweise und die Entwickelung der Nesselkapseln der 
Ceelenteraten.” Bull. Soc. Nat. Moscou, 1896, p. 1. 


JICKELI, Cart F. “Der Bau der Hydroidpolypen. 1. iiber den histologischen Bau von Eudendrium 
Ehrbg. und Hydra L.” Morph. Jahrb. viii, p. 873, 1882. 


JourpAN, Ev. “Recherches Zoologiques et Histologiques sur les Zoanthaires du Golfe de 
Marseille.” (Thése), 1880. 


Kocu, G. von. “Mittheilungen iiber Ceelenteraten.” Jenaische Zeitsch., xi, 1877. 
Ip. “Bemerkungen iiber das Skelet der Korallen.” Morph. Jahrb., v, p. 316, 1879. 
Ip. “Notizen tiber Korallen.” Morph. Jahrb., vi, p. 355, 1880. 


Ip. “Ueber die Entwicklung des Kalkskeletes von Astroides calycularis und dessen morpholo- 
gische Bedeutung.” Mitth. Stat. Neapel., ii, p. 284, 1882. 


Ip. “Mittheilungen tiber das Kalkskelet der Madreporaria.” Morph. Jahrb., viii, p. 85, 1883. 
Ip. “Die Morphologische Bedeutung des Korallenskelets.” Biol. Central., 11, p. 533, 1883. 


Ip. “Uber das Verhaltnis von Skelet und Weichtheilen bei den Madreporen.” Morph. Jahrb., xii, 
p- 154, 1887. 


Ip. “Uber Flabellum.” Morph. Jahrb., xiv, p. 329, 1888. 
Ip. “Uber Caryophyllia rugosa Moseley.” Morph. Jahrb., xv, p. 10, 1889. 


Ip. “Die ungeschlechtliche Vermehrung (Knospung und Stockbildung) von Madrepora.” Verh. 
Natur. Gesellsch. Niirnberg, 1893. 


Ip. “Das Skelett der Stemkorallen.” Festschrift fiir Carl Gegenbaur, Leipzig, p. 251, 1896. 
K6tiickrr, A. von. “Icones Histologice.” Leipzig, 1863. 

Korotnerr, M. pe. “Organes des sens des Actinies.” Archiv. Zool. Exp. Gén., v, p. 203, 1876. 
KWIETNIEWSKI, C. R. “ Actiniaria von Ternate.” Abhandl. Senckenberg. Gesellsch., xxiii, 1897. 


Lacaze Duruiers, HENRI DE. “Développement des Corallaires. Premier Mémoire. Actiniares 
sans Polypier.’ Archiv. Zool. Exp. Gén., 1, p..289, 1872. 


Ip. “Développement des Corallaires. Deuxieme Mémoire. Actiniares & Polypier.” Archiv. Zool. 
Exp. Gén., ii, p. 269, 1873. 


Ip. “Faune du Golfe du Lion. Evolution du Polypier du Flabellum anthophyllum.” Archiv. 
Zool. Exp. Gén., 3° sér., 11, p. 445, 1894. 


Ip. “Faune du Golfe du Lion. Corallaires-Zoanthaires Sclerodermés.” Archiv. Zool. Exp. Gén., 
3° sér., v, p. 1, 1897. 


Ip. “Les Caryophyllies de Post-Vendres.” Archiv. Zool. Exp. Gén., 3° sér., vii, p. 529, 1899. 


LEDENFELD, R. von. “The Functions of Nettle Cells.” Quart. Jour. Micr. Sci., xxvii, p. 393, 
1887. 


Linpstr6m. “Contributions to the Actinology of the Atlantic Ocean.” Kon. Svenska Vet. Akad. 
Handl., xiv, No. 6, p. 17, 1877. 


McMovrricu, J. Phayrarr. “Contributions on the Morphology of the Actinozoa.” 


96. i. “The structure of Cerianthus Americanus.” Jour. Morph., iv, p. 131, 1891. 
97. u. “On the Development of the Hexactiniz.” Jour. Morph., iv, p. 303, 1891. 
98. ui. “The Phylogeny of the Actinozoa.” Jour. Morph., y, p. 125, 1891. 


128 


101. 


102. 
103. 


104. 


105. 


106. 


107. 


108. 


109. 


110. 


111. 
112. 


113. 


114. 


115. 


116. 


117. 


118. 


119. 
120. 
121. 


122. 


PERCY SLADEN TRUST EXPEDITION 


99. iv. “On some irregularities in the number of the Directive Mesenteries in the Hexactiniz.” 
Zool. Bull., 1, No. 3, p. 115, 1897. 


100. v. “The Mesenterial Filaments in Zoanthus sociatus (Hllis).” Zodl. Bull. i, No. 6, 


p. 251, 1899. 


Mincuin, E. A. “Investigations on the Development of Trypanosomes in Tsetse-Flies and other 
Diptera.” Quart. Jour. Micr. Sci., lui, p. 159, 1908. 


Ip. “An Introduction to the study of the Protozoa.’ 1912. 


Mosrus, Kart. “Ueber den Bau, den Mechanismus und die Entwickelung der Nesselkapseln 
einiger Polypen und Quallen.” Abh. Ver. Hamburg, v, p. 1, 1866. 

Mororr, THEopor. “Entwicklung der Nesselzellen bei Anemonia.”’ Archiv. Zellforsch. iv, 
p. 142, 1910. 

MoseEtzy, H. N. “Report on the Deep Sea Madreporaria.” Challenger Reports, Zool., ii, 
Part vu, 1881. 


Moursacu, Lewis. “Beitraige zur Kenntnis der Anatomie und Entwickelung der Nesselorgane der 
Hydroiden.” Arch. Naturg., i, Jahrg. 60, p. 217, 1894. 


Nusssaum, M. “Ueber die Theilbarkeit der lebendigen Materie. ii, Beitriige zur Naturgeschichte ? 


des Genus Hydra.” Archiv. Mikr. Anat., xxix, p. 265, 1887. 


OcitviE, Marta M. “Microscopic and Systematic Study of Madreporarian Types of Corals.” | 


Phil. Trans., clxxxvu, 83, 1896. 


Ip. “Note on the Formation of the Skeleton in the Madreporaria.” Quart. Jour. Micr. Sci., li, 


p. 473, 1907. 


Fr 


y 


ParKer,G.H. “The Mesenteries and Siphonoglyphs in Metridiwm marginatum, Milne Edwards.” 


Bull. Mus. Comp. Zool., xxx, No. 5, p. 259, 1897. 
Prescn, vAN A. J. “The Antipatharia of the Siboga Expedition.” Monogr. xvii, Leiden, 1914. 


Pratt, E.M. “Anatomy of Neohelia porcellana (Moseley).” Willey Zool. Results, Part v, p. 591, — 


1900. 


Satvin-Moorg, J. E., and Breiny, A. “The Cytology of the Trypanosomes, Part 1.” Ann. 
Trop. Med. Parasitology, i, p. 441, 1908. 


SCHNEIDER and ROTTEKEN. “On the Structure of the Actiniz and Corals.” Ann. Mag. Nat. — 


Hist., vu, p. 437, 1871. 

ScHNEIDER, K. C. “Mittheilungen iiber Siphonophoren: v, Nesselzellen.” Arb. Zool. Inst. Wien, 
xu, p. 133, 1900. 

Scrater, W. L. “Ona New Madreporarian Coral of the Genus Stephanotrochus from the British 
Seas, with Notes on its Anatomy.” Proc. Zool. Soc. London, 1886, p. 128. 

Sepewick, A. “On the Inadequacy of the Cellular Theory of Development, and on the Early 
Development of Nerves, particularly of the Third Nerve and of Sympathetic in Klasmobranchi.” 
Quart. Jour. Mier. Sci., xxxvul, p. 87, 1894. 

Ip. “Further Remarks on the Cell-Theory, with a Reply to Mr Bourne.” Quart. Jour. Micr. 
Sci, xxxvil, p. 331, 1895. 

WEISMANN, A. “Die Entstehung der Sexualzellen bei den Hydromedusen.” Jena, 1883. 

Wittem, Victor. “La Digestion chez les Actinies.” Bull. Soc. Médicine de Gand, p. 295, 1892. 

Wison, E. B. “The mesenterial Filaments of the Alcyonaria.” Mitth. Zool. Stat. Neapel, v, 
p. 1, 1884. 

Witson, Henry V. “On the Development of Manicina areolata.’ Jour. Morph., u, p. 191, 
1889. 


MATTHAI—RECENT COLONIAL ASTRAIDA 129 


EXPLANATION OF PLATES* 


. PLATES 1—10: ANATOMY OF THE POLYPS 
Lettering employed. 

Alg. Alga. alg.nu. Nucleusof alga. ax. Axisof nematocystsII. aa. sh. Sheath of axis of nemato- 
eysts II. C. ect. Calicoblastic layer of ectoderm. C. nu. Nuclei of calicoblastic ectoderm. Col. end. 
Endoderm of column-wall. Col. mg. Mesoglea ofcolumn-wall. d.v. Digestive vacuoles. dir.m. Directive 
mesentery. ect. Ectoderm. ect.nw. Nuclei of ectoderm. end. Endoderm. end.nu. Nuclei of endoderm. 
ent. mus. Entoccelic muscle-fibres of mesentery. ent. pl. Kntoccelic mesogleal pleats of mesentery. ent. t. - 
Entoccelic tentacle. ew. pl. Exoccelic mesogleal pleats of mesentery. ea. mus. Exoccelic muscle-fibres of 
mesentery. ex. t. Exoccelic tentacle. g.c. Germ-cells. gr.v. Granular vacuole. in. m. Inner wall of 
gonad. Jat. /. Lateral lobe of mesenterial filament. m. Mesentery. m. end. Mesenterial endoderm. 
m. f. Mesenterial filament. m. mg. Mesenterial mesoglea. m. /. Outer limiting membrane of calico- 
blastic ectoderm. md./. Median lobe of mesenterial filament. mg. Mesoglea. muc.v. Mucous vacuole. 
n,. Type I nematocyst. m,. Type II nematocyst. mn. Type II, nematocyst. n... Type II, nemato- 
cyst. m;. Type III nematocyst. mn». Type III, nematocyst. n./. Nervous layer. n.r. Probable stage 
in the development of nematocyst II,. ov. b. Ovoid bodies. ov. nw. Nucleus of ovum. ov. nul. 
Nucleolus of ovum. py. Pyrenoid of alga. st. end. Stomodzal endoderm. st. mg. Stomodzeal mesoglea. 
st.r. Ectodermal ridge of stomodzum. sub. b. Sub-terminal tentacular battery of nematocysts. 1. end. 
Endoderm of tentacle. ¢. mg. Mesoglea of tentacle. ter. b. Terminal tentacular battery of nematocysts. 
th. Thread of nematocyst III. th. sh. Sheath of thread of nematocyst III. 


Puate 1. 


Hig. 1. Cyphastrea serailia (Forsk.). (Figs. 1—9 all from the same male polyp.) Transverse section 
through inner wall of edge-zone, showing calicoblastic ectoderm, endoderm with varying nuclei, and 
algze with granular nuclei and pyrenoids. (102, Iv, s,4;.)+ x 900. 


Fig. 2. Id. Transverse section through outer wall of edge-zone, showing ectoderm with granular nuclei, 
some with processes to mesoglea. (102, IV, s,3,.) x 900. 


Fig. 3. Id. Longitudinal section through an exoccelic tentacle to show the histology. (102, IV, s,3).) 
x 433. 


Fig. 4. Id. ‘Transverse section through a primary mesentery and part of the stomodzal wall. Note 
that the algee occur principally on the exoccelic side of the mesentery, and that the mesentery is 
contracted in a radial direction. (102, IV, s,5,.) x 210. 


Vig. 5. Id. Transverse section showing a downward extension of the same ridge and of two lateral 
extensions of the ectoderm on either side, these lower forming the median and lateral lobes of 
a mesenterial filament (ep. fig. 6). Semi-diagrammatical. (102, Iv, s,5s.) 


Fig. 6. Id. Transverse section through straight region of the same mesenterial filament. Semi- 
diagrammatical. (102, Iv, s,14.) ; 

Fig. 7. Id. Group of brown vacuoles from the convoluted region of a mesenterial filament towards the 
base of polyp, surrounding a mass of nucleated protoplasm, each sac with a deeper stained spot 
in its inner bluntly-pointed end. (102, Iv, s,1;.) x 400. 


* The mesenterial musculature is marked only in figs. 3, 4, 14 and 27. In all the other sections of the 
meseuteries it is omitted for the sake of clearness. 
+ This refers to the number of the section. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 17 


130 PERCY SLADEN TRUST EXPEDITION 


Fig. 8. Cyphastrea serailia (Forsk.). Nematocyst II (102, 1V, 5:4.) x 1700. 


Fig. 9. Id. Transverse section through a primary mesentery with testis containing triangular-shaped 
spermatozoa. (102, IV, s,2,.) x 350. 

Fig. 10. Id. Transverse section through a primary mesentery of a female polyp, bearing a ripe ovum 
and two groups of germ-cells. (102, vi, 5,25.) x 193. 


Fig. 11. Cyphastrea chaleidicum (Klunz.). Longitudinal section through a primary entoccelic tentacle 
containing a terminal battery and sub-terminal batteries. In the centre of the tentacle lay a 
septum, which has been decalcified, and the endoderm of the tentacle has by contraction come to lie 
in contact with the body-wall (endoderm and calicoblastic ectoderm) over the septum. (200, 1, 
5,3;.) x 180. 

Fig. 12. Cyphastrea suvadive, Gard. Longitudinal section through a primary entoccelic tentacle con- 
taming a terminal battery and a pair of sub-terminal batteries. (102, 1, 5,3,.) x 240. 


PLATE 2. 


Fig. 13. Cyphastrea chalcidicwm (Klunz.). Transverse section through a primary mesentery and part of 
the stomodeeal wall. (200, I, 5,445.) x 180. 


Fig. 14. Cyphastrea suvadive, Gard. Transverse section through a primary mesentery and 1 Par of 
the stomodeeal wall. (102, 1, 5,4;.) x 180. 


Fig. 15. Id. Nematocyst III from convoluted region of a mesenterial filament. (102, 1, 51,.) x 1450. 


Fig 16. Hchinopora hirsutissima, Ed. and H. Nematocyst I11b from convoluted region of a mesenterial 
filament. (107, 11, 8,2..) x 900. 


Fig. 17. Id. Transverse section through convoluted region of a mesentery. Nematocysts IIIb arranged 
in batteries. (107, IL, 8 2;.) x 115. 


Fig. 18. Id. Nematocyst II from convoluted region of a mesenterial filament, its axis extending beyond 
two-thirds the length of its sac. (107, 1, s31,.) x 950. 


Fig. 19. Id. Longitudinal section through an entoccelic tentacle bearing a large terminal battery and 
about six tub-terminal batteries on each side. Alge rare. (107, 1, 5,25.) x 105. 


Fig. 20. Id. Transverse section through a primary mesentery and part of the stomodeal wall. 
(107, I, sgl,.) x 80. 


Fig. 21. Hchinopora lameilosa (Esper). ‘Transverse section through a primary mesentery and part of 
the stomodeeal wall. (107, U1, s;2,.) x 115. 


Fig. 22. Echinopora gemmacea (Iuam.). Transverse section through a primary mesentery and part of — 
the stomodzal wall. The mesentery is somewhat contracted in a radial direction. Alge few. 
(204 11, S345.) x 115. : 

Fig. 28. Galaxea fascicularis (Linn.). Transverse section through a primary mesentery and part of the 
stomodzeal wall. (201, U, s)1,.) x 55. 


Fig. 24. Id. Transverse section through outer wall of edge-zone. Cp. with Pl. 1, fig. 2, and note thick 
ectoderm with many nuclei and ovoid bodies, and swollen endoderm with alge rare. (201, 1, 5.) 
x 600. 


PLATE 3. 


Fig. 25. Galazea fascicularis (Linn.). Transverse section through inner wall of edge-zone near 
attachment of a mesentery to the outer surface of the corallum, showing calicoblastic ectoderm 
greatly thickened with piled up nuclei. (201, U, s;2;-) x 400. 


MATTHAI—RECENT COLONIAL ASTRAIDA 131 


Fig. 26. Galaxea fascicwlaris (Linn.) Longitudinal section through an exoccelic tentacle containing 
a terminal and numerous sub-terminal batteries. Nematocysts I1b mainly forming the terminal 
battery; as in all species only type I in sub-terminal batteries. (201, I, s1,.) x 80. 


Fig. 27. Id. Transverse section of the mesoglea of a primary mesentery in the stomodzal region to 
show the greatly sub-divided condition of the entoccelic pleats. (201, U, 5325.) x 633. 


Fig. 28. Galaxea “musicalis (Linn.). Longitudinal section through an entocelic and an exoccelic 
tentacle. As in PI. 1, fig. 11, the endoderm of the tentacles lies in contact with the endoderm of the 
body-wall. (101, 1, 5,3.) x 766. 

Fig. 29. Id. Transverse section through two mesenteries, each with a ripe egg projecting into the 
exoccele between; the upper part of a succeeding egg partially overlaps one of these. Vacuolated 
appearance of column-wall endoderm and lower ends of mesenterial filaments to be noted. (101, 11, 
S23.) x 766. 

Fig. 30. Leptastrea roissyana, Ed. and H. Longitudinal section through a primary entoccelic tentacle 
with terminal battery only. (202m 11, 5,2;.) x 110. 


Fig. 31. Id. Transverse section through a primary mesentery and part of the stomodzeal wall. Algee 
abundant on the exoceelic side. (202, 1, so4s.) x 120. 


Fig. 32. Diploastrea heliopora (Lam.). ‘Transverse section through a primary mesentery at the level of 
the stomodzeum. In outline to indicate the ento- and exoccelic pleats. (104; 1, 5,1,.) x 60. 


Fig. 33. Leptastrea roissyana, Ed. and H. Nematocyst I1b. (202, 1, 8,25.) x 1700. 


Fig. 34. Diploastrea heliopora (Lam.). Nematocyst II from the convoluted part of a mesenterial fila- 
ment. (104% TI, sly.) x 950. 


PLATE 4. 


Fig. 35. Favia favus (Forsk.). Stages a—g in the development of a nematocyst I. From the ectoderm 
of oral-disc. (202, 1, s,.) x 1500. 


Vig. 36. Id. Transverse section through principal mesentery and part of the stomodzeal wall. (202, 1, 
Sis34-) x 60. 

Rig 37. Id. Stages a—f in the development of a nematocyst II. From the ectoderm of oral-disc 
(202, 1, 8.) x 1500. 

Fig. 38. Id. Nematocyst II from convoluted region of a mesenterial filament. (202, 1, 84424.) x 900. 


Fig. 39. Id. Early stages in the formation of the axis of nematocyst IJ. From nuclei in mesenterial 
filament. (202, 1, si.) x 1500. 


Fig. 40. Id. Nematocyst III with the thread partially extruded. (202, I, 5,52;.) x 900. 


Fig. 41. Pavia doreyensis, Ed. and H. ‘Transverse section through a principal mesentery and part of 
the stomodzeal wall. In the pleatal region algee are more abundant on the exoccelic side, while in 
the non-pleatal region they occur principally on the entocoelic side. (106g 1, sigly.) x 55. 


Fig. 42. Fava hululensis, Gard. Transverse section through a principal mesentery and part of the 
stomodeal wall. (106, 1, 5.2;.) x 80. 


Fig. 43. Faia clouei (Valen.). Transverse section through a principal mesentery and part of the 
stomodeeal wall. (804, 1, s,2;.) x 43. 


PLATE 5D. 


Fig. 44. Favia doreyensis, Ed. and H. Longitudinal section through an entoccelic tentacle. Stages in 
the development of nematocyst I in terminal battery; and vacuolization in ectoderm and endoderm. 
Algee scarce. (106.1, 51,.) x 185. 

17—2 


132 PERCY SLADEN TRUST EXPEDITION 
Fig. 45. Favia doreyensis, Ed. and H. Transverse section of the straight region of a principal mesentery 
and its filament. Note nematocysts I] im endoderm. (106, 1, 52;.) x 150. 


Fig. 46. Favia abdita (Ell. and Sol.). Transverse section through a principal mesentery and part of 
the stomodzeal wall. (1038, 1, 525.) x 55. 


Fig. 47. Pavia bertholleti (Valen.). Transverse section through a principal mesentery and part of the 
stomodzeal wall. (305, 1, s)2,.) x 80. 


Fig. 48. Favia pentagona (Esper). Transverse section through a principal mesentery and part of the 
stomodeeal wall. (106, 111, s;3,.) x 75. 


Fig. 49. Fama hululensis, Gard. Transverse section through a mesentery with three ova. Note germ- 
cells. Cp. Pl. 1, fig. 10 and Pl. 6, fig. 67. (106, 1, S71.) x 153. 


Fig. 50. Favia ananas (Ell. and Sol.). Transverse section through a principal mesentery and part of 
the stomodzeal wall. (202, 11, s;2,;.) x 80. 


Fig. 51. Favia hululensis, Gard. Transverse section through oral-disc. Note ectoderm with a row of 
oval vacuoles. (106, 1, 5,3.) x 600. 


Fig. 52. Fama pentagona (Esper). ‘Transverse section through oral-dise. Nematocysts I more 
abundant im ectoderm than in any other species of Favia. Cp. with fig. 51. (106, 1, 5,3,.) — 
x 650. 


Fig. 53. Id. Transverse section through column-wall near the surface of the corallum. Note wedge- 
shaped processes of attachment in the calicoblastic layer and remarkable regularity in the 
processes to the mesoglea from the nuclei of both layers. Cp. with fig. 52. (106, 1, 5325.) x 650. 


PLATE 6. 


Fig. 54. Goniastrea solida (Ed. and H.). Transverse section through a principal mesentery and part of 
the stomodeal wall. (106; 1, sg2,.) x 80. 


Fig. 55. Goniastrea retiformis (Lam.). Transverse section through a principal mesentery and part of 
the stomodeal wall. Note somewhat regular arrangement of vacuoles in the endoderm in the 
pleatal region. Algze scarce. (105, 1, 5325.) x 110. 


i ee ee ee 


Fig. 56. Gonastrea pectinata (Ehrh.). Transverse section through a principal mesentery and part of 
the stomodzal wall. (203, 1, so1,.) x 80. 


Fig. 57. Favia hululensis, Gard. Transverse section through a nematocyst III. Note membranous wall 
and protoplasmic core. (106, 1, 55.) x 1450. 


Fig. 58. Goniastrea solida (Kd. and H.). ‘Transverse sections a and 6, through two nematocysts I1e 
passing through their axes at different levels. (106; 1, s,2,.) x 1450. 


Fig. 59. Id. Transverse section through the convoluted region of a mesentery. Nematocysts Ic 
arranged to form batteries. Cp. with Pl. 2, fig. 17. (106; 1, s,8,.) x 1450. 


Fig. 60. Favia hululensis, Gard. Nematocyst III. The spiral of thread is only faintly visible in the 
unextruded condition. (106, 1, ss1,;.) x 950. 


Fig. 61. Gonastrea retiformis (Lam.). Nematocyst III. Note narrow sac. Op. with figs. 16 and 60. — | 
(105, U1, s,l3.) x 950. 


Fig. 62. Id. Nematocyst II. Cp. with Pl. 4, fig. 37;, and note narrower sac and shorter axis. 
(105, I, s,3,.) x 950. 


Fig. 63. Galaxea fascicularis (Linn.). Nematocyst I from terminal battery of tentacle. (201, U1, s)1,.) 
x 1600. 


Fig. 64. Id. Nematocyst IIb from terminal battery of tentacle. (201, I, sgl.) x 950. 


a, 


Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


Fig. 
» Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


MATTHAI—RECENT COLONIAL ASTRAXID Ad 133 


65. Favia favus (Forsk.). Fully developed nematocyst I from a terminal battery of a tentacle. 
(202, 1, 2s.) x 1660. 


66. Favia abdita (Hll. and Sol.). Transverse section through a mesenterial filament. (103, I, Sw1,.) 
x 380. 


67. Id. Transverse section through a principal mesentery, containing ova and germ-cells. (103, 1, 


Reon) x 185. 


PLATE 7. 
Transverse sections : 
Cyphastrea chalcidicum (Klunz.). Stomodzal region of giant polyp. (200, 1, s,23.) x 30. 
Favia bertholleti (Val.). Stomodeal region. (805¢ H, Sizl¢.) x 104. 


Goniastrea retiformis (Lam.). Stomodeal region. (301, 1, 8.1y.) x 27. 


Cyphastrea chalcidicwm (Klunz.). Stomodeeal region. (200, 1, 54.) x 45. 


It 

2. 

3 

4. Cyphastrea serailia (Forsk.). Stomodzal region. (102, Iv, s,5,.) x 65. 

5 

6. Cyphastrea microphthalma (Lam.). Stomodeeal region. (200, 1, 8,27.) x 45. 
7 


Cyphastrea suvadive, Gard. Stomodzal region. (102, 1, 5,45.) x 65. 


PLATE 8. 


Transverse sections: 


.1. Leptastrea roissyana, Ed. and H. Stomodeal region, Ceylon polyp. (800, 1, 5.45.) x 27. 
.2. Id. Tentacular region, Donganab polyp. (202, IV, 555.) x35. 
. 3. Id. Stomodeal region, Donganab polyp. (202, 1, S:4u2.) x 25. 


. 4. Galazea fascicularis (Linn.). Stomodeeal region and edge-zone, Donganab polyp. (201, 1, 


Spl;.) x 104. 
5. Echinopora hirsutissima, Ed. and H. Stomodzal region, Hulule polyp. (107, m1, 5,2,.) x 18. 


. 6. Echinopora lamellosa (Esp.). Stomodeal region and edge-zone, Hulule polyp. (107, 11, s,2,.) 


x 25. 
PLATE: 9. 
Transverse sections : 


1. Favia doreyensis, Ed. and H. Stomodzeal region, with tentacles, Hulule polyp. (106, 1, sy1,.) 
x 9. 


2. Fania favus (Forsk.). Stomodeal region, Donganab polyp. (202; 1, s53;.) x 18. 


3. Favia doreyensis Ed. and H. Stomodeeal region, Hulule polyp. (106q 1, s51;.) x 12. 


4, Hchinopora hirsutissima, Ed. and H. Below stomodeal region, Hulule polyp. (107, I, 8192s.) 
x 22: 


5. Fava abdita (Ell. and Sol.), Stomodzal region, Salomon polyp. (108, 1, 822.) x 15. 
6. Favia hululensis, Gard. Stomodeal region, Hulule polyp. (106, 1, s21;.) x 18. 


PuLate 10. 


Transverse sections: 


1. Goniastrea solida (Ed. and H.). Stomodzal region, Amirante (16 fms.) polyp. (106; 1, s,3,.) 
x 20. 


134 


PERCY SLADEN TRUST EXPEDITION 


Fig. 2. Favia ananas (Ell and Sol.). Stomodzeal region, Donganab polyp. (202, mI, s;2,.) x 25. 


Fig. 


3. 


Goniastrea retiformis (Lam.). Stomodzal region, Hulule polyp. (105, 1, ssl.) x 35. 


Fig. 4. Favia ananas (Hl. and Sol.). Below stomodzeal region. (Cp. Pl. 9, fig. 4.) (2025 111, s¢ 1s.) 
x 25. 


Fig. 5. Favia pentagona (Esp.). Stomodzal region, Hutule polyp. (106, 111, s.1;.) x 28. 


Fig. 6. Favia clowei (Val.). Stomodzal region, Ceylon polyp. (804, I, s,1;.) x 14. 


Fig. 


Fig. 2. No. 4 0n p. 46. Typical form. x 3. 
Fig. 3. Typical specimen. x 3. 
Figs. 4—9. Type specimens of Madrepora serailia, Forskal, Copenhagen. Each x 3. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 
Fig. 


Fig. : 


8 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 
Fig. 


PLATES 11—38: CORALLA. 
PuateE 11. 


Cyphastrea serailia (Forskal). 


il, 


= 


SOS OO ESCO Seah i 


Coes Se ON eo 


No. 1 on p. 46. x3. 


i ee ee 


PuaTE 12. 
Cyphastrea chalcidicwm (Klunz.). No. 5 on the list on p. 46. x 3. 
Id. No. 6 on p. 46, showing a giant corallite. x 3. 
Id. Typical form, enlarged view of Pl. 14, fig. 1. x 3. 
Cyphastrea microphthalma (Lam.). No. 15 on p. 47. x 3. 
Id. No. 120np.47. x3. 
Id. No.9 on p. 46. x3. 
Id. - No. 11 on p. 47. x3. 
Id. Typical form. x 3. 


Id. Type of Astrea microphthalma, Lamarck, Paris. x 3. 


h 


PuatE 13. 
Cyphastrea microphthalma (Lam.). Type of Cyphastrea savignyi, Kd. & H., Paris. x 3. 
Id. Type of Madrepora serailia, Forskal, Copenhagen. x 3. 


See aay 


Cyphastrea suvadive, Gard. Type from Felidu of Oyphastrea maldivensis, Gard. x 3. 
Cyphastrea gardineri, n. sp. No. 22 on p. 47. x 3. 

Id. No. 20 on p. 47, typical form. x 3. 

Galaxea lamarcki, Ed. and H. 36 fms., Nilandu, Maldives, typical form. x 3. 
Cyphastrea microphthalma (Lam.). No. 16 on p. 47, hillocky form of growth. x 4. 
Cyphastrea serailia (Forsk.). No. 1 on p. 46, convex form of growth. x $. 


PLATE 14. 
Cyphastrea chalcidicum, Klunz. ‘Typical form, enlarged on Pl. 12, fig. 3. x 4. 
Echinopora lamellosa (Esp.). Typical form, enlarged view of Pl. 16, fig. 6. x 2. 


|| 


MATTHAI—RECENT COLONIAL ASTRAIDA 135 


Fig. 3. Echinopora lamellosa (Esp.). No. 1 on p. BY, 2, 

Fig. 4. Id. No. 2onp. 57. x 2. 

Fig. 5. Id. No.4onp.57. x 2. 

Fig. 6. Id. No.5 onp.57. x 2. 

Fig. 7. Hchinopora hirsutissima, Ed. and H. No.7 on p. 57. Variant towards #. lamellosa (Esp.). 


Fig. 8. Id. Typical form. x 2. 
Fig. 9. Echinopora gemmacea (Lam.). Typical form. x 2. 


PLATE 15. 

Fig. 1. Echinopora lamellosa (Esp.). Corallites near edge. x 2. 
Fig. 2. EHchinopora hirsutissima, Ed. and H. No.8 on p. 57. x 2. 
Fig. 3. Id. No.9 on p.57. x 2. 
Fig. 4. Id. Type of Echinopora hirsutissima, Ed. and H., Paris. x 1. 
Fig. 5. Echinopora gemmacea (Lam.). Type of Echinopora solidior, Ed. and H., Paris. x 1. 
Fig. 6. Id. No.120np.57. x1. 

_ PLATE 16. 
Fig. 1. Galaxzea lamarcki (Ed. and H.). Dredged, Felidu, 25 fms. 
Fig. 2. Galaxea musicalis (Linn.). Enlarged view of fig. 3. 
Fig. 3. Id. Typical form from N. Male. ; 
Hig. 4. Galaxea fascicularis (Linn.). Specimen from which polyps were taken, typical form. 
Fig. 5. Hchinopora gemmacea (Lam.). The fruticulosa type of growth. 
Fig. 6. Echinopora lamellosa (Esp.). Typical form (see Pl. 14, fig. 2). 
Fig. 7. Echinopora gemmacea (Lam.). No. 13 on p. 57. 
Fig. 8. Id. No. 10 on p. 57, irregular mode of growth. 


PLatTE 17. 


Fig. 1. Echinopora hirsutissima, Ed. and H. Tertia facies. x 2. 

Fig. 2. Echinopora gemmacea (Lam.). No. 140n p. 57. x 2. 

Fig. 3. Id. No. 13 on p. 57, enlarged view of Pl. 16, fig. 7. x 2. 

Fig. 4. Leptastrea roissyana, Hd. and H. No.1 onp.71. x 2. 

Fig. 5. Leptastrea ehrenbergana, Ed. and H. Enlarged view of fig. 7. x 2. 
Fig. 6 Id. No.9 on fl. eB. 

Big. 7. Id. Typical form. x 1. 

Fig. 8. Leptastrea solida (Ed. and H.). No. 13 on p.71. x1}. 

Fig.9. Id. No.10onp.71. x14. 


136 PERCY SLADEN TRUST EXPEDITION 


PLATE 18. 

Fig. 1. Leptastrea roissyana, Ed. and H. Typical form. Enlarged view of Pl. 19, fig. 1. x 2. 

Fig. 2. Leptastrea ehrenbergana, Ed. and H. No. 8 on p. 71, region with small corallites. x 2. 

Fig. 3. Leptastrea solida (Kd. and H.). No. 1lonp.71. x13. 

Fig. 4. Id. Typical form, two double corallites. Enlarged view of Pl. 19, fig. 6. x 2. 

Fig. 5. Id. No. 14 on p. 71, showing three giant corallites. x 2. 

Fig. 6. Id. No. 12 on p. 71, also with giant corallites. x 2. 

Fig. 7. Leptastrea ehrenbergana, Ed. and H. No. 8 on p. 71, region with large corallites. Cp. fig. 2. 

x 2. 

Fig. 8. Leptastrea solida (Kd. and H.). Figured type of Cyphastrea bottai, Ed. and H., Paris. x 3. 
PLATE 19. 

Fig. 1. Leptastrea roissyana, Kd. and H. ‘Typical form, enlarged on Pl. 18, fig. 1. x 3. 

Inge 2 dich INO, 2 @ajyos Wile 5< II 

Fig. 3. Leptastrea ehrenbergana, Kd. and H. No.6 onp. 71. x1. 

Fig. 4. Id. No.5 onp.71. x1. 

Fig. 5. Leptastrea solida (Kd. and H.). No.15onp.71. x1. 

Fig. 6. Id. Typical form, enlarged on Pl. 18, fig. 4. x 1. 
PuaTE 20. 

Fig. 1. Favia favus (Forsk.). From Red Sea, var. 2. x 1. 

Fig. 2. Id. From Seychelles, var. 1. x1. 

Fig. 3. Id. From Red Sea, var. 1, deep corallites and weak columella. x 1. 

Fig. 4. Id. From Red Sea, var. 2. x 1. 

Fig. 5. Id. From Red Sea, enlarged view of Pl. 32, fig. 1. x 1. 

Fig. 6. Id. From Red Sea, typical form. x 1. 

Fig. 7. Diploastrea heliopora (Lam.). From Minikoi, edge region. x 2, 

Fig. 8. Id. One of the types of Astrea heliopora, Lam.,= Heliastrea heliopora, Kd. and H. ~ 1. 
PLATE 21. 

Fig. 1. Favia favus (Forsk.). From Red Sea. Corallite rims distinct asin Group I. x 1. 

Fig. 2. Id. From Red Sea. Corallum with distorted growth due to unfavourable conditions. x 1. 

Fig. 3. Id. From Red Sea. Heavy corallum with thickened walls and small corallites, var. 2. x 1. 

Fig. 4. ?7Id. Gardiner’s F. versipora. Cp. Klunzinger’s figure of F. ehrenbergi. x 1. 

Fig. 5. Id. Type of Favia rousseawi, Ed. and H., Paris. x 1. 

Fig. 6. 2Id. Type of F. affinis, Ed. and H., Paris. x 1. 

Fig. 7. Id. One of the types of Fawia jacqwinote, Ed. and H., Paris. x 1. 

Fig. 8. Id. Type of Astrea denticulata, Lam., Paris. x 1. 


MATTHAI—RECENT COLONIAL ASTRAIDA 137 


PLATE 22. 


Figs. 1—4.  Fawa favus (Forsk.). Four types of Madrepora favus, Forskal, Copenhagen. Fig. 3 is a 
typical form ; fig. 4, belonging to var. 2, resembles Pl. 20, fig. 4. x 1. 


Fig. 5. Id. Type of Madrepora cavernosa, Forskal, Copenhagen. Typical form of F. favus. x 1. 
Fig. 6. Favia hululensis, Gardiner. Large specimen from Ceylon. x 1. 


Fig. 7. ? Favia berthollets (Val.). One of the types of Madrepora favus, Forskal, Copenhagen. Perhaps 
only a thin-walled F. favus. x 1. 


Fig. 8. Hawa doreyensis, Ed. and H. Type of Favia doreyensis, Kd. and H., Paris. (Cp. with Gardiner s 
Pl. 1xi, fig. 17.) x1. 


Fig. 9. Id. Specimen in Paris, named both F. amplior and F. ananas. Perhaps Lamarck’s type of 
Astrea ananas, var. amplioribus. x 1. 
PLATE 23. 


Fig. 1. Favia clouer (Val.). ~ One of the types of Fava okeni, Kd. and H., Paris. Corallites larger than 
usual. x 1. 


Fig. 2. Id. Type from Seychelles of Favia clowei, Kd. and H., Paris. x 1. 
Fig. 3. Fava versipora (Lam.). The larger type of Plesiastrwa versipora, Ed. and H., Paris. x 2. 
Fig. 4. Faia bertholleti (Val.). Type of Pavia bertholleti, Ed. and H., Paris. x 1. 


Fig. 5. Fava cloue: (Val.). Four small pieces from Ceylon, from which polyps were taken. Cp. with 
Klunzinger’s figure (PI. 11, fig. 6) of F. tubulifera. x 1. 


Fig. 6. Favia bertholleti (Val.). Four small pieces from Ceylon, from which polyps were taken; the 
lower on the right belongs to var. 1, while the two on the left belong to var. 2. x 1. 


PLATE 24. 
Fig. 1. Favia bertholleti (Val.). From Aldabra. x 1. 
Hig. 2. Favia pentagona (Esper). From Minikow. x 2. 
Fig. 3. Id. From Seychelles. x 2. ; 
Mig. 4. Id. From Minikoi. Gardiner’s Stephanocenia maldivensis. x 2. 
Figs.5 and 6. Fava laa (Klunz,). Two specimens from Red Sea. x 2. 
Mig. 7. Favia hirsuta (Hd. and H.). Gardiner’s Acanthastrwa hirsuta var. megalostoma. x 1. 


Fig. 8. Id. From Red Sea. x1. 


PLATE 25. 
Fig. 1. Favia acropora (Linn.). From Funafuti, Gardiner’s Orbicella orion. x 38. 


Fig. 2. Favia clouei (Val.). From Salomon. Cp. with Klunzinger’s figure (Pl. ii, fig. 4) of 
F. cavernosa. x 1. 


Fig. 3. Favia acropora (Linn.). From Salomon. x 3. 
Fig. 4. Favia wakayana (Gard.). From Rotuma. Gardiner’s Orbicella versipora. x 2. 
Fig. 5. Favia versipora (Lam.). From Rotuma. Gardiner’s Orbicella acropora. x 2. 


Fig. 6. Id. From Minikoi. Gardiner’s Orbicella annuligera, x 2. 
SECOND SERIES—ZOOLOGY, VOL. XVII. 18 


138 PERCY SLADEN TRUST EXPEDITION 


Fig. 7. Favia ananas (Ell. and Sol.). Specimen from Red Sea, from which polyps were taken. x 2. 
Fig. 8. Favia solidior (Ed. and H.). Type of Heliastrea solidior, Ed. and H., Paris. x 1. 
Fig. 9. Favia versipora (Lam.). The type of Heliastrwa laperouseana, Ed. and H., Paris. x 1. 


PLATE 26. 
Figs. 1 and 2. Favia hombroni (Rouss.). Two specimens from Salomon. x 2. 
Fig. 3. Favia halicora (Khr.). From Minikoi. Gardiner’s var. obtusa. x 1. 


Fig. 4. Favia acropora (Linn.). Large type from Indian Ocean of Heliastrea acropora, Kd. and H., 
Paris. x 3. 


Figs. 5—7. Favia halicora (Ehr.). Three specimens from Salomon, showing extent of variation, from 
thin-walled polygonal -corallites without surrounding grooves to oval or circular thick-walled 
corallites with distinct furrows. x 1. 


PLATE 27. 
Figs. 1 and 2. Fawa hemprichi (Ehr.). Two specimens from Seychelles. x 1. 


Fig. 3. Favia vasta, Klunz. From Egmont, Chagos. Cp. with Klunzinger’s figure (PI. iv, fig. 8) of — 
F. vasta var. superficialis. x 1. 

Fig. 4. Favia hemprichi (Ehr.). From Minikoi. Gardiner’s Acanthastrea hirsuta var.microstoma. x iL, 

Fig. Favia vasta, Klunz. From Salomon. Cp. with Klunzinger’s Pl. iv, fig. 12. x 1. 


5 
Fig. 6. Id. From Aldabra. Resembles Klunzinger’s Goniastrewa halicora var. obtusa (Pl. iv, fig. 2). x 1. 


PLATE 28. 
Fig. 1. Fava solidior (Kd. and H.). From Funafuti. Gardiner’s Orbicella coronata. x 1. 
Fig. 2. Favia favosa (Hl. and Sol.). From Red Sea. x 1. 
Gomastrea solida (Kd. and H.). From Hulule. x 3. 


3 

Fig. 4. Id. Specimen from Amirante (16 fms.), from which polyps were taken. x 3. 
5. Goniastrea planulata, Kd. and H. From Turadu. Gardiner’s Goniastrea pectinata. x 2. 
6 


Goniastrea pectinata (Khr.). From Red Sea, enlarged view of Pl. 37, fig. 1. x 2. 


PLATE 29. 


Favia abdita (Ell. and Sol.). Specimens showing extent of skeletal variation; figs. 2—4, Gardiner’s 
Prionastrea fusco-viridis. 


Fig. 1. From Singapore (cp. with Pl. 35, fig. 2, and Ellis and Solander’s Pl. 50, fig. 2). x1. 
Fig. 2. Id. From Goidu. x 1. 

Fig. 8. Id. From Minikoi. «x 1. 
Fig. 4. Id. From Minikoi. x 1. 


PLATE 30. 


Figs. 1—3. Favia complanata, Khr. Three specimens from Red Sea, showing imitation of Favia abdita 
in form of growth and mode of skeletal variation. Cp. with Pl. 29, fig. 1—4. x 1. 


Fig. 
Fig. 


MATTHAI—RECENT COLONIAL ASTRAIDA 139 


PLATE 31. 
1. Goniastrea retiformis (Lam.). From Rotuma. x 3. 


2. Id. From Seychelles. x 3. 


Figs. 3 and 4. Id. Two specimens from Minikoi. x 3. 


Fig. 
Fig. 
Fig. 
Fig. 


Fig. 


Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 
Fig. 


Fig. 
Fig. 


5. Id. From Red Sea, enlargement of Pl. 33, fig. 3. x 3. 
6. Goniastrea solida (Ed. and H.). From Seychelles, enlargement of Pl. 33, fig. 4. x 3. 
7. Goniastrea planulata, Hd. and H. Type of the species, Paris. x 1. 


8. Id. From Egmont, Chagos. x 1. 
PLATE 32. 
1. Fava favus (Forsk.). From Red Sea. x4. (PI. 20, fig. 5, same x 1.) 
2. Favia doreyensis, Kd. and H. From Seychelles. x 2. 
3. Id. From Minikoi, Gardiner’s Orbicella borradailer. x 2. 
4, Id. Type of Astrea rotulosa, Lamarck, Paris. x 1. 
PLATE 383. 
1. Favia acropora (Linn.). From Salomon; typical humpy mode of growth. x 1. 


2. Faia hombroni (Rouss.). From Salomon. x1. 


3. Goniastrea retiformis (Lam.). From Red Sea, typical, even, round or convex mode of growth ; 
enlarged on Pl. 31, fig. 5. x 3. 


4, Goniastrea solida (Ed. and H.). From Seychelles, enlarged on Pl. 31, fig. 6. x }. 


PLATE 34. 


Vertical sections of coralla: 


1. Favia clouei (Val.). Specimens on Pl. 28, fig. 5. x 1. 

2. Galazea lamarcki, Kd. and H. Specimen on Pl. 13, fig. 6. x 2. 

3. Galaxea fascicularis (Linn.). Specimen on Pl. 16, fig. 4. x 2. 

4. Cyphastrea microphthalma (Lam.). No. 14o0n p. 47. x 3. 

5. Cyphastrea gardineri, n. sp. Specimen on Pl. 13, fig. 5. x 3. 

6. Cyphastrea suvadive, Gard. Specimen on Pl. 13, fig. 3. x3. (Cp. with fig. 2.) 

7. LEchinopora hirsutissima, Ed. and H. Specimen on Pl. 17, fig. 1. x2. (Cp. with Gardiner’s 


Pl. lix, fig. 6.) 
8. Leptastrea ehrenbergana, Kd. and H. Specimen on PI. 17, fig. 6. x 2. 


9. Diploastrea heliopora (am.). Second specimen from Minikoi. x 13. 


PLATE 35. 
1. Favia hululensis, Gard. Type of Favia rotulosa, Ehrenberg (Mus. no. 739), Berlin. x 1. 


2. Favia abdita (Ell. and Sol.). Type of Astrea abdita, Ehrenberg (Mus. no. 729), Berlin. (Cp. 
with Pl. 29, fig. 1.) x1. 


18—2 


140 


Fig. 
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PERCY SLADEN TRUST EXPEDITION 


PLATE 36. 
.1. Faria favus (Forsk.). Type of Astrea deformis, Ehrenberg (Mus. no. 731), Berlin. x 1. 
.2. Id. Type of F. favus, Ehrenberg (Mus. no. 693), Berlin. x 1. 

. 3. Favia hemprichit (Khr.). Type of Astrea hemprichi, Ehrenberg (Mus. no. 735), Berlin. x 1. 
.4. Favia pentagona (Esp.). Type of Astrea melicerum, Ehrenberg (Mus. no. 734), Berlin. x 1. 
PLATE 37. 

Goniastrea pectinata (Ehr.). From Red Sea, enlarged on Pl. 28, fig. 6. x 2. 
Favia laza (Klunz.). From Red Sea, rounded form of growth, enlarged on Pl. 24, fig. 5. x 1. 
Fania versipora (Lam.). Type of Heliastreea annuligera, Kd. and H., Paris. x 3. 
Leptastrea roissyana, Ed. and H. Vertical section through specimen on PI. 19, fig. 1. x 2. 
Echinopora gemmacea (Lam.). Vertical section through No. 11 on p. 57. x 2. 
PLATE 38. 
.1. Cyphastrea serailia (Forsk.). Enlarged view of part of fig. 5 below. x 3. 


2. Goniastrea retiformis (Lam.). Enlarged view of part of fig. 4 below. (I have not personally 
examined this nor the next specimen.) x 2. 


3. Goniastrea solida (Ed. and H.). One of the types of Goniastrea favus, Klunz., in the Stuttgart 
Museum. x 2. 


4. Goniastrea retiformis. Another of the types of Goniastrwa favus, Klunz., in the Stuttgart 
Museum (probably Klunzinger’s Pl. iv, fig. 4). x 1. 


5. Oyphastrea serailia (Forsk.). Type of Favia microphthalma, Ehrenberg (ie 3 no. 713), Berlin. 
ale 


6. Galaxea fascicularis (Linn.). One of the types of <Anthophyllum fasciculare, Ehrenberg 
(Mus. no. 624), Berlin. x 1. 


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No. I —COLEOPTERA : CUCUJIDA, CRYPTOPHAGIDA. 
Par A. GROUVELLE. 


(Figures 1—6 dans le texte.) 


AVEC UNE DESCRIPTION DE LA LARVE ET DE LA NYMPHE DE 
PROSTOMINIA CONVEXIUSCULA GROUVELLE [CUCUJID A]. 


Par P. DE PEYERIMHOFF. 
(Figures A—F dans le texte.) 


- (Communiqué par M. Le ProresseuR J. Stantey Garpiner, M.A., E.R.S., F.LS.) 


Lu le 18 juin 1914. 


Létude des Cucujidee et Cryptophagide récoltés par la Perey Sladen Trust 
Expedition a donné 19 espéces reparties entre 10 genres: 9 genres et 17 espéces pour 
les Cucujidee, 1 genre et 2 espéces pour les Cryptophagide. Sur ces 16 espéces 8 sont 
nouvelles, 6 Cucujidee et 2 Cryptophagide; les autres appartiennent & des espéces qui, 
comme cela arrive si souvent pour les insectes de ces familles, ont une aire de dispersion 
considérable. 

Si nous passons en revue les divers genres retrouvés aux Seychelles nous arrivons aux 
constatations suivantes. 


CUCUJIDA. 


Lemophleeus. Genre répandu dans le monde entier. Une des espéces recueillies aux 
Seychelles, L. mirus Grouvelle, appartient 4 la faune tropicale et subtropicale de l'Afrique ; 
elle doit étre transportée dans les fruits et autres matiéres végétales et, d’aprés des 
constatations faites sur des insectes de l’Amérique centrale, semble en voie de dispersion plus 
considérable. La deuxiéme, L. propior, est spéciale au moins jusqu’d ce jour & la faune 
des Seychelles; elle ne peut servir en rien pour caractériser une faune. Elle est trés 
voisine d’especes de |’Inde et des Indes orientales. 

Oryzephilus. Genre créé récemment et & juste titre pour l’ancien Silvanus 
surmamensis Linné, espéce cosmopolite. 

Ahasuerus. Li’espéce rapportée, A. advena Waltl, est cosmopolite. 

Silvanus. Deux espéces; la 1°, S. scuticollis Walker, est répandue dans toutes 
les régions tropicales et subtropicales de l’ancien monde; elle a été retrouvée récemment 
aux Antilles. La 2™°, S. hebetatus Grouvelle, décrite de l Afrique orientale, semble 
appartenir plutot 4 la faune Africaine qu’a la faune Indienne. 

Monanus. Ce genre est plus spéciale & l’ancien monde; une des espéces récoltées, 
M. concinnulus Walker, est devenue cosmopolite; une deuxiéme, M. denticulatus, a été 
décrite sur des insectes provenant des Indes orientales; la troisitme, M. ornatus, est 
nouvelle, elle semble spéciale & la faune des Seychelles. 


142 PERCY SLADEN TRUST EXPEDITION 


Psammeecus. Les espéces de ce genre appartiennent toutes 4 l’ancien continent ; 
elles sont plus nombreuses dans les régions tropicales et subtropicales. Sur les quatre 
espéces récoltées, une en mauvais état semble voisine d’une forme spéciale 4 Madagascar ; 
une deuxiéme, P. semoni Grouvelle, se rencontre depuis les Iles Philippines jusqu’s 
Madagascar ; les deux derniéres, nouvelles, se retrouveront certainement dans les régions 
Inde et des Indes orientales; elles semblent plus éloignées des formes spéciales 4 
Madagascar, que des formes asiatiques. 

Cryptamorpha. L’espéce trouvée, Cryptamorpha desjardinsi Guérin, est cosmo- 
polite; elle est transportée dans les régimes de bananes. 

Inopeplus. Genre répandu dans les régions tropicales et subtropicales de l’ancien et 
du nouveau continent. L’espéce décrite, [. mimetes, est tres probablement spéciale & la 
faune des Seychelles; elle est voisine de I'L. pictus Cast. de Madagascar. 

Prostominia. Ce genre, décrit sur un insecte du Japon, P. lewis: Reitter, est 
représenté aux Seychelles par deux espéces nouvelles: P. scott, trés voisine de P. simont 
Grouvelle de Ceylan ; et P. convexiuscula, forme trés caractérisée au milieu des autres 
par le développement de son épistome. 


CRYPTOPHAGID A. 

Hapalips. Genre répandu dans les régions tropicales et subtropicales de l’ancien 
et du nouveau monde. Les Hapalips ont souvent des aires de dispersion considérables ; 
on ne peut a priori considérer les deux espéces nouvelles décrites comme complétement 
spéciales & la faune des Seychelles. 


Distribution géographique des espéces. 


eels Aldsbea ' 
Groupe Harqu. ar abra (incl. 
P dest Ges ieee nel: a ee Provenance des espéces déja décrites 

epee es Provi- Assomption 

dence) jet Cosmolédo) 
Cucusipm 
Lemophleus mirus ......... _— Afrique équatoriale 
Lzmophleus propior......... — 
Oryzephilus surinamensis — Cosmopolite 
Silvanus scuticollis .....:... — Tend a devenir cosmopolite 
Silvanus hebetatus ......... — Afrique orientale 
Monanus concinnulus _| — Cosmopolite 
Monanus denticulatus ...... —.- Archipel Malais 
Monanus ornatus ............ —- 
Ahasuerus advena ......... — Cosmopolite 
Psammeecus simoni ......... — Manille, Archipel Malais, Madagascar 
Psammecus letulus ......... — 
Psammeecus nitescens ...... — 
Psammicecus Sp. .........2--+- —. 
Cryptamorpha desjardinsi — Cosmopolite 
Inopeplus mimetes ......... = 
Prostominia scotti............ —. 
Prostominia convexiuscula — 
CRYPTOPHAGIDE 

Hapalips scotti ............... — 
Hapalips championi ......... — 


GROU VELLE—COLEOPTERA : CUCUJID®, CRYPTOPHAGIDA 143 


Cucujidz. Uleiotini. 


LamopHiaus Cast., Hist. Nat. Ins. Col., ii, 1840, p. 385. 


1. Lemophleus (Silvanophleus*) mrus, Grouvelle. 
Lemophleus mirus Grouvelle, Bull. Soc. Ent. France, 1905, p. 142. 


Le L. mirus, décrit sur des insectes provenant de Madagascar, a été retrouvé sur la 
edte occidentale de | Afrique, et récemment nous avons eu l'occasion d’examiner des 
exemplaires venant de |’Amérique centrale. Le ZL. mirus doit étre transporté dans les 
matiéres végétales ; il tend 4 devenir cosmopolite. 


2 exemplaires. 
Loc. “Seychelles. Mahé: Cascade Estate, ca. 800 feet, 1908—9.” 


2. Lemophleus propior, n. sp. 


Elongatus, subparallelus, depressus, nitidus, glaber ; capite prothoraceque rufo-piceis, 
antennis pedibusque dilutioribus, elytris piceis, singulo plaga maxima, testacea, vix perspicue 
piceo-infuscata, notato. Antennze apud marem elongatissimee, haud clavatee, apud feminam 
submoniliformes ; clava triarticulata ; 2° et 3° articulis subeequalibus. Caput transversum, 
depressum, tenuissime parceque punctulatum, antice sinuatum, ante antennarum bases 
arcuatim et fronte in longitudinem striatum ; labro sat magno. Prothorax subquadratus, 
utrinque in longitudinem striatus, parcissime tenuissimeque punctulatus ; angulis anticis 
acutis, prominulis, posticis obtusis, haud hebetatis, basi utrinque ad extremitatem sinuata. 
Scutellum subtriangulare, leve. LElytra elongato-ovata, apice subtruncata, ultimum 
segmentum abdominis haud obtegentia; in singulo elytro stria suturali impressa, fere 
integra, et leis punctulatis, vix perspicuis, juxta marginem apicalem magis indicatis. 
Long. 1°2—1°5 mill. 

Allongé, presque paralléle, déprimé, brillant, glabre; antennes et pattes roux testacé 
clair, téte et prothorax plus assombris, élytres brun de poix, marqués chacun d’une grande 
tache testacée, tres légérement teintée de nuance de poix ; dessous du corps brun de poix 
peu foncé, rougedtre dans la partie antérieure. Antennes atteignent chez le male presque 
la longueur du corps, sans massue, plus courtes chez la femelle, submoniliformes, terminées 
par une massue de trois articles lache, allongée, peu accentuée ; 2™° et 3™° article sub- 
égaux. Téte triangulaire, environ deux fois plus large que longue, déprimée sur le front, 
sinuée au bord antérieur, éparsement et trés finement pointillée ; strie interantennaire 
arquée, bien marquée, séparant le front de l’épistome, ce dernier se développant dans un 
plan un peu inférieur 4 celui du front; strie longitudinale du front marquée ; labre trés 
nettement visible. Prothorax subearré, 4 peine rétréci & la base, 4 peine plus large en 
avant que la téte chez le male, un peu plus étroit chez la femelle, strié de chaque cété, 
trés éparsement et trés finement pointillé; angles antérieurs aigus, saillants, postérieurs 
obtus, non émoussés ; base subtronquée au milieu, sinuée de chaque coté vers les extrémités, 
trés finement rebordée entre les stries longitudinales du disque. Ecusson subtriangulaire, 


* Silvanophleus Sharp, Biol. Centr.-Am., Col. 1m. 1, 1899, p. 537: treated as a subgenus of Lemophlaeus.— 
H. Scorr. 


144 PERCY SLADEN TRUST EXPEDITION 


environ deux fois plus large 4 la base que long, lisse. Elytres en ovale allongé, sub- 
tronqués au sommet, un peu plus larges 4 la base que la base du prothorax, subanguleux 
aux épaules, un peu plus de deux fois plus longs que larges ensemble, laissant en partie & 
découvert le dernier segment de l’abdomen, pliés sur les cétés; stries suturales bien 
marquées, effacées vers la base de l’élytre; lignes ponctuées du disque presqu’effacées sur 
le disque, serrées, bien visibles contre la marge apicale formée au sommet par la jonction 
de lintervalle suturale et de la marge latérale. Cavités des hanches antérieures fermées ; 
hanches antérieures, intermédiaires et postérieures écartées. Bord antérieur du dessous 
de la téte tronqué, bordé par une strie arquée. 

Voisin du L. proximus Grouvelle de la région indienne: distinct par sa coloration en 
partie brun de poix, et par la striation de l’extrémité des élytres marquée et serrée qui fait 
contraste 4 cdté de la striation du disque. 


Environ 35 exemplaires. 


Loc. ‘Seychelles. Silhouette: near Mont Pot-a-eau, ca. 1500 feet, VIII. 1908 ; 
forest just above the Mare aux Cochons plateau, over 1000 feet, IX. 1908. Mahé: Mare 
aux Cochons district, ca. 1500 feet, I.—II. 1909. 


In Silhouette most, if not all, the specimens were found in felled heads of an endemic 
palm, Verschaffeltia splendida. They were found among the still unopened central 
(i.e. youngest) leaves. I specially noted at the time that they were not present in 
growing trees, but only in decaying heads of felled trees, in which however the leaves 
were still green. Specimens of certain other species described in this paper were also 
found in these felled palm-heads: Monanus ornatus, Psammacus of perhaps more than 
one species, and Hapalips scottv.H. Scorr.” 


Silvanini. 
ORYZHPHILUS Ganglbauer, 1899, Kiif. Mitteleur., i, p. 584. 


3. Oryzephilus surinamensis (Linné). 


Dermestes surinamensis Linné, 1758, Syst. Nat., ed. X, p. 375. 

Silvanus surinamensis Latreille, 1807, Gen. Crust. Ins., iii, p. 20. 

Oryzephilus surinamensis Ganglbauer, 1899, Kif. Mitteleur., ili, p. 584; Reitter, 
1911, Faun. Germ,, i, p. 46; Grouvelle, 1912, Ann. Soc. Ent. France, Ixxxi, 
p. 318. 

Dermestes sexdentatus Fabricius, 1792, Ent. Syst., i, p. 232. 

Colydium frumentarium Fabricius, 1792, Ent. Syst., i, pars 2, p. 496. 

Silvanus bicornis Erichson, 1848, Naturg. Ins. Deutschl., ili, p. 337. 

Silvanus mercator Fauvel, 1889, Rev. d’Ent. Caen, viii, p. 132. 


Espéce cosmopolite. 3 exemplaires. 


Loc. “Seychelles. Silhouette: from near Mont Pot-d-eau, and from Mare aux 


Cochons, VIIJ.—IX. 1908. Mahé: Port Victoria.” 


GROU VELLE—COLEOPTERA : CUCUJIDAi, CRYPTOPHAGIDA 145 


AHASUERUS Gozis, 1881, Ann. Soc. Ent. France, 6. sér., 1, Bull., p. exxvu. 

4. Ahasuerus advena (Waltl). 

Cryptophagus advena Waltl, 1832, Faunus, p. 169. 

Silvanus advena Erichson, 1848, Naturg. Ins. Deutschl., 11, p. 339; Casey, 1884, 
Trans. Amer. Ent. Soc., xi, p. 73, pl. 4, fig. 6. 

Cathartus advena Reitter, 1876, in Harold, Col. Hefte, xv, p. 127; Ganglbauer, 1899, 
Kiaf. Mitteleur., iii, pp. 587 et 588. . 

Ahasuerus advena Gozis, 1881, Ann. Soc. Ent. France, 6. sér., 1., Bull., p. exxvii. 

Cryptophagus guerint Allibert, 1847, Rev. Zool., p. 12. 

Latridius museorum Ziegler, 1845, Proc. Acad. Philad., ui, p. 270. 

Cryptophagus striatus Rouget, 1876, Ann. Soc. Ent. France, 5. sér., vi, Bull, 
p- cevil. 

Espéce cosmopolite. 3 exemplaires. 

Loc. ‘Seychelles. Mahé: Cascade Estate, ca. 800 feet, 1908—9.” 


Sirvanus Latreille, 1807, Gen. Crust. Ins., ii, p. 19. 

5. Silvanus scuticollis, Walker. 

Silvanus scuticollis Walker, 1859, Ann. Mag. Nat. Hist., 3. ser., 11, p. 53; Grouvelle, 
1908, Ann. Soc. Ent. France, Ixxvii, p. 491. 

Silvanus triangulus Reitter, 1876, in Harold, Col. Hefte, xv, p. 60. 
. Cette espéce tend & devenir cosmopolite; décrite de Ceylan elle a été retrouvée 

dans tout l’Archipel Malais, au Japon, sur la céte d'Afrique, et récemment 4 la 

Guadeloupe. 

7 exemplaires. 


Loc. “Seychelles. Silhouette: near Mont Pot-a-eau, ca. 1500 teet, VIII. 1908 ; 
Mare aux Cochons and forest above, over 1000 feet, IX. 1908.” 


6. Silvanus hebetatus, Grouvelle. 

Silvanus hebetatus Grouvelle, 1912, Ann. Soc. Ent. France, 1xxxi, p. 339. 
Le type de l’espéce provient de l'Afrique occidentale. 

1 exemplaire. 

Loc. Seychelles. Silhouette: Mare aux Cochons, IX. 1908. 


Monanus Sharp, Trans. Ent. Soc. London, 1879, pp. 85, 86; Fauvel, 1903, Rev. d’Ent. 
Caen, xxii, p. 379. 


Emporius Ganglbauer, 1899, Kf. Mitteleur., il, p. 578 et 586. 
Subgen. Monanus, s. str. 
7. Monanus denticulatus, Grouvelle. 
Monanus denticulatus Grouvelle, 1912, Ann. Soc. Ent. France, lxxx1, p. 364. 
Le type de lespéce provient de |’Archipel Malais. 
SECOND SERIES—ZOOLOGY, VOL. XVII. 19 


146 PERCY SLADEN TRUST EXPEDITION 


1 exemplaire. 
Loc. “Seychelles. From Long Island, a small cultivated islet off the coast of 
Mahé, VII. 1908, 1 specimen.” 


8. Monanus concinnulus (Walker). 


Monotoma concinnula Walker, 1858, Ann. Mag. Nat. Hist., 3 ser., i, p. 207; 
(Grouvelle) 1908, Ann. Soc. Ent. France, Ixxvu, p. 489. 

Silvanus signatus Frauenfeld, 1867, Verh. zool.-bot. Ges. Wien, xvii, p. 438, 
Ol, U2, i BS 

Cryptamorpha fascuata Wollaston, 1874, Ent. Month. Mag., x, p. 169. 

Cathartus fascipennis Reitter, 1876, in Harold, Col. Hefte, xv, p. 129. 

Espéce cosmopolite. 10 exemplaires. 

Loc. ‘Seychelles. Silhouette: forest immediately above Mare aux Cochons, over 
1000 feet, IX. 1908. Mahé: Cascade Estate and forest above, ca. 800 feet and over; 
Mare aux Cochons district, ca. 1500 feet, I.—II. 1909.” 


Subgen. Monanops Grouvelle, 1912, Ann. Soc. Ent. France, Ixxxi, p. 344. 
9. Monanus (Monanops) ornatus, n. sp. (Fig. 1, elytra). 


Elongatus, parallelus, modice convexus, nitidus, flavo-pubescens, piceus ; antennarum 
basi, bucca pedibusque pisceo-testaceis, his ultimis paulo obscurioribus, elytris duabus 
maculis transversis ochraceis sectis: 17 macula a basi quam 2* latiore, 2* juxta suturam 
stricte interrupta. Antennz breves; 3° articulo parum elongato. Caput transversum, 
convexiusculum, subparce punctulatum ; oculis magnis, temporibus minutis, haud dentatis. 
Prothorax basi modice angustatus, paulo longior quam in maxima latitudine latior ; 
margine antico arcuato, medio stricte pulvinato; angulis anticis acutis exterius pro- 
minulis; lateribus bisinuatis, tenuissime crenulatis ; angulis posticis acutissimis, basi 
arcuata, marginata; disco dense punctulato. Scutellum transversissimum, apice late 
obtusum. Elytra apice conjunctim rotundata, circiter ter longiora quam simul latiora, 
dense lineato-punctata; intervallis 2°, 4°, et 6° (suturali non numerato) quam aliis 
paulo latioribus. Spatium axillare coxarum posticarum obtuse angulosum. Long. 
3—3°5 mill. 

Paralléle, environ cinq fois plus long que large, modérément convexe, brillant, couvert 
d'une pubescence flave, fine, médiocrement allongée, oblique, assez dense, ne 
masquant pas le tégument, brun de poix avec les antennes sauf les derniers 
articles, et la bouche testacées, teintées de nuance de poix, pattes un peu moins 
claires ; élytres coupés transversalement par deux bandes ochracées; la 1° 
prés de la base trés large, entiére, la 2™° au dela du milieu plus étroite, 
interrompue tres prés de la suture et n’atteignant pas les bords latéraux. 
Antennes plutot courtes, un peu plus allongées chez le male que chez la 


Fig. 1. femelle; 1° article cylindrique, plus long que large, 2™° et 3™° subégaux, 
Monanus or- yn peu plus longs que large, 4™° et 6™° subsphériques, presque plus 
natus, N. sp. » z : os me , A me 
Bem ale étroits que les cuts es TOURS, eS mam pu long que large, 7 subcarré, 

8™e transversal, plus étroit que 7™°, 9™° 4 11™° formant une massue environ 
trois fois plus longue que large, dont les deux premiers articles subégaux sont trans- 


GROUVELLE—COLEOPTERA : CUCUJIDA, CRYPTOPHAGIDA 147 


versaux et dont le dernier environ aussi long que large est émoussé 4 l’extrémité; 
Yensemble des articles 7 4 11 forme presquune massue progressive. Téte triangulaire, 
plus large que longue, un peu convexe, densement pointillée sur le front, saillante en 
forme de trapéze lisse en avant des naissances des antennes; yeux gros, échancrant 4 
peme les marges latérales du front; tempes trés petites. Prothorax médiocrement 
convexe, un peu rétréci a la base, assez fortement bisinué sur les c6tés, un peu plus long 
que large dans sa plus grande largeur, au niveau des angles antérieurs; bord antérieur 
arqué en avant, légérement rebordé en bourrelet au milieu; angles antérieurs algus, 
saillants latéralement; cotés trés finement denticulés surtout en avant; angles postérieurs 
en forme de petite saillie aigue; base arquée, rebordée; ponctuation fine, serrée, inter- 
valles 4 peine visiblement alutacés. cusson trés transversal, en angle largement obtus 
au sommet. Hlytres subtronqués a la base, arrondis aux épaules, arrondis ensemble au 
sommet, environ trois fois plus longs que larges ensemble dans leur plus grande largeur, 
densement ponctués en lignes; intervalles alternes, 4 partir du 3™° en comptant l’inter- 
valle sutural, un peu plus large que les autres; marges latérales pliées ; lignes ponctuées 
atténuées au sommet ; stries suturales bien marquées sur la moitié apicale de la longueur. 
1 segment de l’abdomen plus court que le métasternum. Espace axillaire des hanches 
postérieures s’avangant en angle obtus, largement émoussé sur le premier segment de 
labdomen. 
14 exemplaires. 


Loc. “Seychelles. Silhouette: from near Mont Pot-a-eau, ca. 1500 feet, VIII. 
1908; forest above Mare aux Cochons, over 1000 feet, one specimen being recorded as 
found among the leaves of a felled endemic palm, Verschaffeltia splendida (see under 
Lemophleus propior, p. 144).” 


Psammeecini. 
Psammascus Latreille, 1829, Regne Anim., 2° éd. v, p. 135. 


10. Psammecus sumont, Grouvelle. 

Psammecus simons Grouvelle, 1892, Ann. Soc. Ent. France, Ixi, p. 287; Col. Rég. 
Indienne, in Ann. Soc. Ent. France, Ixxvu, 1908, p. 476. 

Cette espéce a été décrite sur des exemplaires provenant des Iles Philippines; elle 
a été retrouvée depuis dans les Iles Malaises, la région indienne et 4 Madagascar. 

18 exemplaires. 

Loc. ‘Seychelles. Silhouette : Mare aux Cochons and forest above, over 1000 feet, 
TX. 1908. Mahé: from country above Port Glaud and near Morne Blane, a rather dry 
serubby area with a mixture of imported and endemic vegetation, 500—1000 feet, 
XI. 1908.” 


11. Psammecus sp. 
Un exemplaire a coloration incomplete. 


Loc. Seychelles. Silhouette, VIIT. 1908. 
19—2 


148 PERCY SLADEN TRUST EXPEDITION 


12. Psammecus letulus, n. sp. (Fig. 2, elytra). 

Oblongus, vix 2 et } longior quam latior, sat convexus, nitidulus, flavo-pubescens ; 
antennarum articulis 6—10 nigris; capite prothoraceque rufo-testaceis; elytris nigris, 
singulo ochraceo-testaceo bimaculato: 1* macula subbasilari, fere usque longitudinis 
medium extensa, latus subattingente, postice cum alterius elytri macula juncta; 2? 
oblonga, apicali, elongata, suturam fere attingente et usque latus extensa. Caput 
transversissimum. Prothorax basin versus valde angustatus, magis duplo latior quam 
longior, subdense punctatus, lateribus denticulis ex parte longioribus quam latioribus 
armatis. Elytra suboblonga, apice conjunctim rotundata, circiter sesquilongiora quam 
simul in maxima latitudine latiora, striato-punctata; intervallis, extra apicem, quam 
punctis latioribus. Long. 2°7 mill. 

Oblong, un peu moins de deux fois et demie plus long que large dans sa plus grande 
largeur, convexe, légérement déprimé sur le disque des élytres, brillant, 
couvert d’une pubescence flave, fine, assez dense et assez longue ne 
masquant pas la couleur du tégument. Antennes testacées; articles 
6 & 10 noirs; téte et prothorax roux testacés; élytres jaunes-testacés 
étroitement bordés de noir & la base, coupés transversalement vers le 
milieu de la longueur, par une bande noire s’avangant sur la suture, du 


cété de l’écusson, en angle trés aigu et du cdté du sommet en marge 
Fig. 2. étroite* ; bande transversale atteignant le bord latéral en s'élargissant 


Psammecus letu- prooressivement, se réflechissant ensuite contre ce bord pour le border 
lus, n. sp., elytra, 


AG étroitement jusqu’a |’épaule. Antennes allongées, s’épaississant faible- 


ment et progressivement vers lextrémité ; 3™° article environ une fois 
et demie plus long que large, 7™° 4 10™® subcarrés. Téte plus de deux fois et demie plus 
large que longue, faiblement convexe, couverte d’une ponctuation presque serrée, s'effagant 
vers l’épistome, front relevé, striolé de chaque cété entre l’ceil et la base de l’antenne, 
strié entre ces bases; épistome infléchi, environ deux fois plus large que long; labre 
petit; yeux saillants presqu’en forme de demi-cercle, 4 petites facettes. Prothorax 
& peine plus large que la téte dans sa plus grande largeur, fortement rétréci a la base, 
briévement arrondi aux angles antérieurs, d’abord subparalléle, puis convergent vers la 
base, presque deux fois plus large que long, en angle obtus aux angles postérieurs, couvert 
d'une ponctuation semblable a celle de la téte; cdtés armés chacun de six denticules 
étroits irréguliérement espacés, les 2™°, 3™° et 4™° a partir de la base un peu inégaux, un 
peu plus longs que larges; base tronquée, étroitement rebordée. Ecusson moins de 
quatre fois moins large que la base du prothorax. Elytres arqués 2X la base, briévement 
arrondis aux é€paules, alors environ deux fois plus larges que le prothorax 4 la base, 
ovales, faiblement élargis jusque dans les environs du milieu de la longueur, arrondis 
ensemble au sommet, environ une fois et demie plus longs que larges ensemble dans leur 
plus grande largeur, ponctués-striés; stries atténuées vers le sommet; intervalles des 
stries ponctuées plus larges que les points; stries suturales marquées Jusqu’au sommet ; 
marges latérales trés étroitement rebordées. Sillon latéral des hanches antérieures trés 


* The angular production forwards along the suture of the transverse black band is unfortunately not 
shown in the figure.—H. Scorr. 


GROUVELLE—COLEOPTERA: CUCUJIDAI, CRYPTOPHAGIDA 149 


accentué, sa marge antérieure lisse. Métasternum longitudinalement sillonné, ponctué 
sur la région extérieure de la base. Premier segment de l’abdomen subégal au méta- 
sternum; sa saillie entre les hanches postérieures aigue. Hanches intermédiaires 
contigues. 

Voisin de P. reittert Grouvelle, mais trés nettement plus large. 

4 exemplaires. 

Loc. ‘Seychelles. Silhouette: Mare aux Cochons, IX. 1908. Mahé: Cascade 
Estate, ca. 1000 feet, I. 1909.” 


13. Psammecus nitescens, n. sp. (Fig. 3, elytra). 

Ovatus, 2 et 4 longior quam latior, convexus, nitidus, flavo-pubescens ; antennarum 
articulis 6—10 nigris; capite prothoraceque fusco-rufis vel nigris; elytris ochraceis 
transversim nigro-maculatis, sutura strictissime, margine laterali deflexa, apiceque infus- 
eatis; elytrorum fascia nigra transversa postice latissime angulosa antice in suturam 
anguloso-producta. Caput transversissimum. Prothorax basin versus valde angustatus, 
eireiter in maxima latitudine duplo latior quam longior, crebre punctatus, lateribus 
denticulis ex parte longioribus quam latioribus armatis. Elytra suboblonga, apice 
conjunctim rotundata, fere duplo longiora quam simul in maxima latitudme latiora, 
striato-punctata ; intervallis quam punctis latioribus. Long. 2°5—2°7 mill. 

Ovale, environ deux fois et demie plus long que large dans sa plus grande largeur, 
convexe, & peine déprimé sur le disque des élytres, tres brillant sur les 
élytres, un peu moins sur la téte et le prothorax, couvert d’une pubes- 
cence flave, fine, assez longue, ne masquant pas la couleur du tégument. 
Antennes testacées, articles 6—10 noirs; téte et prothorax roux 
enfumé plus ou moins foncé, parfois noirs; élytres jaunes-testacés, 
coupés transversalement, un peu apres le milieu, par une bande noire, 
médiocrement large, atteignant les bords latéraux, s’élargissant de 


chaque cdté de la suture de l’intérieur 4 l’extérieur, formant 4 son bord Fig. 3. 
Psammecus nites- 


postérieur un angle trés obtus et se soudant 4 la tache apicale, saillante Be Fel ay tnt 

en angle aigue sur la suture 4 son bord antérieur*, marges latérales  , 16 | pa 

obseurcies entre les épaules et la bande transversale, trés étroitement 

rembrunies entre cette bande et le sommet, celui-ci assez largement rembruni. Antennes 
allongées, s'épaississant faiblement et progressivement vers l’extrémité ; 3™° article un peu 
moins d’une fois et demie plus long que large, 8™° & 10™® subcarrés. Téte environ deux 
fois plus large que longue, modérément convexe, couverte d’une ponctuation trés serrée, 
seffagant sur lépistome, celui-ci fortement infléchi, séparé du front par un pli arqué, 
briévement relevé de chaque cété entre l’ceil et la base de l’antenne ; épistome testacé, 
subrectangulaire, environ deux fois plus large que long; labre trés-petit ; yeux saillants 
presqu’en forme de demi-cercle, & petites facettes. Prothorax faiblement rétréci en avant, 
fortement 4 la base, & peu prés aussi large dans sa plus grande largeur que la téte au 
niveau des yeux, moins de deux fois plus large dans sa plus grande largeur que long, 
couvert d’une ponctuation trés serrée; bord antérieur arqué en avant dans le milieu, 


sinué de chaque cOté; angles antérieurs obtus; cétés arrondis, armés de six denticules 
* Cf. footnote on preceding page.—H. Scort. 


150 PERCY SLADEN TRUST EXPEDITION 


triangulaires dont le 3™° est un peu plus long que large 4 la base; base tronquée, 
rebordée en bourrelet au milieu. Ecusson environ trois fois moins large que la base du 
prothorax, rembruni. LElytres faiblement arqués 4 la base, brievement arrondis aux 
épaules, alors environ deux fois plus larges que le prothorax a la base, ovales, & peine 
visiblement élargis sur les cédtés, arrondis ensemble au sommet, presque deux fois plus 
longs que larges ensemble dans leur plus grande largeur, ponctués-striés, stries atténuées 
vers le sommet; intervalles des stries ponctuées plus larges que les points; stries 
suturales devenant contigues & la suture avant le sommet; marges latérales fortement 
infléchies, trés étroitement rebordées. Sillon latéral des hanches antérieurs réduit & une 
strie, sa marge antérieure lisse. Métasternum longitudinalement sillonné, ponctué sur la 
région extérieure de la base. Hanches intermédiaires contigues. Premier segment de 
Tabdomen mesure dans sa plus grande longueur plus long que le métasternum. 

18 exemplaires. 

Loc. “Seychelles. Silhouette: from near Mont Pot-a-eau, ca. 1500 feet, VIII. 
1908. Mare aux Cochons and forest above, over 1000 feet, IX. 1908*. Mahé: Cascade 
Estate, ca. 800 feet.” 


CrypraMorPHA Wollaston, 1854, Ins. Mader., p. 156. 


14. Cryptamorpha desjardinsi (Guérin). 

Psammecus desjardinsi Guérin, 1838, Iconog. Regne Anim., Ins., p. 196. 

Cryptamorpha desjardinsi C. O. Waterhouse, 1876, Ent. Month. Mag., xin, p. 122; 
Casey, 1884, Trans. Amer. Ent. Soc., xi, p. 104, pl. 8, fig. 8; Grouvelle, 1908, Ann. Soc. 
Ent. France, Ixxvu, p. 474. 

Dendrophagus suturalis White, 1846, Voy. Ereb. Terr., Ent., p. 18; (Broun) 1880, 
Manual N. Zeal. Col., p. 222. 

Cryptamorpha muse Wollaston, 1854, Ins. Mader., p. 157, of A, fig. 1. 

Pseudophanus signatus Leconte, 1859, Proc. Acad. Philad., p. 85. 

Telephanus fascoatus Redtenbacher, 1867, Reis. Novara, u, p. 41. 

Cosmopolite. Transporté avec les bananes. 

6 exemplaires. 

Loc. “‘Seychelles. Mahé: 5 of the specimens are from country near Morne Blanc 
and above Port Glaud, a rather dry scrubby area with a mixture of imported and endemic 
vegetation, 500—1000 feet, XI. 1908; the sixth is from the district of Mare aux Cochons, 
ca. 1500 feet, I.—II. 1909.” 


Inopeplini. 
Iyopepius Smith, List. Col. British Museum, 1851, p. 4. 


Ino Cast., Etud. Ent., 1835, p. 135. 
Huryplatus Motsch., Etud. Ent., vii. 1859, p. 95. 
Pseudino Fairm., Ann. Soc. Ent. France, 4 sér., ix. 1869, p. 208. 


* In Silhouette certain Psammecus were taken among the leaves of felled Verschaffeltca-palms, but I have 
no record of exactly which species were thus found. See p. 144, under Lemophleus propior.—H. Scort. 


GROU VELLE—COLEOPTERA: CUCUJIDA, CRYPTOPHAGIDA 151 


15. Inopeplus mimetes, n. sp. 


Elongato-ovatus, depressus, nitidus, glaber, nigro-piceus; antennis rufo-piceis, 
preecipue apicem versus modicissime infuscatis; pedibus quam antennis dilutioribus; 
singulo elytro ad basin oblique et juxta apicem stricte testaceo-maculato, callo humerali 
nigro. Antenne subelongatze ; 2° articulo sesquilongiore quam latiore, quam 3° breviore, 
articulis 4°—10° subquadratis. Caput convexiusculum, sat dense punctatum, antice 
juxta antennarum bases transversim truncatum, medio productum et parallelum, apice 
truncatum. Prothorax transversus, cordiformis, plus minusve parce punctatus ; lateribus 
undulatis, obtusissime dentatis. Elytra apicem versus ampliata, longiora quam simul in 
maxima latitudine latiora, levia. Long. 2°5—3°5 mill. 

Ovale, environ quatre fois plus long que large dans sa plus grande largeur, déprimé, 
glabre, brillant, brun noiratre ; antennes roux testacé, lég¢rement assombries, surtout vers 
Yextrémité ; pattes roux testacé; chaque élytre marqué & la base d’une tache testacée, 
oblique, partant de l’épaule, et au sommet dune étroite bordure de méme couleur; calus 
huméraux bruns. Antennes assez allongées chez le male, plus courtes chez la femelle ; 
1 article plus long que large, arqué en dehors, 2™° plus long que large surtout chez le 
male, plus court que le 3™°, 4™° et 5™° subégaux nettement allongés chez le male, un peu 
allongés chez la femelle, 6™° plus long que 5™° et 7™°, 7™° & 11™° progressivement et trés 


x 


faiblement é€paissis, 7™° & 10™° nettement allongés chez le male, suballongés chez la 
femelle, 11™° & peine plus long que le précédent. Téte & peine plus courte que large, 
rétrécie transversalement en avant des naissances des antennes, puis subparallele et 
tronquée au bord antérieur; front légérement convexe, plus ou moins densement ponctué, 
séparé de l’épistome par une strie; épistome légérement infléchi, environ trois fois plus 
large que long, marqué de quelques poimts; labre bien visible, arrondi au sommet, 
pointillé ; yeux latéraux, saillants; tempes allongées. Prothorax cordiforme, plus étroit 
dans sa plus grande largeur que la téte, méme chez la femelle, fortement rétréci 4 la base, 
moins de deux fois plus large dans sa plus grande largeur que long, plus ou moins 
éparsement ponctué; bords latéraux largement et trés obtusement dentés; base sub- 
tronquée, rebordée, denticulée aux extrémités. Hcusson petit, subdemicirculaire. Elytres 
tronqués & la base, arrondis aux épaules, alors & peu prés aussi larges que le prothorax 
dans sa plus grande largeur, s’élargissant presqu’en ligne droite vers le sommet, arrondis 
séparément a l’extrémité, nettement plus longs que larges dans leur plus grande largeur, 
presque deux fois plus larges dans leur plus grande largeur qu’ la base, lisses. Hanches 
antérieures faiblement écartées, postérieures médiocrement. Hspéce voisine comme 
ensemble de T. pictus Cast. de Madagascar. 


10 exemplaires. 


Loc. ‘Seychelles. Silhouette: from high forest, near Mont Pot-a-eau, ca. 1500 
feet, and above Mare aux Cochons, VIIJ.—IX. 1908. Mahé: forest above Cascade 
Estate, ca. 1000 feet, 1908—9.” 


152 PERCY SLADEN TRUST, EXPEDITION 


Prostominini. 


L’étude de la larve du Prostominia convexiuscula Grouvelle, que M. de Peyerimhoff 
a bien voulu donner dans ce mémoire, montre l’opportunité de |’établissement de cette 
nouvelle tribu, qui au milieu des Cucujidse comprend des formes relativement anciennes*. 

La nouvelle tribu des Prostominini nous semble avoir certains rapports avec les 
Xenoscelis. 


ProsrominiA Reitter, 1889, Wien. Ent. Zeit., vili, p. 315. 

16. Prostominia scott, n. sp. (Fig. 4). 

Elongata, subparallela, subdepressa, nitida, glabra, atra ; antennis, bucca pedibusque 
nigro-piceis. Caput fronte fere depressum, in disco parce subtiliterque utrinque fere dense 
punctatum. Prothorax antice vix, postice sat valde angustatus, lateribus arcuatus, vix 
amphatus, paulo longior quam latior, parce subtiliterque punctatus, marginibus lateralibus 
subabrupte inflexus; angulis omnibus obtusis; basi truncata. Scutellum semicirculare, 
leve. Elytra humeris breviter rotundata, apice conjunctim stricte rotundata, circiter 
3 et 4 longiora quam simul latiora, striato-punctata; striis circiter post medium evanes- 
centibus, punctis apicem versus attenuatis ; striarum intervallis in disco quam punctis 
duplo latioribus. Long. 4—4°5 mill. 

Subparalléle, environ cinq fois et demie plus long que large, subdéprimé, brillant, 
glabre, noir avec les antennes, la bouche et les pattes bruns de poix. 
Antennes atteignant environ le milieu de la longueur du prothorax 
chez le male, un peu plus courtes chez la femelle: massue environ 
deux fois plus épaisse que les articles précédents. Téte un peu plus 
longue que large, subdéprimée sur le front, infléchie faiblement en are 
et déprimée en avant des bases des antennes, éparsement et tres 
finement pointillée sur le disque, chargée sur les marges latérales du 
front de points plus forts, un peu allongés, parfois formant presque des 
strioles, dessmant presque des lignes longitudinales, épistome presque 
lisse, trés largement arrondi au bord antérieur ; tempes moins de deux 
fois plus longues que le diametre longitudinal de l’ceil. Prothorax a 


peine rétréci en avant, assez fortement & la base, arrondi sur les cétés, 


Bee surtout dans la partie basilaire, 4 peine élargi, un peu plus long que 
AO ca ee large, & peine plus étroit dans sa plus grande largeur que la téte avec 
n. sp., x 16. 


les yeux, éparsement et trés-finement pointillé, déprimé sur le disque, 
brusquement plié sur les marges latérales ; angles antérieurs obtus, postérieurs encore plus 
ouverts; base tronquée, marge basilaire coupée longitudinalement par un sillon 4 peine 
marqué. Hcusson presqu’en forme de demi-cercle, lisse. Elytres tronqués & la base, briéve- 
ment arrondis aux épaules, & peine arqués et élargis sur les cotés, brievement arrondis au 
sommet, environ trois fois et demie plus longs que larges ensemble dans leur plus grande 
largeur ; ponctués-striés surtout sur la moitié basilaire, puis ponctués en lignes et enfin 
points s'atténuant vers l’extrémité; intervalles des stries presque plans, environ deux 
* Voir étude de M. de Peyerimhoff, p. 156. 


GROUVELLE—COLEOPTERA : CUCUJIDA, CRYPTOPHAGIDA 153 


fois plus larges que les points; stries suturales entitres, bien marquées vers le sommet ; 
marges latérales brusquement pliées. 

9 exemplaires. 

Loc. ‘Seychelles. Mahé: from forest near Morne Blanc, XI. 1908.” 

17. Prostominia conveaxiuscula, n. sp. (Fig. 5, téte). 

Elongata, subparallela, convexiuscula, nitida, glabra, atra ; femoribus tiblisque nigro- 
piceis, antennis tarsisque dilutioribus. Caput fronte subdepressum, in disco parce subtili- 
terque, utrinque densius, punctatum, inter antennarum bases biimpressum; epistomo 
valde producto. Prothorax postice angustatus, lateribus subrectus, transversus ; margine 
antico medio truncato, ad extremitates arcuato, angulis anticis posticisque obtusis ; basi 
arcuata ; disco przecipue ad latera subdense et quam capite densius punctulata. Scutellum 
semicirculare, testaceo-piceum. Hlytra humeris breviter rotundata, apice conjunctim 
rotundata, paulo magis 2 et $ longiora quam simul latiora, striato-punctata ; striis fere 
integris, intervallis in disco quam punctis vix latioribus. Long. 3—3°5 mill. 

Subparalléle, environ quatre fois et demie plus long que large, faiblement convexe, 
glabre, brillant, noir; tibias et fémurs bruns de poix, antennes, tarses et 
écusson, surtout les deux derniers, plus clairs. Antennes courtes méme 
chez le male; massue moins de deux fois plus épaisse que les articles 
précédents. Téte plutot plus large que longue, faiblement convexe sur le 
front, sans infléxion a la base de l’épistome, biimpressionné entre les 
naissances des antennes, éparsement et tres finement pointillée sur le 
disque, chargée sur les marges latérales du front de points plus forts ; 


épistome lisse, trés saillant; front longitudinalement substrié ; tempes Fig. 5. 
au moins égales au double du diametre longitudinal de lceil. Prothorax Prostomania 
goo § : R R vexiuscul 
rétréci 4 la base, presque droit sur les cédtés, assez fortement arqué en = “msm 
n. Sp. 


avant au bord antérieur, arqué 4 la base, environ une fois et demie plus 
large dans sa plus grande largeur que long dans sa plus grande longueur, un peu plus 
étroit dans sa plus grande largeur que la téte avec les yeux ; en angle obtus 4 tous les 
angles, éparsement et finement pointillé, coupé dans la longueur, surtout vers la base, par 
une trés vague impression; marges latérales brusquement pliées, sauf contre langle 
antérieur; bords latéraux vers le c6té droits, fortement arrondis vers la base, celle-ci 
tronqué, tous les deux finement rebordés. Hcusson presque demi-circulaire, lisse. Elytres 
tronqués 4 la base, arrondis aux épaules, subparalléles, arrondis ensemble au sommet, un 
peu plus de deux fois et demie plus longs que larges ensemble, ponctués-striés, presque 
jusqu’a lextrémité ; intervalles des stries 4 peine plus larges sur le disque que les points 
des stries ; marges latérales et apicales assez brusquement infléchies-pliées. 

94 exemplaires. 

Loc. Seychelles. Silhouette, Mahé. 

“The great majority of this long series were found in Silhouette, in the forest 
immediately above Mare aux Cochons plateau, under the bark of fallen twigs of an 
endemic forest-tree, the ‘Bois Rouge’ (Wormia ferruginea): they were taken thus 
several times between 6. IX. and 19. IX. 1908, and on certain occasions the larve and 

SECOND SERIES—ZOOLOGY, VOL, XVII. 20 


154 PERCY SLADEN TRUST EXPEDITION 


pupee discussed on pp. 156—159 of this memoir were found in company with adults. In 
Mahé 9 specimens were collected, all from the high damp forest: Morne Blane and 
Pilot, XI. 1908; between Trois Fréres and Morne Seychellois, 1500—2000 feet, XII. 
1908 ; above Cascade Estate, I. 1909.—H. Scort.” 


Cryptophagide. Xenoscelini. 


Hapauies Reitter, 1877, Verh. nat. Ver. Briinn, xv, p. 122 (Rhizophagide) ; 
Gorham, 1898, Biol. Centr.-Amer., Col. vu, p. 250 (Erotylidee); Grouvelle, 1908, Ann. 
Soc. Ent. France, xxvii, p. 58 (Cryptophagide); Champion, 1913, Trans. Ent. Soe. 
London, p. 96 (Cryptophagidee). 


18. Hapalips champion, n. sp. 


EHlongato-ovatus, modice convexus, nitidus, glaber, fulvo-testaceus antennis pedi- 
busque dilutioribus. Caput transversum, convexiusculum, subdense punctulatum, utrinque 
ad antennz basin oblique tenuiterque striolatum, temporibus retrorsum divergentibus, 
minimis, angulis posticis vix hebetatis. Prothorax modice transversus, lateribus extra 
extremitates vix arcuatus ; angulis anticis rotundatis, posticis acutis; basi medio arcuato- 
producta, utrinque sinuata; lateribus basique tenuiter marginatis. Elytra basi subparallela, 
dein apicem versus attenuata, apice separatim breviter rotundata, striato-punctata ; 
punctis apicem versus attenuatis et evanescentibus; striis suturalibus paulo ante 
apicem magis impressis, intervallis striarum in disco quam punctis paulo latioribus. 
Mesosternum subparce punctulatum. Metasternum in longitudinem vix perspicue 
. Sulcatum, subdense punctulatum. Segmenta abdominis parce et tenuissime punctulata. 
Long. 3°5 mill. 

Ovale, un peu moins de cing fois plus long que large dans sa plus grande largeur, 
modérément convexe, brillant, glabre, testacé, temté de nuance de poix; antennes et 
pattes plus claires. Antennes atteignant 4 peine le milieu du prothorax; 3™° article 
subcarré, 4™°, 6™* et 8™° plus courts que les articles voisins, 7™® et 8™° un peu plus larges 
que les articles précédents ; massue brusque, environ deux fois plus large dans sa plus 
grande largeur que les articles 7 et 8. Téte subtriangulaire, environ deux fois plus 
large que longue, faiblement convexe, couverte d’une ponctuation fine, presque dense, 
satténuant en avant, infléchie, arquée en avant des bases des antennes, trés-largement 
arrondie au bord antérieur, yeux plus de deux fois plus longs que larges; tempes petites 
convergentes en avant, angles postérieurs & peine émoussés. Prothorax médiocrement 
transversal, subparalleéle, trés-faiblement arqué sur les cétés, sauf aux extrémités, briéve- 
ment arrondi aux angles antérieurs, briévement arqué, sinué en avant des angles 
postérieurs, ceux-ci aigus, presque saillants; base arquée en arriére dans le milieu ; 
bords latéraux et base étroitement rebordés. Ecusson subpentagonal, plus de deux fois 
plus large que long. Elytres un peu plus larges & la base que la base du prothorax, 
subdentés aux épaules, subparalléles sur plus de la moitié basilaire, puis atténués vers 
Yextrémité, bri¢vement et séparément arrondis au sommet, environ trois fois et demie 
plus longs que larges ensemble 4 la base, ponctués-striés ; stries atténuées, puis effacées 
vers le sommet; stries suturales mieux marquées vers l’extrémité, trés atténuées sur 


GROUVELLE—COLEOPTERA: CUCUJIDM, CRYPTOPHAGIDA 155 


lextréme marge apicale; intervalles des stries discoidales plus larges que les points; 
bords latéraux bordés par une marge concave, étroite & la base, sélargissant faiblement 
vers le sommet et s'effacant un peu avant de l’'atteindre. Mésosternum & peine visiblement 
alutacé, subéparsement ponctué. Métasternum A peine visiblement sillonné dans la 
longueur, presque densement ponctué. Segments de l’abdomen éparsement et trds fine- 
ment ponctués surtout vers l’extrémité. Lignes fémorales du premier segment courtes, 
divergentes. 


3 exemplaires. 


Loc. ‘Seychelles. Praslin: from between the leaf-bases of a growing endemic 
palm-tree, a ¢ Lodoicea sechellarum (‘Coco-de-mer’), in the Vallée de Mai, Cotes d’Or 
Estate, 28, XI. 1908.” 


19. Hapalips scotti, n. sp. (Fig. 6). 


Elongatissimo-ovatus, modice convexus, nitidus, glaber, piceus, antennarum basi 
et ultimo articulo pedibusque dilutioribus. Caput transversum, convexiusculum, parce 
punctulatum, punctis apicem versus attenuatis, temporibus parallelis, minimis, angulis 
posticis hebetatis. Prothorax subquadratus, capite paulo latior, subparce punctulatus ; 
lateribus basique tenuiter marginatis. Elytra apicem versus attenuata, tenuiter punctato- 
substriata, punctis ad apicem attenuatis; stria suturali ad apicem magis impressa, 
intervallis striarum in disco quam punctis latioribus, juxta apicem haud separatis et 
calloso-elevatis. Mesosternum alutaceum, subparce punctatum. Metasternum alutaceum, 
subparce punctulatum. Segmenta abdominis tenuiter alutacea et parce tenuissime 
punctulata. Long. 3—4 mill. 

Ovale, plus de cing fois plus long que large dans sa plus grande largeur, modérément 
convexe, brillant, glabre, brun de poix avec la base et le dernier article 
des antennes et les pattes plus clairs. Antennes atteignant 4 peine le 
milieu de la longueur du prothorax; 3™° article subcarré; 4™°, 6™° et g™me 
plus courts que les articles voisins; 7™° et 8™° trés faiblement plus larges 
que les précédents ; massue brusque, un peu moins de deux fois plus large 
que larticle 8. Téte modérément transversale, faiblement convexe, 
couverte d’une ponctuation presque dense, trés fine & la base, s’atténuant 
vers l’épistome, celui-ci infléchi, tronqué au bord antérieur ; yeux plus 
de deux fois plus longs que larges ; tempes paralleles, courtes; angles 
postérieurs émoussés. Prothorax subcarré, trés légérement arqué sur les 
cdotés, couvert d'une ponctuation dense et trés fine, entremélée de points 
encore plus fins visibles 4 un fort grossissement, légérement relevé au 


te pe ss 3 Fig. 6. 
milieu de la base ; angles antérieurs et postérieurs faiblement obtus, les 2 
: : 2 p Hapalips scotti 
premiers & peine émoussés; cdtés et base trés finement rebordés. Ecusson a a a 
spe : 


environ deux fois plus large que long, subpentagonal. Elytres bri¢vement 
arrondis aux épaules, progressivement et faiblement atténués vers lextrémité, trés briéve- 
ment arrondis séparément au sommet, environ quatre fois plus longs que larges 4 la base, 
fnement ponctués-striés ; stries suturales enfoncées vers le sommet; stries discoidales 
anténuées vers lextrémité, effacées au sommet; marges latérales fortement infléchies, 


20—2 


156 PERCY SLADEN TRUST EXPEDITION 


bordées par une strie bien marquée, limitant en dedans une marge explanée trés étroite, 
un peu plus large au sommet; disque de chaque élytre terminé au sommet par un calus 
allongé, subacuminé 4 lextrémité, limité en dedans par le fort enfoncement de la strie 
suturale, se raccordant en dehors et au sommet en pente trés brusque avec le rebord 
marginal. Mésosternum et métasternum finement alutacés, éparsement et finement 
ponctués, le premier plus fortement que le second. Segments de labdomen 4 peine 
visiblement alutacés, éparsement et trés finement pointillés. Lignes fémorales du premier 
segment légérement divergentes, atteignant 4 peine le milieu de la longueur du segment. 

28 exemplaires. 

Loc. Seychelles. Silhouette, Mahé. 

“Tn Silhouette (VITI.—IX. 1908) a number of specimens were beaten from a dead leaf 
of a coconut-palm still hanging to the tree, at the edge of the Mare aux Cochons plateau ; 
others were taken in the forest above Mare aux Cochons, one being recorded as found 
among leaves of a felled Verschaffeltia-palm (see under Lemophleus propior, p. 144); 
another was found between leaf-bases of a growing endemic palm (species unrecorded) in 
the high damp forest near Mont Pot-d-eau. In Mahé two specimens were taken between 
leaf-bases of a growing endemic palm (Stevensonia) near Morne Blanc, X.—XI. 1908; 
several others were collected in the forest above Cascade Estate.” 


DESCRIPTION DE LA LARVE ET DE LA NYMPHE DE PROSTOMINIA CONVEXIUSCULA 
GROUVELLE*, [COLEOPTERA, CUCUJIDA]. PAR P. DE PEYERIMHOFF. 
(FIGURES A—F DANS LE TEXTE) 


Matériel étudié: (1) quatre larves de méme Age, conservées dans l’alcool faible ; 
toutes ont les téguments distendus et sont proches d'une mue: la cuticule déja soulevée 
laisse apercevoir le stade suivant, & peine chitinisé.—(2) une nymphe déformée par le 
séjour dans lalcool.—(3) un imago. 

Provenance (d’aprés une étiquette manuscrite jointe) : “Seychelles. Silhouette : Mare 
aux Cochons, from under bark of fallen twigs of ‘ Bois Rouge’ (Wormia ferruginea), IX. 
1908 (see p. 153).” 

Longueur de la larve la mieux conservée: 3°5 mill.—Largeur: 0°4 mill. 

Corps (fig. A) trés allongé, paralléle, déprimé, luisant, presque lisse, jaunatre ; 
mandibules et dernier segment d’un roux foncé.  Ocelles noirs. Soies claires, peu 
nombreuses et courtes. 

Téte & peine enchassée dans le prothorax, légérement élargie en arriére, puis un peu 
rétrécie & la base. Sutures craniennes jalonnées par une assez large dépression en V. 
Epistome défini par une ligne postérieure nette. Labre (fig. C) membraneux, saillant 
en are de cercle, et portant de fines soies. 

Antennes relativement courtes, basées sur une articulation membraneuse trés 
détachée. 1° article carré, 2™° subcylindrique et sensiblement plus long, 3™° deux fois 
plus court et plus mince que le précédent, prolongé par une longue soie; cone sensitif 
de méme longueur et de largeur presque égale. 

* Voir pp. 152—154. 


GROUVELLE—COLEOPTERA: CUCUJIDA, CRYPTOPHAGIDA Wa7 


Mandibules (fig. C) larges et courtes, saillantes, bidentées au sommet, avec une 
mola distincte, denticulée de profil. 

Pleures céphaliques (fig. FZ) non contigues, creusées dans leur moitié antérieure en 
forme de rectangle émoussé, pour recevoir les piéces buccales dont l'ensemble est plat. 

Mazilles formées d’un stipe robuste, peu incliné, prolongé sans suture par un lobe 
presque rectangulaire, armé & l’intérieur et surtout au sommet d’un double peigne de 
spinules. Palpes mamillaires dépassant beaucoup le labium, 4 articles diminuant en 


Figs. A—F. Prostominia convexiuscula Grouv. 


Larve, vue en dessus. 

Nymphe, vue en dessus. 

Mandibule droite (légerement détachée) et labre, vus en dessus. 
Les deux derniers segments de l’abdomen, vus de profil. 

Téte et prothorax, vus en dessous. 


S&SQh6h 


Derniers segments de l’abdomen, vus en dessous. 


largeur et croissant en longueur, les deux premiers aussi longs que larges, le dernier 
presque une fois et demie plus long que le précédent. Cardo visible en partie, enraciné 
sous la membrane articulaire qui est longuement ovale et rejoint le menton vers la 
moitié. 

Labiwm composé d’avant en arriére: (1) d'un palpigére cordiforme portant sur des 
socles assez détachés un palpe court, 4 2™° article cylindrique terminé par une papille ; 


158 PERCY SLADEN TRUST EXPEDITION 


au milieu, une languette arrondie, portant deux soies terminales.—(2) d’un menton 
subearré, orné de deux soies.—(3) d’un sous-menton en forme d’enclume, entiérement 
contigu 4 la membrane articulaire des maxilles et s'appuyant en arriére sur une gula de 
contour semblable, mais plus large, qui fait le pont entre les gene. 

Segments thoraciques & cdtés dorsaux peu curvilignes, présentant chacun en leur 
milieu un sillon longitudinal trés prononeé, et munis latéralement d'une assez longue soie 
insérée sur la pleure. Pronotum en rectangle transversal de méme largeur que la téte, 
portant quatre soies en avant et en arriére. Mésonotum et métanotum légérement plus 
courts et plus larges, portant deux soies antéro-latérales et quatre soies postérieures. 

Segments abdominaux transversaux, a cotés eurvilignes subanguleux, de longueur 
croissante du 1° au 5™ inclusivement ; 6™¢ et 7™° subégaux, 8™° plus long et moins large, 
en trapeze renversé; chacun portant des soies & peu prés disposées comme sur le méta- 
notum. 9™ segment (fig. /’) carré dans son ensemble, trés chitinisé, rugueux, convexe sur 
sa moitié antérieure, puis profondément déclive, chevronné sur les cdtés, et terminé par 
quatre processus chitineux, les latéraux bidentés a dent externe tres relevée (fig. D), les 
médians en forme de crochets affrontés vers la ligne médiane, ot ils enclosent une espace 
circulaire. 

Dessous du corps déprimé, luisant et de méme consistance que le dessus. Pleures 
détachées. Sternite du 9™ segment trés saillant en dessous, débordant en avant sur 
le 8™e, armé de chaque cdté vers la base de deux papilles chitineuses aigues. Anus 
faisant suite vers l'arriére, précédé et suivi de quatre soies. 

Stigmates normaux, la premiére paire grande, saillant entre les deux premiers 
segments du thorax; les autres paires latérales, non saillantes, situées au premier tiers 
des pleures, sur les 8 premiers segments de l’abdomen. 

Pattes distantes, courtes, portant quelques rares spinules. Sanches globuleuses, 
trochanters plus courts que les fémurs, égaux en longueur aux tebias, qui sont plus 
minces, trongoniques, et terminés par un ongle aigu a base membraneuse. 

Cette larve a le facies et la structure de toutes celles connues dans la famille des 
Cucwjide: corps long, aplati, terminé par des processus compliqués; organes buccaux 
inférieurs, mais non saillants, et relativement réduits, c’est-a-dire laissant, entre le trou 
occipital et le niveau du cardo, une gula et des genz étendues. 

La forme épanouie et la structure membraneuse du lobe maxillaire rappellent 
beaucoup le type Znopeplus (cf. Peyerimhoff, in Annales de la Société entomologique de 
France, 1902, p. 715). Mais Prostominia s'en distingue profondément par le développe- 
ment plus accentué des piéces buccales, le 8™° segment plus court, le 9™° beaucoup 
plus long, terminé par un systéme chitineux totalement différent. Tous ces caractéres 
marquent a priori une moindre différenciation. Au cas ot la structure de Vimago 
inclinerait 4 rapprocher ces deux genres, c'est en tous cas Prostominia qui représenterait 
la forme phylétiquement la plus ancienne. 

Nymphe (fig. B) membraneuse, nue, pale, terminée par des cerques, surtout 
caractérisée par ses trés longues soles motrices. De chaque coté, la téte en porte 6, 
savoir: une sur |’épistome, 2 sur le front, 2 au-dessus de Vceil, et une sur le vertex,—le 
pronotum 10, savoir: 4 au bord antérieur, 4 au bord latéral, les deux postérieures rappro- 


GROUV ELLE—COLEOPTERA: CUCUJIDA, CRYPTOPHAGIDA 159 


chées de langle, et 2 discales proches de la base,—les segments dorsaux de 1l’abdomen 
4 (sauf le dernier qui n’en porte que 2), savoir: une latérale sur le tubercule de la pleure, 
2 de taille différente sur le rebord interne de la pleure, et une dorsale,—au genou, les 
fémurs antérieurs 2, les médians et les postérieurs une. 

La forme de la téte et du pronotum (dont le disque est trés déprimé), ainsi que la 
striation des élytres si accentuée chez l’imago, sont déja trés sensibles. Les antennes 
sont chargées de tubercles. 


728 


No. I1.—MALLOPHAGA, APHANIPTERA, AND DIPTERA PUPIPARA. 


By Hueu Scort, M.A., F.L.S., F.ES., 


Curator in Entomology in the Uniwersity of Cambridge. 
(Text-figures 1—4.) 
Read 18th June 1914. 


MALLOPHAGA. 


Only one species of bird-louse was obtained by the Percy Sladen Trust Expedition. 
Through the kindness of Professor G. H. F. Nuttall it was submitted to Professor L. G. 
Neumann of Toulouse, to whom its determination is due. 


Lrpzurus, Nitzsch. 
1. Lnpeurus subsignatus, Giebel. 
Inpeurus subsignatus Giebel, Ins. Epiz., 1874, p. 232. 


Loc. Aldabra: a number of specimens were collected by Fryer from the flamingo 
Phenicopterus minor, 1908—9. Professor Neumann stated (in. litt.) that this species 
was already known as a parasite of Phamcopterus: Kellogg, in Gen. Ins., Fascic. 66, 
1908, p. 45, gives it “from Phenicopterus antiquorum.” 


APHANIPTERA. 


A single species of flea is included in the collections of the Expedition. I am 
indebted to the Hon. N. C. Rothschild for its determination. 
OTENocEPHALUS, Kolenati. 
2. Ctenocephalus felis (Bouché). 
Pulea felis Bouché, Nova Acta Acad. Leopold., xvii. 1835, p. 505. 


Loc. Seychelles. A number of specimens were obtained in Silhouette, IX. 1908, in 
the hut at Mare aux Cochons: there were no other human habitations near, but the 
thick palm-leaf thatch of the hut was tenanted by many rats. A single specimen was 
also collected in Mahé (Cascade Estate, II. 1909). 


DIPTERA PUPIPARA. 
Hippoboscide. 


Only two species of this family are represented in the collections formed by the 
members of the Expedition. Both are found all over the warmer parts of the globe. I am 


SECOND SERIES—ZOOLOGY, VOL. XVII. 21 


162 PERCY SLADEN TRUST EXPEDITION 


much indebted to Mr E. E. Austen of the British Museum for examining and determining 
the material. 
PSEUDOLFERSIA, Coquillet. 
3. Pseudolfersia spinifera (Leach). 
Feronia spinifera Leach, Eprobose. Ins., ii. 1818, p. 557, pl. 26, figs. 1—3. 
Pseudolfersia spinifera Speiser, Zeitschr. Hym. Dipt., 1. 1902, p. 146; Austen, Ann. 
Mag. Nat. Hist., ser. 7, xu. 1903, p. 265. 


It is not attempted here to give the complete synonymy of this very wide-spread 
species. Its association with the Frigate-bird, which is found on the islands and coasts 
of all the warmer parts of the world, both tropical and subtropical, is discussed by Speiser 
(i.c.), and mentioned again by Austen (/.c.). The latter writer also refers to some other 
species of birds on which the fly has been found. 

The collections of the Percy Sladen Trust Expedition: include seven specimens. 
Four examples from Aldabra are very much larger than three from the Cargados Islands, 
but the disparity in size is no greater than that visible among the specimens of the 
British Museum series. 

Loc. Aldabra: 1907 (Thomasset). Cargados Carajos Islands, 26 and 28. VIII. 1905. 
Hosts unrecorded. 

Lyncuia, Weyenbergh. 

4, Lynchia maura (Bigot). 

Olfersia maura Bigot, Ann. Soc. ent. France, ser. 6, v. 1885, p. 237. 

Lynchia maura Speiser, Zeitschr. Hym. Dipt., i. 1902, pp. 155, 163. 


A single specimen has been determined by Mr Austen as belonging to this species, 
which, he informs me, is a parasite of domestic pigeons in all subtropical and tropical 
parts of the world. 


Loc. Seychelles: Mahé, 1908—9 (no further data recorded). 


Nycteribiide. 


No members of this family of bat-parasites were found in the Seychelles, nor, so far as 
I am aware, have any previously been recorded from the group. It must be confessed, 
however, that very little search was made for Nycteribudze. A specimen of the flying-fox 
Pteropus edwardsi was shot in Silhouette in August 1908, and one of Coléura seychellensis 
was shot at night at Cascade, Mahé, by Fryer, on March Ist, 1909. Neither of these 
specimens bore any Nycteribiidze: but no further search was made, and the resting-places 
of bats were not discovered. This branch of the entomology of the Seychelles, there- 
fore, should decidedly be further investigated. 

As regards Aldabra and the neighbouring coral-islands the case is different. There 
Fryer did search for Nycteribiidee, and he succeeded in finding one interesting species. 
The bat-fauna of those islands is mentioned in his work on the ‘‘formation of Aldabra,” 
ete. (Trans. Linn. Soe. London, ser. 2, Zool., vol. xiv. 1911, pp. 416—7). There is 
an endemic flying-fox, Pteropus aldabrensis True, contined to Aldabra, i.e. not occurring 


SCOTT—MALLOPHAGA, APHANIPTERA, AND DIPTERA PUPIPARA 163 


in the neighbouring islands of Assumption, Cosmoledo, or Astove. Fryer tells me that 
he examined 12—16 specimens of this bat for Nycteribiide, but found none, which is 
remarkable, since in Ceylon and the East Indies bats of this genus appear often to harbour 
these parasites. The single species of Nycteribiidee collected was taken in Assumption 
Island on a wide-ranging bat, Taphozvus mauritianus. The following particulars of its 
capture have been given me. The bats were clinging to the stem of a big coconut-palm 
—the only big one in the island—just below the crown of leaves. A boy was sent up the 
tree, but though it was broad daylight the bats were very active and dodged him round 
the trunk. Two of them were, however, killed with a stick, and on these were found the 
18 specimens of the Nycteribiid, which were also very active. 

Of the other family of Dipterous bat-parasites, the Streblidee, none were obtained in 
any of the islands visited by the Percy Sladen Trust Expedition, nor have any been 
previously recorded therefrom. 


NycreripiA, Latreille. Subgenus Acrocuonip1a (Kolenati). 


The single species belongs to this genus and subgenus: that is, it is entirely without 
eyes ; its tibize are not ringed, nor are they broad and flattened; the anal segment of 
the $ is without dorsal appendages. 

This Nycteribiid is quite distinct from any species of which I have seen named 
specimens, and it does not agree with any published description. It is therefore described 
here as new, and dedicated to its finder under the name Nycteribia (Acrocholidia) fryer. 
Tt has not been possible for me to see the types or named specimens of a number of species 
of the subgenus. Had this been possible, it would certainly have been more satisfactory 
to have had examples of all species for comparison, particularly as the descriptions in some 
cases are far from complete. According to Speiser’s descriptions (Arch. Naturg., 67.1.1901, 
pp. 31—36) both WN. (A.) stichotricha Speiser from Nias Island, and N. (A.) bellardii 
Rondani from South America, seem to have certain points of resemblance with JN. (A.) 
Jryeri, but both appear to differ from it in other respects. The most striking feature of 
N. (A.) fryeri lies in the form of the dorsal abdominal segments of the ?; and in no 
description have I found reference to anything like its remarkable two-processed 2nd 
tergite. 

Tn any case, NV. (A.) fryer is not confined to the region where it was found by Fryer, 
since there is in the British Museum an old, dried (and previously unnamed) ? specimen, 
undoubtedly referable to this species, from the widely remote island of Labuan. 

5. Nycteribia (Acrocholidia) fryert, sp. nov. (figs. 1—4). 

Length of body (without head) between 24 and 24mm. Colour of chitinous parts 
yellowish, abdomen of 3 darker. Head with about 6 bristles on the vertex in front 
(the complete absence of eyes has been demonstrated by mounting the head of a specimen 
in balsam and examining it under a high power). Thorax beneath (fig. 2) a little broader 
than long: of three specimens (preserved in spirit) in which its dimensions were measured 
with the help of a drawing apparatus, in 1 $ the breadth is about 14 times the length, 
in 29 it appears about 12 times the length: the median longitudinal line is slightly 

21—2 


164 PERCY SLADEN TRUST EXPEDITION 


broadened just behind the middle, and is depressed rather abruptly just before its 
posterior end: the oblique sutures dividing meso- from metasternum appear as rather 
broad white lines: the surface bears extremely fine short bristles and there are no very 
long bristles on the hind margin. Legs very long and slender, front coxee over 4 the 
length of the femora. 

g¢ ABDOMEN (figs. 3, 4): at the extreme base dorsally is a number of short erect 


rate 


S 


a , 


Fig. 1. Mycteribia (Acrocholidia) fryeri, sp. nov., ¢ , dorsal view. 


bristles marking the hind margin of the true tergite 1. Tergites 2, 3, 4 have their 
surfaces clothed with short bristles, and their hind margins set with alternating short and 
moderately long bristles, rather spaced out. Tergites 5 and 6 have their hind margins 
set with alternating very long and shorter bristles, the very long ones nearest the middle 
being strongly divergent: tergite 5 has a few short bristles on its surface, tergite 6 has 
its surface bare. Anal segment rather strongly tapering, its breadth at the apex a little 
less than 3 that at the base; in fig. 3 its length appears about equal to its breadth at the 


SCOTT—MALLOPHAGA, APHANIPTERA, AND DIPTERA PUPIPARA 165 


base, but when the segment is viewed from a slightly different angle its length appears a 
little greater than its breadth at the base: the anterior half of the segment is bare, but 
the posterior half bears a number of short erect bristles: there are numerous rather 
longer erect bristles on its sides, and there is a long stout bristle at either apical angle. 
Ventrally (fig. 4), the basal sternite is bare towards its base, but has several irregular 
series of short bristles on its surface behind, and rather longer bristles, directed outwards, 
at the sides: teeth of the ctenidium rather long. Sternites 2 and 3 have alternating 
moderately long and shorter bristles on their hind margins, and at the hind angles 
several much longer bristles, of which one is very long and outstanding: the surface 
of sternite 2 is almost hidden under the ctenidium, but some very short bristles can 
be discerned on it: the surface of sternite 3 is bare except for a few very short bristles 
just in front of the hind margin: both sternites have a few rather longer bristles 
on their surfaces towards the lateral margins. The next sternite (4+ 5) is a plate as long 
as, or longer than, the two foregoing taken together; its margin is curved at either side, 
so that it forms roughly a trapezoidal figure: its surface is bare except for a narrow area 
immediately before the margin: the apical part of its margin bears a number of stout 
black thorn-like bristles, immediately in front of which is a series of very fine bristles, 
long and short alternating: the sloping parts of the margin on either side bear very long 
bristles, immediately in front of which (on the surface) are short erect bristles, and in 
front of these again some very short bristles. The anal segment is rather densely covered 
with stiff erect bristles: the claspers lie parallel but not contiguous, taper from base 
to apex, and are black-pigmented not only at the apex but throughout their length almost 
to the base. 

The 2 ABDOMEN dorsally (fig. 1) is very remarkable. Basal tergite small and narrow, 
frequently appearing much foreshortened owing to distension of the abdomen, narrower at 


S 
DORIAN 
veal CN 


STILE NS 
/({U EE a) 
ann 1 fais x 


Figs. 2—4. Mycteribia (Acrocholidia) fryeri, sp. nov. 2. @, thorax and abdomen, ventral. 3. g, abdomen, 
dorsal. 4. $, abdomen, ventral. 


the base and broader behind, with the hind angles rounded ; divided in the middle of the 
hind margin by a triangular cleft extending forwards about half its length, in which cleft 


166 PERCY SLADEN TRUST EXPEDITION 


pale whitish connexivum is exposed; the hind margin on either side of this cleft is 
broadly, and the lateral margins are rather more narrowly, black-pigmented ; the hind 
margin on either side of the cleft bears 5 or 6 very long strong blackish bristles set close 
together; on either side of these, at the outer angles, the margin bears 2 or 3 short 
blackish bristles and the dise of the tergite bears a number of short sub-erect bristles 
in its lateral portions. Tergite 2 large, yellowish, chitinous, occupying nearly half the 
length of the abdomen: divided longitudinally by a pale line which is not always very 
evident: the hind margin slopes obliquely backwards on either side to the middle line, 
immediately on either side of which the segment is produced into a remarkable blunt- 
pointed process: the two processes are slightly divergent, and separated from one 
another by an elongate triangular cleft. A streak of darker brown pigment extends 
along the outer side of each process some way forward into the segment ; the two streaks 
diverge slightly from behind forwards, and each becomes broader and more dilute anteriorly 
until it is gradually lost in the surrounding yellow colour of the segment. ach of the 
processes bears several bristles; a long stout divergent bristle rises from its inner 
side just before the apex, there is a shorter bristle at the extreme apex and 
one or two still shorter ones near it. The hind margin of the segment bears 3 
or 4 rather long bristles, spaced out, on either side towards the sides of the body. 
The disc of the segment bears scattered short erect bristles laterally, but the median 
portion, especially in front, is bare except for a very few short bristles on either side 
of the median longitudinal line. Behind this 2nd tergite is an area of whitish con- 
nexivum, bare in the middle, at the sides covered with numerous scattered minute and 
exceedingly short bristles. Behind this connexivum is a small, more strongly chitinised, 
yellowish portion, with a slightly rounded hind margin bearing 6 bristles, 3 on either 
side separated by a median interval*. The anal segment is short, tapering slightly, 
but rather broad apically: quite bare above, but with one or two bristles at either side, 
and a longer and a shorter one at each hind angle, 

Ventrally (fig. 2), the basal sternite is as in the g. Sternite 2 consists of connexival 
membrane ; surface rather closely covered with fine short bristles, becoming slightly 
longer behind ; hind margin indicated by a transverse series of longer and stouter bristles, 
those towards the sides longer than the median ones. Sternite 3 consisting of a very 
short area of connexivum, its surface bare, its hind margin indicated by a series of bristles 
similar to that on sternite 2. Sternite 4 also short and with its surface bare: its hind 
margin has, at either lateral extremity, a small yellowish chitinous area: the marginal 
series of bristles is nearly similar to those on the two preceding sternites, except that 
on each of the two small chitinous areas two of the bristles are very long and directed 
outwards. Sternite 5 bears two much larger yellowish chitinous areas, which are 
separated in the middle line by only a very narrow space of white connexivum : each 
chitinous area bears a transverse series of short bristles in front of the hind margin, 
and a series of much longer bristles on the hind margin: in the marginal series longer 


* In the British Museum specimen from Labuan this small chitinous area is much smaller and bears only 
two bristles. The specimen has long been preserved dry, and it is possible that other bristles may have been 
broken off. In other respects the Labuan example corresponds closely with those from Assumption. 


SCOTT—MALLOPHAGA, APHANIPTERA, AND DIPTERA PUPIPARA 167 


and shorter bristles alternate, and one or two near the outer extremities are very long 
and directed outwards. Subgenital plate roughly trapezoidal, with hind angles much 
rounded : divided by a narrow, pale, median area into two yellowish chitinous lateral 
parts, each of which bears on its posterior half two transverse series of bristles, the 
anterior consisting of about 5 short bristles, the posterior consisting of about 3 very 
long outstanding bristles ; the apical part of the margin bears about 9 bristles, of which 
the two at the rounded hind angles are the longest. 


Loc. Assumption. Labuan. 

Assumption: IX. 1908 (Fryer); 33, 15? (14, 129, in spirit: 23, 39, preserved dry), 
collected from Taphozous mauritianus (for circumstances of capture see above in the 
general remarks on Nycteribiidee). 


Labuan: 1 in British Museum, labelled “78.6. From Taphozous saccolaimus.” 


No. IV.—MEDUS4 FROM THE INDIAN OCEAN. 


(Collected by Pror. StanLEY GarpinErR, in H.M.S. “Sealark,” in 1905.) 
By Epwarp T. Browne, M.A., F.L.S. 
Plate 39. 


Read 17th June, 1915. 


INTRODUCTION. 


The collection of Medusz, made by Prof. Stanley Gardiner during the voyage of 
H.M.S. “Sealark” in the Indian Ocean, between Chagos, Mauritius and Seychelles, in 
1905, was sent to me for examination. I heartily thank Prof. Gardiner for allowimg 
me the privilege of writing this report upon the specimens, and must also express to 
him my regret for the delay over the work. 

The chief interest in the collection centres in the geographical distribution -of species 
as this part of the Indian Ocean had scarcely been explored for medusze. It should be 
borne in mind that the collecting of medusee was only a subsidiary part of the expedition’s 
work. Nevertheless Prof. Gardiner was able to collect not less than thirty-five different 
genera, but none proved to be new to Science. The species were slightly more numerous 
than the genera, three new ones have received names, and others could have been added 
if the specimens had been in better condition. 

The Anthomedusze show a much better list of genera and species than in the 
. previous collections from the Maldives and Ceylon. The poor list of species belonging 
to the Leptomeduse is partly due to my failure to identify the specimens. There are 
always a certain number of bad specimens in every collection, and it so happened that the 
Leptomedusze got more than their fair share of these. 

The Anthomedusze and- Leptomedusz are associated with the littoral fauna and are 
rarely found far from land or shallow water. Many of their genera are known to be 
connected with hydroids, so that their geographical distribution depends upon their 
hydroids finding a suitable habitat. Some are widely distributed, others have a very 
limited range. It is certainly strange that some of the commonest species have so far 
evaded the search for their hydroids and leave us still without a clue, but it is ditiicult to 
presume that some have no alternation of generations. 

It is unfortunate that systematists on meduse still hold divergent views over the 
question of species, but nevertheless progress is slowly being made. There has been 
a good advance since the day that saw the appearance of Haeckel’s monograph which 
was by no means a success, except from an artistic point of view. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 22 


170 PERCY SLADEN TRUST EXPEDITION 


The medusze which are known to be, or expected to be, connected with hydroids 
may require a finer discrimination of specific characters than those which have direct 
development. The linking together of species which very closely resemble each other 
under a common name, but are found in localities very far apart, may lead later on to 
further confusion and erroneous ideas on geographical distribution of marine animals. 
We are not yet certain that hydroids which are universally recognised as distinct species 
of a genus may not have medusze so much alike that there is the possibility of their 
medusze being regarded as belonging to the same species. We know that hydroids 
belonging to families far removed have medusze closely related according to our present 
system of classification, but we do not know what surprises are in store when all the 
hydroid species of Syncoryne, Bougainvillia, Perigonimus, Obelia, Campanulina, ete., 
have had their medusz reared and traced to their adult stages. 

A different treatment of specific characters appears to me to be required for those 
medusze which have direct development (without any hydroid stage), especially those 
belonging to certain genera, such as Liriope and Pelagia, which are found throughout 
the warm regions of the oceans. There are many genera which have a vast geographical 
range, but how far their species are valid is still a debatable subject. At one time 
a formidable list of species was being piled up, but the tendency is now to reduce the 
species to about one per genus. I think this latter process is being carried too far, it 
makes work easy for the systematist, but is bad for Science. It appears to me that 
a species belonging to this class of medusze shows a far greater range of variation than 
those belonging to the Anthomedusze and Leptomeduse. We have formerly been 
basing the specific characters upon a too limited number of specimens, and have described 
the characters of an individual rather than those of a race. There are certainly distinct 
races of Liriope and Pelagia, and some of them are good species. 

The following is a classified list of genera and species found on the cruise of the 


““Sealark ” : 


HYDROMEDUSi. 

ANTHOMEDUS. 
Euphysora bigelowi, Maas. Amphinema sp. 
Steenstrupia normani, n. sp. Leuckartiara gardinert, n. sp. 
Zanclea orientalis, n. sp. Pandea juv. 
Zanclea juv. Heterotiara minor, Vanhoffen. 
Cyteis tetrastyla, Eschscholtz. Proboscidactyla tropica, Browne. 
Bougainvillia fulwva, Agassiz and Mayer. Proboscidactyla sp. 
Turritopsis nutricula, McCrady. : 

LEPTOMEDUS. 

Tiaropsis rosea, Agassiz and Mayer. Mesonema pensile (Modeer). 
Phaalidium sp. Equorea macrodactyla (Brandt). 


Irene sp. 


BROWN—MEDUSA FROM THE INDIAN OCEAN We 


Olindias singularis, Browne. 


Rhopalonema velatum, Gegenbaur. 


Sminthea eurygaster, Gegenbaur. 
Pantachogon rubrum, Vanhoffen. 
Halicreas papillosum, Vanhoften. 


Solmaris sp. . 
gina citrea, Eschscholtz. 


Charybdea sp. : 
Nausithie punctata, Kolliker. 
Atolla wyville, Haeckel. 


TRACHOMEDUSZ. 


Aglaura hemistoma, Péron et Lesueur. 
Amphogona apsteini (Vanhéffen). 

Liriope tetraphylla (Chamisso et Eysenhardt). 
Liriope sp. 

Geryonia proboscidalis (Forskal). 


NARCOMEDUS&. 


Solmundella mediterranea (Miiller). 
Cunina sp. 


SCYPHOMEDUSi. 


Pelagia flaveola, Eschscholtz. 
Pelagia sp. 
Rhizostoma andromeda var. maldivensis, Browne. 


Pelagia panopyra (Péron et Lesueur). 


Lists OF SPECIES FOUND WITHIN DEFINITE AREAS DURING THE CRUISE OF 


THE ‘‘ SEALARK.” 


I have not prepared lists of medusze taken in every haul of the nets nor for every 
station, but have grouped together the stations under convenient geographical headings. 
The longest list belongs to the Chagos Archipelago, and that may be probably due to more 
collecting of plankton having been done within that area, than in other areas, such as at 
Mauritius. Professor Gardiner made serial hauls at different depths and also hauls at 
definite intervals of time at several of the stations. I made records of the meduszx 
found in every haul, but failed to obtain any reliable results owing to the scarcity of 


specimens. 


CHAGOS ARCHIPELAGO. 


Steenstrupia normant. 
Zanclea orientalis. 
Cyteis tetrastyla. 
Bougainvillia fulva. 
Turritopsis nutricula. 
Pandea juv. 
Heterotiara manor. 
Phialidium sp. 
Mesonema pensile. 
Aiquorea macrodactyla. 
Olindias singularis. 
Rhopalonema velatwm. 
Sminthea eurygaster. 


Pantachogon rubrum. 
Aglaura hemistoma. 
Amphogona apsteint. 
Liriope tetraphylla. 
Geryonia proboscidalis. 
Solmaris sp. 

Afgina citrea. 
Solmundella mediterranea. 
Cunina sp. 

Nawsithie punctata. 
Atolla wyviller. 
Pelagia panopyra. 
Pelagia sp. 


SAYA DE MALHA BANKS. 


Cyteis tetrastyla. 
Irene sp. 
fEquorea macrodactyla. 


Aglaura hemistoma. 
Liriope tetraphylla. 
Pelagia flaveola. 


LS 
bo 
{ 


172 PERCY SLADEN TRUST EXPEDITION 


NAZARETH BANK (Cargados Carajos). 


EHuphysora bigelow?. Amphogona apsteint. 
Irene sp. Liriope tetraphylla. 
MAURITIUS. 

Tiaropsis rosea. Liriope tetraphylla. 

Rhopalonema velatum. Geryonia proboscidalis. 

Aglaura hemistoma. Cunina sp. 
FARQUHAR GROUP. 

Proboscidactyla sp. Solmaris sp. 

Khopalonema velatum. Solmundella mediterranea. 

Halicreas papillosum. Cunina sp. 

Aglaura hemistoma. Charybdea sp. 


LInriope tetraphylla. 


BETWEEN PROVIDENCE AND ALPHONSE ISLES 
(lee. & 16’ S., Meme, Sly 2G O10.) 


Pantachogon rubrum. Liriope tetraphylla. 

Halicreas papillosum. Atolla wyvillei. 
ALPHONSE ISLAND. 

Euphysora bigelow?. Liriope tetraphylia, 

Zanclea juv. Cunina sp. 


Aglaura hemistoma. 


AMIRANTE GROUP. 


Bouganvillia fulva. Sminthea eurygaster. 
Amphinema sp. ; Halicreas papillosum. 
Leuckartiara gardinert. Aglaura hemistoma. 
Proboscidactyla tropica. Liriope tetraphylla. 
Laodice % Solmundella mediterranea. 
Phialidium sp. Nausithoe punctata. 


Rhopalonema velatum. 


SEYCHELLES GROUP. 


Cassiopea andromeda var. maldivensis. 


Some of the results obtained by this expedition for the geographical distribution of 
medusze are rather interesting. 

Turritopsis nutricula had not before been found in the Indian Ocean. It is a well- 
known species on the North Atlantic coast of the United States, south of Cape Cod, and 
it also has quite recently been recorded from Japan. 

Tiaropsis rosea and Euphysora bigelowi have had their range extended westwards 
from the Malay area. 

Proboscidactyla tropica is the most interesting find in the collection; it has not 
been recorded since Huxley first found it in Louisiade Archipelago (Malay area). 

Olindias singularis occurs at Chagos, which is not so far from the Maldives, where 
it was first discovered. Since its discovery it has been found far away in the middle of 
the Pacific among the Paumotu Isles. 

Sminthea eurygaster is new to the Indian Ocean; its old records are for the 
Mediterranean, the Bay of Biscay, and in the Atlantic as far south as Brazil. 


BROWNE—MEDUSAi. FROM THE INDIAN OCHAN 173 


HYDROMEDUS Ai. 


ANTHOMEDUS&. 
Genus Eupuysora, Maas, 1905. 


Hartlaub (1907) and Mayer (1910) agree in uniting the genera Huphysa and 
Euphysora with Corymorpha. (Mayer uses the name Steenstrupia in the place of 
Corymorpha; the former is a medusoid generic name and the latter a hydroid generic 
name.) I am in agreement with them so far as Huphysa is concerned, but am not, at 
present, inclined to follow them in the case of Huphysora. 

Both Steenstrupia rubra and Huphysa aurata have one fully developed perradial 
tentacle and three perradial marginal bulbs. The latter are called rudimentary tentacles 
by the above authors, but it must be clearly understood that the tentacles have com- 
pletely disappeared on the three bulbs. Far remote ancestors may possibly have had 
four fully developed tentacles, but now only one tentacle remains, the others having their 
former positions marked by the bulbs. 

Tt is known that Steenstrupia rubra is the medusa belonging to the hydroid 
Corymorpha nutans. Hartlaub believes that Huphysa aurata is probably connected 
with Corymorpha nana Alder, but at present there is no definite proof, though Mayer 
records the connection as a definite fact. We have no clue to the hydroids connected 
with Huphysora. 

The genus Huphysora was established by Maas for those Codonide having four fully 
developed tentacles, but one tentacle unlike the others being larger and different in 
external appearance. 

There are three species belonging to this genus. The first was described by Bigelow 
(1904, p. 251, pl. 1) from the Maldives under the name of Huphysa tetrabrachia, and the 
second by Maas from the Malay Archipelago under the name of Huphysora bigelowi, and 
this is the type species of the genus. Bigelow was doubtful about the generic position of 
his new species, but Maas placed it in the new genus Huphysora and at the same time 
pointed out the characters by which the two species could be recognised. 

Without going into every minute detail the two species can be easily distinguished 
by the structure of their tentacles. Huphysora tetrabrachia has annular rings of nemato- 
cysts on all the tentacles, but more on the longest tentacle. Huphysora bigelowi has 
large globular clusters of nematocysts on the longest tentacle only. Mayer is inclined to 
regard the two species as being identical, for in his monograph (p. 37) he has written : 
“Future studies will probably show that these distinctions are not of specific value, but 
merely changes due to growth and variation, and that the two medusz are identical and 
should be called Steenstrupia tetrabrachia.” 

The third species, Huphysora valdivie Vanhoften (1911), has lateral branches on its 
principal tentacle. This species I shall refer again to under Steenstrupia norman. 


174 PERCY SLADEN TRUST EXPEDITION 


1. EupHysorA BIGELOWI, Maas, 1905. 


Euphysora bigelowi, Mass, 1905, p. 7, Taf. 1, figs. 1—3. 

Huphysora bigelowi, Maas, 1906, p. 84, pl. 2, figs. 1—2. 
Huphysora bigelowi, Miller, 1908, p. 59. 

Steenstrupia bigelowt, Mayer, 1910, vol. i, p. 36, fig. 9. 

Euphysora bigelowi, Vanhoffen, 1911, p. 197. 

ELuphysora bigelowi, Vanhoffen, 1912, p. 7, Taf. 1, fig. 3. 


Localities. Cargados Carajos, Surface. 30 Aug. 1905,1. 2 specimens. Alphonse 
Is., Surface (Temp. 80° F.). 7 Oct. 1905, dd.* 1 specimen. 

The specimens in the ‘“Sealark” collection belong to the species Huphysora bigelown, 
and only a brief description of them is necessary as the species has been fully described 
and well figured by Maas. The smallest specimen is about 1°5 mm. in length, and has 
seven globular clusters of nematocysts upon the principal tentacle. The next in size is 
about 3 mm. in length, and the principal tentacle is also provided with seven batteries of 
nematocysts. Neither of these specimens shows gonads. The largest specimen is about 
4 mm. in length and 2 mm. in width, and has a well-developed spermary which surrounds 
the stomach and extends along its whole length, just leaving the circular mouth free. 
Upon the principal tentacle there are eleven globular clusters of nematocysts forming 
a half loop round the tentacle. The three other marginal tentacles are quite different in 
shape and appearance. They taper to a point and have a smooth external surface, the 
nematocysts being scattered. In the largest specimen the tentacle opposite to the 
principal tentacle is much shorter than the others, but in the smaller specimens the three 
tentacles are about equal in size. None of the specimens show the apical prolongation of 
the stomach in the shape of a canal, which was present in many of the medusz seen by Maas. 

Distribution. Indian Ocean; Pacific Ocean, Malay Area. 


2. STEENSTRUPIA NORMANI nova, species. 


(Plate 39, fig. 1.) 


Locality. North of Chagos, Lat. 4° 16’S., Long. 71° 53’ EZ. 100—O fms. 17 May, 
1905, B. 1 specimen. 

Description of the Species :—Umbrella about twice as high as broad, with a conical 
apex, and thin walls. Ex-umbrella with twelve longitudinal streaks of nematocysts 
extending from the margin to the apex, having lateral branches and forming a kind of 
network on the lower part of the umbrella. Stomach large, cylindrical, with a tube-like 
mouth extending beyond the velum, and with an endodermal cellular prolongation into 
the apex of the umbrella. Four thick radial canals and a very thick circular canal. 
One perradial tentacle, with a long, hollow, sac-like basal bulb, and terminating with a 
Jarge globular ball containing nematocysts, and also three other globular clusters of 
nematocysts probably unilaterally arranged. Three small perradial bulbs without 
tentacles on the margin of the umbrella. 

Size. Umbrella 1:25 mm. in width and 2°5 mm. in height. 

The specimen is in very good condition, but a few more showing later stages would 


* For List of stations see rans. Linn. Soc. Ser. 2. Zool. xii. (1907), p. 170. 


BROWNE—MEDUS4 FROM THE INDIAN OCEAN 175 


have been welcomed. The stomach is very much swollen out with food and it almost fills 
up the whole of the cavity of the umbrella. It is evidently a young stage as I cannot 
detect gonads upon the stomach, which has fairly transparent walls. The stomach 
occupies the whole of the top of the cavity of the umbrella and even extends above 
it, as the radial canals leave the stomach laterally. The radial canals and circular 
canal have a very cellular appearance and look as if composed of a solid chord of very 
large cells. The solitary tentacle has a large sac-like basal bulb with rather thin walls, 
covered with large flat ectoderm cells and practically free from nematocysts. The tentacle 
itself is contracted and coiled up. It has three lateral clusters of nematocysts projecting 
from the tentacle and a large globular terminal cluster. The lateral clusters are 
globular and vary in size; the smallest at the top and the largest at the bottom. 
Owing to the coiling of the tentacle their exact position on the tentacle is doubtful. 

In the three perradii without tentacles there are only small bulbs projecting from the 
margin. They are solid in appearance, covered with nematocysts, and show no signs of 
developing tentacles. It is not easy to trace out completely the tracks of nematocysts 
upon the ex-umbrella. There are clear indications of twelve longitudinal tracks running 
right up to the apex of the umbrella and some of these tracks unite near the apex. The 
main tracks have also short lateral branches which join on to branches from adjacent tracks 
and form a kind of network on the lower part of the umbrella. There are no signs of any 
ocelli on the marginal bulbs or tentacle. 

The characters selected for distinguishing the species are the shape and structure of 
the tentacle and the numerous tracks of nematocysts upon the ex-umbrella. 

This new species is named after my friend and neighbour the Reverend Canon 
A. M. Norman, whose name is well known to all marine zoologists. 

Euphysora valdiwie, which Vanhéffen (1911) described as a new species from a 
single specimen taken near Siberut Island off the west coast of Sumatra (west of Padang), 
has certain characters in common with Steenstrupia norman. Both species have similar 
tracks of nematocysts upon the ex-umbrella; the same type of large stomach, and large 
radial and circular canals. The principal tentacle of Huphysora valdivie has lateral 
branches, and is evenly covered with nematocysts without any arrangement of rings or 
knobs. In addition to the principal tentacle there are three other smaller tentacles 
without branches. 

As Steenstrupia normani is not half the size of Euphysora valdivie and is at 
an immature stage, one has to consider the question of Steenstrupia norman: being an 
early stage of EHuphysora valdivie. There is no evidence that three marginal bulbs 
of Steenstrupia normani will develop tentacles, though there is a possibility of their 
doing so. ‘The principal tentacle of Steenstrupia normant has very conspicuous globular 
clusters of nematocysts. These clusters project from the tentacle, but they do not appear 
to be upon stalks or lateral branches. _ But if such stalks or lateral branches were strongly 
contracted, then possibly the clusters would come alongside the tentacle. Vanhéffen 
states that the principal tentacle of Huphysora valdime is without knobs or rings 
of nematocysts and his figures do not show them. Their absence is in favour of 
Steenstrupia normani being a distinct species. 


176 PERCY SLADEN TRUST EXPEDITION 


3. ZANCLEA ORIENTALIS nova species. (Pl. 39, figs. 2 and 3.) 


Locality. North of Chagos, Lat. 4°16’ 8., Long. 71° 53’ E. 500 fms. (Wolfenden’s 
closing net). 18 May, 1905, G. 1 specimen. 

Description of the Species :—Umbrella a little higher than broad, with a rounded 
summit and rather thin walls. Velum very narrow. Stomach cylindrical, about half 
the length of the cavity of the umbrella. Mouth circular. Gonads extending nearly the 
whole length of the stomach and forming interradial swellings. Two large opposite 
perradial tentacles armed with globular batteries of nematocysts upon contractile stalks. 
No tentacles in the other two opposite perradii, but only two very rudimentary internal 
bulbs. Four perradial patches of nematocysts upon the margin of the umbrella, without 
any groove or streak leading to them. 

Size. Umbrella about 1:5 mm. in width and 2 mm. in height. 

The character selected to distinguish this new species is the presence of rather broad 
perradial patches of nematocysts upon the margin of the umbrella. These patches (fig. 3) 
are situated directly on the margin and have no tracks or grooves leading to them. 
There are no longitudinal bands of nematocysts upon the ex-umbrella, but its whole 
surface is sprinkled with small isolated nematocysts. The tentacles are in a contracted 
condition and so also are the stalked batteries of nematocysts. The exterior of the 
battery is covered with a thin layer of ectoderm, and the nematocysts are packed 
away inside, practically filling the whole of the interior of the ball. The stalked 
batteries of nematocysts are along only the outer side of the tentacles. 

The depth at which this specimen was taken is not trustworthy, as the messengers 
working the net were not acting properly. 


4. ZANCLEA juv. 


Locality. Alphonse Is., Surface. 7 Oct. 1905, ee. 1 specimen. 

The umbrella is nearly 2°5 mm. in height and 1:75 mm. in width, with thick walls 
and a rounded summit. The stomach is short, about 0°5 mm. in length, and with 
a circular mouth. Four perradial canals. There are two opposite perradial tentacles 
armed along their outer side with stalked batteries of nematocysts, which are oval in 
shape, about twice as long as wide, and about twice the size of those in the batteries of 
Zanclea orientalis. The stalks have the appearance of rather fine filaments, studded with 
small isolated nematocysts. In the other two opposite perradii there is the merest trace 
of a marginal bulb, which has not the appearance of ever developing a tentacle. Upon 
the ex-umbrella there are four perradial tracks of nematocysts. They are very short, just 
curling over the margin and terminating with a small cluster of nematocysts. 

This is evidently a young medusa, as there are no signs of gonads upon the 
stomach. 

5. ZANCLEA ?. 


Locality. Chagos, Peros Atoll. 75—0 fms. 30 June, 1905, M. 1 specimen. 
The umbrella is thick but contracted and torn; about 2°5 mm. in width and 2 mm. 
in height. The stomach is badly damaged, but there are cells at its base, interradially 


BROWNE—MEDUSAD FROM THE INDIAN OCEAN 177 


situated, having the appearance of ova. Two large opposite perradial basal bulbs, 
globular in shape, from which tentacles have been broken off. In each of the other two 
opposite perradii there is a minute rudimentary bulb. There are four perradial tracks of 
nematocysts upon the ex-umbrella, about 0°5 mm. in length. The tracks lead to an oval 
patch of nematocysts, situated upon a slight prominence of the ex-umbrella. 

This specimen probably belongs to the genus Zanclea, as it has tracks of nematocysts 
upon the ex-umbrella, and only two tentacles. 


6. ZANCLEA ?. 

Locality. Lat. 8° 16’ S., Long. 51° 26’ E. (between Providence and Alphonse). 
900—0 fms. 6 Oct. 1905, aa. 1 specimen. 

The umbrella is thick, with a rounded summit, about 2°5 mm. in height and slightly 
less in width. The stomach at its base is cross-shaped and tapers down to a slender tube 
terminating with a circular mouth, which is about on a level with the margin of the 
umbrella. Upon the upper half of the stomach are situated four gonads, which occupy 
the whole of the spaces between the four perradii. There are four perradial basal bulbs 
all about the same size, from which tentacles have apparently been broken off. Upon the 
margin of the ex-umbrella there are four perradial tracks of nematocysts, one above each 
of the basal bulbs. They have the appearance of straight, narrow canals, less than 0°5 mm. 
in length. | 

Owing to the absence of tentacles it is impossible to be sure of the correct 
determination of the genus. It is probably a Zanclea with four tentacles. 


Genus Cyrais, Eschscholtz, 1829. 

There have been several species of Cyteis described at intervals, and located in 
different regions of the world. Vanhoffen, who has examined a very large number of 
Cyteis collected by different expeditions in the Atlantic, Indian and Pacific Oceans, 
definitely comes to a conclusion that there is only one species in this genus, and to it 
he assigns the oldest specific name, Cytevs tetrastyla, Eschscholtz. 

There is, however, a feeling of doubt in my mind as to there being only one species 
of Cytwis. I am inclined to regard Cytais vulgaris, Agassiz and Mayer, from the Fiji 
Isles as a distinct species, and also Cyteis pusilla, Gegenbaur, from the Mediterranean. 
I think that as Vanhéffen has taken as a type the somewhat vaguely defined and badly 
drawn Cytais tetrastyla of Eschscholtz, it will be best to follow him by grouping under 
its name at all events the following :— 

Cyteis nigritina, Haeckel, 1879, p. 74, Taf. 6, figs. 2—). 

Cyteis macrogaster, Haeckel, 1879, p. 74, Taf. 6, fig. 1. 

Cyteis nigritina, Maas, #904, p. 8, pl. 1, fig. 3. 

Cyteis herdmant, Browne, 1905, p. 135, pl. 1, fig. 1. 

Cytwis vulgaris, Bigelow, 1909, p. 190, pl. 6, fig. 3, pl. 40, fig. 2 and fig. 5, pl. 43, figs. 4—5. 
Cyteis tetrastyla, Vanhoffen, 1911, p. 204, Text-fig. 6. 

Cytwrs tetrastyla, Vanhoften, 1912, p. 8, Taf. 1, fig. 5. 

The above-mentioned figures show specimens with large triangular basal bulbs 
extending some way up the umbrella, and the tentacles should have a pigmented 
endoderm. 

SECOND SERIES—ZOOLOGY, VOL. XVII. : 23 


178 PERCY SLADEN TRUST EXPEDITION 


7. Cy?Ta&Is TETRASTYLA, Eschscholtz, 1829. 


Localities. North of Chagos Archipelago, Surface (Temp. 82° F.). 18 May, 1905, D. 
1 specimen. South of Saya de Malha Banks. 50—O fms. 4 Sept. 1905,m. 2 speci- 
mens. North of Saya de Malha Banks, Surface (Temp. 77°—80° F.). 8 Sept. 1905, n. 
1 specimen. 

The specimens collected by the “Sealark” are not in good condition. The largest is 
about 2 mm. in diameter. Its stomach is on a short peduncle and the mouth surrounded 
with at least twelve capitate tentacles. The four perradial marginal tentacles are 
internally pigmented with a very dark reddish colour along their whole length, and 
covered with a very thick ectoderm. The specimens have a few medusa-buds upon the 
stomach. The structure of the tentacles and the shape of the basal bulbs are similar to 
those of Cytais herdmani. 

Some of the specimens reported upon by Vanhdffen were collected by the “ Valdivia’ 


) 


in the Indian Ocean, one station being north of Chagos. 


8. BouGAINVILLIA FULVA, Agassiz and Mayer, 1899. 


Bougainvillia fulva, Agassiz and Mayer, 1899, p. 162, pl. 2, fig. 6; 1902, p. 145, pl. 2, fig. 8. 
Bougainvillia fulva, Maas, 1905, p. 10, Taf. 1, fig. 8, Taf. 2, figs. 9—10; 1906, p. 87, pl. 2, figs. 4—5. 
Bougainvillia fulva, Bigelow, 1909, p. 195, pl. 6, fig. 7, pl. 44, figs. 5—7. 

Bougainvillia fuleva, Hartlaub, 1909, p. 448, Taf. 19, figs. 1—4. 

Bougainvillia fulva, Mayer, 1910, p. 160. 

Localities. Chagos Archipelago, Salomon Atoll. 180—O fms. 30 June, 1905, O. 
11 specimens. Amirante Isles, Desroches Atoll. 400—O fms. 16 Oct. 1905, mm. 
1 specimen. 

Except for the contraction of the margin of the umbrella the specimens are in very 
good condition. The smallest one measured 6 mm. in height and 4mm. in width, and 
the largest 13 mm. in height and 10 mm. in width. 

The umbrella is very thick, about as thick at the top as on the side. Its natural 
shape is altered owing to the great contraction of the margin. The contraction produces 
deep perradial furrows and interradial ridges or lobes which meet in the centre of the 
umbrella opening and close it up completely. The specimen figured by Maas (1906, pl. 2, 
fig. 5) resembles this very closely in the shape of the contracted umbrella, but the jelly 
above the sub-umbrella cavity is thicker than in my specimens. 

The stomach varies considerably in shape. When empty and uncontracted it hangs 
down as a thin cruciform sac, but when full of food it becomes nearly globular. The four 
oral tentacles are dichotomously branched, four times in the smallest specimen and seven 
times in the largest one. The gonads form swellings upon the wall of the stomach and 
have the appearance of eight distinct sacs, isolated perradially and interradially. 

The marginal tentacles are more or less contracted, and are very small and slender 
when compared with the size of the umbrella. In the smallest specimen there are twelve 
tentacles on each compound bulb; and fourteen on a bulb was the maximum number 
counted in the largest specimen. An ocellus is situated on the inner side of every 


BROWNE—MEDUSA FROM THE INDIAN OCEAN 179 


tentacle, very close to the bulb; it is a narrow band of dark pigment on the surface of 
the ectoderm. 

One specimen is infested with Cwnina buds in different stages of development. The 
buds are attached to the sub-umbrella close to the base of the stomach. The largest 
Cunina shows three rows of otoporpe on each lappet, and is about ready for 
liberation. 

Bougainvillia fulva is widely distributed over the Pacific and Indian Oceans, and 
lives within the tropical belt. 


Genus Turritropsis, McCrady, 1857. 


There are differences of opinion amongst specialists on medusee as to the number 
of species that should be recognised as belonging to the genus Turyitopsis. Mayer (1910) 
and Bigelow (1913) clearly consider that the American Turritopsis nutricula, McCrady, 
is identical with the European Turritopsis polycirrha, Keferstein. Maas (1909) is 
practically of the same opinion, but he suggests that the species should be kept apart, 
as local forms or varieties. Acting upon his own suggestion, Maas described a new 
Turritopsis from Japan as a new variety of Turritopsis nutricula. 

Hartlaub (1911) on the other hand seems fairly convinced that the American and 
European Turritopsis belong to distinct species. I may here say that I agree with 
Hartlaub in placing Turris neglecta, Lesson, as a synonym of Turritopsis polycirrha. 
Sometime before the appearance of Dr Hartlaub’s publication I had already come 
to the same conclusion. 

With regard to the medusa described by Maas (1909) under the name of Turritopsis 
nutricula, var. pacifica from Japan, I think it should be regarded as a distinct species, 
and under the name Turritopsis pacifica. Maas clearly states that the ocelli are abaxial, 
that is to say, on the outer side (ex-umbrella side) of the basal bulbs, whereas both 
Turritopsis nutricula and Turritopsis polycirrha have their ocelli on the inner side 
of the basal bulbs. Even if Maas, as Bigelow (1913) has hinted, has accidentally 
recorded and figured the ocelli on the wrong side of the bulb, still, in my opinion, it 
remains a distinct species. Maas’s figures show that the formation of endoderm above 
the stomach is similar to that of Turritopsis polycirrha, but it can be distinguished from 
Turritopsis polycirrha by having a much larger number of tentacles (120—150) arranged 
in several rows round the margin of the umbrella. 

Since I have seen the specimens of Twrritopsis in the “Sealark” collection and 
compared them with specimens of Turritopsis polycirrha from the English Channel and 
the North Sea, I am strongly in favour of Hartlaub’s views and believe the reasons given 
by him for separating the American and Huropean species are sound. 

In Turritopsis nutricula the radial canals, by a considerable thickening of their walls, 
and coalescence, form a kind of endodermal peduncle upon which the stomach hangs. 

In Turritopsis polycirrha a further development of endoderm takes place. It is no 
longer confined to the walls of the radial canals, but grows across the top of the umbrella 
cavity and forms a homogeneous mass of cells. This extension also grows downwards and 
takes in the radial canals in the uppermost part of the sub-umbrella. The extension 


23—2 


180 PERCY SLADEN TRUST EXPEDITION 


of the endoderm is recorded by the leaving of the outer half of the radial canal wall 
in situ alongside the umbrella. In adult specimens the radial canals run alongside this 
mass of endoderm nearly to the top of the sub-umbrella and there they curve sharply 
inwards to the stomach. 

Turritopsis nutricula has been well described and figured by American naturalists, 
and there is no evidence to show that its endodermal peduncle is converted by further 
development into a mass of cells which block up the upper part of the cavity of the 
sub-umbrella. Maas (1909) has already said that the differences between the two species 
may be defined as stages in development, and that is the case, but up to the present 
I have failed to find any proof that Turritopsis nutricula develops into Turritopsis 


polycurrha. 


9. TURRITOPSIS NUTRICULA, McCrady, 1857. 


Oceania (T'urritopsis) nutricula, McCrady, 1857, p. 55, pl. 4. 

Turritopsis nutricula, McCrady, 1858, p. 127, pl. 8, fig. 1. 

Modeeria multitentacula, Fewkes, 1881, p. 149, pl. 3, figs. 7—9. 

Turritopsis nutricula, Brooks, 1886, p. 388, pl. 37. 

Turritopsis nutricula, Brooks and Rittenhouse, 1907, pp. 429—460, pls. 30—35. 
Turritopsis nutricula, Mayer, 191C, p. 143, pl. 14, figs. 10—13, pl. 15, figs. 10—13. 
Turritopsis nutricula, Bigelow, 1913, p. 8. 

Locality. Chagos Archipelago, Salomon Atoll, Surface. 5 July, 1905, Q. 
2 specimens. 

The umbrella is about as high as broad (2°5 mm.), with a rounded summit; its 
sides are slightly curved inwards about the middle and the walls are rather thin. Velum 
moderately broad. The stomach is large and cross-shaped. The mouth has four very 
short perradial lips and its whole margin is lined with isolated, globular, clusters of 
nematocysts. Four fairly broad radial canals. The radial canals on leaving the stomach 
proper are very wide and have very thick walls, which coalesce, so that the stomach 
appears to hang from a thick cellular peduncle, about one-third its length. The gonads 
are interradial upon the walls of the stomach; one specimen has ova and the other 
spermatozoa. The tentacles are arranged im a single row on the margin of the umbrella 
(one specimen with 56, the other with 45 tentacles). They have a smooth, even surface, 
but are densely covered with minute nematocysts, and terminate with a bulbous 
enlargement. The basal bulbs of the tentacles vary in size, especially in length. They 
adhere on the outer side to the margin of the umbrella, and the basal half of the inner 
side is covered with a semicircular loop of nematocysts. There is a small reddish-looking 
ocellus on the inner side of every bulb, situated close to the tentacle. 

The specimens in the “Sealark” collection are similar to the published figures of 
Turritopsis nutricula. The formation of the endodermal peduncle is identical with that 
of Turritopsis nutricula, and not at all like that of Turritopsis polycirrha. 

Bigelow (1913) in his description of Turritopsis nutricula from Japan calls attention 
to a distinct terminal dilatation of the tentacles, and he points out that similar dilatations 
also oceur in specimens taken on the American coast at Newport. It is strange that these 
dilatations have not been previously noticed by the American naturalists. The specimens 


BROWNE—MEDUSA FROM THE INDIAN OCEAN ; 181 


in the “‘Sealark” collection also show a large hollow enlargement of the distal end of 
some of the tentacles. I am rather inclined to regard these dilatations in some way 
due to the action of the killing re-agents used. I can find no such dilatations in 
Turritopsis polycirrha ; their tentacles in a semi-contracted condition are rather club- 
shaped, as shown in the figure of Turritopsis nutricula by Mayer (1910, pl. 14, fig. 18). 

The hydroid of Turritopsis nutricula is known under the name of Dendroclava, 
which is probably identical with Tubielava of Allman. 

The medusoid genus has been previously recorded from the Indian Ocean. Vanhéffen 
(1911) found a damaged specimen of Turritopsis between the Chagos Islands and the 
Seychelles. Bigelow (1904) found a young Turritopsis in Felidu Atoll, Maldives. 

Distribution. North Atlantic along the coast of United States. Pacific Ocean, 
Japan. 

10. AMPHINEMA sp. ? 


Locality. Amirante Islands, Desroches Atoll. 200—0O fms. 16 Oct. 1905, kk. 
1 specimen. 

This little medusa was preserved in alcohol and is in rather a fragile condition. 

The umbrella is thin, a little less than 2 mm. in length and width, with apparently 
a small apical projection. The stomach is not situated upon a peduncle. The mouth 
has four small lips. The whole manubrium projects about half its length beyond the 
margin of the umbrella. There are four inconspicuous radial canals, which apparently 
leave the stomach a little way down its sides, and slight traces of very small incon- 
spicuous mesenteries, but as it is difficult to estimate the amount of contraction and 
shrinkage the presence of mesenteries remains doubtful. The gonads form eight adradial 
folded bands along the upper half of the manubrium, and each contains a few fairly 
large ova. 

There are two large, opposite, perradial tentacles, with large tapering basal bulbs, 
slightly laterally compressed. The basal bulbs are very opaque and of a dark brownish 
colour ; but probably of quite a different colour when alive. On each half of the margin 
of the umbrella, between the two tentacles, there are five very small bulbs, little longer 
than broad, and evidently do not develop large tentacles. No trace of an ocellus could 
be seen on any of the bulbs. 

This is an adult medusa probably belonging to a Perigonimus-like hydroid. 
I cannot determine the species with any degree of certainty, as it does not show 
clearly any well-marked character. A few more specimens were wanted, and in better 
condition. 


Genus LEUCKARTIARA, novum nomen, Hartlaub, 1914. (T1aRa, preoccupied name.) 


11. LeUCKARTIARA GARDINERI, nova species (Pl. 39, fig. 4). 


Locality. Amirante Islands, Surface. 10 Oct. 1905, gg. 1 specimen. 

Description of the Species:—Umbrella conical, about twice as high as broad, with 
rather thin wails. Four very conspicuous perradial canal-like bands projecting from the 
surface of the ex-umbrella, and extending from the tentacles nearly to the summit of 


z \ 


LIBRARYi= 
woe | 


182 PERCY SLADEN TRUST EXPEDITION 


the umbrella. Stomach large, occupying more than half the cavity of the umbrella. 
Mouth large, and its margin in folds. Four fairly broad perradial canals, without any 
lateral processes, and with a slit-like union with the stomach along its whole length ; 
thus forming the so-called “mesenteries” which attach the stomach perradially to the 
wall of the sub-umbrella. Four gonads, isolated perradially, and arranged in eight 
adradial bands, which are composed of bifurcated transverse folds and united interradially 
by a transverse fold. Four long perradial tentacles, with laterally compressed basal 
bulbs clasping the margin of the umbrella. Also very minute interradial and adradial 
tentacles, and a few marginal bulbs without tentacles. A blackish ocellus on the outer 
side of all basal bulbs of the minute tentacles and marginal bulbs. 

Size. Umbrella about 3°5 mm. in width and 6 mm. in height. 

The collection contains only one specimen, and it is in a splendid state of preser- 
vation, but with the margin contracted imwards. 

This new species, which I have great pleasure in naming after the leader of the 
“Sealark” expedition, has a character well-marked by the presence of four perradial 
canal-like bands upon the ex-umbrella. These bands are probably brightly coloured 
in the sea. They extend beyond the margin on to the outer edge of the basal bulbs 
of the large tentacles and.contain nematocysts. Although ocelli are clearly visible on 
the basal bulbs of the minute tentacles, still they cannot be seen upon the basal bulbs 
of the large perradial tentacles. I am inclined to take the view that this species has not 
more than four large tentacles and that the minute tentacles remain in a rudimentary 
condition. The gonads show ova fairly well advanced, so that the specimen is not an 
early stage. 


12. PANDA juv. 


Locality. Chagos Archipelago, Salomon Atoll. 10—0O fms. 1 July, 1905, P. 
1 specimen. Surface. 5 July, 1905, Q. 2 specimens. 

The specimens are young stages in a contracted and crumpled condition. The smallest 
is. about 2°5 mm. in diameter, its umbrella has conspicuous longitudinal ridges carrying 
nematocysts. There is a nematocyst track corresponding to every tentacle and bulb. 
The perradial ridges extend to the summit of the umbrella and the other ridges or tracks 
are shorter, their length and size being correlated with the age of the tentacle. The 
development of the gonads is just commencing and their position is marked by small 
isolated pits in the wall of the stomach. There are seven tentacles and a few adradial 
bulbs, the latter are at different stages of growth. A conspicuous black ocellus is situated 
on the outer side of all the basal bulbs of the tentacles and on some of the larger adradial 
bulbs. 

The second specimen is a little older and about twice the size of the smallest specimen. 
In this specimen the stomach has flattened out and the mouth expanded to its extreme 
limit. The four perradial canals leave the stomach close to the four corners of the mouth. 
There are nine tentacles and seven tentacular bulbs, each one having a black ocellus. 

The third specimen closely resembles the last one. It shows very clearly that the 
adradial tentacles do not all begin to develop at the same time. 


BROWNE—MEDUS4 FROM THE INDIAN OCEAN 183 


These early stages are either identical with or closely related to Pandea conica 
(Quoy et Gaimard) which is a common species in the Mediterranean. Wanhdéffen (1911) 
records the occurrence of Tiara (Pandea) conica, Lesson, in the Agulhas Current, Indian 
Ocean. 


Genus HetTrrotiarA, Maas, 1905. 


There are two species’ belonging to the genus Heterotiara, namely Heterotiara 
anonyma, Maas (1905) and Heterotiara minor, Vanhéffen (1911). A complete description 
of Heterotiara anonyma could not be given by Maas as the specimens had lost their 
tentacles. Vanhéffen on the cruise of the “ Valdivia” obtained Heterotiara anonyma as 
well as Heterotiara minor, and again Heterotiara anonyma was taken without tentacles. 

Bigelow (1909) found, in the material collected by the “ Albatross” in the Humboldt 
Current off Peru, two specimens of a medusa which he has fully described and figured 
under the name of Heterotiara anonyma. These specimens have their tentacles 
tapering to a point, and therefore are not like those of Heterotiara minor terminating 
with a bulb. 

_ Bigelow (1913) also records Heterotiara anonyma from the Bering Sea, where the 
“ Albatross” obtained many specimens. Their tentacles, however, terminate in a spherical 
knob much the same as in Heterotvara minor. On the same cruise Heterotiara minor was 
plentifully found at the Philippmes, but the specimens have not yet been fully described. 
Bigelow states that Heterotiara munor and Heterotiara anonyma are undoubtedly distinct, 
“They are separated by the number of tentacles correlated with size, H. anonyma having 
twelve tentacles (or less) when adult, and reaching a height of 20 mm., whereas H. minor 
has about twice as many tentacles, though much smaller (only about 10 mm. high).” 

Bigelow has described and figured Heterotiara anonyma from off the coast of Peru 
with tentacles tapering to a point, and from the Bering Sea with tentacles terminating in 
a bulb, as in Heterotiara minor. It seems to me that either the Peruvian specimens have 
been inaccurately described or that they belong to another genus, and I am inclined to 
take the latter view. 


13. HetTERoTIARA MINOR, Vanhdoffen, 1911. 


Heterotiara minor, Vanhoffen, 1911, p. 212, Taf. 22, fig. 5, Text-fig. 8. 
Heterotiara minor, Bigelow, 1913, p. 25. 

Locality. Lat. 4° 16’S., Long. 71° 53’ E. (North of Chagos). 1200—0 fms. 17 May, 
1905, C. 1 specimen. 

This single specimen is in fairly good condition. Its umbrella measures 13 mm. in 
height and 12 mm. in width. The stomach is contracted into folds and hangs free inside 
the-cavity of the umbrella. There are four radial canals and no centripetal canals. The 
tentacles are transparent, hollow, and terminate with a large hollow bulb, thickly covered 
with nematocysts. There are nineteen tentacles, varying slightly in size, the differences 
being due to age. The tentacles have no definite basal bulbs, but their basal portion is 
partly embedded in the jelly and situated in grooves on the margin of the umbrella. The 
specimen resembles Vanhéffen’s text-figure 8. 


184 PERCY SLADEN TRUST EXPEDITION 


14. PRoBoscIDACTYLA TROPICA, Browne, 1904. 
Willsia sp. Huxley, 1877, p. 120, fig. 17. 
Proboscidactyla tropica, Browne, 1904, p. 727. 

Locality. Amirante Isles, Desroches Atoll. 100—0O fms. 16 Oct: 1905, kk. 
1 specimen. 

Huxley, when off the Louisiade Archipelago in 1849, found a medusa which he briefly 
described and figured under the name of Willsta. In my revision of the Williadz (1904) 
I recognised this medusa as a distinct species and called it Proboscidactyla tropica. 

Bigelow (1909, p. 220) considers Proboscidactyla tropica to be identical with 
Proboscadactyla ornata McCrady, 1857, var. stolonifera, Maas, 1905. 

One of the characters, which I made use of to distinguish Proboscidactyla tropica 
from all the other species of the genus, was the form of a blastostyle bearing a cluster of 
nematocysts at its free end, and upon the sides of this blastostyle the medusa-buds develop. 
This blastostyle is similar in structure to the blastostyle of the hydroid Lar sabellarum, 
upon which medusa-buds develop, which when set free belong to the medusoid genus Willa. 

Maas (1905) in his description of the variety stolonifera from the Malaysian region 
makes no mention of this peculiar blastostyle, and if it had been present in his specimens 
he could not have failed to have noticed it. 

Bigelow (1909, p. 219) states that Proboscidactyla gemmifera (Fewkes) is only the 
budding form of Proboscidactyla ornata, McCrady, 1858. I cannot very well link 
Proboscidactyla tropica to Proboscidactyla ornata by means of Proboscidactyla gemmifera, 
as Brooks (1880) states clearly that the stolons have branches and each branch terminates 
with a medusa-bud. There is here also no mention made of any blastostyle-like stolon 
with a terminal cluster of nematocysts. Mayer's figures (1910, Pl. 21) confirms Brooks's 
statement. Under these circumstances I prefer to retain Proboscidactyla tropica as a 
distinct species until at least we know more about the life-histories of the other species 
of the genus. 

The specimen in the “‘Sealark” collection is about 2 mm. in diameter, and has lost 
its natural shape owing to contraction. The velum is very narrow. The stomach has 
four perradial lobes, from which the four main radial canals leave to join the tentacles. 
As in other species of the genus there is no circular canal. Each main radial canal has 
the appearance of being twice dichotomously branched, each terminal branch going to a 
tentacle. At the juncture of the first branch of each canal is situated a blastostyle-lke 
stolon. This blastostyle has rather the appearance of a tentacle, for at its free end there 
is a large cluster of nematocysts. The medusa-buds are at different stages of development. 
Two to four on each stolon, and the largest is nearly ready for liberation. They are 
arranged round the sides of the stolon. Huxley, however, states that the buds are uni- 
lateral. He saw them alive, whereas I have only a contracted specimen. In one of the 
perradii at the junction of the second branch there is a second blastostyle-like stolon with 
a small bud developing. So apparently the medusa has stolons on the junction of the 
second branch as well as on the first. 

There are sixteen tentacles, rather thick, in a contracted condition, with a thick semi- 
circular band of nematocysts on the inner side of the basal bulbs, which project into 


BROWNE—MEDUS FROM THE INDIAN OCEAN 189 


the substance of the umbrella. Between every two tentacles are isolated clusters of 
nematocysts, which extend over the ex-umbrella from the margin up to the summit, and 
these clusters are similar to those described in the other species of the genus. Though 
the specimen is in fairly good condition, it is not suitable, owing to contraction, for the 
making of a good drawing. 


15. PROBOSCIDACTYLA sp. ! 


Locality. Farquhar Group, Providence Is., Surface. 3 Oct. 1905, x. 1 specimen. 

Umbrella about 1:25 mm. in diameter. Five main radial canals, each with a lateral 
branch. Ten tentacles. No medusa-beds. 

The specimen is not in very good condition. The number of radial canals shows that 
it is abnormal, and as each canal has only one branch it is evidently a young stage. 


16. ANTHOMEDUSA, Genus ?. 


Locality. Mauritius. 300—0 fms. 22 Aug. 1905, C. 1 specimen. 

The umbrella is in a contracted condition, about 5 mm. in width and probably when 
alive about the same in height, rather thick. It has on its exterior surface four conspicuous, 
prominently projecting, perradial ridges, which extend from the-margin to, or nearly to, the 
top of the umbrella. These ridges have rather a blunt rounded edge, with what appears 
to be a narrow shallow groove running along the middle of the edge. From the shape and 
general appearance I do not think that these ridges were formed artificially either by 
preservation or by contraction, but existed in the living specimen. I am not so sure, 
however, about the groove along the middle of the ridge, it might be due to shrinkage. 
I cannot recall seeing a figure of any Anthomedusa with ridges, such as appear in this 
specimen. The velum is very narrow, and the sub-umbrella is well covered with a layer’ 
of fine muscles. The stomach is rather contracted, and has a plain circular mouth. In its 
contracted condition it is rather broad, and sac-shaped. It is not likely to extend beyond 
the margin of the umbrella even when fully stretched out. There are four conspicuous 
radial canals. The wall of the stomach has the appearance of being covered with male 
gonads, and if so, then the gonads are at an early stage as there is no definite swelling. 

The medusa has only two opposite perradial tentacles, both of which are unfortunately 
broken off at the basal bulbs and leaving not the slightest clue as to their structure. The 
basal bulbs are rather large and globular, and are attached to the margin of the umbrella, 
but do not project into the substance of the umbrella. The other two opposite perradii 
are naturally without any tentacles and also without any rudimentary bulbs, and the 
whole margin is also without minute rudimentary bulbs. There is apparently a patch of 
nematocysts on the margin of the umbrella in each of the four perradii, and there is also 
evidence of a short extension of them along the ridges of the ex-umbrella. No ocelli could 
be found. 

With the complete loss of the tentacles it is impossible to determine the genus. It 
has certainly only two opposite tentacles, so it may be a Dicodoniwm, a Zanclea, or even 
a new genus. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 24 


186 ~ PERCY SLADEN TRUST EXPEDITION 


LEPTOMEDUS A. 
17. Laopice ?. 


Locality. Amirante Isles, Surface. 9 Oct. 1905, go. 1 specimen. 

This specimen is probably a young Laodice, but the absence of gonads, which have 
not yet begun to develop, makes the determination of the genus uncertain. The umbrella 
is saucer-shaped, about 4 mm. in diameter, and moderately thick. The tentacles, not 
exceeding sixty in number, are very slender and short, with small basal bulbs partly 
extending over the margin of the umbrella and without a well-defined spur. There are 
a few marginal bulbs, from some of which tentacles have been broken off. Between every 
two tentacles.or bulbs there is generally a single cordylus. Cirri are also present, but not 
many remain. A conspicuous black globular ocellus is situated on the inner side of most 
of the basal bulbs of the tentacles.and marginal bulbs. 


18. Traropsis RosEA, Agassiz and Mayer, 1899. 
Tiaropsis rosea, Maas, 1905, p. 30, Taf. 7, figs. 45—47. 
Tiaropsis rosea, Mayer, 1910, p. 260. 

Locality. Mauritius. 125—0 fms. 22 Aug. 1905, a. 1 specimen. 

Description of the specimen :-—Unmbrella, about 3 mm. in diameter, moderately thick, 
and hemispherical. Velum broad. Stomach cross-shaped. Four radial canals. Gonads 
upon the radial canals, forming globular sacs near the stomach. Four large perradial 
tentacles, with pigmented basal bulbs. Seven marginal bulbs between every two tentacles. | 
Eight adradial sensory vesicles, with a large black roundish ocellus, situated at the base 
of the vesicle and on the inner side. 

This specimen, though much smaller in size, agrees very closely with a Tiaropsis 
described by Maas from Damar, Malay Archipelago. Maas’ description and figures are 
based upon a single adult specimen about 15 mm. in diameter, and he considers it to be 
the adult stage of Tiaropsis rosea of Agassiz and Mayer. It has yet to be definitely 
proved that Traropsis rosea of Agassiz and Mayer found in Suva Harbour, Fiji, is an 
early stage of Tiaropsis rosea, Maas. 

The arrangement of the marginal bulbs may help to determine this species. There 
are seven in each quadrant, situated as follows :—three in the interradial portion, and 
two on the perradial side of the two sense organs. The gonads do not extend along the 
radial canals as figured by Maas, but the specimen has the appearance of being immature 
and has probably not reached its maximum growth. 


PHIALIDIUM sp. ? 
There are apparently two species of Phialidium in the collection, but the specimens 
are not in a condition suitable for the determination of species. 
19. PHTaALiprum sp. A. 


Locality. North of Chagos, Lat: 4° 16’ S., Long. 71° 53’ E., Surface (Temp. 86° F.). 
17 May, 1905, B. 5 specimens. 
Umbrella hemispherical, 3—5 mm. in diameter. Stomach quadrangular, on a slight 


BROWNE—MEDUSA FROM THE INDIAN OCEAN 187 


thickening of the sub-umbrella. Mouth with four lips. Gonads close to the margin of 
the umbrella and extending over about one-quarter the length of the radial canals. 
Tentacles, 14—16, with globular basal bulbs. One or two sensory vesicles between every 
two tentacles, very minute and probably have contained only one otolith. 


20. PHIALIDIUM sp. B. 


Locality. Amirante Isles, Surface (Temp. 79° F.). 9 Oct. 1905, ge. 6 specimens. 
Surface. 17 Oct. 1905, 00. 3 specimens. 

Umbrella hemispherical, 5—6 mm. in diameter. Gonads close to the margin of the 
umbrella, linear or oval sacs extending over about one-quarter the length of the radial 
canals. Tentacles, 30—40, with globular basal bulbs. One to three sensory vesicles 
between every two tentacles. 


21. IRENE sp. ? A. 


Locality. Nazareth Bank, Cargados Carajos, Surface. 30 Aug. 1905, g. 
1 specimen. 

This is an early stage without gonads. The umbrella is about 6 mm, in diameter 
and very thin. The stomach is small and upon a short peduncle. There are eight 
tentacles, with globular basal bulbs, and three to four marginal bulbs between every two 
tentacles. All the bulbs have a pair of cirri adjacent to them, and excretory pores are 
visible. A few sense organs were found, and there is probably one between every two 
marginal bulbs and tentacles. 


22. IRENE sp. ? B.- 


Locality. Saya de Malha Banks. 58—0 fms. 6 Sept. 1905, C, 14. 11 specimens. 

These specimens are all in bad condition, so that only a general description can be 
given just to indicate that the Irene-like medusa occurs in that district. Their umbrellas 
measured 30 to 40 mm. in diameter and are moderately thick. From the centre of the 
umbrella hangs down a broad thick peduncle, about 20 mm. in length. Upon it a broad 
stomach is situated, and its mouth has four lips with the margin closely folded. It 
is a large mouth even in a contracted condition. There are four radial canals. 
The gonads, some containing well-developed ova, form laminar bands along the canals 
upon the sub-umbrella, but they do not extend down the peduncle. Some of the 
specimens have the above-mentioned organs in fairly good condition, but all have the 
margin of the umbrella, either completely denuded of its appendages or in such a frag- 
mentary condition that not even one quadrant is complete. I consider there should have 
been fifty or more tentacles, about three marginal bulbs between every two tentacles, and 
at least one small sensory vesicle between every two bulbs. Hxcretory pores are visible 
in connection with the basal bulbs of the tentacles and the marginal bulbs. I can find no 
definite indication of marginal cirri, The margin is not only broken but has rather 
a macerated appearance, therefore it is not advisable to describe it in detail or base 


drawings upon it. 
24 —9, 


188 PERCY SLADEN TRUST EXPEDITION 


23. MesoneMA PENSILE (Modeer) 1791. 


Mesonema pensile, Browne, 1904, p. 733, pl. 55, fig. 4, pl. 57, figs. 2—9. 
Mesonema pensile, Browne, 1905, p. 147, pl. 2, figs. 11—15. 

Mesonema pensile, Maas, 1905, p. 42, Taf. 8, fig. 52. 

Mesonema pensile, Maas, 1909, p. 26. 

Locality. Chagos Archipelago, Diego Garcia. 10—14 fms. 10 July, 1905. 
1 specimen. 

The genus Mesonema of Eschscholtz has undergone several changes in its definition. 
In 1904 I defined it as follows :—‘ Aquoride with numerous, simple, unbranched radial 
canals. Stomach circular, with lower wall quite rudimentary. Mouth nearly as large as 
the diameter of the stomach and cannot be closed.” The type species of the genus is 
Mesonema pensile (Modeer) 1791, and specimens similar to the type were described and 
figured by me in 1904. Both Mayer (1910) and Bigelow (1909) have raised objections 
to my distinguishing marks of the genus, and by which I separated it from Afquorea. 
They rightly say that owing to the great contractibility of the lower wall of the stomach 
it is difficult to assign to the correct genus certain species of the Aiquoride, and therefore 
they consider it is advisable to recognise Hquorea only, and place Mesonema in its list of 
synonyms. I clearly foresaw this difficulty, but had hopes that further researches on 
the Aiquoride might lead to the permanent retention of Mesonema by adding other 
characters so as to definitely separate it from A/quorea. 

Vanhéften (1911) also rejects my definition of Mesonema and brings forward a new 
definition as follows :— 

Mesonema. Leptomedusze with numerous radial canals, and with large tentacles 
considerably less in number than the canals. Tentacles with triangular and especially 
heart-shaped bulbs. 

Aiquorea. Leptomedusze with numerous radial canals, with fully developed 
tentacles about as many or more than radial canals. Tentacles with conical basal 
bulbs. 

Vanhoffen’s characters for separating the two genera may be an improvement on 
mine. It is quite easy to isolate the extremes as separating characters, whether one uses 
the size of the mouth, the number of canals in relation to the number of tentacles, or the 
shape of the basal bulbs, but the difficulty is to assign to the right genus those specimens 
which come near the border line. We have a very imperfect knowledge of the species 
belonging to the genus Aquorea, using the term in its widest sense. Some of the 
species exist on paper only, as their descriptions are too vague for any further use, a 
few have been fully described and figured, and many disputed as to their validity 
by systematists. 

Vanhoffen (1911) considers that Mquorea macrodactyla, Brandt, Aiquorea maldi- 
vensis, Browne, and Afquorea parva, Browne, are all identical with Mesonema celum 
pensile of Modeer. It is still, however, my firm opinion that 4quorea macrodactyla and 
Mesonema pensile are quite distinct species. Although the original descriptions of these 
two old species are rather vague and their figures more like rough sketches, still their 
names are now associated with medusee which have been described and figured according 


BROW NE—MEDUS4 FROM THE INDIAN OCEAN 189 


to modern requirements. At present, I have no valid reasons for ranking Avquorea 
parva aS a synonym. 

It appears to me, in spite of the descriptions and figures given by Maas and myself 
of Mesonema pensile and Aquorea macrodactyla, that Vanhéffen has failed to see clearly 
the difference between the two species. In his sketch (1911, p. 223, text-fig. 21) of the 
margin of umbrella showing the basal portion of the tentacles of Mesonema pensile, they 
appear to be very much like the basal bulbs of Aiquorea macrodactyla ; they are not at 
all like the basal bulbs of a genuine Mesonema pensile. 

In the “Sealark” collection there is only one specimen, which is in very bad 
condition. By the shape of the basal bulbs I think it is Mesonema pensile. The 
umbrella is about 15 mm. in diameter. There are eight tentacles. The radial canal 
system is practically destroyed, just a few canals left. 

Distribution. Tropical Pacific and Indian Oceans. Maas (1909) records it from 
Sagami Bay (35° N., 139° 37’ E.), Japan. 


24, AXQUOREA MACRODACTYLA (Brandt) 1834. 


Aiquorea maldivensis, Browne, 1904, p. 732, pl. 56, figs. 4—12. 
Mesonema macrodactylum, Maas, 1905, p. 40, Taf. 8, figs. 51 and 54. 
quorea macrodactyla, Bigelow, 1909, p. 37, pl. 36, figs. 5—10. 

Localities. North of Chagos. 1200—0O fms. 17 May, 1905, C. 1 specimen. 
Chagos Archipelago, Salomon Atoll. 180—0 fms. 30 June, 1905, O. 1 specimen. 
Saga de Malha. 55—0 fms. 6 Sept. 1905, C. 14. 8 specimens. 

Aiquorea macrodactyla of Brandt received a description according to modern 
requirements by Maas in 1905, and more recently Bigelow has published an account 
of some specimens taken in the Eastern Tropical Pacific. As the two best specimens 
in the “‘Sealark” collection agree very well with the descriptions given by Maas and’ 
Bigelow, I have no doubt that we have all seen the same species. 

In one of my earlier publications on Medusz I pointed out that the shape of the 
basal bulbs of the tentacles was a useful aid in the determination of the species belonging 
to the Aiquoride. There are distinct types of basal bulbs, and for each type an allowance 
must be made for variation and also for contraction or distortion due to preservation. 
Aiquorea macrodactyla has the basal bulbs curling over or clasping a thickening of the 
margin of the umbrella, and the character is clearly seen when a tentacle is cut out from 
the umbrella and viewed sideways. 

Maas, when he re-described Aquorea macrodactyla, pointed out that its basal bulbs 
belonged to the same type as that found in .Hquorea maldivensis, which had then not 
long been described by me as a new species. Bigelow, however, has gone a step 
further and considers Aquorea maldivensis to be the same species as Aiquorea macro- 
dactyla. It would now be very hard for me to keep the two species apart except by 
hair-splitting differences which, as far as possible, should be avoided. 

The basal bulbs of the ‘‘Sealark” specimens are more like those figured by Maas and 
Bigelow for Aquorea macrodactyla than those figured by me for Aquorea maldivensis, 
but the type is the same and the differences would come within the range of variation. 


190 PERCY SLADEN TRUST EXPEDITION 


The oral lips are similar in shape and structure as those figured for Aquorea maldivensis, 
but are more numerous. Iam not sure that it would be advisable to take the number 
into consideration for one of the specific characters. Maas found the oral lips to be less 
than half as numerous as the radial canals. Bigelow does not mention them in his text, 
but a figure indicates fewer lips than canals. One of the ‘“‘Sealark” specimens has about 
the same number of lips as radial canals. 

All the specimens from the Saya de Malha Banks are in bad condition, and it is only 
just possible to determine the species in one specimen which has a few tentacles left. 
Their stomachs measured from 23—34 mm. in diameter, and radial canals numbered 
72 to 100. 

A few notes can be given on the specimens from the other stations. 

A, The umbrella measures 42 mm. in diameter and is moderately thick. The 
stomach measured 27 mm. and the mouth 23 mm. in diameter. Even allowing for the 
lower wall of the stomach being considerably contracted, I doubt very much if it could 
expand sufficiently so as to allow the mouth to close up. There are 92 radial canals, 
a few have developed much later than the rest and bear no gonads. Seventeen large 
tentacles and seven much smaller in size at different stages of development. There are 
between every two tentacles 5—10 marginal bulbs with excretory pores. This specimen 
shows in a marked manner the radial muscle bands extending from the proximal end of 
the radial canals, across the lower wall of the stomach, to the oral lips. There is a muscle 
band in connection with every radial canal. 

B. The umbrella is shaped like a biconvex lens and measures 45 mm. in diameter. 
The stomach is about 30mm. and the mouth about 20 mm. in diameter. Around the 
margin of the mouth are numerous lips, about as many radial canals, and of the latter 
there are seventy-nine. The gonads show ova and extend as laminar bands along nearly 
the whole length of the canals. There are 16 large tentacles and four smaller in size. 
Between every two tentacles are situated 8 to 10 marginal bulbs, the central one is 
usually larger than the other and shows signs of developing a tentacle. The sense organs 
are very numerous, usually 2 to 5 between every two bulbs. 

The excretory pores in both these specimens are very conspicuous on the inner side 
of the circular canal. They stand out like papillae. There is one corresponding to and 
opposite every tentacle and large marginal bulb. 

Distribution. Indian Ocean and Tropical Pacific. 


25. LepromepusA, No. 1. 


Localities. North of Chagos. 75—0 fms. 16 May, 1905, A. 1 specimen. Far- 
quhar Group, Providence Is., Surface. 5 Oct. 1905, y. 1 specimen. 

The specimens are young stages, without gonads, and they probably have not yet 
developed their generic characters. At all events, I cannot assign them to a definite 
genus. Mayer (1910) gives figures (Pl. 25, fig. 8, and Pl. 27, fig. 1) of early stages of 
a Dupleurosoma which somewhat resembles these specimens. 

Description :—The umbrella is thick, not quite so high as broad, about 4 mm. in 
diameter, without any apical projection. Velum narrow. The stomach is cross-shaped 


BROWNE—MEDUSA FROM THE INDIAN OCEAN 191 


when contracted, and quadrangular when expanded. It is fairly large and occupies the 
greater part of the top of the cavity of the umbrella. The mouth is fairly large, and has 
four lips. There are four radial canals, without any branching. Eight tentacles (4 per- 
radial and 4 interradial) with thick conical basal bulbs. Between every two tentacles 
there are three marginal bulbs, very small and short, a little longer than broad, and from 
their general appearance they evidently do not develop tentacles. A conspicuous roundish 
dark brownish ocellus is situated on the outer side of all the basal bulbs and on all the 
small marginal bulbs. No trace of any cordyli, cirri, or marginal sensory vesicles could be 
seen. One of the specimens is a little smaller and younger than the other. It has only 
two marginal bulbs between every two tentacles, and one of its interradial tentacles is 
missing. Both specimens are badly contracted, but in a fairly good state of preservation. 


26. LEepromepusa, No. 2. 


Localities. Farquhar Group, Surface. 2 Oct. 1905, w. 1 specimen. Amirante Isles, 
Desroches Atoll. 50—O fms., 200—0O fms. 16 Oct. 1905, kk. 2 specimens. 

Description :—Umbrella hemispherical, fairly thick, 5—8 mm. in diameter. Stomach 
small, quadrangular when expanded. Mouth with four large lips having a crenate 
margin. Four narrow radial canals. Gonads extending over the outer half of the radial 
canals. Tentacles 16 (perhaps more), small, with small globular basal bulbs. Three to 
seven (usually five) elongated marginal bulbs (?) between every two tentacles. Two 
sensory organs between every two tentacles. 

I cannot determine the genus to which these specimens belong owing to the condition 
of the marginal bulbs. These bulbs are elongated, and vary in length. They may 
be cirri in a state of contraction, or minute tentacles unlike the large ones, or genuine 
bulbs. I am in favour of their being contracted cirri. The sense organs are small, about 
large enough to contain a couple of otoliths, and are probably open sensory pits; if so, 
the specimens would belong to the Mitrocomide. The tentacles are not always equi- 
distant apart, hence the irregular number of bulbs between them. One specimen has 
a quadrant containing six tentacles. 


27. Lepromepusa, No. 3. ° 


Localities. Nazareth Bank, Cargados Carajos, Surface. 30 Aug. 1905, i. 1 specimen. 
Amirante Isles, Surface. 18 Oct. 1905, 00. 3 specimens. 

Description :—Umbrella rather flat, about 3 mm. in diameter. Stomach small and 
flat, similar to that of a Phialidiwm. Four radial canals (one specimen with five canals) 
without branches. Gonads sausage-shaped, along nearly the whole length of the radial 
canals. One specimen has conspicuous ripe ova. Tentacles small and slender, about 
40, and terminating with a small cluster of nematocysts. No signs of cirri, or rudi- 
mentary marginal bulbs between the tentacles. No sense organs of any kind could 
be found. 

All the specimens have rather a macerated appearance, and consequently it is not 
safe to rely upon the absence of sense organs. 


192 PERCY SLADEN TRUST EXPEDITION 


TRACHOMEDUS. 


28. OLINDIAS SINGULARIS, Browne, 1904. 


Olindias singularis, Browne, 1904, p. 737, pl. 56, fig. 2, pl. 57, fig. 1. 
Olindras singularis, Bigelow, 1909, p. 109, pl. 4, fig. 1, pl. 31, figs. 1—10, pl. 32, fig. 8. 
Olindias singularis, Mayer, 1910, p. 357. 


Locality. Chagos Archipelago, Diego Garcia. 10—14 fms. 10 July, 1905. 
4 specimens. 

This species was first described by me from a single specimen, which Professor Stanley 
Gardiner found on his expedition to the Maldive Archipelago. 

The chief specific character was based upon the presence of a single sense organ at 
the base of the primary tentacles. All the other species of the genus having a pair of 
sense organs in that position. 

Bigelow fortunately found no less than 23 specimens in Managreva Harbour, 
Paumotu Archipelago in the Tropical Pacific, and he has given a more adequate 
description of the species, accompanied by excellent figures. In that series of specimens 
Bigelow noticed that the presence of a single sense organ did not hold good, for in the 
largest specimens (30—60 mm. in diameter) single and paired sense organs were found 
in the proportion of about four to one. | 

On searching the margin of the umbrella of the specimens from Diego Garcia I could 
find only a single sense organ at the base of the primary tentacles in three specimens, 
but the fourth specimen showed the presence of pairs. In quadrant (A) thirteen singles 
occurred, in quadrant (B) twenty-three singles and two pairs, in quadrant (C) twelve 
singles and five pairs, in quadrant (D) thirteen singles. It will be noticed that one 
quadrant is more prolific in pairs than the others, and that two quadrants are 
without pairs. 

In these specimens the umbrella is somewhat flattened out, and measures 17 to 
22 mm. in diameter. There are about seven centripetal canals in each quadrant. All 
the specimens show a fair quantity of primary tentacles, but the secondary tentacles 
have all but disappeared. It was not until a special search was made for them that 
a few, at a very early stage of development, were found. The fully grown secondary 
tentacles were probably broken off in the net, and now only their stumps remain. All 
the four specimens have gonads well developed. 

In my report on the Maldive medusze I stated that the primary tentacles of Olindias 
did not possess adhesive pads or suckers, and included their absence amongst the generic 
characters. Mayer and Bigelow have, however, proved the presence of adhesive pads 
in Olindias tenuis, and no doubt they occur in all the species of the genus. The error 
on my part arose from using sections of material which had not been specially preserved 
for that purpose. I failed to distinguish the adhesive cells, and on finding plenty of 
nematocysts came to the conclusion that the primary tentacles terminated with a battery 
of nematocysts. 

Distribution. Tropical Pacific and Indian Oceans. 


BROWNE_MEDUS FROM THE INDIAN OCEAN 193 


29. RHOPALONEMA VELATUM, Gegenbaur, 1856. 


Rhopalonema velatum, Maas, 1893, p. 14, Taf. 1, figs. 5, 9—11. 

Rhopalonema velatum, Vanhoffen, 1902, p. 59, Taf. 10, fig. 16, Taf. 11, fig. 32. 
Rhopalonema velatum, Lo Bianco, 1904, p. 55, Taf. 34, fig. 137. 

Rhopalonema velatum, Maas, 1905, p. 50, Taf. 10, fig. 69. 

Rhopalonema velatum, Bigelow, 1909, p. 129. 

Rhopalonema velatum, Mayer, 1910, p. 378, text-figs. 214, 216, 218, 219. 
Rhopalonema velatum, Vanhoffen, 1912, p. 29. 

Rhopalonema velatum, Vanhoffen, 1912, p. 371. 


Localities. Chagos Archipelago; Mauritius; Farquhar Group; Amirante Isles. (The 
collection contained about sixty specimens taken at about twenty different stations.) 

The specimens collected by the ‘“‘Sealark” are about 2 to 5 mm. in diameter, and 
in different stages of development; the larger specimens have quite ripe gonads. The 
gonads form elongated swellings, not exceeding one millimetre in length and are usually 
situated about the middle of the radial canals. In shape these gonads are very similar 
to those of Rhopalonema velatum from Naples. 

Rhopalonema velatum and Rhopalonema ceruleum have one character in common, 
namely, a conical top-knot on the summit of the umbrella. Bigelow found the top-knot 
to be very constant in his Pacific specimens. Although, at first, the top-knot was 
not regarded as of any importance, later its absence became useful for distinguishing 
Rhopalonema funerarvum, Vanhétten, from the other species of the genus. Nearly all 
the specimens in the “Sealark” collection show clearly the top-knot on the summit 
of the umbrella, but a few do not. The top-knot when properly formed stands out 
as a kind of conical projection on the top of the umbrella, and is usually marked off 
from the rest of the umbrella by a transverse circular furrow or depression. In shape 
and size it varies considerably, and is scarcely recognisable in extreme cases, as only 
a slight depression in the contour of the upper part of the umbrella is present. In this 
collection there are a few specimens which show no signs of a depression. These specimens 
are about 5 mm. in diameter, with well-developed gonads, and have rather a conical-shaped 
umbrella. Although certain specimens have not the characteristic top-knot, still there 
is not sufficient evidence to connect them with Rhopalonema funerarium. This latter 
species has a differently shaped umbrella and apparently does not begin to develop its 
gonads until about 6 mm. in diameter, and the gonads extend over the outer two-thirds 
of the radial canals. It lives at a greater depth than Rhopalonema velatum, and belongs 
to the mesoplankton. 

According to Vanhéffen Rhopalonema velatum has eight perradial tentacles, eight 
interradial cirri, sixteen adradial cirri, and eight sense organs adjacent to the interradial 
cirri. Certain authors, however, have described and figured Rhopalonema velatum with 
sixteen sense organs. Maas (1893) gives the number of sense organs as sixteen, and 
figures two octants with three sense organs in each, thus showing that the number may 
even exceed sixteen. Mayer (1910, p. 380) gives a figure of Rhopalonema velatum 
drawn by himself at Naples, and it shows clearly sixteen sense organs, eight adjacent 

SECOND "SERIES—ZOOLOGY, VOL. XVII. 25 


194 PERCY SLADEN TRUST EXPEDITION 


to the perradial tentacles, and eight adjacent to the interradial cirri. Bigelow (1909) 
could only find eight sense organs in specimens collected in the Eastern Pacific. 

There should be no difficulty in seeing sense organs in living specimens, but after 
specimens have been preserved for some time the sense organs frequently have a 
marvellous way of either disappearing or so changing their appearance that it is not an 
easy matter to recognise them. Among the specimens in the “Sealark” collection 
I have found three with sense organs adjacent to the perradial tentacle and in one of 
the specimens the root of the adradial cirrus was visible. The tentacles and cirri are 
broken off in most of the specimens. When any are present it is the interradial cirrus. 
Only about three specimens have any perradial tentacles left and the adradial cirri have 
either not developed or else broken off at the base. 

Rhopalonema velatum is widely distributed throughout the warm regions of all the 
oceans. It is generally found at or near the surface. 

Bigelow (1909) after an examination of a large series of specimens collected by the 
“Albatross” in the Eastern Tropical Pacific, has come to the conclusion that Rhopalonema 
velatum, Gegenbaur, and Rhopalonema ceruleum, Haeckel, are identical. In 1906, when 
I reported upon specimens of Rhopalonema caruleum collected in the Bay of Biscay, 
I certainly felt sure that Rhopalonema velatum and [hopalonema cerulewm were 
distinct species. The distinguishing character was based upon the shape of the gonads, 
which in Rhopalonema velatum are either globular or oval, but in the Biscayan 
Rhopalonema ceruleum the gonads form narrow bands, which occupy the central 
third of the radial canals. Although I have re-examined the Biscayan specimens, still 
I am not yet convinced that they are identical with Rhopalonema velatum. 

Mayer (1910, p. 380) regards Rhopalonema ceruleum, Haeckel, as a distinct and 
good species. Under its name he has placed the following :—Rhopalonema caruleum, 
Maas (1905); Browne (1906). Rhopalonema funerarvum, Vanhoffen (1902); Bigelow 
(1909). Bigelow considers the Rhopalonema ceruleum, Haeckel and Browne, to be 
identical with Rhopalonema velatum, Gegenbaur. Mayer, on the other hand, regards 
it as identical with Rhopalonema funerarium. I cannot agree with Mayer's synonymy, 
for, I believe, that he has mixed up two distinct species; Rhopalonema caruleum, Haeckel 
and Browne, belongs to one species, and Rhopalonema funerarium, Vanhéffen (1902), 
Bigelow (1909); Rhopalonema ceruleum, Maas (1905) belong to another species. 


30. SMINTHEA EURYGASTER, Gegenbaur, 1856. 


Sminthea eurygaster, Gegenbaur, 1856, p. 245, Taf. 9, figs. 14—15. 
Trachynema eurygaster, Haeckel, 1879, p. 260. 

Trachynema mammeforme, Haeckel, 1879, p. 262, Taf. 17, figs. 13—15. 
Smainthea eurygaster, Metschnikoff, 1886, p. 244, Taf. 1, figs. 18 —20. 
Trachynema euryguster, Maas, 1893, p. 12. 

Trachynema eurygaster, Browne, 1906, p. 171. 

Sminthea ewrygaster, Mayer, 1910, p. 383, text-figs. 226—227. 


Localities. North of Chagos, Lat. 4° 16’S., Long. 71° 53’ KE. 125—0 fms. 17 May, 
1905, B. 1 specimen. 50—O fms. 18 May, 1905, F. 2 specimens. Chagos Archipelago, 


BROWNE—MEDUSA FROM THE INDIAN OCEAN 195 


Peros Atoll. 75—0 fms. 30 June, 1905, M. 1 specimen. Amirante Isles, Desroches. 
100—O fms. 16 Oct. 1905, kk. 1 specimen. 

Mayer (1910) has revived the old generic name Sitiathea of Gegenbaur and defines 
the genus as follows :—‘‘ Trachymedusze with only eight tentacles, one at the foot of each 
of the eight radial canals. In other respects this genus is similar to Rhopalonema.” 
The chief advantage in the use of the name Sminthea is that it has a good type species, 
known as Sminthea eurygaster. It is quite easy to recognise this species when the 
gonads are present as they form globular swellings on the radial canals adjacent to the ~ 
margin of the umbrella. In the early stages with the gonads undeveloped, it is necessary 
to rely upon the absence of any intercanal tentacles or cirri, and take the risk of their 
being early stages belonging to another genus. 

The specimens in the ‘‘Sealark” collection are between 2 to 3 mm. in diameter and 
some have gonads. The tentacles are all broken off at their base. 

This is rather a rare medusa. It has been previously recorded from the Mediterranean 
and the Atlantic. Its southernmost record in the Atlantic being in the South Equatorial 
Current off the north coast of Brazil. 


31. PANTACHOGON RUBRUM, Vanhoffen, 1902. 


Pantachogon rubrum, Vanhéffen, 1902, p. 63, Taf. 9, fig. 9, Taf. 10, figs. 19—20. 
Pantachogon rubrum, Maas, 1905, p. 55, Taf. 10, fig. 66. 
Pantachogon rubrum, Mayer, 1910, p. 389, text-figs. 240—241. 

Localities. Chagos Archipelago, off Peros Banhos, 600—0 fms. 30 June, 1905, N. 
1 specimen. Between Providence Is. (Farquhar Group), and Alphonse Is., Lat. 8° 16’S., 
Long. 51° 26’ KE. 900—0 fms. 6 Oct. 1905, aa. 1 specimen. 

Unfortunately both specimens are in bad condition, but it is just possible to determine 
the species. The umbrella is dome-shaped, with an evenly rounded summit, about 7 mm. 
in width and 5 mm. in height. The stomach is about 2 mm. long, without a peduncle, and 
its mouth has four short lips. The gonads are evidently just beginning to develop on the 
lower half of the eight radial canals. The tentacles are indicated by their stumps, and 
there are evidently eight in each octant. 

The second specimen is of about the same size and shape as the first one. It is, 
however, in far worse condition, but useful as it shows a certain amount of bright 
reddish coloration on the sub-umbrella. 

Distribution. Widely distributed in the warm regions of the Atlantic and Indian 
Oceans, and belonging to the mesoplanktonic zones. 


32. HALICREAS PAPILLOSUM, Vanhéffen, 1902. 


Halicreas papillosum, Vanhoffen, 1902, p. 68, Taf. 9, figs. 7—8, Taf. 11, fig. 30. 
Halicreas papillosum, Maas, 1905, p. 57, Taf. 10, fig. 70, Taf. 11, fig. 71. 
Halicreas papillosum, Bigelow, 1909, p. 138, pl. 3, fig. 3, pl. 33, figs. 8—9, pl. 34, figs. 1—3, 5, 8, 10, 11. 
Halicreas papillosum, Mayer, 1910, p. 391, figs. 242—243. 

Localities. South of Farquhar Group, Lat. 10° 27’8., Long. 51°17’ EK. 1000—0 fms. 
27 Sept. 1905, q. 3specimens. North of Farquhar Group, Lat. 8° 16’S., Long. 51° 20’ E. 


25—2 


ry 
196 PERCY SLADEN TRUST EXPEDITION 


900—O0 fms. 6 Oct. 1905, aa. 2 specimens. Amirante Isles. 750—0O fms. 16 Oct. 
1905, ll. 3 specimens. 16 Oct. 1905,mm. 400—0 fms. 2 specimens. 

All the specimens are in very bad condition, and but little better than clear lumps 
of jelly. 

This species has been very rarely taken near the surface, and it evidently belongs to 
the mesoplanktonic zone. 

Distribution. Throughout the tropical and sub-tropical regions of all the oceans. 


33. AGLAURA HEMISTOMA, Péron et Lesueur, 1809. 


Aglaura hemistoma, Haeckel, 1879, p. 275, Taf. 16, figs. 3—4. 
Aglaura hemistoma, Maas, 1893, p. 25, Taf. 1, figs. 12—13. 
Aglaura hemistoma, Vanhoffen, 1902, p. 78. 

Aglaura hemistoma, Lo Bianco, 1904, p. 55, Taf. 34, fig. 138. 
Aglaura hemistoma, Browne, 1906, pp. 176, 184. 

Aglaura hemistoma, Bigelow, 1909, p. 119, pl. 2, fig. 6. 

Aglaura hemistoma, Mayer, 1910, p. 398, pl. 46, figs. 4—5, pl. 49, figs. 3—7, pl. 50, fig. 11, text-figs. 250—251. 
Aglaura nausicaa, Haeckel, 1879, p. 274, Taf. 16, fig. 1. 

Aglaura hemistoma var. nausicaa, Maas, 1893, p. 26. 

Aglaura hemistoma var. nausicaa, Mayer, 1910, p. 400, fig. 252. 
Aglaura laterna, Haeckel, 1879, p. 274, Taf. 16, fig. 2. 

Aglaura hemistoma var. laterna, Maas, 1893, p. 25, Taf. 1, fig. 14. 
Aglaura hemistoma var. laterna, Mayer, 1910, p. 400, fig. 253. 
Aglaura prismatica, Maas, 1897, p. 24, Taf. 3, figs. 4—9. 

Aglaura prismatica, Agassiz and Mayer, 1899, p. 165, pl. 4, fig. 13. 
Aglaura hemistoma var. prismatica, Mayer, 1910, p. 400. 

Aglaura octogona, Bigelow, 1904, p. 257, pl. 2, fig. 9. 

Aglaura hemistoma var. octogona, Mayer, 1910, p. 401. 


For further synonyms and references see Mayer, 1910, pp. 397—401. 


Localities. North of Chagos; Off Mauritius; North of Saya de Malha Bank; 
Farquhar Group; Alphonse Is.; Amirante Isles. (The collection contained about 120 
specimens taken at 21 different stations.) 

After the eruise of the “ Valdivia” Vanhoffen came to the conclusion that only one 
species of Aglaura existed. Bigelow on the “ Albatross” cruise in the Kastern tropical 
Pacific paid special attention to Aglawra by examining specimens alive, and he also has 
decided in favour of a single species. Mayer is of the opinion that only one species exists, 
but he retains in his monograph the names of the varieties. 

Aglaura hemistoma belongs to the epiplanktonic fauna, and is widely distributed 
throughout the warm regions of all oceans and seas. In the Biscayan Plankton (Browne, 
1906) it was most plentiful at about 50—100 fms, scarcest at the surface. It did not 
occur in closing nets below 100 fms. On the “Sealark” expedition Professor Gardiner 
used the Wolfenden closing-net at only three stations and at each Aglawra hemistoma was 
taken. It occurred once at 250 fms, and twice at 500 fms. The serial hauls taken with 
open nets from different depths gave no reliable clue, owing to the paucity of specimens, 
as to the depth at which the species was most abundant. On each occasion the nets used 
within 50 fms of the surface contained specimens, and the numerous surface tow-nettings 
showed that it was not uncommon at the surface. 


BROWNE—MEDUSAi FROM THE INDIAN OCKAN 197 


Most of the specimens are in good condition and resemble figures given by Maas 
(1893, Taf. 1, figs. 12—14). The umbrella having its length greater than its width, but 
in a few the length and width are about equal. None exceeded 3 mm. in length. The 
peduncle showed great variability in its length. In some specimens it is scarcely visible 
owing to contraction, whilst in others it reaches halfway down the cavity of the umbrella. 
The gonads at an early stage are globular, and become cylindrical or sausage-shaped when 


fully grown. 


34, AMPHOGONA APSTEINI (Vanhoffen) 1902. 


Pantachogon apsteint, Vanhoffen, 1902, p. 65, Taf. 10, fig. 18, Taf. 11, fig. 28. 

Amphogona apsteint, Browne, 1904, p. 740, pl. 54, fig. 5, pl. 56, fig. 1, pl. 57, figs. 1O—15. 
Amphogona apsteini, Bigelow, 1909, p. 126, pl. 2, figs. 1—2, pl. 34, figs. 12—15, pl. 45, fig. 10. 
Amphogona apsteint, Mayer, 1910, p. 405, text-fig. 257. 

Localities. North of Chagos, Surface. 18 May, 1905, H. 1 specimen. Cargados 
Carajos, Surface. 30 Aug. 1905, 1. 3 specimens. Farquhar Group, Surface. 2 Oct. 
1905, x. 5 specimens. 

Hermaphroditism occasionally occurs in this species and as such a feature is very rare 
amongst the Hydromedusz it breaks the monotony of the usual descriptions. LI first 
noticed (1904) in specimens collected by Professor Gardiner from the Maldives Islands ; 
the male and female gonads alternating with one another in the radial canals. 

Bigelow on the cruise of the “ Albatross” in the Eastern Pacific found six specimens 
in Acapulco Harbour. These specimens turned out to be unisexual, all the gonads of any 
given individual being either male or female. 

Hartlaub (1909, p. 462, Taf. 21, fig. 27) has described a new species under the name 
of Amphigona pusilla from Dyibuti in the Gulf of Aden. One specimen showed herma- 
phroditism, and the other was unisexual and female. 

In the “Sealark” collection one specimen shows three gonads, similar in shape and 
size on adjacent canals, and they have every appearance of being males, so it may be 
presumed that this specimen is unisexual. Another specimen shows large and small 
gonads alternating with one another, as figured by Vanhdéffen, but they are too immature 
for the determination of the sex. The other specimens have either lost or nearly lost 
all their gonads and are useless for this purpose. 

The specimens are about 2°5 mm. to 4°5 mm. in diameter, and have their stomach 
upon a peduncle, which varies in length according to the size of the specimen and the 
amount of contraction. The smallest specimen has about six tentacles, and the largest 
eight tentacles in each octant. The sense organs have all disappeared. 

Distribution. Previously recorded from Indian Ocean, Maldive Is.; West coast of 
Sumatra. Pacific Ocean; Acapulco Harbour (Mexico). 


Genus Lrriopr, Lesson, 1843. 


This is a difficult genus for finding reliable specific characters. The character which 
has usually been selected is the shape of the gonads. If the gonads be taken for the sole 
character, then it is possible to connect together specimens with linear gonads, through 


198 PERCY SLADEN TRUST EXPEDITION 


a series of oval and ovoid forms, on to specimens with heart-shaped and triangular gonads, 
providing that the specimens are sufficiently numerous and taken over a large area. This 
method of determination leads to a considerable reduction in the number of described 
species, and practically it means that the genus should have only a single species. 

Another character which has been used as an aid to the determination of species is 
the number of centripetal canals. These increase in number by age and by the growth 
of the umbrella, so that taken alone they are not very reliable. Other characters have 
also been used, such as the length of the peduncle and the shape of the umbrella. By 
taking each character separately one can connect together specimens which have every 
appearance of being quite distinct species. 

Some of the species have no doubt been described from single specimens, every 
character has been used for the purpose, especially the shape of the gonads and no 
allowance made for variation or growth. Apparently a description and figure based upon 
a single or just a few individuals are not sufficient in the case of Liriope. It has, however, 
occurred to me that by taking a sufficient number of adults at the same place and time, 
and using all the possible characters, one would be able to obtain an impression of the 
type of that particular lot of specimens, and produce a sketch not of an individual but 
of the characters of the type. By adopting this method I think there is a chance of 
isolating species or at all events local races. : 

The number of specimens in the “Sealark” collection was not sufficient for finding 
out the exact number of local races or species living within the area covered by the voyage. 
After rejecting early stages and bad specimens the number left for the purpose was small. 
I am able, however, to isolate two races or species, and got on to the track of a third one. 


35. LIRIOPE TETRAPHYLLA (Chamisso et Hysenhardt) 1821. 


The specimens with triangular gonads I have placed under the old name of Lariope 
tetraphylla, and they are very similar to Vanhoffen’s figure of Lnriope tetraphylla (1902, 
Taf, 10, fig. 14). 

The umbrella is thin, 5—8 mm. in diameter. The length of the peduncle is about 
twice the diameter of the umbrella. Velum very broad. In each quadrant there are three 
centripetal canals, the interradial the longest extending up to or nearly to the top of the 
gonads, the adradial canals about half the length of the interradial. The gonads are of 
the triangular type, about equilateral, either with angular or rounded corners and do not 
extend down to the margin of the umbrella. 


36. LIRIOPE, sp. ? 


The second species or race I leave without a name, as I have not been able to find 
a published figure showing the characters of the race, and the condition of the specimens 
is not suitable for drawing. 

The umbrella is thin, about 5 mm. in diameter. The peduncle is a little longer than 
the diameter of the umbrella. Only one centripetal canal (interradial) in each quadrant. 
The gonads are longer than they are broad, either oval or elongated shield-shaped. 

The great differences between the two species are in the shape of the gonads and 


BROWNE—MEDUS FROM THE INDIAN OCEAN 199 


in the number of centripetal canals. The specimens with triangular gonads have always 
three centripetal canals in each quadrant, even in the intermediate stages when the gonads 
are beginning to develop. The specimens with elongated oval gonads have only one 
centripetal canal in each quadrant and the gonads begin as linear enlargements of the 
radial canals. 

Liriope was found at 27 stations. A good haul was made north of Chagos (17 May, 
1905, C.) when 78 specimens were taken. These were mostly early stages with a few 
adults of Ziriope sp. Another large haul of the same species was made at the Farquhar 
Isles (1 Oct. 1905, v.). Except at these two stations Liriope was rather scarce. 


37. GERYONIA PROBOSCIDALIS (Forskal) 1776. 


Geryonia proboscidalis, Vanhoften, 1902, p. 84, Taf. 10, fig. 15. 

Geryonia proboscidalis, Bigelow, 1909, p. 116. 

Geryonia proboscidalis, Mayer, 1910, p. 425, pl. 53, figs. 1—3, pl. 54, fig. 10, text-fig. 282. 
Geryonia hexaphylla, Maas, 1897, p. 26, Taf. 3, fig. 6. 

Carmarina hastata, Haeckel, 1865, p. 74, Tafs. 1, 4, 5. 

Carmarina hastata, Lo Bianco, 1904, p. 56, Taf. 35, fig. 140. 

Geryones mexicana, Agassiz and Mayer, 1902, p. 149, pl. 4, fig. 17. 

Carmaris rosea, Agassiz and Mayer, 1902, p. 149, pl. 4, fig. 18. 


For further synonyms and references see Mayer, 1910, p. 425. 


Localities. North of Chagos, 1200—0 fms. 17 May, 1905, C. 1specimen. Chagos 
Archipelago, Salomon Atoll, 10 fms. 1 July, 1905, P. 1 specimen. Mauritius, 300—0 fms. 
22 Aug. 1905, c. 1 specimen. 

The specimens are early stages and not in good condition. The smallest, about 3 mm. 
in diameter, has one centripetal canal in each interradius, and the perradial hollow tentacles 
are just beginning to develop. The largest specimen, about 8 mm. in diameter, has three 
centripetal canals in each interradial sextant. The gonads are beginning to develop. Two 
of the sense organs have two otoliths instead of the usual single one. 

Distribution. At the surface in the tropical and warm regions of all the Oceans, and 
in the Mediterranean. 


NARCOMEDUS. 


38. SOLMARIS sp. ? 


Localities. North of Chagos, Surface (Temp. 82° F.). 18 May, 1905, F. 1 specimen. 
Chagos Archipelago, Salomon Atoll, Surface. 4 July, 1905, Q. 3 specimens. Farquhar 
Is., Surface. 29 Sept. 1905, u. .1 specimen. 

Out of the five specimens taken only one is in moderately good condition, and it is 
the largest. 

Deseription:—The umbrella is about 5mm. in diameter, slightly curved, and 
moderately thin. The marginal lappets are about as broad as long, and curved on the 
outer edge. Velum narrow. The stomach is circular, its lower ,wall rather loose, and 
the mouth closed in the form of projecting lips. There are broad marginal canals, but not 


200 PERCY SLADEN TRUST EXPEDITION 


in the form of a solid chord of cells. The gonads are developing on the outer third of the 
lower wall of the stomach in a continuous ring, which is without any projecting pouches. 
There are eight tentacles, about as long as the diameter of the umbrella; they are not 
very stiff, and taper to a fine point. Below each tentacle is a well-marked peronial 
groove, lined with nematocysts. On each of the eight marginal lappets are five sense 
organs, rather ovate in shape, showing a clear circular vesicle, and without any external 
hairs. Otoporpe are present as narrow bands just curling over the margin of the 
umbrella and situated on ridges. 

The other specimens are smaller in size and very much contracted. Two of them 
have nine tentacles. 

It is difficult to assign a specific name to these specimens, as some are certainly quite 
young stages and the largest has not reached maturity. They come nearest to Haeckel’s 
Solmaris lenticula from the Indian Ocean. . This species was only briefly described and 
never figured by Haeckel. Mayer thinks that it was probably only an immature form. 


39. ANGINA cCITREA, Eschscholtz, 1829. 


digina citrea, Maas, 1905, p. 71, Taf. 11, fig. 72, Taf. 13, figs. 79—82. 
gina citrea, Bigelow, 1909, p. 73, pl. 1, fig. 5, pl. 14, fig. 5. 

Locality. Chagos Archipelago, Peros Atoll. 600—O fms. 30 June, 1905, N. 
1 specimen. 

Tam not certain about the correct determination of this specimen, which is far from 
being perfect. The umbrella has a rather rounded summit, and measures 14 mm. in width 
and 7 mm. in height. There are eight stomach-pouches, one of which shows signs of 
a small notch in the middle of the lower edge; the other pouches are either without 
a notch or in an imperfect condition. Four tentacles about 30 mm. long. The margin of 
the umbrella is imperfect, but four sense organs were seen in one octant. 


Genus SOLMUNDELLA, Haeckel, 1879, ex Maas, 1904. 


This genus is found in all the oceans from the tropics to the icy waters of the poles. 
It is quite easy to determine the genus even from very bad specimens, but one has yet to 
settle the exact number of species, and this delicate point is not so easily disposed of. 

Vanhoffen considers that the genus has only a single species, which he calls 
Solmundella bitentaculata, after the oldest specific name. The doubtful species is 
known as Solmundella mediterranea. Whether this is a good species, or a variety of 
Solmundella bitentaculata or identical with it, is a point which, in my opinion, requires 
further researches. 

Maas and Bigelow evidently recognise two species; Mayer is very doubtful, and only 
ranks Solmundella mediterranea as a variety. As in the case of Liriope, I think that 
there are at least local races of Solmundella. The difficulty is to make sure of their 
status. Are they distinct species or varieties? If it should be finally decided that there 
is only one species, then I think that local races should be recognised by names for the 


sake of geographical distribution. 


BROWNE—MEDUS FROM THE INDIAN OCEAN 201 


The Antarctic race’has a character which has not been described in specimens from 
other places. Both Vanhoffen and myself note the occurrence of clusters of nematocysts 
upon the ex-umbrella, especially near the margin. If these clusters are not found in 
specimens from other localities they could be used to isolate the Antarctic race. The 
English and German expeditions to the Antarctic found Solmundella very plentifully at 
their respective winter quarters, and it apparently stays there and breeds there throughout 
the year. Though we both found the same race, still we called it by different names. 
Vanhoffen regarded it as Solmundella bitentaculata on account of his recognising only 
a single species, whereas I tried to separate it from that species and called it Solmundella 
mediterranea on account of its possessing only eight sense organs. The number of sense 
organs has generally been used to separate the two species. Solmundella mediterranea 
usually with eight sense organs and not more than sixteen, whereas Solmundella biten- 
taculata may have double that number. The sense organs alone are hardly sufficient to 
separate the two species because Solmundella bitentaculata passes through stages with 
eight and sixteen sense organs. The shape of the umbrella may be useful when one is 
examining living specimens or specimens preserved in perfect condition; the latter, 
however, are not often found in collections from. abroad. 


40. SOLMUNDELLA MEDITERRANEA (Miiller) 1851. 


Solmundella mediterranea, Maas, 1906, p. 12, Taf. 1, fig. 5, Taf. 3, figs. 2324. 
Solmundella mediterranea, Browne, 1910, p. 38. 


Solmundella bitentaculata var. mediterranea, Mayer, 1910, p. 456, pl. 54, figs. 1—3, pl. 55, fig. 4. 


Localities. North of Chagos, Surface (Temp. 84° F.). 16 May, 1905, A. 2 speci- 
mens. 50—0fms. 18 May, 1905,F. 1 specimen. Chagos Archipelago, Salomon Atoll, 
Surface. 5 July, 1905, Q. 4 specimens. Farquhar Group, Surface. 3 Sept. 1905, x. 
3 specimens. Amirante Isles, Desroches Atoll. 50—0 fms. 16 Oct. 1905, kk. 2 speci- 
mens. Amirante Isles, Surface. 18 Oct. 1905, 00. 1 specimen. 

I have decided to place the “Sealark” specimens under the name of Solmundella 
_ mediterranea as the best and largest specimen comes nearest to it. The umbrella is 
highly arched, about 3 mm. in diameter, with isolated nematocysts scattered over the 
ex-umbrella. The gonads are confined to the pouches and show ripe ova. The tentacles 

are 13 mm. in length. There are nine sense organs, with otoliths, and four interradial 
_ marginal bulbs. The other specimens are of little value for specific determination, being 
either early stages or damaged. One specimen has Cunina-buds inside the stomach. 


41. CUNINA sp. ? 


Localities. North of Chagos. Chagos Archipelago. Farquhar Alphonse Is. 
Amirante Is. 

The collection contains about 50 specimens taken at nine different stations. 

Nearly all the specimens of Cunina belonged to early stages, about 3 mm. in 
diameter. Most of them have 8 tentacles, and a few have 7, 9 or 10 tentacles. They are 
in rather bad condition and probably represent more than one species. . 

At a station, North of Chagos, a series larger in size was taken, and had the 

SECOND SERIES—ZOOLOGY, VOL. XVII. 26 


202 PERCY SLADEN TRUST EXPEDITION 


specimens been in better condition I might have succeeded in determining the species. 
In general appearance they resemble Gegenbaur’s figure of Cunina lativentris (Gegenbaur, 
1856, Taf. 10, fig. 2). The largest specimen measures 8 mm. in diameter, and its 
umbrella is moderately thick. It has twelve tentacles, about 3 mm. long, tapering to 
a fine point; at their base is a semicircular band of nematocysts. The gastric pouches 
are about as broad as long, with gonads in the course of development, and in shape 
somewhat similar to Gegenbaur’s figure. The otoporpe take the form of short narrow 
lines, situated on ridges, four to five on each lappet. 


SCYPHOMEDUS. 


CHARYBD AIDA 
42. CHARYBDEA sp. 4 


Locality. Farquhar Group, Surface. 2 Oct. 1905, x. 1 specimen. 

This single specimen is immature, and at a stage which makes identification so Nem 
uncertain that it is perhaps best not to give it a specific name. 

The umbrella is 15 mm. wide and 23 mm. high, and tapers very slightly towards the 
summit, which is slightly rounded. The ex-umbrella is free from warts and clusters of 
nematocysts. The stomach is very short and rather flat ; the mouth has four lips. The 
phacellze or gastric filaments are neither arranged in groups nor branched, but each 
interradial set is composed of simple filaments. The velarium has six unbranched canals 
in each quadrant. There are four tentacles, with pedalia about 7 mm. in length, and 
having a flat spatula-like expansion on the inner side. The gonads are immature, about 
11 mm. in length, and do not extend so far down as the plane of the sense organs. The 
four perradial sense organs are situated about 4mm. above the margin of the umbrella, 
and the sensory clubs are suspended by a stalk in a deep pit, which projects out on the 
wall of the sub-umbrella. The sensory clubs have two ocelli, one of which is very large 
and nearly terminal, the other is above it, close to the stalk, and is very much smaller. 
These are the median ocelli, but no lateral ocelli could be detected. 


CORONAT A. 
43. NausiTtH6e puncTATA, Kolliker, 1853. 


Nausithie punctata, Bigelow, 1909, p. 35, pl. 12, fig. 5. 
Nausithoe punctata, Mayer, 1910, p. 554, pl. 60, figs. 4—5 
Nausithée punctata, Bigelow, 1913, p. 85. 

Localities. North of Chagos. Chagos Archipelago. Amirante Isles. 

The collection contains 16 specimens taken at eight different stations. 

The specimens are rather small, not exceeding 5 mm. in diameter. Nearly all the 
specimens have globular gonads, about equidistant apart. Two specimens, however, have 
oval gonads, about twice as long as broad. 

This medusa inhabits the tropical and warm regions of all the oceans, and belongs to 
the surface fauna. 


BROWNE—MEDUSAi FROM THE INDIAN OCEAN 203 


Genus AtottaA, Haeckel, 1880. 


After reading the work of Bigelow (1909 and 1913), Mayer (1910) and Broch (1913) 
on Atolla, it seems to me that the genus has only two species, namely, Atolla wyviller 
and Atolla chuni. The latter is easily distinguished by the presence of warts on the 
marginal lappets. All the other species which have been described by various authors 
apparently fall under the oldest name of Atolla wyviller. The presence and shape, or the 
absence, of radial furrows on the central disc, formerly relied upon for distinguishing 
species, are apparently worthless as specific characters. 

Broch, after examining over 200 specimens of Atolla collected by the “‘ Michael Sars” 
in the North Atlantic, was able to separate the material into three groups. 

“ First, those furnished with distinct radial furrows all over the central disc; second, 
those with incomplete radial furrows, in many cases visible only at the margin of the 
central disc; and third, those with a perfectly smooth central disc, showing no trace 
whatever of radial furrows.”...“‘The intermediate group contains every transition stage 
from Atolla bairdw (with a smooth central disc) to Atolla verrillia (with narrow radial 
furrows on the central disc), and we are therefore compelled to consider Atolla verrilli as 
a synonym of Atolla bairdw.” 

Out of this large number of Atolla Broch was able to isolate a single specimen, 
which he has placed under the name of Atolla wyvillec. “The broad and conspicuous 
radial furrows of the central dise and the strongly-marked longitudinal furrows of the 
pedalion distinguish this species from the other Atlantic species.” Broch apparently 
found no connecting link between Atolla wyvillei and Atolla verrillw. 

Bigelow, according to his reports on collections from the Eastern and North-Western 
Pacific regions, has experienced the difficulty of separating Atolla wyvillec (with broad 
radial furrows) from Atolla verrillw (with narrow radial furrows). He says: “In the 
Eastern Pacific specimens there was considerable variation in the breadth of the furrows, 
which were usually broad in large, narrow in small specimens.” In the North-Western 
Pacific specimens “the furrows vary so much that no sharp line can be drawn between 
specimens in which they are broad and those in which they are narrow.” 

Bigelow records no specimens in these collections with a perfectly smooth disc 
(Atolla bairdii). It is clear from Broch’s researches that the absence of or presence of 
radial furrows are not good characters for the determination of species, and from Bigelow’s 
researches that the width of the radial furrows is also useless. 

Broch also gives tables which show that there is no good evidence for assuming that 
the smooth-dise form of Atolla and the furrow-disc form are separate geographical races, 
or that they live at different depths, or that differences are due to growth. The two 
kinds are found together at the same stations, depth, and of the same size. 


44, ATOLLA WYVILLEI, Haeckel, 1880. 
Localities. N.E. of Chagos, Lat. 4° 30’S., Long. 71°15’ E. (Depth about 2000 fms.) 
2 specimens caught in Fowler’s self-closing net. 1000—500 fms. 18 May, 1905, J. 
Between Providence and Alphonse, Lat. 8° 16’ S., Long. 51° 26’ E. 1 specimen caught 


in a large square net. 900—0 fms. 6 Oct. 1905, aa. 
26—2 


204 PERCY SLADEN TRUST EXPEDITION 


The specimens are in bad condition. The smallest measured 7 mm. across the circular 
muscle band, and it is an early stage, with 20 tentacles. The largest is 30 mm. in 
diameter, across the circular muscle band and has 24 tentacles. The radial grooves on the 


margin of the central dise are well marked and belong rather to the type associated with 
Atolla verrillu than to Atolla wyvillei. 


SEMAOSTOME. 


Genus PEnacta, Péron et Lesueur, 1809. 


Pelagia is one of the genera which has its species in a state of confusion. There has 
been a general reduction in the number of species, and some of those which have been left 
are still uncertain. The characters selected for the determination of the species have not 
produced satisfactory results. 

The classification based upon the warts on the ex-umbrella failed owing to their being 
treated too minutely, but I believe that they will yet prove useful for distinguishing races, 
if not species. Warts vary considerably in shape and size in each individual, but there is 
a predominating type which becomes characteristic of the race or species, and belongs to 
that particular race only. The predominance of high conical warts is characteristic of 
Pelagia flaveola. This type of wart does not occur in Pelagia panopyra, which has low 
oval-shaped warts ; but minute roundish warts are common to both. 

The marginal lappets in all the specimens which I have seen are too much alike in 
shape and size to be of any definite use for distinguishing species. The position and density 
of the warts on the lappets may be useful for distinguishing races. 

Attempts have been made to distinguish species by the length and size of the 
manubrium and oral arms, but one never knows when examining preserved specimens 
how much the oral arms have contracted. 

In the “Sealark” collection there are three distinct kinds of Pelagia. Pelagia 


Jlaveola 1 am in favour of recognising as a distinct species on account of its peculiar 
warts. oh ee 


45. PELAGIA PANOPYRA (Péron et Lesueur) 1807. 


Pelagia panopyra, Bigelow, 1909, p. 43. 
Pelagia panopyra, Mayer, 1910, p. 575. 
Pelagia panopyra, Kishinouye, 1910, p. 9. 
Pelagia panopyra, Bigelow, 1913, p. 88. 


Locality. South West of Chagos, Surface. 31 July, 1905. 5 specimens. 

The specimens from this station have warts on the ex-umbrella marked by ridges 
and furrows. The warts form low mounds varying in shape from round to oval. The 
oval-shaped warts predominate and are characteristic. The ridges and furrows are formed 
in the jelly, and are clearly revealed where the ectodermal covering has been rubbed off. 
An oval-shaped wart has a longitudinal ridge with lateral ridges and furrows running 


BROWNE—MEDUSA FROM THE INDIAN OCEAN 205 


down at right-angles to the main central ridge. A roundish wart has ridges and furrows 
radiating out from the centre. 

The specimens are from 20 to 25 mm. in diameter; the largest measured 25 mm. in 
width and 15 mm. in height. The umbrella is hemispherical in shape with a slightly 
flattened top, and moderately thick. 

The marginal lappets are about as long as broad, with rounded corners. They are 
thin and partly covered with small roundish or oval patches of nematocysts. The oral 
arms are rather contracted, and have a firm solid appearance, about 15 mm. in length. 
The manubrium, oral arms, and frills, are covered with warts, usually oval in shape, but 
more variable in shape and size on the frills. On the margin of the umbrella, opposite 
every sense organ, there is a small shallow conical pit. The largest specimen has ova in 
the genital sacs. 


46. Prnacia sp.?. A. 


Locality. South of Saya de Malha, Surface. 4 Sept. 1905, m. 1 specimen. 

One specimen found amongst those of Pelagia flaveola by the shape of its warts 
clearly belongs to another species. It has a number of long and oval-shaped warts upon 
the ex-umbrella, which is in an excellent state of preservation and completely covered with 
ectoderm. The warts are very variable in shape and size, some are roundish. On the top 
of the warts there is a patch of nematocysts corresponding to the general shape of the 
raised warts. There are no clear indications of ridges and furrows in these warts. I am 
rather inclined to think that the appearance of ridges and furrows on the surface of the 
warts, which have been denuded of ectoderm, has some connection with a shrinkage of 
the jelly, and that they probably would not be noticed if the warts were completely covered 
with ectoderm. | 

The specimen measures about 12 mm. in diameter. The marginal lappets are loosely 
covered with small roundish or oval warts, which are confined to the areas occupied by the 
canals on the lappets. A shallow, roundish pit exists on the margin of the ex-umbrella, 
opposite each sense organ. This single specimen comes nearest to Pelagia panopyra on 
account of the presence of the oval-shaped warts. 


47, PELAGIA FLAVEOLA, Eschscholtz, 1829. 


Pelagia ftaveola, Eschscholtz, 1829, p. 76, Taf. 6, fig. 3. 

Pelagia papillata, Haeckel, 1880, p. 509. 

Pelagia tahitiana, Agassiz and Mayer, 1902, p. 158, pl. 8, figs. 34—35. 
Pelagia flaveola, Mayer, 1910, p. 575, text-fig. 364. 


Locality. South of Saya de Malha, Surface. 4 Sept. 1905, m. 24 specimens. 

The specimens of Pelagia taken at this station can easily be distinguished from those 
taken at the other stations by the shape of the warts upon the ex-umbrella. The warts 
are formed of conical lumps of jelly capped with a small cluster of nematocysts, and they 
stand up conspicuously as figured by Agassiz and Mayer (1902, PI. 8, fig. 34) for Pelagia 
talitiana. The largest warts are upon the upper half of the umbrella and they decrease 


206 PERCY SLADEN TRUST EXPEDITION 


in size towards the margin. There is a considerable variation, both in shape and size of 
the warts, due either to pressure or to a shrinkage of the jelly. The latter produces strange 
effects, such as a wart having the appearance of being surmounted by a small capitate 
tentacle, or the ex-umbrella being covered with tentacular-like papillee clearly visible to 
the naked eye. It is quite easy to distinguish this species by the shape of the warts 
provided that the specimens are in good condition, but there is a good chance of a failure 
if the warts are squeezed out of shape by pressure. 

The description of Pelagia tahitiana by Agassiz and Mayer agrees so well with my 
specimens that I cannot find anything of importance to add to it, and as Mayer considers 
that Pelagia tahitiana is identical with Pelagia flaveola of Eschscholtz I have followed 
his synonymy. 

The size of the specimens in the “ Sealark” collection is from 10 to 25 mm. in diameter 
with the umbrella flattened out. A few large specimens have immature gonads hanging 
down from the sub-umbrella. Al the specimens are colourless in formaline, but they should 
have been yellowish when alive. The oral arms of all the specimens are in bad condition. 
They are apparently only very thinly covered with small patches of nematocysts upon slight 
elevations of jelly. There is a small pit on the margin of the umbrella opposite each sense 
organ. 

Some of the specimens have small barnacles attached to the ex-umbrella. 

Distribution. Tropical Pacific and Indian Oceans. 


48. PELAGIA sp.?. B. 


Locality. North of Chagos, Lat. 4° 16’S., Long. 71° 53’E. 75—0 fms. 17 May, 
1905, B. 1 specimen. Surface. 18 May, 1905, B. 6 specimens. 75—0Ofms. 18 May, 
1905, B. 2 specimens. 

The warts on the ex-umbrella of these specimens are low and roundish, with a broad 
patch of nematocysts in the centre. These warts show no traces of ridges and furrows, 
and none are oval-shaped as seen in Pelagia panopyra. In one specimen in a depression 
at the top of the umbrella some of the warts are rather taller than the others, suggesting 
a resemblance to the warts of Pelagia flaveola. 

The specimens measured from 7 to 13 mm. in diameter. The umbrella has a flattish 
top, nearly twice as broad as high and moderately thick. The marginal lappets are about 
as long as broad, with rounded corners, and well covered with rounded patches of nema- 
tocysts. The manubrium and oral arms are rather longer than in Pelagia panopyra and 
thickly covered with roundish patches of warts. There is a shallow pit on the ex-umbrella, 
opposite every sense organ. The gonads are only just beginning to develop in the larger 
specimens. } 


BROWNE—MEDUS4 FROM THE INDIAN OCEAN 207 


RHIZOSTOM 4. 


49. CASSIOPEA ANDROMEDA var. MALDIVENSIS, Browne, 1905. 


Cassiopea andromeda var. maldivensis, Browne, 1905, p. 962. 
Cassiopea andromeda var. maldivensis, Mayer, 1910, p. 963. 


Locality. Seychelles Group. Praslin Reef. 2 specimens. 

These specimens agree fairly well with the description of the variety which I described 
from the Maldives. 

The umbrella of the largest specimen measured 110 mm. when flattened out, and the 
smaller one measured 80 mm. in diameter. The ex-umbrella is quite smooth, without any 
trace of a circular band, and without any colouration markings. The margin of the umbrella 
is too imperfect for counting the number of lobes. The largest specimen has at least 
18 sense organs, which are very irregular in position. The oral arms are provided with 
appendages very similar to those described in Maldive specimens. The small appendages 
adjacent to the oscula are either leaf-shaped or cylindrical. In addition to these appendages 
there are a few others very much larger. One in the very centre of the oral dise measured 
22 mm. in length and 7 mm. in width; others near it are shorter and more eylindrical. 
Both specimens are thickly covered with Green Cells or Zooxanthelle. 


208 PERCY SLADEN TRUST EXPEDITION 


LIST OF BOOKS AND MEMOIRS QUOTED IN THE TEXT. 


Agassiz, A., and A. G. Mayer, 1899. “Acalephs from the Fiji Islands.” Bull. Mus. Comp. Zool., 
Harvard College, vol. 32, pp. 157—189, 17. pls. 

+— 1902. “Reports on the Scientific Results of the Expedition to the Tropical Pacific...in the 
‘Albatross’.” Part 11. Meduse, Memoirs Mus. Comp. Zool., Harvard College, vol. 26, pp. 139— 
176, pls. 1—13. 


BicEtow, H. B., 1904. “ Medusze from the Maldive Islands.” Bull. Mus. Comp. Zool., Harvard College, 
vol. 39, pp. 245—269, 9 pls. 


—— 1909. “Reports on the Scientific Results of the Expedition to the Eastern Tropical Pacific... 
in the ‘ Albatross’.” Part xvi. The Medusze, Memoirs Mus. Comp. Zool., Harvard College, vol. 37, 
243 pp., 48 pls. 


—— 1913. “Medusz and Siphonophore collected by the ‘Albatross’ in the North-western Pacific, 
1906.” Proc. U.S. Nat. Mus., vol. 44, pp. 1—119, pls. 1—6. Smithsonian Instit., Washington. 


Brocu, H., 1913. “Scyphomeduse” in Report, Scientific Results of the “Michael Sars,’ North 
Atlantic Deep Sea Expedition, 1910. Vol. 3, part 1. (Zool.), 23 pp., 1 pl. Bergen. 


Brooks, W. K., 1886. “The Life-History of the Hydromeduse.” Mem. Boston Nat. Hist. Soe., vol. 3, 
pp. 359—480, pls. 37—44. 


Brooks, W. K. and S. RrrrenHouse, 1907. “On Turritopsis nutricula (McCrady).” Proc. Boston Nat. 
Hist. Soc., vol. 33, pp. 429—460, pls. 30—35. 


Browne, E. T., 1904. “Hydromeduse with a Revision of the Willadz and Petaside.” Fauna and 
Geography of the Maldive and Laccadive Archipelagoes, vol. 2, pp. 722—749, pls. 54—57. Cambridge. 


—— 1905. “Scyphomeduse.” Fauna and Geography of the Maldive and Laccadive Archipelagoes, 
vol. 2, suppl. 1., pp. 958—971, 1 pl. Cambridge. 


—— 1905. “Meduse,” in Herdman’s Pearl-Oyster Fisheries, Ceylon. Part Iv., pp. 131—166, 4 pls. 
Royal Soc. London. 


— 1906. Biscayan Plankton collected during a Cruise of H.MLS. “Research,” 1900. Part x. “The 
Medusae.” Trans. Linn. Soc. London, vol. 10, pp. 163—185, pl. 18. 


— 1910. National Antarctic Expedition, “Discovery.” “Medusex.” Vol. v., Part v., 62 pp., 7 pls. 
British Museum, London. 


EscuscHo.tz, F., 1829. System der Acalephen. Berlin. 


Frewxes, J. W., 1881. “Studies of the Jelly-fishes of Narraganseett Bay.” Bull. Mus. Comp. Zool., 
Harvard College, vol. 8, pp. 141—176, pls. 1—10. 


GEGENBAUR, C., 1856. “Versuch eines Systemes der Medusen.” Zeitschr. wiss. Zool., Bd. 8, pp. 202— 
273, Taf. 7—10. 


Haercke, E., 1865. “Die Familie der Russellquallen (Geryonida).” 194 pp., 6 Taf. Leipzig. (Abdruck. 
Jenaische Zeitschr. natw. Bd. 1—2.) 


—— 1879—1880. Das System der Medusen. Jena. 


Harriavus, C., 1907. “Craspedote Medusen. Codoniden und Cladonemiden.” Nordisches Plankton 
X11, 135 pp., 126 figs. 


—— 1909. “Uber einige von Ch. Gravier in Djibuti gesammelte Medusen.” Zool. Jahrb. Jena. 
Bd. 27, pp. 447—476, Taf, 19—23. 


BROWNE—MEDUSA FROM THE INDIAN OCEAN 209 


HarrLavs, C., 1914. “Craspedote Medusen. Tiaride.” Nordisches Plankton x11, pp. 237—363. 
Huxtey, T. H., 1877. A Manual of the Anatomy of Invertebrated Animals. London. 


KisHinovuve, K., 1910. “Some Medusx of Japanese Waters.” Journ. Coll. Sci. Tokyo, vol. 27, Art. 9, 
35 pp., 5 pls. 


Lo Branco, S., 1904. “Pelagische Tiefseefischerei der ‘Maja’ in der Umgebung von Capri.” Bd. 1, 
91 pp., 42 Taf. Jena. 


Maas, O., 1893. “Die Craspedoten Medusen.” Ergebnisse der Plankton-Expedition “National.” Bd. 11, 
K.C., 107 pp., 8 Taf. 


— 1897. “Reports on an Exploration off the West coast of Mexico, Central and South America, and 
off the Galapagos Islands.” “Albatross,” 1891, xx1. “Die Medusen,’ Memoirs Mus. Comp. Zool., 
Harvard College, vol. 23, pp. 1—92, 15 Taf. 


— 1905. “Die Craspedoten Medusen der Siboga-Expedition.” Monograph x, 84 pp. 14 Taf. 
Leyden. 


— 1906. “Méduses d’Amboine.” Rev. Suisse Zool. Geneve, Tom. 14, pp. 81107, pls. 2—3. 


—— 1909. “ Beitriige zur Naturgeschichte Ostasiens. Herausgegeben von Dr F. Déflein. No. 8. 
Japanische Medusen.” Abh. Akad. Wiss. Miinchen. Suppl. Bd. 1, pp. 1—52, 3 Taf. 


McCrapy, J., 1857. “ Description of Oceania (Turritopsis) nutricula nov. spec., and the embryological 
history of a singular Medusan Larva, found in the Cavity of its Bell.” Proc. Elliott Soc., vol. 1, 
pp. 55—90, pls. 4—7. 


—— 1858. “Gymnopthalmata of Charleston Harbour.” Proc. Elliott Soc., vol. 1, pp. 103—221, 
pls. 8—12. 


Mayer, A. G., 1910. Medusz of the World. 3 vols, 4to. . Washington. 


METSCHNIKOFF, E., 1886. Medusologische Mitteilungen. Wien. Arb. Zool. Instit.' Univ. Bd. 6, 
pp. 287—266, Taf. 1—2. 


Murer, H., 1908. “Untersuchungen tiber Eibildung bei Cladonemiden und Codoniden.” Zs. wiss. 
Zool., Leipzig. Bd. 89, pp. 28—80, Taf. 3—5. 


VANHOFFEN, E., 1902. “Die craspedoten Medusen der deutschen Tiefsee-Expedition, ‘ Valdivia, 
1898—1899.” Wissen. Ergebnisse, Bd. 3, pp. 53—86, Taf. 9—12. 


— 1911. “Die Anthomedusen und Leptomedusen der Deutschen Tiefsee-Expedition, ‘ Valdivia, 
1898—1899.” Bd. 19, pp. 193—288, Taf. 22. 


—— 1912. “Die craspedoten Medusen des ‘ Vettor Pisani’.” Zoologica, Heft 67, 33 pp., 2 Taf. 


—— 1912. “Die craspedoten Medusen der deutschen ‘Sudpolar’ Expedition, 1901—1903.” Bd. 13, 
Zool. v, pp. 353—395, Taf. 24—25. 


SECOND SERIES—ZOOLOGY, VOL, XVII. 27 


210 PERCY SLADEN TRUST EXPEDITION 


DESCRIPTION OF PLATE 39. 


Fig. 1. Steenstrupia normani, n. sp. x 25. 
Fig. 2. Zanclea orientalis, n. sp. x 25. 


Fig. 3. Zanclea orientalis. A perradial rudimentary bulb (B) with a patch of nematocysts (N) on the 
margin of the ex-umbrella (Ex.). Circular Canal (CC). Velum (Y). 


‘Fig. 4. Leuckartiara gardineri, n. sp. x 12. 


Cambridge University Press. 


TRANS. LINN. SOC. SER. 2. ZOOL. VOL. XVII .PL.389. 


MEDUSAE FROM THE INDIAN OCEAN. 


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: ; V 


No. V.—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 
COLLECTED BY H.M.S. “SHEALARK” IN THE INDIAN OCEAN. 


By Artaur Denpy, D.Sc., F.R.S., F.L.S8., Professor of Zoology in the 
Unwersity of London (King’s College). 


(Plates 40—43.) 
Read 17th June, 1915. 


The “Sealark” collection contains only three species of Hexactinellid Sponges, but 
these are all extremely interesting forms, and I venture to hope that the somewhat 
detailed study that I have been able to make of them may add materially to our 
knowledge of this remarkable group. 

Aulocalyx serialis is a new species of a genus hitherto known only by fragments of the 


expedition and described by 


type species (A. irregularis) obtained by the “ Challenger’ 
Schulze. 

Heterorete pulchra is the type species of a new genus related to Dactylocalyx, and 
remarkable for the entire absence of special dermal and subdermal spicules. Fortunately 
the material of this species was much better preserved than is usually the case with 
hexactinellids, so that I am able to give some particulars as to the soft tissues and canal 
system. This sponge is further interesting on account of the presence in it of a commensal 
or parasitic hydroid ramifying through the substance of the wall, as well as of numerous 
Anthozoa attached to the surface. 

Sarostegia oculata is a very beautiful and remarkable sponge, first described by 
Topsent from deep water off the Cape Verde Islands. 

The preparation of this Report has been greatly facilitated by a grant from the 
Trustees of the Perey Sladen Memorial Fund to enable me to pay an assistant. I desire 
to express my thanks to the Trustees for their generous action, and I wish also to express 
my indebtedness to Miss Hilda Lucy Deakin, the assistant appointed, by whom most of 
the microscopical preparations and drawings of skeletal structures have been made, for 
the skill and care with which she has carried out the work entrusted to her. 

The drawing’s of external forms were made for me by Mr T. P. Collings. 


Genus AuLocaLyx Schulze [1887]. 


This genus was founded by F. E. Schulze in 1887 for the reception of several 
fragmentary specimens obtained by H.M.S. “Challenger” from a depth of 310 fathoms 
off Marion Island, south-east of the Cape of Good Hope, and between Marion Is. and the 
Crozets at a depth of 1600 fathoms, and thus practically within the area of the Indian 

27—2 


212 PERCY SLADEN TRUST EXPEDITION 


Ocean. To these specimens the specific name irregularis was assigned. Schulze’s 
diagnosis of the genus [1887, p. 380] is as follows :—“ A thin-walled cup, much folded, 
extended into lateral diverticula, and also continued into short laterally projecting tubes. 
The cup is fixed by a firm irregular base. The connected framework of beams consists 
of much curved hexacts, partly united by synapticula, partly soldered together. The 
parenchyma contains loose discohexasters with short or with medium-sized principal rays, 
bearing §-shaped terminals disposed in perianth-like fashion. Under the skin there are 
large hexasters in which each of the short principal rays bears six long diverging 
terminals, which gradually increase in thickness towards the round outer end, and are 
beset all round with backward bent pointed hooks. The dermalia and gastralia are 
rough medium-sized oxypentacts.” 

Schulze placed his new genus in the family Rossellidee of the then still accepted 
sub-order Lyssacina. I am not aware that the genus has again been met with excepting 
by the “ Sealark” expedition in the Indian Ocean ; but in his report on the Hexactinellida 
of the Valdivia (p. 180) Schulze [1904] accepts a new arrangement proposed by [Tjima 
[1903], in accordance with which it is placed in the latter’s new family Dactylocalycide, 
all the other genera of which are more or less typical dictyonine forms. 


1. Aulocalyx serialis n. sp. 
(Plate 40, figs. 1—10 a.) 


The largest and most perfect specimen (R.N. v. 1) has the form of a narrow, 
obconical goblet, terminating above in a slightly contracted mouth without any sieve- 
plate (cf. fig. 1). The wall of the goblet is longitudinally folded, and the prominent 
ridges on the outer surface appear to be made up each of a single row of short tubes 
fused together. The principal rows of tubes extend longitudinally from the constricted 
base of the sponge to the margin of the mouth; as they extend upwards the intervals 
between adjacent rows become wider and new rows appear between them. The tubes 
have conspicuous external openings and lead right through the wall of the goblet into the 
central cavity. Their internal openings lie in the bottom of longitudinal depressions of 
the inner surface. Altogether there are some eight or nine rows of these tubular openings 
in the wall of the sponge. In this specimen, unfortunately, the base of attachment has 
been broken off ; but two other specimens (R.N. v. 2 and R.N. lxxxi.), which are in other 
respects much less perfect, show how the body of the sponge gradually contracts below 
and then expands into a small flattened disc of attachment (cf. fig. 1). The central 
cavity of the sponge is continued downwards practically as far as the basal disc, and 
there is no distinct stalk. The sponge is very fragile, the greater part of the body soft 
and compressible, almost woolly, but gradually becoming much more rigid below, owing 
to the stronger development of the dictyonal framework. All the soft tissues seem to 
have disappeared, and the best specimen (R.N. v. 1) contains a considerable quantity of 
fine white sand, consisting largely of foraminiferan and radiolarian skeletons. The 
height of the largest specimen, without the basal disc, is about 70 mm., and the greatest 
width, close to the top of the goblet, 29 mm. 


DENDY—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 213 


The skeletal framework (fig. 2) consists of a very lax reticulation of large, smooth 
hexacts, with very long, slightly curving rays soldered together and joined by synapticula 
in a most irregular, but often ladder-like, fashion. A characteristic feature of this frame- 
work is that it sends out free ends, which project into the gastral cavity in the form of 
large hooks with very varying degrees of curvature (fig. 3). 

The following kinds of separate spicules occur in the sponge :— 

(1) Large and small parenchymal hexacts, having straight, slender rays with slightly 
roughened ends (figs. 4, 5). 

(2) Large dermal and gastral pentacts; rays slightly roughened at the ends 
(fig. 6). 

(3) Occasional small pentacts, with knob-lke vestige of the sixth ray ; rays slightly 
roughened (fig. 7). 

(4) Discohexasters, varying somewhat as regards the curvature and arrangement of 
the terminal rays, which end in small dises with toothed margins (figs. 8, 9). 

(5) Large hexasters with terminal rays ending in small knobs, and provided with 
‘rather long, backwardly pointing spines, the most distal of which may arise in a whorl 
from the terminal knob. These spines are very easily broken off, and this has evidently 
been the case to a large extent in the specimen figured (fig. 10). It is very difficult to 
find a specimen of this spicule perfect enough to draw, though a fair number of them 
occur in my preparations. 

The only other species of Aulocalyx hitherto described is Schulze’s A. arregularis, 
the type of the genus. To judge from the description and figures given in the 
“Challenger” report the chief difference between the two lies in the external form. In 
the “ Challenger” species “the general form was that of a broadly expanded cup with 
complex, much folded or diverticulated wall.” The generic diagnosis mentions the occurrence 
of “short laterally projecting tubes.” These are not mentioned in the specific description 
but the general form is compared to that of Periphragella elise, the figure of which 
(Pl. LXXX) shows numerous short tubular projections, with open mouths, scattered 
irregularly over the outer surface of the cup. Schulze’s general figure of the external 
form of A. wregularis (Pl. LX, fig. 1) does not show any distinct lateral tubes, but the 
specimen was evidently much injured. He gives a separate figure (fig. 2), however, 
of the macerated skeleton of a lateral tube, which seems to show that these structures 
are of the same nature as the tubular openings in A. serials. There is no indication, 
however, that they are arranged in longitudinal rows as in the latter species, and the 
form of the cup in the two cases appears to be quite different, being very much narrower 
in A. serialis. 

Another difference is found in the shape and size of the large, spiny-rayed hexasters 
or rosettes. Those of A. serialis are only about 0°24 mm. in diameter, while those of 
A. wregularis measure about 0°'4mm. Moreover, in the latter the ends of the long 
terminal rays are ‘“‘ simply convex, or more rarely somewhat knobbed,” while in A. serzalis 
they are, usually at any rate, distinctly knobbed, with a terminal whorl of backwardly 
pointing spines coming off from the knob as shown in fig. 10 a. 

How far the differences between the two forms can be regarded as really specific 


214 PERCY SLADEN TRUST EXPEDITION 


cannot be decided in the absence of better preserved specimens of A. irregularis. The 
two are evidently closely related and the localities from which they were obtained 
suggest that intermediate forms may occur at great depths in the intervening area of 
the Indian Ocean. 

Register Nos., Localities, dc. Vv. 2, 4 (fragment), Saya de Malha, 7.9.1905, C. 20, 
3—500 fathoms; LxXxx1., Saya de Malha, 8.9.1905, C. 21, 450 fathoms. 


Genus HETERORETE™ n. gen. 


The sponge consists of thick-walled, branching, cylindrical tubes, of stony hardness. 
The main skeleton is a stout dictyonal framework with very irregular meshes, in which 
slender-rayed hexacts gradually become incorporated by fusion, especially in the inner 
part of the tube-wall, where the stout dictyonal framework gives place to a much finer 
network formed by union of the slender-rayed hexacts. The nodes of the reticulation 
are not provided with spiny warts or with lychnisks. There are no uncinates, no scopulze 
nor clavule, and no pentacts. There are discohexasters scattered in the parenchyma, 
especially in the inner portion of the wall. The canal system is complicated by strong 
folding of the chamber-layer and the flagellate chambers are comparatively small, oval 
or thimble-shaped. 

This genus evidently falls into Ijima’s family Dactylocalycide. It perhaps comes 
nearest to Dactylocalyx itself, but is remarkable for the complete absence of pentacts and, 
indeed, of special dermal or subdermal spicules of any kind. 


2. Heterorete pulchra n. sp. 
(Plate 41, figs. 11—18.) 


This species is represented in the collection by one good-sized fragment and a few 
small pieces evidently belonging to the same specimen. ‘The sponge consists of irregularly 
branching tubes (fig. 11), about 8 mm. in diameter, and with walls about 2 mm. 
in thickness, so that there is a wide lumen of about 4 mm. diameter. There is some 
evidence of anastomosis between adjacent tubes, but this cannot be regarded as definitely 
established. Both imner and outer surfaces of the tubes are marked with numerous 
small, thickly-scattered pits, representing the openings of the exhalant and inhalant 
canals. The texture is rigid and stony, but brittle; the colour in spirit (with the soft 
tissues preserved) is opaque yellowish white, in the macerated condition it is glassy and 
transparent. Numerous minute Anthozoa (? Zoanthids) are attached to the outer surface 
of the tubes, but at wide intervals and apparently without any connection with one 
another. Apparently they have no effect on the growth of the sponge, which does not 
seem to respond in any way to their presence. 

The main skeleton (figs. 12, 17) is a stout dictyonal framework of cylindrical trabeculee, 
with irregular meshes. The bars are somewhat stouter and the meshes smaller at the 
dermal surface, where also short, conical processes are given off from the trabeculze towards 
the dermal membrane, which they help to support. The whole of the dictyonal framework 


* This name is proposed in allusion to the two kinds of skeletal net-work. 


DENDY—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 215 


thus constituted is roughened with small conical spines, but these are more strongly 
developed on the subdermal trabecule than deeper down. 

At various points rather small, spimy-rayed hexacts are seen to be undergoing 
incorporation in the general framework by fusion of their rays with the trabecule. To 
what degree the framework really grows by this incorporation of originally separate hexacts 
it is impossible to say, but the followmg observations rather suggest that it extends 
centripetally in this manner. 

For some little distance beneath the gastral surface, in the region occupied by the 
inner trabecular layer of soft tissues, the main dictyonal framework is absent and its place 
is taken by scattered hexacts quite irregularly arranged. These hexacts (figs. 12, 17, hew.) 
are far more numerous here than anywhere else in the sponge, and it is at the junction of 
this layer with the chamber-bearing layer that the incorporation of hexacts in the dictyonal 
framework is chiefly seen (figs. 12, 18). 

Possibly growth of the main skeleton also takes place by the formation and subsequent 
fusion of outgrowths from the trabecule themselves, as in Sarostegia oculata, but it is often 
difficult to distinguish between such outgrowths and the projecting rays of partially 
incorporated hexacts. 

The hexacts of the subgastral layer also frequently unite with one another by fusion 
of rays, and thus tend to form a very irregular dictyonal framework with much smaller 
meshes and more slender trabecule than those of the main skeleton (fig. 12, hez.). 

Sometimes even discohexasters may be incorporated in the skeletal framework, 
giving rise to very curious appearances (fig. 13). 

The parenchymal spicules are as follows :— 

(1) Spimy-rayed hexacts (figs. 12, hex., 14); with rays straight or slightly curved, 
and varying a good deal in length and thickness ; sometimes sharply pointed at the ends 
and sometimes more or less clubbed. These spicules occur chiefly in the subgastral layer, 
where they are united together by fusion of rays into an irregular, loose network, while 
the outer ones are also united in the same manner with the inner portion of the main 
dictyonal framework. It is difficult to find a single hexact lying entirely free in the 
parenchyma. 

(2) Oxyhexasters (fig. 15); with long, slender, sharp-pointed rays. These appear to 
be extremely rare and are perhaps not a normal constituent of the spiculation. 

(8) Discohexasters (fig. 16); with slender, curved rays terminating in toothed discs 
and varying much in length in different specimens. There are usually about five terminal 
rays to each principal. These spicules are very abundant in the subgastral portion of the 
sponge wall. Occasionally they become incorporated in the skeletal framework (fig. 18). 

This sponge is sufficiently well preserved to enable me to give some account of the 
structure of the soft parts (figs. 17, 18). A very delicate dermal membrane (d.m.) is 
stretched over the outer surface of the main skeletal framework. This framework is 
interrupted at frequent intervals by the rather wide inhalant canals (7.c.). The dermal 
membrane still extends over the outer ends of these canals in some cases as a thin net 
pierced by the inhalant pores, but for the most part it is absent from the openings in 
question, perhaps owing to abrasion or shrinkage. Beneath the dermal membrane comes 


216 PERCY SLADEN TRUST EXPEDITION 


a very thin external trabecular layer (0.¢./.), followed immediately by the chamber layer, 
which occupies by far the greater part of the thickness of the sponge wall. Then, on the 
inside of the chamber layer, comes a fairly thick subgastral or inner trabecular layer 
(z.2.1.), bounded internally by a thin gastral membrane (g.m.). 

The wide inhalant canals run inwards approximately at right angles from the dermal 
surface and extend throughout the greater part of tle thickness of the sponge wall. They 
branch more or less and interdigitate with similar exhalant canals, also branched and also 
extending through the greater part of the thickness of the wall, which open by wide 
apertures on the gastral surface (whether the gastral membrane ever extends over these 
apertures as a delicate net I am unable to say definitely, but I think it highly probable 
that 1t sometimes does). Owing to the obliquity and irregularity of their direction a 
transverse section of the sponge wall (fig. 17) shows the inhalant and exhalant canals 
sometimes cut across and sometimes cut lengthwise. 

The flagellate chambers (f:c.) are very small for a hexactinellid sponge, only just 
about as large as those of a typical Leucandra, such as L. phillipensis [Dendy 1893], and 
much smaller than those of L. australiensis [Dendy 1893]. Indeed the whole canal system 
very closely resembles that of a Leucandra, except that the trabecular layers are represented 
in the latter by more continuous mesogleeal tissue. The large exhalant and inhalant canals 
may be regarded as formed by folding of the chamber-bearing layer, and it is probable that 
in reality there is only a single, much folded layer of chambers. Such appears to be the 
case at any rate in many places (e.g. in part of the section represented in fig. 17), but 
usually the arrangement has become greatly confused. 

The chambers come close up to the surfaces of both inhalant and exhalant canals, but 
the actual surface in both is probably formed by a very thin, net-like trabecular layer. 
Most of the chambers communicate with the large exhalant canals, but some of them 
open into the irregular spaces in the subgastral trabecular layer (fig. 18). 

The chambers themselves (figs. 17, 18, fc.) are oval or thimble-shaped. I have 
measured them up to 0°2 mm. in length, but usually they appear a good deal shorter than 
this. The collared cells are very small and indistinct and it is impossible to make out any 
satisfactory histological details. Such histology as I have been able to observe is repre- 
sented in fig. 18. 

In addition to the numerous small Anthozoa attached to the outer surface, the sponge 
wall is penetrated in various directions by the branching stolons of a hydroid colony. The 
hydranths (fig. 17, hyd.) are elongatedly club-shaped, with few tentacles (two or three ?) 
arranged in a single whorl springing from a short distance beneath the mouth. They are 
only sparsely scattered at long intervals on the hydrorhiza and appear to be capable of 
protrusion sometimes from the outer and sometimes from the inner surface of the sponge 
wall, though all now in a state of complete retraction. There is no distinct horny perisare, 
though sometimes a very thin layer can be discerned which may represent the last vestige 
of such a structure. The hydranths occupy definite tubular cavities which run inwards 
from the surface of the sponge and appear to be lined by a continuation of the dermal or 
gastral membrane as the case may be. This hydroid is probably closely related to Ampha- 
brachium euplectellé, described by Schulze [1880] as occurring in the soft tissues of 


DENDY—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 217 


Euplectella aspergillum. The latter species, however, has only two tentacles to each 
hydranth, while our form certainly seems to have at least three in some cases. Moreover 
the tentacles in Schulze’s species are much longer than in ours, assuming both to be 
retracted to approximately the same extent. Schulze also speaks of a delicate, annulated 
perisarc tube in his species. 

It seems probable therefore that the commensal hydroid of Heterorete belongs to a 
distinct species from that of Euplectella and, provisionally at any rate, it may be named 
Amphibrachium infestans. 

Register No. and Locality. cxv., Salomon, 3.7.1905, C. 120—150 fathoms. 


Genus SarosrEcra Topsent [1904]. 


The sponge forms a coral-like colony of stony hardness, the more or less cylindrical, 
tubular or solid branches ramifying chiefly in one plane and sometimes anastomosing. 
The rather close dictyonal framework of the skeleton is made up of stout trabecule. The 
separate spicules consist of (1) dermal and gastral hexacts, in which one ray is frequently 
more or less completely reduced, (2) spinose hexacts, which tend to become incorporated 
with the dictyonal framework, (3) dermal sarule, (4) uncinates, (5) oxyhexasters, 
(6) discohexasters. . 

This well-characterised genus was founded by Topsent in 1904 for a remarkable 
sponge obtained by the “‘ Princess Alice” and the “Talisman” in deep water off the Cape 
Verde Islands, and named Sarostegia oculata. 

In the same year Schulze [1904] proposed the genus Ramella for fragments of 
a similar sponge collected by the “ Valdivia” expedition near the Cape Verde Islands 
and Sumatra respectively. There can be little doubt that the specimen from the Cape 
Verde Islands at least is both generically and specifically identical with Topsent’s. 

Curiously enough, in the same year again, H. V. Wilson [1904, p. 84] proposed 
a genus, Sclerothamnopsis, for some fragments collected by the “ Albatross” expedition 
in the Eastern Pacific, which may very well be generically identical with Topsent’s and 
Schulze’s specimens. In neither of these two latter cases, however, was the material 
sufficiently well preserved to afford the basis of a satisfactory generic diagnosis. In 
the ‘“ Valdivia” material the only separate spicules found were the uncinates. In the 
“ Albatross” fragments the following are described, although in the generic diagnosis it 
is stated that the free spicules are not known with certainty :— 

(1) Spinose hexacts (similar spicules occur in the “Sealark” material). 

(2) Slender, smooth oxydiacts, always broken (probably broken uncinates, which, 
in the “Sealark” material, may have the spines very feebly developed, so that they 
resemble smooth oxydiacts). 

(3) Oxyhexasters (similar spicules occur in the “ Sealark” material). 

(4) Pinnules of peculiar form, with the distal ray enormously swollen and beset with 
very short spines. Wilson remarks that these pinnules are very few in number but so 
peculiar that it seems likely that they belong to the sponge. On the other hand, in the 

SECOND SERIES—ZOOLOGY, VOL, XVII, 28 


218 PERCY SLADEN TRUST EXPEDITION 


same work he figures practically identical pinnules for Hurete erectum, and I am strongly 
inclined to the opinion that they occur only as foreign bodies in Sclerothamnopsis. 

(5) *Scopulee, few in number and also very likely foreign. 

Neither pinnules nor scopulz occur in the specimens of Sarostegia examined by 
Topsent and myself, but, in place thereof, the peculiar dermal spicules which Topsent 
termed “sarules” and which I propose to call “sarule.” 

On the whole there seems to be a strong probability that Wilson’s Sclerothamnopsis 
is generically identical with Topsent’s Sarostegia and Schulze’s Ramella. If so, the genus 
is a very widely distributed one, occurring in the Atlantic, the Pacific and the Indian 
Oceans. 

Under these circumstances the question of priority naturally arises, and we have to 
determine whether the genus is to be known as Sarostegia, Ramella or Sclerothamnopsis. 

The date of publication of Topsent’s paper is May 1904, that of Wilson’s July 
1904. Schulze’s report on the “ Valdivia” Hexactinellids is marked “ Eingegangen den 
16 Dezember 1903,” but it was not published till some time in 1904; I have been 
unable to obtain further information as to the exact date. 

Clearly Sclerothamnopsis may be eliminated, unless it should prove necessary to 
retain it on account of some generic peculiarity of the “ Albatross” specimens, which 
seems improbable. As between the other two names I choose Sarostegia, on the ground 
that Schulze’s genus Ramella, based upon very imperfect material, in which the 
characteristic spicules were entirely wanting, was quite insufficiently diagnosed. 

Topsent placed Sarostegia in the family Farreide. This family was merged by 
Schulze [1904] in the Euretidz, in which he also included his Ramella. The reason ~ 
for the union of the two families was the breaking down of the distinction between 
“Scopularia” and “Clavularia” by the discovery of the genus Claviscopulia, described 
by Schulze in the “ Amerikanische Hexactinelliden” [1899, p. 76 &c.]. Claviscopulia 
intermedia, the type species of the genus, in addition to clavulee, possesses also spicules 
of a peculiar kind intermediate between clavule and scopule. These spicules, which 
Topsent terms ‘“‘sarules,” have the distal extremity club-shaped and beset with long 
spines, so that the whole comes to resemble somewhat a besom. It is very interesting 
to observe that sarules (or sarulee) occur also in Sarostegia, although of a somewhat 
different form from those of Claviscopulia. 

In his diagnosis of the family Euretidee, Schulze [1904, p. 177] says that the, 
coherent supporting framework is composed of dictyonal hexacts, which for the most 
part are united in a regular manner by the enclosure of the parallel apposed rays in 
layers of silica, so as to form a scaffolding with predominantly rectangular meshes. 
This is probably the primary arrangement of the framework in Sarostegia, but it is 
much obscured, at any rate in the “Sealark” specimen, by the formation of secondary 
trabecule subdividing the primary meshes into triangular areas. 


DENDY—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 219 


3. Sarostegia oculata Topsent. 


(Plates 42 and 43, figs. 19—36.) 


Sarostegia oculata Topsent [1904]. 

Ramella tubulosa Schulze [1904]. 

2? Sclerothamnopsis compressa Wilson [1904]. 

Topsent’s account of this beautiful species is not very detailed. The figures of 
external form are very fragmentary and only one kind of spicule, the sarula, is figured. 
It seems desirable, therefore, to give a complete account, with illustrations, of the 
“Sealark ” material. 

A large number of pieces were obtained, apparently in a single haul of the dredge, 
from a depth of 450 fathoms at Saya de Malha. It seems highly probable that they all 
formed part of a single specimen, which, owing to its brittle character, was broken into 
fragments in the dredge. The largest piece, drawn of the natural size in fig. 19, does 
not represent more than about one-sixth of the total material. Unfortunately the base is 
missing, but the sponge was doubtless attached in an erect position to a hard substratum 
by a somewhat expanded basal plate, as in the type. The branching is dichotomous 
(figs. 19a, 196), and seems to have taken place mostly m one plane, though with 
occasional deviations into a plane even at right angles to the principal one. The 
branches are approximately circular in transverse section, there being no conspicuous 
flattening. The thickest branches measure about 10 mm. in diameter, while the most 
slender, terminal, branches may measure as little as 2mm. Anastomosis of the branches 
appears to take place only occasionally. 

The branches are partly tubular and partly solid. The tips may sometimes bear 
a small terminal opening, but at other times they appear to be solid, while considerable 
cavities may occur in the older portions. These cavities open to the exterior by very 
irregularly distributed oval apertures (figs. 19 ¢, 19d, ap.) in the wall of the tube. They 
are sometimes occupied by polychzte worms, and I suspect it is the presence of these that 
keeps them open by preventing the ingrowth of tissues which takes place elsewhere. 
Certainly the solidification may take place while the branch is still very young. Topsent 
appears to regard the apertures in question as oscula, but I am very doubtful whether 
they can be correctly interpreted as such. 

The surface of the sponge has a finely granulated character. The texture is hard but 
brittle, and the colour in alcohol and formalin is very pale brown™. 

The whole of the surface of the sponge is more or less thickly studded with com- 
mensal or parasitic polyps (pol.), presumably zoanthids related to Palythoa. These are 
attached to the surface by expanded bases and can be picked off like scabs, leaving 
shallow depressions behind. It is extremely interesting to observe that the sponge 
responds to the presence of these polyps by enclosing each one in a delicate upgrowth 
of the dermal membrane, forming a thin translucent collar (figs. 19, 19 f, 19g, col.), 
supported by the characteristic dermal spicules of the sponge, with a marginal fringe 


* Topsent describes the sponge in life as being “‘semi-transparente, de teinte délicate, jaunatre-rosée, émaillé 


d’Actinies commensales d’un orangé assez vif.” 
28—2 


220 PERCY SLADEN TRUST EXPEDITION 


of sarulee. These collars, however, are only preserved where the surface of the sponge 
has been protected from rubbing, as shown in fig. 19. In one case (fig. 19 f) two polyps, 
apparently formed by fission of a single one, were observed within the same collar. The 
polyps appear to be connected with one another by a network of stolons ramifying in the 
thickness of the sponge wall. 

Similar polyps were described by Topsent in his specimen, and Schulze [1904] speaks 
of the macerated fragments obtained by the “ Valdivia” as showing “dellenartige Ober- 
flichenvertiefungen von ovaler oder doch rundlicher Form mit schwach erhabenem 
Rande.” It seems probable that these are the shallow depressions left after the removal 
of the polyps. The association between sponge and polyp thus appears to be a 
constant one. 

The main skeleton is a close framework of usually stout trabeculee (up to about 
0°07 mm. in thickness), with triangular meshes. As many as six bars of this frame- 
work may radiate from a common centre in approximately the same plane*, like the 
spokes of a wheel, connected at their outer ends by the spokes of similar adjacent 
systems, giving the whole framework a very characteristic appearance, as shown in 
fig. 33. 

Apparently the whole framework grows not so much by incorporation of new 
hexacts as by the outgrowth of secondary trabecule, whose ends meet and fuse to form 
systems similar to those just described. This process is certainly responsible for the 
inward extension of the framework by which the original central cavity becomes more 
or less obliterated. 

The structure of the dictyonal framework thus agrees very closely with Schulze’s 
description and figure of Ramella tubulosa, except that what I may perhaps term the 


” 


“rotulate” character, due to the formation of triangular meshes, appears to be more 
strongly pronounced (fig. 33). I have no doubt, however, that this character is a very 
variable one. In the older parts of the skeleton the trabecule are smooth, but in the 
younger parts, adjacent to the dermal and gastral surfaces, where the trabeculz are 
more slender, they are often roughened with minute projections (fig. 34). 

The manner in which the dictyonal framework spreads into the central cavity is 
well shown in figs. 35 and 36. Slender, more or less radially arranged outgrowths 
are given off from the superficial trabeculee of the gastral surface (fig. 35, pr.), and their 
ends, coming in contact with one another, fuse to form a new node of the skeleton 
(fig. 36, pr.). Doubtless these slender processes, which are at first minutely roughened, 
are thickened and become smooth later on by the addition of concentric layers of silica. 
Growth seems to take place at the dermal surface of the dictyonal framework in 
precisely the same manner, and probably some, at any rate, of the freely projecting, 
minutely spiny knobs, which occur on this surface, are the immediate agents concerned 
therein. I have seen just the same fusion of such outgrowths to form a new node on the 
dermal as on the gastral surface. Some of the minutely spiny knobs on the dermal 
surface, however, appear to be the reduced centrifugal rays of the outermost fused hexacts 
of the original framework, as described by Topsent. 2 


* Of course similar bars radiate from the same centre in other planes. 


DENDY—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 221 


The dermal skeleton consists, in the first place, of stout hexacts (figs. 20, 21, 22), 
with the outwardly projecting ray much shortened and the inwardly projecting ray 
lengthened. The rays are bluntly rounded at their extremities and usually quite smooth, 
although a little roughening may sometimes be detected, especially towards the end 
of the longest ray, which may also be more sharply pointed (fig. 21). 

Between the dermal hexacts occur numerous radially arranged sarule of the form 
shown in figs. 238, 24, 25. These remarkable spicules consist of a straight shaft, some- 
times slightly roughened, and terminating at the inner end in a blunt point, while the 
outer end is oval club-shaped and covered with stout, sharp, forwardly-directed spines 
more or less fused together. 

The dermal skeleton is evidently very easily rubbed off and is only well preserved in 
places, especially in the membranous collars surrounding the polyps, where, as already 
stated, the sarule form a marginal fringe. 

The gastral skeleton consists of hexacts (figs. 26, 35, hea.) of the same general type 
as the dermal hexacts, but the rays are more slender, generally less unequal in length 
(though usually one still seems to be very short), and frequently knobbed at the 
extremities. In the invasion of the original central cavity by the skeleton these spicules 
become very irregularly arranged (fig. 35, hewx.), but they may still be seen, crowded 
together in the middle of the sponge, even after the central cavity has been completely 
obliterated. They seem to retain their independence for a long time, but it is quite 
possible that they ultimately become incorporated in the dictyonal framework. 

The parenchymal spicules are as follows :— 

(1) Very long and slender uncinates, sharply pointed at the two extremities and 
with feebly developed teeth (fig. 32). 4 

(2) Small spiny hexacts (fig. 27) lying between the trabecule of the dictyonal 
framework, with which their rays may become fused. (It is sometimes dittcult to 
distinguish these from new nodes formed by outgrowth and fusion of spiny processes 
from the trabecule (cf. fig. 36, pr.).) 

(3) Oxyhexasters (figs. 28, 29, 30). Very variable, with smooth, slender, sharp- 
pointed rays. 

(4) Discohexasters (fig. 31). Rather compact, with not very long terminal rays, 
about five to each principal. 

- Although a good deal of the soft tissues of the sponge still remains, I have not been 
able to make out the flagellate chambers. 

I do not think there can be any reasonable doubt that the ‘Sealark” specimen is 
specifically identical with Topsent’s type from the Cape Verde Islands, in spite of the 
difference in locality. There do, it is true, appear to be some minor differences in 
spiculation, as indicated by Topsent’s account of the dermal hexacts and _pentacts, 
which, unfortunately, he does not figure, and by the absence of all mention by him of 
the spiny hexacts (No. 2 above). It is probable, however, that these apparent differ- 
ences would disappear if it were possible to make a direct comparison of the specimens, 
and the agreement in other respects is so close that I feel justified in making an 
identification. 


222 PERCY SLADEN TRUST EXPEDITION 


Previously known Distribution. Near Cape Verde Is. (Topsent and Schulze) ; 
Sumatra (Schulze); 4 Hastern Pacific (Wilson). 


Register Nos., Locality, de. 1., U., UL, 1v., Iva., Saya de Malha, .8.9.1905, 
C. 21, 450 fathoms. 


LIST OF LITERATURE REFERRED. TO. 


1893. Dernpy, A. “Studies on the Comparative Anatomy of Sponges, V. Observations on the Structure 
and Classification of the Calcarea Heteroccela.” (Quart. Journ. Micro. Sci., vol. 35.) 


1903. Isima, 1. “Studies on the Hexactinellida. Contribution III.” (Journ. Coll. Sci. Imp. Univ. 
Tokyo, vol. 18.) 


1880. ScHunzE,F.E. “On the Structure and Arrangement of the Soft Parts in Huplectella aspergillum.” 
(Trans. Royal Soc. Edin., vol. 29.) 


1887. Id. Report on the Hexactinellida of the “Challenger” Expedition. 
1899. Id. “ Amerikanische Hexactinelliden nach dem Materiale der ‘ Albatross ’-Expedition.” 
1904. Id. Hexactinellida of the “ Valdivia” Expedition. 


1904. Topsent, E. “Sarostegia oculata. Hexactinellide nouvelle des iles du Cap-Vert.” (Bull. Mus. 
Océanographique de Monaco, No. 10, 20 Mai 1904.) 


1904.. Witson, H. V. Sponges of the “ Albatross” Expedition of 1891. 


Fig. 
Fig. 
Fig. 


Fig. 
Fig. 
Fig. 
Fig. 


DENDY—REPORT ON THE HEXACTINELLID SPONGES (TRIAXONIDA) 223 


DESCRIPTION OF PLATES. 


PuatE 40. 


Figs. 1—10a. Aulocalya serialis n. sp. 


1. Restoration of entire sponge (based mainly on R.N. v. 1). x 2. 

2. Part of skeletal framework (R.N. v. 1). x 110. 

3. Part of skeletal framework ending in hooks which project into the gastral cavity (R.N. v. 1). 
x 110. 

4, Large parenchymal hexact (R.N. v. 1). x 100. 

5. Small parenchymal hexact (R.N. v.1). x 320. 

6. Large pentact (R.N. v.1). x 100. 

7. Small pentact with vestige of sixth ray (R.N. v. 1). x 320. 


Figs. 8,9. Discohexasters (R.N. v.1). x 320. 


Fig. 
Fig. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 
Fig. 


Fig. 


Fig. 


10. Characteristic large hexaster with spiny rays (R.N. v. 1). x 320. 
10a. End of ray of large hexaster more highly magnified. 


PLATE 41. 


Figs. 11—18. Heterorete pulchra n. gen. et sp. 


11. Part ofa colony. x 2. 

pol. parasitic or commensal Anthozoa. 

12. Part of dictyonal framework, as seen in transverse section, showing the transition to the layer 
of scattered and partially fused hexacts (hew.) in the inner portion of the sponge wall. x 100. 

13. Portion of skeletal framework from inner part of sponge wall, showing incorporation of 
discohexasters. x 360. 

14. Spiny-rayed hexact. x 540. 

15. Oxyhexaster. x 540. 

16. Discohexaster. x 540, 

17. Transverse section of sponge wall, showing soft tissues and canal system. (Combined drawing.) 
x 50. 

d.m. dermal membrane; e.c. exhalant canal; fc. flagellate chambers; g.m. gastral membrane ; 
hyd. hydroid polyp; 2.c. inhalant canal ; 7.t./. inner (subgastral) trabecular layer; o.t.l. outer 
(subdermal) trabecular layer. 

18. Part of inner portion of a transverse section showing soft tissues, more highly magnified 
(Zeiss D. oc. 2). 
disc. discohexaster ; hex. hexacts. Other lettering as before. 


PLATE 42. 


Figs. 19-31. Sarostegia oculata Topsent. 


19. The largest piece of the sponge. Nat. size. 
col. collars of sponge tissue surrounding parasitic or commensal polyps (pol.). 


Figs. 19a, 19b. Two other pieces, showing ends of branches. Nat. size. 
Figs. 19¢, 19d. Portions of two hollow branches showing apertures (ap.). Nat. size. 


224 PERCY SLADEN TRUST EXPEDITION 


Fig. 19¢. A single polyp as it appears when removed from the sponge. x 3. 
Fig. 19 f Two polyps enclosed in the same collar (col.). x 3. 

Fig. 199. Side view of collar (col.) enclosing polyp. x 3. 

Figs. 20, 21, 22. Stout dermal hexacts, from collar around polyp. x 175. 
Figs. 23, 24, 25. Sarulz, from collar around polyp. x 175. 

Fig. 26. Gastral hexact, from central cavity of skeletal framework. x 175. 
Fig. 27. Spiny parenchymal hexact. x 175. 

Figs. 28, 29, 30. Oxyhexasters. x 540. 

Fig. 31. Discohexaster. x 540. 


PLatE 48. 


Figs. 32—36. Sarostegia oculata Topsent. 


Fig. 32. Uncinate. x 102. ; 

Fig. 38. Part of dictyonal framework as seen in longitudinal section through end of branch. x 60. 

Fig. 34. Part of dictyonal framework as seen in tangential section just below surface. x 175. 

Fig. 35. Skeleton surrounding central cavity, as seen in transverse section near tip of young branch. 
x 60. 

c.c. central cavity; hex. hexacts; pr. spiny processes of trabecule. 

Fig. 36. Skeleton surrounding central cavity, as seen in transverse section, to show the union of spiny 

processes (pr.) to form a new node of the network. x 60. 


PgRcy SLADEN TRUST EXPEDITION. TRANS. LINN. Soc. SER.2.Z00L VoL XVII Pr. 40. 


T.P Collings & HL,Deakin del. Cambridge University Press. 


AULOCALYX SERIALIS n sp. 


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AD. TPC. &H.L.D, del. Cambridge University Press. 


HETERORETE PULCHRA nisp. 


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Percy SLADEN TRUST EXPEDITION. TRANS.LINN. Soc. SER.2.Zoon VoL.XVIL .Pu.42. 
(DENDY ) 


TP Collings & H.L.Deakin del, Cambridge University Press. 


SAROSTEGIA OCULATA Topsent. 


a 


TRANS. LINN. SOc. SER 2.Z00L VOL.XVIL .PL.43. 


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PERCY SLADEN TRUST EXPEDITION 


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No. VI—REPORT ON THE HOMOSCLEROPHORA AND ASTROTETRAXONIDA 
COLLECTED BY H.M.S. “SEALARK” IN THE INDIAN OCEAN. 


By Arruur Denpy, D.Sc., F.R.S., F.LS., Professor of Zoology im the 
Unwwersity of London (King’s College). 


(Plates 44—48.) 


Read 17th June, 1915. 

The present instalment of my Report on the Sponges collected in the Indian Ocean 
by the “Sealark” expedition deals with a portion only of the Tetraxonida, viz. the 
Homosclerophora and Astrotetraxonida, leaving the Sigmatotetraxonida—a very large 
group—to be dealt with subsequently. It will be noticed that the families Spiras- 
trellidze and Latrunculiide, hitherto included by general consent in the Astrotetraxonida, 
are omitted from the present communication. The reason for this is that I have con- 
vinced myself—largely through investigation of the «“Sealark” material—that the 
so-called asters of these two families are really pseudasters, and that the groups im 
question are of desmacidonid origin and must therefore be included amongst the 
Sigmatotetraxonida. I hope to discuss the question at length in a future instalment of 
my Report. . 

The classification of the Astrotetraxonida is as yet by no means in a satisfactory 
condition, but a comprehensive revision of the group will be necessary before attempting 
any radical reform. Such a revision I have already commenced in conjunction with my 
colleague Mr R. W. H. Row, but it would not be desirable to delay the publication of 
this report until it is completed, as it must necessarily take a long time. I may be 
allowed to state, however, that, although I retain the family Pachastrellide as a matter 
of convenience, I cannot regard that family, as generally understood, as constituting 
a natural group—it probably contains forms on the up-grade from the Homosclerophora, to 
the Stellettidee and others which are nothing but degenerate Stellettids. The latter 
appear to be analogous to the so-called ‘“Epipolaside,” but differing from these in that 
it is only the rhabdome of the trizene that has undergone reduction. The “ Epipolasidee ” 
I no longer accept as a family, those which are present in the “Sealark ” collection will be 
found amongst the Stellettidze. 

SECOND SERIES—ZOOLOGY, VOL. XVII. : 29 


226 PERCY SLADEN TRUST EXPEDITION 


Twenty-five species are dealt with in the present contribution, of which nine appear 
to be new. The list is as follows :— - 


Order TETRA XONIDA. 


Sub-order HomoscLEROPHORA. 


Family Plakinide. 


1. Dercitopsis minor n.sp. 
Sub-order AsTROTETRAXONIDA. 


Family Pachastrellide. 


Pachastrella tenuilaminaris (Sollas). 


ow bo 


Yodomia perfecta n.sp. 


Family Stellettide. 
Myriastra parva (Row). 
Myriastra cavernosa un. sp. 
Dragmastra lactea (Carter) var. mauritiana nov. 
Lthabdodragma (n. gen.) conulosa (Kieschnick). 
Heionemia carteri Dendy. 


co I TF 


Hevonemia laviniensis Dendy. 
10. Aurora providentice n. sp. 
11. Aurora cribriporosa n. sp. 
12. Aurora rowi n. sp. 

13. Astéropus simplex (Carter). 
14. Jaspis johnstonii (Schmidt), 


Family Geodiide. 


15. Geodia auroristella n. Sp. 


Family Erylide. 
16. Hrylus lendenfeldi Sollas. 
17. Hrylus proximus n. sp. 


Family Donatiide. 
18. Donatia lyncuriwm auctorum. 
19. Donatia japonica (Sollas). 
20. Donatia ingalli (Bowerbank). 
21. Donatia seychellensis (Wright). 
22. Donatia stella-grandis nu. sp. 


Family Chondrosiide. 
23. Chondrilla australiensis Carter. 
24. Chondrilla miata Schulze. 
5) 


5. Chondrilla sacciformis Carter. 


I am again indebted to the Trustees of the Percy Sladen Fund for financial assistance 
in the preparation of illustrations, &c., and to Miss Deakin for much valuable help 
rendered possible thereby, especially for her faithful and painstaking drawings of spicules. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 227 


I refrain from publishing a reference list of literature with the present instalment, 
as it will be quite sufficient to publish one list later on for the whole of the Tetraxonida. 
In the meantime the dates given after authors’ names may afford sufficient clue to the 
memoirs referred to. 


Order TETRAXONIDA Dendy [1905]. 


Sponges with siliceous spicules whose fundamental form is tetraxonid and tetracti- 
nellid. (This fundamental form is often obscured by secondary modifications, and the 
spicules may even disappear completely in some degenerate forms.) 

In 1905 I proposed to arrange the order Tetraxonida as follows :— 


Grade TETRACTINELLIDA. 


Sub-order Homosclerophora. 
Astrophora. 


3? 


ee Sigmatophora. 
Grade LiTHISTIDA. 


Grade MoNAXONELLIDA. 


Sub-order Astromonaxonellida. 
Sigmatomonaxonellida. 


9 


The Sub-orders Astrophora and Sigmatophora were, of course, adopted from Sollas 
[1888] and the Homosclerophora replaced his Microsclerophora. 

The Astromonaxonellida were regarded as being derived from the tetractinellid 
Astrophora and the Sigmatomonaxonellida from the tetractinellid Sigmatophora [ Dendy, 
1905, p. 133], 

The views thus expressed as to the phylogeny of the Tetraxonida have been accepted 
by Hentschel [1909, 1911 4] in his work on the Tetraxonida of S.W. Australia. He has, 
however, proposed a modification of my arrangement which gives clearer expression to 
these views and which I gladly accept. He divides the order Tetraxonida directly into 


three sub-orders :-— 


Sub-order 1. Homosclerophora Dendy. 

2. Astrotetraxonida Hentschel (= Astrophora + Astromonaxonellida 
Dendy). 

3. Sigomatotetraxonida Hentschel (=Sigmatophora + Sigmatomonaxo- 
nellida Dendy). 


We are thus, I hope, finally rid of the old artificial distinction between “ Tetracti- 
nellida” and “Monaxonida,” introduced by Zittel [1878 4], which the “Challenger” 
Reports unfortunately did so much to emphasise. 

With regard to the Lithistida and Ceratosa Hentschel remains in some doubt, 
suggesting that they may have to be added as two separate sub-orders to the three 
above mentioned. This is possibly the best thing to do with the Lithistida in the 
present stage of our knowledge, though I should not like to commit myself to a definite 

29—2, 


29 


228 PERCY SLADEN TRUST EXPEDITION 


opinion as yet. With regard to the Ceratosa I adhere to the views I have already 
expressed [1905]. Only the “‘ Pseudoceratosa” have any claim to be included in the 
Tetraxonida, and whether these can be constituted into a distinct sub-order is extremely 
doubtful. Some of them appear to me to be certainly Chalinine in origin, while others 
are very possibly Ectyonine. 


Sub-order 1. HomosctrropHora Dendy [1905]. 


Tetraxonida in which microscleres and megascleres have not yet become sharply 
differentiated from one another and no triznes are as yet developed. 

I cannot agree with Hentschel in including [1909] the Oscarellide in this sub-order, 
which seems to me to be a distinctly retrograde step. I adhere to the opinion which 
I expressed in 1905, that Oscarella must be placed in a separate order, Myxospongida, 
which represents the common ancestors of all the siliceous sponges, both Triaxonida and 
Tetraxonida, and also of the Euceratosa. 


Family Plakinide. 


With the characters of the sub-order. 

This family was proposed by Schulze in 1880 for the reception of the ee genera 
Plakina, Plakortis and Plakinastrella. References to its history between 1880 and 1900 
are given by Lendenfeld [1903, p. 118]. In addition to the three original genera 
Lendenfeld includes in the family Corticium and Thrombus. In 1905 I removed 
Plakinastrella from the Plakinide on account of the presence of short-shafted trizenes, 
and placed it in the Pachastrellide, an arrangement to which I must adhere. 


Genus Dercrtoprsis Dendy [1905]. 


Plakinidee with calthrops, oxea and sometimes triods, but no candelabra. All 
spicules smooth. 

When I proposed this genus in 1905 I unfortunately overlooked the existence of 
two species which must certainly be taken into account in discussing its affinities, viz. 
Plakinastrella clathrata, described by Kirkpatrick [1900 8] from Funafuti, and P. oxeata, 
described by Topsent [1904 4] from the Azores. More recently Lendenfeld [1906] has 
described a species under the name Plakinastrella mammillaris, from the west coast 
of Australia, which must also be considered in the same connection. ‘That all these three 
species are closely related to my Dercitopsis ceylonica there can be no doubt, but I am 
not disposed to agree with Lendenfeld [1906] that Dercitopsis ceylonica should be 
associated with them in the genus Plakinastrella. On the contrary I think that all 
three should be removed from Plakinastrella and placed in cP raeopels, and that for 
the following reasons. 

The type species of Plakinastrella is P. copiosa, described and figured by Schulze 
[1880]. That species possesses well-differentiated, short-shafted trizenes, definitely 
orientated beneath the surface of the sponge, and, as already stated, it was mainly for 
that reason that in 1905 I placed it in the Pachastrellidee. Not one of the species placed 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 229 


by Kirkpatrick, Topsent and Lendenfeld in the genus Plakinastrella possesses trizenes, 
but, on the contrary, all of them belong to the Homosclerophora. _ 

The genus to which Dercztopsis is most closely allied is not, in my opinion, Plakina- 
strella, but Plakortis [Schulze 1880]. Indeed it is perhaps not easy to separate the two 
satisfactorily, but Schulze, in his original diagnosis of the genus Plakortis, emphasised the 
fact that tetract spicules were wanting, only triacts and diacts being developed. We may 
take this as the basis of the generic distinction. Plakortis also seems to be more primitive 
than Dercitopsis as regards canal system, but in this respect the gap seems to be bridged 
over to some extent by Dercitopsis clathrata (Kirkpatrick). 

The presence of radially arranged small oxea at the surface must be abandoned 
as part of the generic diagnosis of Dercitopsis, for, though such a layer is present in 
D. ceylonica Dendy, D. clathrata (Kirkpatrick) and D. minor n. sp., it is absent in 
D. oxeata (Topsent) and (apparently) in D. mammullaris (Lendenfeld). The last named 
species is remarkable in another respect, for the spiculation includes only oxea and 
calthrops, triods bemg completely absent. 


1. Dercitopsis minor n. sp. 
(Plate 44, fig. 1; Plate 45, fig. 1.) 


The sponge (Plate 44, fig. 1) forms irregular, rounded, cushion-like masses, encrusting 
pieces of rock, &c. The margins are broadly rounded and may project considerably beyond 
the base of attachment, and become tucked in, thus tending to envelope the support. The 
maximum dimensions of the largest specimen (R.N. xuit. 6) are as follows:—Length 78 mm., 
breadth 57 mm., thickness about 16 mm. ‘The surface is smooth but rather uneven ; 
subglabrous and minutely punctate. The colour of the surface (in spirit) varies from 
light brown to dark slate grey ; internally it is pale yellowish. Vents of moderate size, 
up to about 3 mm. in diameter, each with a prominent, membranous collar ; few in number 
and scattered singly on prominent portions of the upper surface. Inhalant pores closely 
scattered all over the surface. Texture firm and compact. 

The skeleton is a dense feltwork of loose spicules, quite irregularly arranged except 
at the surface, where very small oxea are placed more or less at right angles to the surface 
to form a dermal layer. 

Spicules:—(1) Calthrops (Plate 45, fig. 1 a), with smooth, sharp-pointed rays measuring 
about 0°037 by 0:005 mm. in a well grown specimen, but varying a good deal. 

(2) Triods (Plate 45, fig. 16), differing from the calthrops in the absence of one ray. 
Perhaps, on an average, the rays of the triods are more slender than those of the calthrops, 
but slender-rayed forms of both occur. 

(3) Oxea (Plate 45, fig. 1c), fusiform, slender, slightly curved, gradually sharp- 
pointed, almost always with a kink or enlargement in the middle. Size variable, averaging 
in the deeper parts of the sponge, say, about 0°1 by 0:004 mm., though often much more 
slender. The largest seen, and that only once, measured only about 0°19 mm. in length. 
The small dermal oxea measure up to ‘about 0:04 mm. long and are of proportionate 
thickness. Intermediate sizes between these and the deeper oxea are abundant. The 
oxea are far more numerous than both triods and calthrops together. 


230 PERCY SLADEN TRUST EXPEDITION 


The ectosome and the outer part of the choanosome contain numerous small, brown, 
granular pigment-cells. I have not examined the canal system in detail, but it appears 
to agree closely with that of Dercitopsis ceylonica [Dendy 1905}. 

Dercitopsis minor would appear to be a common species in the Indian Ocean. At 
first I thought it must be specifically identical with the Ceylon form, but the fact that the 
oxea never, in any of the specimens, seem to reach half the size that many of them attain 
in D. ceylonica, renders it, in my opinion, desirable to recognise a specific distinction. 

The external appearance of the sponge strongly recalls that of a Chondrilla, with 
which genus it may readily be confounded until examined microscopically, especially when, 
as in the case of R.N. xxxuir. 1 a, the two are growing together. 

Register Nos., Localities, de. Xxx. 1A, XXxul. 2, and xu. 3, Cargados Carajos, 
30.8.05, B. 13, 30 fathoms ; xxi. 6, Cargados Carajos, 30.8.05, B. 9, 30 fathoms; ci. 24 
(encrusting Hrylus lendenfeldi), Amirante, 18.10.05, E. 25, 44—20 fathoms; cxi. 7 and 
cx. 12, Egmont Reef. 


Sub-order 2. AstRoTETRAXxoNIDA Hentschel [1909]. 


Tetraxonida with astrose microscleres (except when these have been lost secondarily) ; 
without sigmata or their derivatives. 


Family Pachastrellide. 


Astrotetraxonida with calthrops and (or) short-shafted trizenes, usually scattered 
irregularly in the interior of the sponge, though some of the short-shafted trizenes may be 
definitely orientated, with the cladi supporting the ectosome. Without typical long- 
shafted triznes and without sterrasters. 

’ As already stated, this family is only retained provisionally. It is probably of poly- 
phyletic origin, containing both primitive forms and degenerate stellettids. 


Genus PacHAsTRELLA Schmidt [1868]. 


Pachastrellidze with oxea and calthrops and (or) short-shafted trizenes for megascleres ; 
without mesotrizenes ; with microrhabds and various forms of streptaster for microscleres, 
but without spherasters. 

I must agree with Lendenfeld [1903] in merging Sollas’s genus Peecillastra, which 
I was at first strongly disposed to retain, in Pachastrella. There appears to be really 
nothing but the plate-like form and the distribution of pores and oscula to distinguish the 
two, and these characters vary so much that they cannot, at any rate in this case, be 
regarded as of generic import. 


2. Pachastrella tenuilaminaris (Sollas). 


(Plate 45, fig. 2.) 


Normania tenuilaminaris Sollas [1886]. 
Pecillastra tenwilaminaris Sollas [1888]. 
Pachastrella crassiuscula Lendenfeld [1903]. 
Pachastrella tenwilaminaris Lebwohl [1914]. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 231 


I identify with this species a plate-like fragment of considerable size, about 58 mm. 
in length, 45 mm. in breadth and 8 mm. in thickness. The plate is slightly curved and one 
surface bears small, thickly scattered pore-sieves, while the other bears numerous small, 
scattered oscula, less than 1 mm. in diameter and with their margins level with the general 
surface. The margin of the plate, where intact, is broadly rounded, but the incurrent face 
ends sharply above in a well-marked edge. Part of this edge only is provided with a 
fringe of long projecting oxea and both surfaces are also hispid in places. 

The broken edges show the narrow inhalant and exhalant canals running through the 
plate more or less at right angles to the two surfaces. 

The colour in spirit is dull, pale yellow; texture firm and harsh, but rather friable. 

The skeleton is an extremely confused feltwork of large oxea. There are also 
numerous very long and very slender oxea which are chiefly arranged in loose fibres or 
wisps, which run towards the surface and in places project therefrom in long loose 
bundles. The comparatively few tetract spicules appear to be quite irregularly scattered 
through the sponge. 

Spicules. (1) Calthrops and short-shafted trizenes (Plate 45, fig. 2a), not sharply 
distinguishable from one another. Rays sharp-pointed or rounded, about 0°5 mm. long by 
0-05 mm. in diameter at the base. The rays are sometimes slenderer. These spicules are 
not numerous and they rarely show any reduction of rays, but irregularly branched forms 
are occasionally found. 

(2) Stout, fusiform, slightly curved and sharply pointed* oxea (fig. 2 b), measuring 
about 3°0 by 0:05 mm. Sometimes reduced, by rounding off of one or both ends, to styl 
or strongyla (figs. 2c, 2d). Shorter oxea also occur. 

(3) Long, hair-like oxea (fig. 2e), of about the same length as the largest but only 
about 0°008 mm. thick. Very numerous and commonly arranged in loose wisps. 

(4) Slender-rayed metasters (fig. 2), with about six or eight long rays or spines. 
Greatest length of entire spicule usually about 0°016 mm. 

(5) Microxea (fig. 2g); slender, slightly curved, sharply pointed; with very slight 
indications of roughening ; size about 0°15 by 0:004 mm. Rather scarce. 

I have in my possession several of Sollas’s original preparations of Pachastrella 
(Pecillastra) tenmlaminaris and a careful comparison of these with the “Sealark” 
Specimen seems to me to justify a specific identification. The chief apparent differences 
are as follows. 

(1) The “Challenger” specimen had no special hispidating fringe at the margin. 
This is of little importance. 

(2) The long hair-like oxea are not mentioned in Sollas’s description. They are 
certainly very rare m the “Challenger” material, but I have seen a few. 

(3) The tetract spicules are less numerous in the “Sealark” specimen; they show no 
tendency (so far as observed) to regular arrangement at the margin, as in the “Challenger” 
specimen, 

(4) Reduced tetracts with only one ray are common in the “Challenger” material. 
T have never seen them in the “ Sealark ” specimen. 


* Sollas’s statement that the oxea are not sharply pointed is not borne out by his preparations. 


232 PERCY SLADEN TRUST EXPEDITION 


(5) The metasters are decidedly smaller in the “Sealark” specimen and do not show 
so strong a tendency to pass into plesiasters. 

(6) The microxea are much fewer in the “Sealark” specimen and the tendency to 
roughening of the surface is less pronounced, but it is very slight even in the “Challenger” 
specimen. 

(7) The plate of which the sponge is composed is more than twice as thick in the 
“‘Sealark ” specimen as in the type of P. tenuilaminaris, but in spite of the specific name, 
somewhat unfortunately chosen, this character cannot be regarded as of great importance. 
Lendenfeld [1903] regards P. tenwilaminaris Sollas as a synonym of P. crassvuscula 
Sollas, but I doubt whether this is justifiable in the present state of our knowledge. 
The chief distinguishing feature of the species appears to be the absence of short-spined 
spirasters. 

Previously known Distribution. South of Japan, 775 fathoms (“ Challenger”); Japan 
(Lebwohl). 

Register No., Locality, dc. Uxxit. 2, Amirante. 


Genus Yopom1a Lebwohl [1914]. 


Pachastrellidee in which the principal megascleres are calthrops (or short-shafted 
trizenes), mesotriznes and oxea, with various derivatives of these often exhibiting very 
abnormal characters. The microscleres consist of amphiasters (or possibly some other form 
of aster) and microrhabds. 

This genus has recently been proposed by Lebwohl [1914] for the reception of a 
remarkable Japanese species, Yodomia yimai, which agrees with Triptolemus in the 
presence of mesotriznes but differs in the possession of calthrops (or trizenes) and oxea as 
well, thereby approximating to the more typical Pachastrellide. The presence of abnormal- 
looking derivatives of the megascleres, sometimes forming spheres, appears also to be very 
characteristic. . 

Lebwohl gives the following diagnosis of his new genus “ Pachastrellidze mit lang- 
schiiftigen Triznen; mit radial orientirten Plagiotriznen und Mesotriznen an der Ober- 
fliche.” It appears to me that he has here laid undue emphasis on the long-shafted 
trieenes, which are far from being typical long-shafted trizenes and are said to be relatively 
scarce. In the new species discovered by the “Sealark” expedition they do not occur 
at all. 

The genus Triptolemus, proposed by Sollas in 1888, includes small encrusting forms, 
and may possibly be regarded as having been derived from Yodomia by reduction of the 
spiculation. 

Schmidt’s Stelletta pathologica [1868] from the coast of Algiers, redescribed by Sollas 
in 1888, also includes mesotrizenes in its spiculation and is perhaps a nearly related form. 


3. Yodomia perfecta n. sp. 


(Plate 44, figs. 2, 2a; Plate 45, fig. 3.) 


The external form of the sponge is irregular and variable. Thus R.N. x. 1 (Plate 44, 
fig. 2) forms a flat, spreading crust about 8 mm. in thickness, with an uneven, nubbly 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 233 


surface and strongly hispid in places, especially along part of the margin, where dense 
tufts of oxea project for as much as 3 mm.; the entire crust is about 55 mm. in length 
and 27 mm. in greatest breadth; irregularly oval in shape. R.N. x. 2 and x. 3 are 
essentially similar to R.N. x. 1. R.N. vr. (Plate 44, fig. 2a), on the other hand, has the 
form of a thick, vertical, wall-like plate, attached without any spreading base to a stone. 
The two sides of the plate are flattened but rather uneven, the margin broadly rounded, 
the surfaces alike, coarsely granular in appearance but at the same time slightly hispid in 
places. The entire specimen measures about 50 mm. in length, 30 mm. in height and 
18 mm. in maximum thickness. R.N. rx. 1 closely resembles R.N. v1. 

None of the specimens show any oscula. The texture is compact, firm and very 
harsh to the touch. Colour in spirit pale, dull yellow. 

The skeleton is a dense, confused mass of large and small calthrops and small meso- 
trizenes, penetrated here and there by loose wisps or bundles of oxea, running towards 
the surface. The relative numbers of the large and small calthrops vary greatly in 
different specimens. The microrhabds are abundantly scattered all through the sponge 
but are accumulated in an especially dense layer at the surface. 

Spicules. (1) Large calthrops or short-shafted trizenes (Plate 45, figs. 3a—3 a”). 
Rays generally about equal in length, sometimes straight and sometimes curved or 
crooked ; sometimes sharp-pointed and sometimes rounded off, sometimes (in R.N. 1x. 1) 
reduced to rounded knobs, so that the whole spicule may become almost spherical 
(fig. 8 a’); occasionally divided into two short branches at the extremity (figs. 3a, 3 a’) ; 
size very variable; rays measured up to about 1°5 by 0°15 mm. 

(2) Small calthrops or short-shafted trivnes (figs. 3b, 30’). Rays usually straight, 
gradually sharp-pointed ; sometimes differing a good deal in length, while more frequently 
the three which are alone fully visible at the same time appear to be about equal; length 
very variable, say about 0°2 mm., with a diameter of about 0°03 mm. Numerous inter- 
mediate sizes between these and the large calthrops also occur. 

(3) Mesotrizenes (figs. 3c—3c”). The three cladi spring from about the middle of 
a short shaft. The cladi are always branched, usually bifurcating once only but occasionally 
twice. All the cladi and both ends of the shaft are gradually sharp-pointed. Size very 
variable, say about 0°2 mm. across the cladome, from tip to tip of cladi, in R.N. x. 1, but 
may be at least twice this size in R.N. vi. Each half (ray) of the straight, unbranched 
shaft (rhabdome) is about as long as the cladi, An abnormal form with four cladi has 
been met with and one with only one ray of the rhabdome developed. 

(4) Oxea (fig. 3d). Very long, straight and slender; tapering very gradually to 
each extremity; measured up to 7°77 by 0°04 mm. 

(5) Amphiasters (fig. 3e). The shaft between the two whorls of rays is so short 
that the spicule looks like an oxyaster, especially when seen obliquely or end on, but 
I think it is really an amphiaster. The rays are long, slender and sharply pointed, 
altogether about 10 in number. Total diameter of spicule about 0-016 mm. These 
spicules are abundantly scattered through the sponge. 

(6) Smooth microrhabds (fig. 3/). Oval, measuring about 0°012 by 0-006 mm. 
Extremely abundant throughout the sponge, but especially so at the surface. Sometimes 

SECOND SERIES—ZOOLOGY, VOL. XVIL. 30 


234 PERCY SLADEN TRUST EXPEDITION 


varying to more slender forms as shown in the figures, and even passing into the next 
form. 

(7) Spined microrhabds (fig. 3g). Slender; covered with minute short spines ; 
measuring about 0°022 by 0:002 mm. (exclusive of spines), but variable. This spicule is 
extremely scarce in some specimens, though plentiful in others, so that it may very easily 
be overlooked. It seems to be a characteristic feature of the genus Triptolemus and is 
probably proper to Yodomia also. It may be a reduced streptaster. 

The condition of the material and the character of the skeleton make it impossible 
to prepare satisfactory paraftin sections, but investigation by this method reveals the 
presence of an enormously thick, gelatinous ectosome. This tissue appears to be of the 
nature of that termed ‘“‘chondrenchyme” by Sollas [1888]. Imbedded in the clear, 
gelatinous, faintly stainmg matrix are numerous oval, granular cells, about 0°02 mm. 
in diameter, each with a small nucleus and each surrounded by a well-defined shrinkage 
cavity or lacuna in the matrix, usually much larger than itself. The relative proportion 
of cells and matrix varies in different parts, but generally the cells lie pretty close 
together. The outermost portion of the ectosome is occupied by the thick layer of oval 
microrhabds. 

This species in many respects resembles Lebwohl’s Yodomia yimai from Japan, but 
it differs in important details of spiculation. Thus the mesotrizenes of Y. yumaz have 
simple cladi while in Y. perfectus they are branched; Y. perfectus has no long-shafted 
triznes and Y. yimai appears to have none of the smooth oval microrhabds which are so 
abundant in our species. 

Register Nos., Locality, de. vi., 1X. 1, x. 1, 2, 3, 4, all from Saya de Malha, 4.9.05, 
C. 1, 150 fathoms. 


Family Stellettide. 


Astrotetraxonida with long-shafted triznes; without calthrops, sterrasters and 
aspidasters. (In a number of genera and species with reduced spiculation, constituting 
the so-called family Epipolaside, the tetract megascleres have completely disappeared, 
while in certain forms the astrose microscleres seem to have vanished.) 

The original Stellettids appear to have arisen by the development of long-shafted 
triznes and the radial arrangement of the megascleres in some primitive pachastrellid 
ancestor. It is a very remarkable and interesting fact that along a number of more or 
less distinct lines of descent within the stellettid family the power to produce tetract 
megascleres seems to have become exhausted and these lines have passed over into the 
monaxonellid condition. It was for such lines that Sollas [1888] proposed the family 
Epipolasidze, which he placed as an “Appendix” to his Astrophora Euastrosa (= Stellettidee). 
In the Epipolasidee he recognised three genera, Amphius, Asteropus and Coppatias. His 
diagnosis of the family fully recognises the principle of the loss of tetract megascleres, 
it runs as follows :—‘“ Euastrosa (?) without trizenes, possessing oxeas and one or more 
forms of aster. The oxeas arranged partly in radiating fibres, partly scattered loosely 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 235 


in the choanosome; in the ectosome they lie tangentially. The chamber system (so far 
as investigated) diplodal* ” (p. 177). 

Since 1888 a considerable number of species have been assigned to the “ Epipolasidze’ 
and it has become increasingly evident that the group is of polyphyletic origin. It is 
perhaps not even certain that all Epipolasids are reduced Stellettids, for it is quite con- 
ceivable that similar reduction may have taken place in primitive Pachastrellids and given 
rise to Epipolasids which never passed through a stellettid stage in their ancestry. Apart 
from this question, however, it is by no means difficult to find, amongst the known 
Stellettidee, genera, or even species, which seem to represent very closely the ancestors 
of certain Epipolasids. Sollas himself pointed out that “If Asteropus is a reduced 
- Stellettid it is to Stryphnus that we must look for its nearest alliance” (/.c. p. 206). 
In another part of the same work, however, he expresses the strong opinion that “ Asteropus 
has resulted from an Algol by the loss of trizenes”’ (p. exlii). 

In 1905 I pointed out that the epipolasid genera Coppatias (now sunk in Jaspis) . 
and Cryptotethya are evidently very closely related to Stelletta, and said that “Crypto- 
tethya may be regarded as derived from some such form as Séelletta herdmani by further 
reduction of the triznes and by the outgrowth of the ectosome into finger-like processes ” 
(p. 110). 

Most remarkable, however, is the existence of three epipolasid species, viz. 
Rhabdastrella distincta Thiele, Diastra sterrastrea Row and Aurora cribriporosa, n. sp. 
each of which is represented by a closely related, trizene-bearing species in the stellettid 
genus Aurora. The relationships of these species will be discussed later on. 

In view of these facts it seems to me that the time has now arrived when we may 
conveniently abandon the family Epipolasidze altogether and distribute its members as 
best we can amongst the Stellettidee, and, if subsequent research should render it 


necessary, amongst the Pachastrellidze also. 


b} 


Genus Myrtiastra Sollas [1886]. 


Stellettidee with or without a distinct fibrous cortex and with only one form of 
microsclere, a chiaster. 

This genus has been sunk by Lendenfeld [1903] in Stelletta, but it appears to me 
convenient to retain it, at any rate pending a much needed revision of the Stellettide. 
It seems impossible, however, to draw a real distinction between Myrvastra and Pilochrota, 
for all degrees of development of the fibrous cortex occur in different species of these 


genera. I therefore propose to merge Pilochrota in Myriastra. 


-4, Myriastra parva (Row). 
Pilochrota parva Row [1911]. 
This species is represented in the collection by a small fragment about 6 mm. in 


greatest diameter and of a pale, dull yellow colour. 


* On p. 141 he says “aphodal.” 
30—2 


236 PERCY SLADEN TRUST EXPEDITION 


The skeleton is dense, radially arranged, and the spiculation very typical, as 
follows :— 

(1) Orthotrizenes ; with simple unbranched cladi; shaft straight, tapering gradually 
to a sharp point, measuring about 0°76 by 0°026 mm.; cladi gradually sharp-pointed, 
measuring about 0°15 by 0-026 mm. 

(2) Anatriznes; numerous and frequently projecting beyond the surface. Cladi 
strongly recurved. Shaft very long and slender, measuring about 0:9 by 0°0086 mm. 
Cladi gradually sharp-pointed, about 0°034 mm. long. 

(3) Oxea; straight or nearly so, fairly gradually and sharply pointed, measuring 
up to about 0°9 by 0°02 mm. Considerably shorter and more slender forms also occur. 

(4) Chiasters (tylasters); very minute, with very slender rays and very small 
heads ; total diameter about 0°008 mm. Scarce. 

The cortex is very feebly developed and not sharply differentiated from the choano- 
some; say about 0°12mm. thick. It contains very little fibrous tissue and the large 
subcortical crypts push their way through it to within a short distance of the surface, 
lying between the distal portions of the bundles of large orthotrizenes, whose cladi are 
extended actually at the surface. The inhalant pores seem to open singly by short, 
narrow canals into the subcortical crypts. An inner zone of smaller orthotriznes extend 
their cladi beneath the subcortical crypts. 

The “‘Sealark” fragment agrees closely with the type of the species from the Red 
Sea, as described by Row. I have examined one of Mr Row’s preparations of the type 
and can find no important difference. I cannot find the slender, hair-lhke oxea which 
he describes and figures, but which I cannot regard as of any taxonomic importance. 

The species is evidently closely related to Sollas's Myriastra sumplicifurca [1888] 
from Torres Strait; differing, however, in the much smaller size of the spicules. It 
also comes near to Hentschel’s Stelletta tuberosa [1909] from S.W. Australia, from which 
it differs in the form of the cladome of the anatrizene, the cladi being, usually at any rate, 
much more strongly recurved. ‘Hentschel also mentions the occurrence of small, slender 
oxea here and there in the choanosome in his species. They are probably merely young 
individuals of the large oxea. Probably all three forms will have to be united as varieties 
of one and the same species, but it would be premature to do this at present. 

Previously known Distribution. Red Sea (Row). 

Register No., Locality, dc. Lv. 1, Coetivy. 


5. Myriastra cavernosa n. sp. 
(Plate 44, figs. 3, 3a; Plate 46, fig. 1.) 


Sponge (Plate 44, figs. 8, 3a) massive, irregularly subspherical; without definite 
points of attachment but more or less thickly encrusted with nullipores and Orbitolites. 
Surface uneven, granular, occasionally hispid where well protected. A few rounded 
openings, say about 3 mm. in diameter, irregularly scattered between the débris on 
the surface, and without prominent margins, probably represent the vents. They com- 
municate with the extensive system of wide canals which ramify all through the interior 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 237 


of the sponge and give it its characteristic cavernous appearance when cut open. In the 
outer part of the sponge these canals often approach very close to the surface, being 
covered in only by a thin membrane. ‘Texture compressible, resilient, fairly compact 
between the wide canals. Colour in spirit light yellowish grey. There are four specimens 
in the collection ; three of these have each a diameter of about 40 mm., while the fourth 
is much smaller. ; 

The skeleton, at any rate towards the surface, is radially arranged, consisting of 
large, stout oxea and orthotrienes grouped to some extent in loose, very ill-defined 
bundles. Most of the triznes are in the outermost portion and have their cladi 
extended at or very near the surface. 

Spicules :— (1) Orthotrizenes (Plate 46, fig. 1 a); with stout, straight or nearly straight 
shaft, usually tapering very gradually to a fine point. Cladi simple, stout, nearly 
straight ; extended nearly at right angles to the shaft but inclined slightly forward 
and then slightly recurved (may be slightly inclined forward again towards the apex). 
Shaft measuring up to about 2°0 by 0°066 mm., with cladi about 0:27 by 0:066 mm. 

(2) Oxea (fig. 1b); long, stout, fusiform, slightly and gently curved, usually gradually 
and sharply pointed at each end ; measuring up to about 2°6 by 0°07 mm. 

(3) Oxea (fig. 1c); short, slightly curved, fairly sharply pointed at each end; 
measuring about 0°155 by 0:0086 mm. A few of these occur scattered through the 
choanosome; they are possibly foreign. 

(4) Chiasters (fig. 1d); small, with slender, slightly tylote rays; total diameter 
about 0-013 mm. . 

There is a good deal of fibrous tissue in the thin ectosome (only about 0-085 mm. 
thick), but the ectosome is not very sharply differentiated from the underlying choano- 
some and it is impossible to speak of a distinct cortex. 

This species seems to differ from most species of Myriastra in the absence of the 
anatrizne, of which I have found no trace. Its curious cavernous character and general 
habit are also probably very distinctive. 

Register No., Locality, dc. vu. 5, Saya de Malha, 6.9.05, C. 15, 55 fathoms. Four 
specimens. 


Genus Dracmastra Sollas [1888] emend. 


Stellettidee in which the microscleres consist of euasters and trichodragmata. 

Sollas restricts this genus to corticate species in which the middle or collenchymatous 
layer of the cortex is crowded with trichodragmata. I have already pointed out that the 
degree of development of the cortex in the Stellettidze is so variable, and so many transi- 
tions occur, that it does not form a satisfactory generic character. I therefore omit all 
reference to the cortex from the diagnosis of this genus. 

Sollas [1888] expressly excluded Carter’s Stelletta lactea from the genus Dragmastra, 
on the ground that “the orthodragmas [= trichodragmata], as stated by Carter, are 
confined to the choanosome, and there is no necessity therefore to assign it to Drag- 
mastra, with which it is evidently not nearly related.” 

I cannot, myself, see why it is not nearly related to Dragmastra, and I find, in the 


238. PERCY SLADEN TRUST EXPEDITION 


variety about to be described, that the trichodragmata occur in the cortex as well as in 
the choanosome. I therefore think that Dragmastra is the genus to which Stelletta 
lactea must be assigned. 


6. Dragmastra lactea (Carter) var. mauritiana nov. 


(Plate 46, fig. 7.) 

Stelletta lactea Carter [1871 a]. 

Stelletta lactea Norman [Bowerbank 1882]. 

Pilochrota (?) lactea Sollas [1888]. 

Pilochrota lactea Topsent [1894 a]. 

Stelletta lactea Lendenfeld [1903]. 

Mr Carter originally described this species from the coast of Devonshire, and stated 
that it is “massive, spreading, fixed, following and filling the cavities of deciduous 
small boring shells (Saaicave) and Annelids, which confine themselves to the surface of 
the sandstone rock in which they live, almost entirely concealed by overgrowths of small 
Cirripedes and Fuci, and communicating with the exterior only through the openings 
of the cavities mentioned.” 

It is very interesting to find a closely related form, obviously a variety of the same 
mode of life at Mauritius. 


? 


species, adopting a similar “ cryptozoic’ 

The single specimen forms a thin crust, growing upon a horny sponge (R.N. cxxvi. 4) 
beneath a specimen of Latrunculia (R.N. oxxv1. 4c). The main skeleton consists of 
dichotrizenes and oxea; the dichotrizenes being mostly arranged in loose brushes with 
their cladomes just beneath the surface, while others are irregularly scattered in the 
deeper parts of the sponge. <A few of the oxea are radially arranged, but most of them 
seem to be irregularly scattered, singly or in bundles. , 

Spicules :—(1) Dichotrizenes (Plate 46, fig. 7 «); shaft short and stout, gradually 
sharp-pointed, characteristically bent somewhat to one side at about one quarter of its 
length below the cladome, measuring about 0°4 by 0°0258 mm ; cladi very short, each 
bifurcating into two short, sharply conical branches about equal in length to the main 
branch; total diameter of cladome about 0°1mm. <A few much more slender trizenes, with. 
unbranched cladi, also occur; these I take to be young forms of the dichotrizenes. 

(2) Oxea (fig. 7 6); nearly straight, fusiform ; gradually and fairly sharply pointed ; 
measuring about 0°75 by 0°02 mm., but frequently more slender. 

(8) Oxyspherasters (fig. 7c); very minute, sometimes with well-developed centrum 
and numerous very slender rays about as long as the diameter of the centrum, but the 
proportions are variable; total diameter about 0°008 mm. LEspecially abundant in a 
superficial layer. A few (fig. 7 d) occur as much as 0-016 mm. in diameter, usually 
with long, slender rays and relatively small centrum. 

(4) Trichodragmata (fig. 7); very numerous in some parts of the sponge, rare 
in others ; measuring about 0°02 by 0:004 mm. 

The small size and general condition of the sponge do not allow of my saying much 
about its minute anatomy or histology. There is no distinct fibrous cortex, but there 
appears to be a thick gelatinous ectosome (about 0°86 mm. thick), distinguished by its 
lighter colour and clearer appearance from the underlying choanosome. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 239 


This variety differs from the type of the species chiefly in the much smaller size of 

the megascleres and in the fact that all the trizenes when full-grown seem to be dichotrizenes. 
Previously known Distribution. Devonshire coast (Carter) ; French coast (Topsent). 
Register No., Locality, de. Oxxvi. 4 5, Mauritius, 23.8.05. 


Genus RHABDODRAGMA nh. gen. 


Stellettidee with (always) a very strongly developed, partly fibrous cortex. The 
microscleres include asters, microrhabds and trichodragmata. 

This genus stands in the same relation to Sollas’s Psammastra as that im which the 
same author's Dragmastra stands to Stelletta. Stelletta includes forms without micro- 
rhabds and without trichodragmata, while Dragmastra includes forms with trichodrag- 
mata but no microrhabds. Psammastra includes forms with microrhabds but without 
trichodragmata, and is perhaps indistinguishable from Hcionemia, while Rhabdodragma 
includes forms with both microrhabds and trichodragmata, in addition, of course, to the 
asters. 

Topsent’s genus Sanidastrella [1892 D] cannot, in my opinion, be distinguished from 
Psammastra, for the so-called sanidaster merges into the microrhabd type of spicule, as 
the figures given by Topsent [1894 G] clearly show. 

We have here a group of usually corticate Stellettidee which are evidently all closely 
related to one another but in which the microscleres show great variation from genus to 
genus. The presence or absence of such distinct types of microsclere as microrhabds and 
trichodragmata appears to me to afford good ground for generic distinction, and the same 
may be said of the characteristic spherasters of the genus Aurora and the sterrasters of 
the Geodiide and Erylide, but we cannot attribute a like value to the extremely 
variable oxyasters (and chiasters and other related forms), as Lendenfeld [1903] has 
done in attempting to differentiate subgenera of Stellettide. 

Kieschnick’s Psammastra conulosa from Ternate, first adequately described by 
Thiele [1900], is the type and so far only known species of the genus, and it is extremely 
interesting to meet with this little-known sponge again at Cargados Carajos. 


7. Rhabdodragma conulosa (Kieschnick). 
(Plate 44, fig. 7; Plate 47, fig. 1.) 


Psammasira conulosa Kieschnick [1896]. 
Psammastra conulosa Thiele [1900]. 


There are in the collection three specimens of this remarkable sponge, two 
large and one small, all from Cargados Carajos. The type specimens, from Ternate, 
were only about 1 cm. in diameter, but the smallest of the “Sealark” specimens has 
a diameter of about 2cm., while the largest has a diameter of about 6 cm., 
being nearly as large as a cricket ball. The external appearance (Plate 44, fig. 7) 


240 PERCY SLADEN TRUST EXPEDITION 


is very characteristic, the surface beg beset with irregularly arranged, more or less 
elongated conuli. These conuli are stiff and supported by spicule bundles. In addition 
to the conuli there are short, root-like attaching processes on the lower surface. Between 
the conuli the surface is in many places minutely reticulate, with pore-sieves in the meshes 
of the reticulation. There are no conspicuous vents, and it appears probable that the 
exhalant openings are covered over by sieve-nets. The smallest specimen is almost 
spherical, but the two larger ones are both slightly flattened dorsoventrally, and the 
largest (fig. 7) has a rather prominent equatorial ridge, dividing the more flattened upper 
from the more convex lower surface ; on this ridge the conuli are especially numerous. 
The colour in spirit is dark purplish-brown. All the specimens show clearly the small 
white specks due to the accumulation of trichodragmata just beneath the surface, as 
described by Thiele. The texture is firm and compact. 

The cortex, in the largest specimen, is about 3 mm. thick. It is divided into two 
very distinct layers, an outer, soft, pigmented one, to which the colour of the sponge is 
due, and an inner, very dense, fibrous one with only occasional pigment cells. The inner, 
fibrous layer is very much more ‘strongly developed than represented in Thiele’s figure, 
the difference being no doubt correlated with the much larger size and presumably greater 
age of the specimen. It makes up considerably more than half the thickness of the 
cortex and is composed of an extremely dense interlacement of fibre-tracts running in all 
directions. From its outer surface fibre-tracts run into the outer layer of the cortex, 
where they form a loose network, concentrated, however, towards the surface and around 
the radially arranged bundles of megascleres. In the meshes of this network lie the large 
vesicular pigment cells described by Thiele, the pigment being most strongly developed 
in the deeper part of the layer. The outer layer of the cortex also contains great rounded 
masses of trichodragmata. 

The inner layer of the cortex is pierced by narrow canals, which may branch and 
anastomose with one another and frequently unite to form wide chones, and whose course 
is clearly marked out by the microrhabds lying in their walls. In the outer layer of the 
cortex the canals are not easy to follow, but they appear to be still narrow and no doubt 
lead inwards from the dermal pores. 

Beneath the fibrous layer of the cortex there is a much thinner collenchymatous layer 
containing numerous subcortical crypts. This layer also contains a good many pigment 
cells. 

I am unable to say anything definite about the exhalant canal system, but there is 
good reason to believe that it is similar to that described by Topsent [1894 G] for his 
Sanidastrella coronata, a sponge which is obviously nearly related to our species. In 
that sponge the exhalant canals open at the sides of the much elongated dermal 
appendages (corresponding to the conuli of Rhabdodragma), not by large openings but 
through sieves. At the same time it seems probable that in our species the exhalant 
openings are not confined to the conull. 

The skeleton of the choanosome is very dense, consisting of pretty closely packed, 
radially arranged, large oxea and plagiotrizenes. Most of these terminate below the cortex, 
but here and there, at wide intervals, dense bundles of oxea and plagiotrizenes penetrate 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 241 


the cortex and either reach the surface of the sponge between the conuli or are continued 
into the conull. 

Spicules :—(1) Plagiotrizenes (Plate 47, fig. 1 a); with stout shaft and short, stout, 
often somewhat incurved cladi; shaft sharply and very gradually pointed, cladi sharp or 
blunt; shaft about 2°5 by 0:068 mm. ; cladi about 0:14 by 0°04 mm. 

(2) Oxea (fig. 1b); stout, fusiform, straight or nearly so, gradually and finely pointed 
at both ends or with ends somewhat blunted; measuring sometimes as much as 4 by 
0:07 mm. 

(3) Chiasters (fig. 1 ¢); with about eight rather slender, usually slightly roughened 
rays each ending in a small oval knob (I have seen one specimen with spiny instead of 
knobbed ends, as described and figured by Thiele); usually about 0°02 mm. in total 
diameter but sometimes nearly twice as much. 

(4) Slender oxea (fig. 1d); straight, gradually sharp pointed at each end, with very 
faintly roughened surface ; measuring about 0°266 by 0:005 mm. 

(5) Microrhabds (fig. 1e); short, thickly covered with small, short spines ; varying 
in diameter; say about 0-012 by 0°004 mm., including spines. Especially abundant in the 
dermal membrane. . ; 

(6) Trichodragmata (fig. 1f); each dragma about 0:03 by 0°012 mm.; readily 
breaking up into hair-like raphides. Enormously abundant and sometimes collected in 
large oval or spherical masses, especially in the outer part of the cortex. 

A careful comparison with Thiele’s excellent description and figures leaves no doubt 
as to the specific identity of the “Sealark” with the Ternate specimens. Yet Lendenfeld 
[1903] speaks of the asters as ‘‘oxyasters” (Thiele simply calls them asters), while in our 
specimens they are chiasters with knobbed ends, the difference between the two being 
extremely minute; and Thiele regards the spiny microrhabds as sanidasters! These facts 
show the utter hopelessness of basing generic distinctions upon oxyasters and sanidasters 
as distinct from chiasters and microrhabds. I think a comparison of my figures of the 
spicules with those given by Thiele will fully justify the identification. 

Previously known Distribution. Ternate (Thiele). 

Register Nos., Locality, &c. ULxxvitt. 1A, B, 3, Cargados Carajos, 28.3.05, B. 2, 

30 fathoms. 


Genus Ecronemra Bowerbank [1862 c]. 


Stellettidee in which the microscleres include microrhabds in addition to euasters ; 
the former are commonly minutely spined or roughened and usually form a dermal layer. 
There are no trichodragmata. 

I accept this genus in the same sense as that in which I employed it in my Ceylon 
Pearl Oyster Report [1905]. I cannot at present enter into the very difficult and complex 
question of its relationship to Ancorina and other stellettid genera and sub-genera, but 
I may say that neither the arrangement of Sollas [1888] nor that of Lendenfeld [1903] 
appears to me satisfactory. There can be no doubt that the species which I include in’ 
Ecionemia are closely related to H. acervus, Bowerbank’s type of the genus. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 31 


242 PERCY SLADEN TRUST EXPEDITION 


8. Ecwonemia carterr Dendy. 

Ecionema carteri: Dendy [1905]. 

This species is represented in the collection by a number of specimens, varying in — 
shape from almost spherical to irregularly massive, and in colour from light to dark brown. 
They agree very closely with the Ceylon form and I need add nothing to my original 
description except the statement that granular brown pigment cells occur in very varying 
numbers in the outer parts of the sponge, both in the Ceylon and im the “Sealark” 
specimens. 

Previously known Distribution. Ceylon (Dendy). 

Register Nos., Localities, dc. 11. 3, 6, Coetivy ; uxxi. 2, Amirante, 17.10. 05, 1Dy, aL. 
30 fathoms; LXXvitl. 18, Cargados Carajos, 28.3.05, B. 2, 30 fathoms. 


9. EHecvonemia laviniensis Dendy. 


(Plate 44, fiz. 6; Plate 46, fig. 5.) 


Ecionema laviniensis Dendy [1905]. 


There is a single specimen (Plate 44, fig. 6) of this species in the collection, con- 
siderably larger than the type; irregularly massive, potato-lke, in form, and measuring 
46 mm. in longer diameter. The surface is marked here and there by very irregular, 
strongly hispid grooves, some of which contain openings which may be vents. Elsewhere 
the surface is rather uneven, granular and minutely porous. Only a small quantity of 
calcareous débris is attached to the surface. The texture is hard, incompressible. The 
colour in spirit is light purplish brown. 

In spiculation this specimen agrees closely with the type, but the cladi of the . 
dichotrizenes (Plate 46, figs. 5 a, 5 a’) are stouter, while the microstrongyla (fig. 5 g) appear 
on an average to be somewhat smaller. Minute chiasters with stout cylindrical rays 
(fig. 5d) and small, slender-rayed oayersiiang (fig. 5e) are both present, together with 
intermediate forms (figs. 5e, 5/). | 

Previously known Distribution. Ceylon (Dendy). 

Register No., Locality, &c. xt. 2, Saya de Malha, 7.9.05, C. 19, 29 fathoms. 


Genus Aurora Sollas [1888]. 


Stellettidee in which the principal microscleres are large spherasters (or sterro- 
spherasters), accumulated especially in a cortical layer. 

This genus was proposed by Sollas [1888] for the sac Sam of Carter’s Stelletta 
globostellata and S. reticulata. Lendenfeld [1903] has again merged it in Stelletta, and 
in this respect he has been followed by Hentschel [1909], but the large spheraster (or 
sterrospheraster) forms such a characteristic and well-defined feature, and so many species 
are now known, that it seems to me desirable to retain Sollas’s genus. To Carter's species 
must be added, as typical members of the genus, Hentschel’s Stelletta awrora [1909] and 
Isops membranacea [1909], and two new ones to be described presently under the names, 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 243 


Aurora providentie and A. rowi. In addition I propose to include in this genus three 
reduced or ‘epipolasid’ species, viz. Coppatias (Rhabdastrella) distinctus Thiele [1900], 
Diastra sterrastrea Row [1911] and Aurora cribriporosa n. sp. 

Aurora globostellata remains the type of the genus, and I therefore call special 
attention to the fact mentioned later on (p. 247), that it probably does not possess tricho- 
dragmata*, as described by Sollas. 

The genus is of great phylogenetic interest as representing almost certainly the 
starting point for the evolution of the Geodiide, as well as for other reasons. To work 
out this problem as fully as it deserves would require more time and material than I at 
present have at my disposal, but the following considerations appear to me to leave little 
doubt as to the results which such an investigation would yield. 

The Geodiide are, as is well known, distinguished by the possession of a very 
peculiarly modified spheraster, known as the sterraster, and these spicules form a dense 
cortical crust. In the more typical forms of Aurora, such as A. globostellata and A. prow- 
dentia, the characteristic microsclere is a large spheraster with conical rays, very like that 
of Donatia or Chondrilla, and these also are arranged in a cortical layer. Now in certain 
species of Aurora, such as A. membranacea (Hentschel), A. sterrastrea (Row) and A. rowi, 
n. sp., the typical spheraster is either associated with or replaced by a peculiarly modified 
_spheraster resembling a sterraster, which I propose to call a sterrospheraster (Plate 46, 
fig. 4c). What appears to be this type of spicule was indeed actually described as a 
sterraster by Hentschel [1909] in his Isops (Aurora) membranacea. The same type of 
spicule was described by Row [1911] in his Diustra sterrastrea, which may be looked 
upon as an Aurora with reduced spiculation. Row pointed out the resemblance which it — 
bears to a sterraster. He also noted the absence ofa “hilum,” and it is possible that this 
may prove to be a distinctive feature of the sterrospheraster, though I think it hardly 
likely. 

Row gave some account of the development of the sterrospheraster in Diastra, and 
I have been able to work it out somewhat more fully in the case of Aurora rowi (vide 
mfrd and Plate 46, fig. 4). It certainly resembles pretty closely the development of a 
typical geodiid sterraster, but at the same time it passes through a stage im which it is 
a fairly typical spheraster, identical, in fact, with the spheraster of Awrora aurora, which 
has no sterrospherasters. There can be no doubt that the typical spheraster of Aurora, the 
sterrospheraster, and the sterraster of Geodia, are all closely related spicule-forms, the 
sterrospheraster being in some respects intermediate between the other two. At the same 
time we must not forget that a ‘“sterrospira,” practically; indistinguishable in the adult 
condition from the sterraster of the Geodiide, has arisen independently in the spirastrellid 
genus Placospongia, as shown by Vosmaer and Vernhout [1902 |. 

' The sterrospheraster, however, actually occurs in certain undoubted Geodiide, as, for 
example, Geodia carter Sollas [1888, p. 247], associated with true sterrasters. Carter, 
who originally described that species from the south coast of Australia under the name 
Geodia canaliculata, Sdt., and figured the spicules [1883 B], regarded the sterrospheraster 


* Sollas [1888] uses the term “orthodragma,” but the spicules in question are identical with the 
spicules described by Ridley and Dendy [1887] as “trichodragmata.” 


31—2 


244 PERCY SLADEN TRUST EXPEDITION 


as an abnormal form of spicule*. That this is not the case I have convinced myself by 
examination of Mr Carter’s preparations now in my possession. Schmidt [1868] certainly 
figured a very similar spicule in his Geodia canaliculata, but it is not clear, from the 
descriptions given by him and by Sollas [1888], whether or not it is associated in this case 
with a typical sterraster, 2.e. whether the species is a Geodia or an Aurora. 

It is obvious from what has been said that the genus Aurora bridges over to a very 
large extent the gap between the Stellettide and Geodiide, and the fact that Hentschel 
described as a geodiid (Isops) a species which I feel constrained to place in the genus 
Aurora, affords eloquent testimony to the close relationship of the two families}. If asked 
exactly where we ought to draw the line between the two, I should say that further 
minute anatomical investigation is needed before the question can be answered. Pro- 
visionally we may take the typical, hilum-bearing sterraster as a distinctive feature of 
the Geodiide. The sterrospheraster occurs in both families and cannot be regarded as 
distinctive of either; indeed we cannot even regard it as affording the basis of a generic — 
separation from Aurora, because of its close relationship to the typical spheraster, the 
gap between the two being completely bridged over by the adult spheraster of Aurora 
aurora and A. reticulata and the developmental stages of the sterrospheraster. There is 
just the possibility that the spheraster of Awrora aurora is not a true spheraster but 
represents a case of convergence, and that true spherasters do not develop in the same 
manner as sterrospherasters, but this does not seem very likely, and in any case more 
information is wanted before we can settle the point. 

The close relationship between the Geodiidz and Stellettidee was recognised many 
years ago by Czerniavsky [1879], who proposed the sub-genus Stello-geodia for a species 
(Geodia stellosa) which he regarded as intermediate between the two, with the following 


diagnosis :—‘‘Membrana sarcodea superticialis, corticem tegens, stellulas minimas 
numerosas breviradiatas continens. Parenchyma preeter globulos siliceos stellas majores 
numerosas pauciradiatas continet.” The author evidently regarded the presence of 


? 


euasters in addition to the “siliceous globules” as the distinguishing feature of his 
* sub-genus, but such euasters as he figures of course occur abundantly in the genus 
Geodia itself and are by no means distimctive. On the other hand, it seems probable 


? 


from his figures and description that the “siliceous globules” are not true sterrasters, 
and that the sub-genus may be identical with Aurora. ‘The description and figures, 
however, are not sufficiently accurate to enable me to decide this point, and it hardly 
seems necessary to abandon the generic name Aurora in favour of Stello-geodia, especially 
as the type species of the latter appears to contain tylostyles, which are not met with in 
Aurora, though possibly these spicules are only abnormal forms of the oxea. 

As already indicated above, the transition from the tetractinellid to the epipolasid 
condition by loss of the trizne megascleres appears to have taken place at least three 
times in the genus Aurora. At any rate there are three epipolasid species each of which 


* This is also the view taken by Lendenfeld [1910] of the spicules termed by him “sterroids” (e.g. in 
Geodia variospiculosa), which seem to be identical with the sterrospherasters. 

+ It is not impossible that several other species at present included in the Geodiide, such, for example, 
as Hentschel’s Geodia punctata [1909], may be shown by future research to be Auroras. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 245 


is evidently closely related to a corresponding species of Aurora which still retains its 
trienes. Aurora (Rhabdastrella) distincta (Thiele), with large smooth spherasters but 
without trizenes, is closely related to Awrora globostellata (Carter) with large smooth 
spherasters and triznes. Aurora cribriporosa, n. sp., with large rough spherasters but 
without trizenes, is closely related to Aurora providentia, n. sp., with large rough spher- 
asters and triznes ; and Aurora (Diastra) sterrastrea (Row), with sterrospherasters but 
without triznes, is closely related to Aurora rowi, n. sp., with sterrospherasters and 
trieenes. 

Aurora (Rhabdastrella) distincta was first described by Thiele [1900] as a Coppatias, 
but he subsequently [1903] separated it from Coppatias under the new generic name 
_Rhabdastrella. 

Aurora sterrastrea was described by Row in 1911 and also recognised as an 
epipolasid, for which the new genus Diastra was instituted. If these genera were to 
be retained it would, I think, be necessary also to propose a new genus for Awrora 
eribriporosa, but all three genera would have to be based upon very inadequate characters. 
We cannot include all three species in a single genus distinct from Aurora unless we are 
willing to retain that single genus merely as an artificial one of polyphyletic origin, and 
any argument in favour of doing this would apply equally to the case of the so-called 
family Epipolaside. There seems nothing for it, then, but to abandon the genera 
Rhabdastrella and Diastra and place all three species in Aurora. It may be said that 
we ought also to abandon the genera Amphius, Asteropus and Jaspis (Coppatias), and 
this I shall be quite prepared to do when I feel as certain about their origin from 
particular tetractinellid genera as I do in the case of Diastra and Rhabdastrella. 

In this connection it is also necessary to say something about Sollas’s genus Magog 
[1888], which was proposed for the reception of Carter’s Chondrilla sacciformis [1879 B]. 
This species was described by Carter as having “acerates” (oxea) and “ globostellates” 
(spherasters). Sollas accepted this as correct after examination of one of Carter's slides, 
and diagnosed the genus Magog thus :—‘“Tethyide in which the rhabdus spicule is an 
oxea, which is confined to the choanosome.” 

Were the spiculation of Chondrilla sacciformis really as described I think we should be 
obliged to accept that species also as an epipolasid Aurora. J have in my possession, how- 
ever, in Mr Carter’s cabinet, two microscopical preparations of his Chondrilla sacciformis, 
one consisting of teased fragments mounted in balsam and the other of a number of fair-sized 
fragments put up dry in a cell. I have examined sections and boiled-out spicules from 
one of these fragments, but neither in these preparations nor in Carter’s own balsam slide 
can I find any oxea at all. The species is a genuine Chondrilla, identical with Thiele’s 
Chondrilla grandistellata [1900]. As to how the mistake arose I can only surmise that 
the preparation from which Carter’s original description was taken, and which is now 
presumably lost, must have contained, as accidental inclusions, oxeote megascleres of 
Rhaphidhistia spectabilis, which Carter described as a thinly encrusting sponge growing 
on the same mass as Chondrilla sacciformis. That the preparations now in Mr Carter’s 
cabinet, and labelled by himself “ Chondrilla sacciformis,” were made at a later date than 
the original description seems certain, as they bear the date 1881. That they were taken 


246 PERCY SLADEN TRUST EXPEDITION 


from one or more of the original types (there were several specimens on the same mass) 
seems also certain, for one of the slides is labelled ‘‘ Bk. Coll. 701. Mauritius,” and this 
number and locality are quoted in the original description. Moreover the size and form 
of the large spherasters are extremely characteristic (vide infrd, p. 269 and Pl. 48, fig. 8). 

The genus Magog therefore must fall to the ground, (1) as being based upon a serious 
misconception as to the characters of the type species, which is a genuine Chondrilla, 
and (2) because even were the type species as represented it would have to be regarded 
merely as a reduced Aurora. 

It seems quite probable that the genus Chondrilla itself has been derived from 
Aurora by complete loss of ald the megascleres, and not only of the trizenes. 


10. Aurora providentie n. sp. 
(Pl. 46, fig. 2.) 


This species is represented by a thin crust, about 25 mm. in greatest breadth, which 
has apparently been pared off some other object, or possibly sliced off from the surface of 
a larger specimen of the same species, but if the latter hypothesis be correct it is remark- 
able that the specimen in question is missing from the collection*. The shape of the 
fragment is quite irregular. The upper surface is fairly smooth and shows two sieve-like 
groups of openings, of very varying size, which are evidently oscula. There are a large 
number of small calcareous foreign bodies scattered over the surface. Colour in spirit 
pale, dull yellow. Texture cavernous. 

There is a thin, fibrous cortex, about 0°1 mm. thick, containing numerous large and 
small spherasters in its outer portion, and pierced at wide intervals by inhalant chones. 
Each chone is divided into ectochone and endochone by a thin diaphragm perforated by 
a pore. The ectochone terminates externally in a single (?) dermal pore, the endochone 
merges indistinguishably into a subcortical crypt. : 

In the deeper part of the sponge the skeleton consists of loose bundles of oxea 
running towards the surface, with many loose oxea scattered between. As these bundles 
approach the surface they spread out into brushes composed mainly of slender oxea and 
orthotrizenes, the cladi of the latter being extended in or beneath the fibrous cortex, 
below the layer of large spherasters. 

Spicules -—(1) Orthotrizenes (Plate 46, fig. 2a). Shaft varying from gradually and 
finely pointed to rounded or even knobbed at the extremity. Cladi usually simple, 
gradually and sharply pointed ; rarely bifid for a short distance at the extremity. Shaft 
measuring about 0°5 by 0-017 mm., but variable ; cladi about 0°14 by 0°013 mm., but also 
variable. 

(2) Anatrizenes (figs. 26, 20’, 26”). Minute, with long, hair-like shaft and widely 
extended cladi curving backwards almost on ares of a sphere. Shaft about 0°37 by 
0001 mm. Cladome about 0:0164 mm. across from tip to tip of cladi; cladi about 


* The fact that Aurora (Isops) membranacea [Hentschel 1909] forms a thin crust suggests that A. pro- 
videntie not improbably has a similar form. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 247 


0-001 mm. in diameter at base, but may be reduced to a mere knob (fig. 2b”). These 
spicules are very rare in the boiled out preparations, but paraffin sections show them in 
considerable numbers piercing the cortex and usually projecting freely from the surface, 
where their cladomes are generally broken off. 

(3) Oxea (fig. 2¢, 2c’). Very slightly curved, usually gradually and finely saat 
at each end; measuring up to about 0°84 by 0:02 mm., but varying a good deal, both in 
length and diameter, and often very slender. 

(4) Large spherasters (fig. 2d); with large centrum and generally sharp-pointed, 
conical rays, usually more or less roughened on their sides, especially towards the apex ; 
total diameter about 0°04 mm., with rays about 0:008 mm. long. Very numerous, 
especially in the cortex. Occasionally the rays are all blunted and roughened at the 
ends, very much as in Hentschel’s Aurora (Stelletta) aurora [1909]. 

(5) Small spherasters and oxyasters (fig. 2¢); total diameter about 0:008 to 
0:02 mm., variable. Some of these are probably early stages of the large spheraster, 
but I doubt if this is the case with all. 

(6) Oxyasters (fig. 2f); rather large, with straight, moderately stout rays, about 
6 or 8 in number and usually more or less roughened. Sometimes there is a small but 
distinct centrum. Total diameter of spicule about 0°07 mm. 

This species is evidently related not distantly to Carter's Aurora (Stelletta) globo- 
stellata from Ceylon [18838], which has never been satisfactorily described. I have two 
_ preparations in Mr Carter’s cabinet, evidently made from the type, ‘though labelled 
“ Stelletta globostellifera.” From an examination of these I find that the principal points 
in which Carter’s species differs from Aurora providentia are as follows:—(1) The large 
spheraster has a somewhat smaller centrum and the rays are rather longer, smooth, and 
often blunted at the ends; (2) The large oxyaster is represented by a smaller spicule with 
long and very slender rays (apparently the “chiaster” of Sollas’s description [1888] but 
reaching a total diameter of 0°045 mm., whereas Sollas gives 0°015 mm.). I have found 
no anatrizenes, but these are so minute and the cladomes are so easily broken off in the 
“Sealark” species that I cannot attribute any great importance to their apparent absence 
in A. globostellata. Sollas gives an “ orthodragma” as a constituent of the spiculation in 
A. globostellata, but I can find none in Carter’s slides nor does Carter himself mention it, 
so we must suppose that Sollas has made a mistake in this respect. He does not say how 
he obtained his information. é 

It is also evident that our species is related to Hentschel’s Aurora (Stelletta) aurora, 
from §.W. Australia, but the details of spiculation afford quite sufficient differences to 
separate the two. 

Register No., Locality, dc. LxXxxvi. 2, Providence, 4.10.05, D. 4, 50—78 fathoms. 


11. Aurora cribriporosa n. sp. 


(Plate 44, fig. 4; Plate 46, fig. 3.) 


The single specimen (Plate 44, fig. 4) has a broad, oval base of attachment, from 
which it rises up on all sides like a hillock, culminating in a somewhat excentrically 


248 PERCY SLADEN TRUST EXPEDITION 


placed, rounded apex bearing a group of between twenty and thirty minute, sphinctrate 
vents of varying sizes. The surface is smooth and only slightly uneven; minutely 
punctate under a pocket lens owing to the thickly and for the most part uniformly 
scattered pore-sieves. The texture is rather leathery owing to the well-developed cortex, 
internally somewhat cavernous owing to the more or less radially arranged canals. Colour, 
throughout, brown, not very dark (in spirit). A small amount of calcareous débris 1s 
attached to the surface but there seems to be little or none internally. The longer 
diameter of the base measures 34 mm., the shorter diameter 17 mm., and the maximum 
height from base to apex about 17 mm. 

The cortex is fibrous and about 0°35 mm. thick. It is much excavated, however, 
by the large inhalant chones, each divided into ectochone and endochone. The ectochone 
is conical in shape, with the broad base turned outwards and formed by the thin sieve- 
membrane pierced by numerous inhalant pores. The ectochone is separated from the 
endochone by a moderately thick diaphragm of fibrous tissue pierced by a single pore 
which occupies the apex of the cone. The endochones merge insensibly into the large, 
irregular, subcortical crypts, which unite together in groups and give rise to the wide 
inhalant canals which penetrate to the interior of the sponge. 

The mesogloea of the choanosome is finely granular, penetrated by numerous narrow, 
branching canaliculi. The chamber system cannot be satisfactorily made out in my 
preparations. 

The main skeleton is an irregular feltwork of moderate-sized oxea. These are - 
present abundantly in the fibrous cortex, as well as in the choanosome. Indeed, they 
almost seem to form a special cortical layer, being arranged paratangentially, while 
beneath the cortex there is a tendency towards radial arrangement. The radial ones 
also frequently penetrate the cortex with their outer ends. There are also in the cortex 
numerous very much smaller, slender oxea, arranged radially and with their outer ends, 
now mostly broken off, projecting slightly beyond the surface. 

The outermost part of the cortex and the thin, pore-bearing membrane which covers 
over the inhalant chones are densely charged with minute spherasters ; beneath this layer 
lie numerous large spherasters, practically confined to the outer half of the cortex, where 
the fibrous tissue is less strongly developed. Slender-rayed oxyasters, together with 
a few of the other kinds, occur scattered through the choanosome. 

Spicules -—(1) Oxea (Plate 46, fig. 3a); very slightly curved, fusiform, usually 
gradually and finely pointed at each end, occasionally blunted or even becoming stylote, 
and occasionally with short abnormal branches. Measuring up to about 0°7 mm. in length. 
Thickness very variable, up to about 0°025 mm. but usually more slender. 

(2) Slender cortical oxea; slightly curved ; ends varying from sharply and gradually 
pointed to rounded off Size about 0-2 by 0:004 mm. 

(3) Large spherasters (fig. 3b); with large centrum and conical rays beset with 
short spines. Total diameter about 0°0287 mm. 

(4) Spherasters (fig. 3c); with small centrum and numerous long, slender rays ; 
diameter about 0°016 mm. Fairly common in the cortex; possibly young forms of the 
large spheraster. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 249 


(5) Minute spherasters (fig. 3d); with moderately large centrum and numerous 
short, strongylote rays; total diameter about 0°008 mm. 

(6) Oxyasters (fig. 3¢); with no centrum and about 6 or 7 long, slender, slightly 
roughened rays, usually strongylote. Total diameter about 0:04 mm. Very abundant 
in the choanosome. 

This species must, as already stated, be regarded as derived from some form closely 
related to Aurora providentie by loss of the triznes. It is also nearly related to Aurora 
(Rhabdastrella) distincta Thiele [1900] from Ternate, differing mainly in its rough spher- 
asters. 

Register No., Locality, ée. cx1x. 12, Salomon. 


12. Aurora rowi n. sp. 
(Plate 44, fig. 5; Plate 46, fig. 4.) 


The single specimen (Plate 44, fig. 5) is an irregularly rounded, massively lobose 
sponge, attached to a calcareous nodule very much smaller than itself. (There are 
indications that at least one other large foreign body has been removed from the surface.) 
The specimen measures about 36 mm. in height, 41 mm. in breadth, and has a very 
varying thickness up to about 25mm. The surface is subglabrous ; minutely punctate 
under a pocket lens, and has a curious crumpled appearance, with irregular meandering 
grooves of varying depth. The oscula are of moderate but variable size, measured up 
to 2mm. in diameter; few in number, scattered singly; without prominent margins. 
The inhalant pores are scattered singly on the surface of the sponge, each in the middle 
of a small polygonal area. 

The colour in spirit is a uniform chocolate brown throughout. The texture is 
compressible but resilient; fairly compact, but with a tendency towards the inclusion 
of rather large foreign bodies. 

The skeleton is not very dense and somewhat confused, but on the whole shows 
a marked radial arrangement, consisting principally of very loose wisps of large oxea 
running towards the surface. In the more superficial part of the sponge occur numerous 
radially arranged orthotrienes with cladi extended beneath a dermal crust of sterro- 
spherasters and small spherasters. 

Spicules :—(1) Orthotrizenes (Plate 46, fig. 4a); shaft well developed, straight, 
gradually sharp-pointed, measuring about 0°74 by 0°023 mm. Cladi unbranched, shghtly 
recurved, sharply pointed, about 0°16 mm. long. 

(2) Oxea (fig. 4b); rather slender, straight or very slightly curved; gradually and 
sharply pointed at each end, measuring about 1:1 by 0°023 mm. 

(3) Sterrospherasters (figs. 4c—4h). The full-grown spicule (fig. 4c) is spherical, 
with a very large, solid centrum whose surface is covered with close-set, irregular, 
flattened protuberances, incompletely separated from one another by narrow grooves 
which form a reticulate pattern. The diameter of the spicule is about 0:04 mm. The 
development of this spicule is very interesting. ‘The first stage (fig. 4d) is a slender- 
rayed oxyaster, but apparently the rays may sometimes be truncated (fig. 4d’). A distinct 

SECOND SERIES—ZOOLOGY, VOL, XVII, 32 


250 PERCY SLADEN TRUST EXPEDITION 


centrum is then developed between the basal portions of the rays, and the whole spicule 
at the same time increases in size (figs. 4¢, 4e’). As the centrum grows larger the rays 
grow longer and increase in number. They may still be either sharp-pointed (fig. 4) or 
truncated (fig. 4f’), but remain slender, so that their bases stand far apart from one 
another on the surface of the spherical centrum. As growth proceeds the rays begin to 
thicken and become bluntly conical. The radius of the centrum is now growing more 
rapidly than the length of the rays, so that the intervals between the latter become to 
a large extent filled up, the ends of the rays forming blunt conical projections on the 
surface of a sphere (fig. 4g). Short spines are now developed at the ends of the rays 
(fig. 4h). The spheraster has now reached what appears to be the adult condition in 
Aurora (Stelletta) aurora Hentschel [1909]*. The adult condition in our species (fig. 4 ¢) 
appears to be arrived at by further development of the terminal spines and by their 
fusion with one another to form the enlarged extremities of the rays, and with those of 
adjacent rays to form bridge-like connections between these extremities. There seems to 
be little doubt that we have here a very interesting case of the recapitulation of phylo- 
genetic history, the penultimate stage in the development of the sterrospheraster, with 
terminally spined rays, representing an ancestral condition which is still retained as the 
adult form in A. aurora. The oxyaster with which the development commences, and 
the various intermediate stages, no doubt also represents ancestral adult forms. 

The sterrospheraster and its developmental stages are abundantly scattered through 
the choanosome. The adult form is especially abundant in the cortex. 

(4) Small, irregular spherasters without any distinct rays but merely a nubbly 
surface (fig. 47). Varying in diameter up to about 0°0082 mm., but usually a good deal 
smaller. These spicules look as if they might owe their origin to the separation of the 
enlarged extremities of the rays of the sterrospherasters. They resemble these extremities 
closely in shape, and usually also in size, though some of them are larger. I have, 
however, found no evidence of such an origin, and they are probably independent spicules, 
homologous with the strongylasters of Aurora aurora. They are scattered abundantly 
throughout the sponge, but are especially numerous in the cortex. 

(5) Oxyasters (fig. 44); minute, slender-rayed, with no distinct centrum and about 
5 or 6 rays; total diameter about 0°0165 mm. It is very doubtful whether these can 
be sharply distinguished from the early stages of the sterrospheraster. 

The condition of the specimen does not permit of my giving a detailed account of the 
canal-system and histology, but the following particulars may be noted. There is a thin, 
fibrous cortex, about 0°2 mm. thick, densely packed, especially in its outer part, with 
sterrospherasters and spherasters. This cortex is pierced at intervals by chones. Each 
chone is divided into ectochone and endochone and the two are connected by a very 
narrow and fairly long canal. The ectochone terminates at the surface of the sponge in a 
single inhalant pore. The endochone opens into a subcortical crypt, from which, indeed, 
it is not sharply distinguishable, and the subcortical crypts unite together to form large 
irregular spaces from which the incurrent canals take their origin. The entire arrangement 
is very similar to that figured by Keller for Stelletta siemensi [1891, fig. 56]. 


* Compare especially Hentschel’s figure of 4. awrora var. arenosa (loc. cit., p. 363, fig. 7). 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 251 


The choanosome contains numerous large, oval, thin-walled, vesicular pigment-cells, 
about 0°04 mm. in longer diameter and containing a granular reticulum of brown or yellow 
pigment to which the colour of the sponge seems to be chiefly due. 

This is an extraordinarily interesting species, evidently closely related to Aurora 
(Stelletta) awrora, Hentschel, Aurora (Isops) membranacea (Hentschel) and Aurora 
(Diastra) sterrastrea Row. The sterrospherasters are identical with those of Row’s 
species, only somewhat larger*. In A. sterrastrea, however, there are no triznes, and 
Row therefore very naturally included that species amongst the so-called Epipolaside. 
Aurora vow: undoubtedly represents very closely the actual stellettid species from which 
Aurora (Diastra) sterrastrea was derived by loss of trizenes. The resemblance between 
the two is remarkably close, even to the characteristic chocolate-brown colour. Unfortu- 
nately Row’s description of his species is faulty in certain respects. His account of the 
cortex is very misleading. He has evidently included as part of the cortex a considerable 
portion of the choanosome. The special layer of oxea which he describes in the supposed 
deeper parts of the cortex, 0°5 mm. from the surface of the sponge, does not belong to the 
cortex at all; moreover it is not a properly defined layer but part of a confused general 
skeleton. The mistake seems to have arisen through the lack of stained paraffin sections. 

Register No., Locality, dc. cxxxt1., Seychelles, F. 9, 37 fathoms. 


Genus AstTEROPUS Sollas [1888]. 


Reduced Stellettidee which have completely lost their tetractinellid megascleres. The 
main skeleton is a confused feltwork of large oxea. The microscleres are oxyasters and 
sanidasters. 

I have already pointed out that Sollas regarded this genus as probably being derived 
from either Stryphnus or Algol by loss of trizenes.. 


13. Asteropus simplex (Carter). 


(Plate 46, fig. 6.) 


Stellettinopsis simplex Carter [1879 8]. 

Asteropus simplex Sollas [1888]. 

Asteropus haeckeli Dendy [1905]. 

Asteropus simplex Hentschel [1909]. 

Asteropus simplex Dendy [1915]. 

The single specimen in the collection is a cake-shaped mass about 90 mm. in maximum 
diameter, flattened below and convex above. A few mammiform projections scattered 
here and there on the upper surface are penetrated each by several narrow oscular tubes 
and bear at their summits the rather small vents, most of which appear to be closed. 
The surface is uneven and encrusted by a good deal of calcareous débris, together with a 
species of Gelliodes. Texture firm, compact, coarse. Colour in spirit rather dark, purplish 
brown. 

* Row gives the diameter of the full-grown sterrospheraster in D. sterrastrea as 0°36 mm. This is 
obviously a mistake. I find it to be 0-028 mm. by actual measurement from the type. 


32—2 


252 PERCY SLADEN TRUST EXPEDITION 


The main skeleton is a confused, dense feltwork of large oxea, with numerous 
oxyasters and sanidasters scattered between. 

Spicules :—(1) Oxea (Plate 46, fig. 6 a); fusiform, more or less curved ; for the most 
part gradually and sharply pointed, but may be rounded off at one or both ends, 2.e. stylote 
(fig. 6 a’) or strongylote. Size up to about 2°1 by 0:065 mm. 

(2) Oxyasters (fig. 6b); rays slender, sharply pointed, slightly roughened, usually 
about 10 in number, often springing from a small but distinct centrum ; total diameter of 
spicule about 0°05 mm. 

(3) Sanidasters (fig. 6¢); with approximately straight axis, usually bifurcate or 
trifurcate at the ends, and with a number of rather short, strongylote spines coming off 
irregularly along its length or in two principal whorls; total length about 0°02 mm. 

In deseribmg my Asteropus haeckeli from Ceylon in 1905 I pointed out its close 
relationship to Carter’s Stedlettinopsis simplex from Western Australia, and suggested that 
the future discovery of intermediate forms might justify us in uniting the two. This 
union has since been effected by Hentschel [1909] on the basis of Australian material 
collected by the Hamburg South-West Australian Expedition. I have no doubt that he 
is right, nor have I any doubt that the ‘“‘Sealark” specimen falls within the same species. 

Mr Carter, when he first described the species in 1879 referred to a specimen from 
Hayti which he believed to belong to the same species, but which differs in small details 
of spiculation. In 1905 I expressed doubt as to this identification, but a careful examina- 
tion of Carter's original preparation, now in my possession, convinces me that he was right, 
and that the differences in spiculation, such as the very slightly inflated ends of the rays 
of the aster, are not of specific value. 

Previously known Distribution. Fremantle, Western Australia (Carter); Port 
Philip Heads, Victoria (Carter 1886); South-West Australia (Hentschel); Ceylon 
(Dendy); Okhamandal (Dendy) ; Hayti (Carter). 

Register No., Locality, &c. Lxxvut. 2, Cargados Carajos, 28.3.05, B. 2, 30 fathoms. 


Genus Jaspis Gray [1867 F]. 


Stellettidze (7). with oxeote megascleres irregularly interlaced to form a confused 
skeleton, and with microscleres in the form of euasters (and perhaps microxea which cannot 
be sharply distinguished from the megascleres). 

This genus was proposed by Gray [1867 Fr] for the reception of Schmidt’s Vioa john- 
stonit, with the diagnosis ‘‘Spicules of two kinds:—1. Fusiform. 2. Stellate.” 

Sollas [1888] proposed the genus Astropeplus for the reception of his Astropeplus 
pulcher. Lendenfeld [1896] showed that Astropeplus pulcher Sollas is synonymous with 
Viow johnston Schmidt, but unfortunately placed that species in the genus Xenospongia, 
which had been proposed by Gray [1858] for a totally different sponge. Topsent [1898 B] 
pointed out that Vioa johnstonii cannot be a Xenospongia and relegated it to Sollas’s 
genus Coppatias, which had been defined by Sollas [1888] as comprising “ Epipolasidee in 
which but one form of aster, and that a euaster, is present.” Sollas’s Astropeplus thus 
became for Topsent a synonym of the same author’s Coppatias. In his Monograph on the 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 253 


Sponges of France [1900] Topsent fully discussed the synonymy of Vioa johnstonit (up to 
date) and decided upon the name Coppatias Johnstoni, rejecting Gray’s generic name 
Jaspis on the ground that the diagnosis of that genus had no scientific value and could 
not be accepted. A few years later, however, he had changed his opinion, and in his work 
on the Sponges of the Azores [1904 4] he accepted Gray’s name. 

If we are to accept others of Gray’s generic names, proposed in the same paper as 
Jaspis [1867 F], as has lately been done in the case, for example, of Mycale, I certainly do 
not see how we can refuse to accept Jaspis ; indeed, according to the laws of nomenclature, 
I believe we are bound to accept it, for not only did Gray give a diagnosis but he 
mentioned a type, and there can be no possible doubt as to what he meant. 

The character of the oxea, and especially of the centrotylote microxea of Jaspis 
johnston, suggests that this genus may have originated directly from some pachastrellid 
ancestor without having passed through a stellettid stage, but until we have more 
evidence that this is the case it seems better to include it with the other “ epipolasid ” 
genera in the Stellettide. 


14. Jaspis johnston (Schmidt). 
(Plate 47, fig. 2.) 


Vioa Johnstonii Schmidt [1862]. 

Jaspis Johnstonia Gray [1867 F]. 

Viow Johnstonii var. Schmidt [1868]. 
Viow Schmidtit Carter [1882 a]. 
Astropeplus pulcher Sollas [1888]. 
Dorypleres incrustans Topsent [1892 c]. 
Coppatias inconditus Topsent [1892 p]. 
Xenospongia johnstonii Lendenfeld [1896]. 
Asteropus incrustans Lendenfeld [1896]. 
Coppatias Johnstonz Topsent [1898 8]. 

1 Dorypleres biangulata Lindgren [1898]. 
Coppatias Johnstoni Topsent [1900]. 
tJaspis biangulata Thiele [1903 8]. 

Jaspis Johnstoni Topsent [1904 a]. 

Jaspis johnstoni var. incrustans Topsent [1904 a]. 
*Coppatias albescens Row [1911]. 


The specimen forms a thin crust of a light pinkish-brown colour, spreading over and 
cementing together a mass of Siliquaria shells and other calcareous débris. The vents 
are minute and scattered. The soft tissues contain numerous granular brown pigment- 
cells. 

The skeleton is a more or less dense feltwork of small oxea and microxea. 

Spicules :—(1) Short, stout, fusiform oxea (Plate 47, fig. 2a); slightly curved; usually 
very gradually and finely pointed, but.may be a little blunted and may also occasionally 
become stylote, but styli are rare. Size when full-grown about 0-46 by 0-024 mm. 

(2) Microxea (fig. 2b); sharply pointed, curved, usually centrotylote, closely 
resembling the microxea of Erylus; size very variable, say about 0-08 by 0-004 mm., 


254 PERCY SLADEN TRUST EXPEDITION 


but numerous smaller ones occur and also numerous forms connecting with the larger 
oxea (1). 

(3) Oxyasters (fig. 2c); with no distinct centrum, or quite a small one, and slender 
rays, usually about eight in number ; total diameter of spicule about 0°016 mm. 

Topsent has pointed out the variability in the size of the spicules in this species, and 
also mentions that the smaller oxea are frequently centrotylote. I have therefore no 
hesitation in identifying the “Sealark” specimen with the European form. I have 
thought it desirable, however, to give measurements and figures of the spicules of the 
“Sealark” specimen as a contribution to our knowledge of the range of variation. Topsent 
[1898 B] has also expressed the opinion that Lendenfeld’s Asteropus incrustans is identical 
with Jaspis johnstonw, an opinion with which I agree. It appears to me very probable 
that Row’s Coppatias albescens from the Red Sea [1911] may belong to the same species, 
and possibly also Lindgren’s Dorypleres biangulata from Java [1898], also recorded by 
Thiele [1903 8B], under the name Jaspis biangulata, from Ternate. 

Previously known Distribution. Adriatic (Schmidt, Lendenfeld); Mediterranean 
coast of France (Topsent); St Iago, Porto Praya, Cape Verde Is. (Sollas); Azores 
(Topsent) ; ?Red Sea (Row) ; ? Java (Lindgren) ; ?Ternate (Thiele). 

Register No., Locality, dc. wuxxvi. 11, Cargados Carajos, 28.3.05. B. 2, 
30 fathoms. 

Family Geodiide. 


Astrotetraxonida with trizene megascleres, a cortical layer of sterrasters, and various 
forms of euasters, to which microrhabds may be added. 

The question of the probable origin of this family from the genus Aurora has 
already been discussed. The classification of the. Geodide is an extremely difficult 
problem and one which cannot be properly tackled without a comprehensive re-investiga- 
tion of the group. I may state, however, that I have the gravest doubts as to the value 
of the character and arrangement of the inhalant and exhalant openings for the dis- 
crimination of genera, upon which so much stress has been laid by Sollas and subsequent 
writers. I therefore use the old genus Geodia in a far less restricted sense than is 
customary, but, laying the greatest stress as usual upon skeletal characters, I have framed 
the diagnosis thereof so as to exclude Pachymatisma, Caminus and Geodinella, which may 
conveniently be kept distinct. 

I have followed Lendenfeld [1910 8] in referring the genus Erylus to a separate family. 


Genus Gropi1a Lamarck [1815 a]. 


Geodiidee with well-developed triznes arranged radially at or near the surface ; 
without microrhabds and without spherules. 


15. Geodia auroristella n. sp. 
(Plate 47, fig. 3.) 


The single specimen, in its present condition, has the form of a thin crust, which has 
evidently been sliced off either from a larger specimen of the sponge or from some foreign 
object which it was encrusting. As there is no larger specimen of the sponge in the 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 255 


collection the latter explanation is probably correct. The specimen measured about 
25 mm. in length by 8 mm. in breadth and 3 mm. in thickness. The colour on the 
surface is almost white, internally more yellowish. The surface is almost smooth, very 
minutely hispid. A group of some 20 or 30 minute, inconspicuous openings, situate close 
to the margin, but not in any special depression, represent the vents. 

Paraffin sections show that each of the small vents is really covered in by a thin 
pore-sieve, which roofs over a funnel-shaped chone that penetrates the cortex. The 
inhalant pores are very difficult to make out, but I conclude from the study of sections 
that they are scattered over the surface, probably in groups, and lead into narrow canals 
which penetrate the cortex, anastomosing as they go. The cortex is about 0°7 mm. thick, 
densely charged with sterrasters and with a superficial layer of minute spherasters. It 
contains a good deal of fibrous tissue connecting the sterrasters with one another and 
especially strongly developed round the inner ends of the exhalant chones, where it 
apparently forms sphincters. 

The skeleton in the deeper part of the sponge is confused, but loose bundles of 
diactinal megascleres radiate towards the surface, terminating distally in tufts of trizenes 
which penetrate the cortex. The stout orthotrizenes for the most part extend their cladi 
in the outermost part of the cortex, just beneath the surface ; beyond them again loose 
tufts of slender spicules project freely and are almost invariably broken off short ; some of 
the latter are very slender oxea (‘styli), others are reduced anatriznes with vestigial 
cladome. 

Spicules :-—(1) Orthotrizenes (Plate 47, fig. 3a); with long, straight or slightly curved 
shaft, gradually and finely pointed at the end or somewhat blunted, measuring about 1:7 
by 0:°034 mm. ; with short, stout, sharp-pointed cladi extended almost at right angles KO 
the shaft, measuring about 0°086 by 0:034 mm. 

(2) Anatrizenes (fig. 3b); with long, slender, almost hair-like shaft, tapering away 
to a very fine point, and sharply recurved cladi, rather long, slender and gradually and 
finely pointed. Shaft about 1:7 by 0:008 mm.; cladi about 0:07 by 0:006 mm. The 
cladome may be reduced to a mere knob. 

(3) Oxea (fig. 3c); long, slender, sometimes slightly curved ; may be sharply pointed 
at each end, but more often one or both ends is more or less rounded off, giving rise to 
stylote (fig. 3c’) and strongylote (fig. 3 c”) forms; measuring about 1-4 by 0-023 mm. The 
long, very slender oxea (? styli) projecting from the dermal surface may be regarded as 
modifications of these. 

(4) Sterrasters (figs. 3d—8 d’’”); of typical oval form, with distinct hilum and surface 
reticulation formed by the stellate ends of the ftised rays; size about 0°123 by 0:1 mm., 
but many smaller ones occur which appear to be fully developed. Developmental stages 
are shown in figs. 3d’/—3d’”. 

(5) Spherasters (fig. 3 e); with stout, smooth, conical but somewhat irregular rays, 
resembling an Aurora or Donatia spicule ; total diameter about 0°05 mm. Choanosomal 
(subcortical). 

(6) Spherasters (fig. 3); with relatively longer, more slender and minutely spined, 

conical rays; total diameter about 0:037 mm. Choanosomal. 


2956 PERCY SLADEN TRUST EXPEDITION 


(7) Minute spherasters (fig. 3 g); with small centrum and short, strongylote (perhaps 
sometimes tylote) rays; total diameter about 0°008 mm. Choanosomal and dermal. 

(8) Oxyasters (fig. 3); with usually about 8 or 9 slender, minutely roughened rays 
and inconspicuous centrum ; total diameter about 0°02 mm. 

In spiculation this species approaches Lendenfeld’s Sidonops oxyastra [1910 a] pretty 
closely, but differs in the possession of the large spherasters and perhaps in other details. 
It also comes near to Carter's Geodza globostellifera [1880 B], from the Gulf of Manaar, 
which has a similar spheraster (‘‘globostellate”), but is only 0°021 to 0°028 mm. in 
diameter, instead of 0°05 mm. as in our species. Carter’s species, however, appears 
to have protriznes and no anatriznes, and the oxyasters seem to be absent. 

Register No., Locality, &c. Uxxxvi. 1, Providence, 4.10.05, D. 4, 50—78 
fathoms. 


Family Erylide. 


Astrotetraxonida with a cortex containing aspidasters. The typical megascleres are 
triznes and oxea (or strongyla). The microscleres include microrhabds and choanosomal 
euasters. 

The remarkable genus Erylus was included by Sollas [1888] in the family Geodiide, 
as it had been by Gray [1867 F], and this procedure has since been generally followed. 
In 1910, however, Lendenfeld proposed to remove Erylus to a separate family by itself, 
to which he gave the name “Erylidz.” He pointed out that Erylus differs very con- 
siderably from Geodia and proposed the name “‘aspidaster” for the characteristic cortical 
spicule, which had previously been regarded as a sterraster. This spicule is undoubtedly 
the most characteristic feature of the Erylidee, and it is quite conceivable that, like the 
sterrospira of Placospongia, its resemblance to the Geodia sterraster may be due simply 
to convergence. This view is supported by the fact that Erylus differs from Geodia also 
in other respects, such as the presence of the very characteristic microrhabd and the 
absence of anatrieenes. The triznes that are present seem to be less differentiated than 
those of the Geodiide and may resemble the short-shafted trizenes of the Pachastrellidee, 
from primitive members of which family it seems quite possible that the Erylidz have 
been independently and directly evolved, while the Geodiide have almost certainly arisen 
through the Stellettidee. 

It is customary at the present time to recognise only a single genus in this family, 
but it seems possible that we may before long be able to sub-divide it according to the — 
form of the aspidaster. Even if we are able to do this, however, I fear that Ferrer’s 
recently proposed genus Scutastra [1912] will have to be regarded as a synonym of 
Erylus. 


Although a good many species have been described these are for the most part 
evidently very closely related to one another. 


Genus Eryztus Gray [1867 F]. 
With the characters of the family. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 257 


16. Hrylus lendenfeldi Sollas. 
(Plate 47, fig. 4.) 


Stelletta euastrum Carter [1880 8, p. 136], not Schmidt. 
Lrylus lendenfeldi Sollas [1888]. 


I am not aware that this species has been met with since it was first described by 
Carter in 1880 from Fremantle, Western Australia, and Carter’s description was taken 
from a single dry specimen. It seems therefore worth while to describe it again. 

The single specimen in the ‘‘Sealark” collection is cushion-shaped, subspherical, 
about 16 mm. in maximum diameter. It has been torn off from some attachment below. 
It bears a single circular vent, about 2 mm. in diameter, not far from the centre of the 
upper surface, without any oscular collar or fringe of any kind. The inhalant pores are 
distinctly visible under a pocket lens, scattered irregularly and singly over the general 
surface, each with a narrow whitish margin ; sometimes widely open and sometimes closed 
to varying extents. The surface is smooth and of a dark, greyish-brown colour; the 
interior is yellow. Texture firm and fairly compact, but with a wide oscular tube 
running up to the vent and appearing (broken across) on the lower surface, which 
suggests that there may have been a good deal more of the sponge below. 

Skeleton confused internally, radially arranged towards the surface, consisting 
principally of large oxea but with a fair number of orthotriznes typically orientated 
beneath the cortex of aspidasters. 

Spicules :—(1) Orthotrizenes (Plate 47, fig. 4a); with well-developed cladi, typically 
unbranched but sometimes irregularly bent and even forked; shaft and cladi tapering to 
more or less sharp points (often blunted). Size variable, a typical specimen gave the 
following measurements :—shaft 0°5 by 0:034 mm.; cladi 0°24 by 0:034 mm. 

(2) Oxea (fig. 46); slightly curved, tapering at each end to a sharp or blunt point, 
measuring about 1:0 by 0°026 mm. 

(3) Aspidasters (figs. 4c—4c””); much flattened, oval or irregular in outline, 
surfaces beset with numerous small, scattered, simple or slightly stellate tubercles ; 
dimensions about 0°17 by 0°082 mm. Young forms radially striate, formed by fusion 
of slender rays proceeding from a central mass ; intermediates smooth, without tubercles. 

(4) Large oxyasters (fig. 4d); with no distinct centrum. and long slender rays, 
minutely spined and not quite sharply pointed; rays usually 4 in number, measuring 
each up to about 0:07 by 0:004 mm. 

(5) Small. oxyasters (fig. 4); similar to the last but with rays only about 0°01 mm. 
long; perhaps young forms ; abundant. 

(6) Polyactinose oxyasters (fig. 4); with minutely spined, slender rays and no 
distinct centrum. Total diameter up to about 0°028 mm. 

Intermediate forms between (4), (5) and (6) occur; and some of the small asters may 
have slightly tylote rays. 

(7) Microrhabds (fig. 4g); smooth, curved, often centrotylote and usually bluntly 
pointed at each end; measuring, say, about 0°07 by 0°006 mm., but variable. 

SECOND SERIES— ZOOLOGY, VOL. XVII. 33 


258 . PERCY SLADEN TRUST EXPEDITION 


The cortex is about 0°2 mm. thick and is sharply differentiated into two layers—an 
outer layer forming about four-fifths of the whole and so densely charged with tangentially 
placed aspidasters, arranged in several layers, that it is impossible to make out its 
histological characters; and an inner layer, forming the remaining fifth, composed of 
fibrous tissue. Both layers contain numerous pigment cells, which, in the fibrous layer, 
are elongated in the same direction as the fibres, 7.e. parallel to the surface. A few 
pigment cells also occur in the outer part of the choanosome. 

The scattered inhalant pores lead each into a very well-defined chone, subcylindrical 
in shape but widening out somewhat as it penetrates the cortex, the bottom of the chone 
being at about the junction of the outer and inner layers of the cortex. A rather thick 
sphincter diaphragm, formed from the inner, fibrous layer of the cortex, separates the 
chone from the subcortical crypt into which it opens. The subcortical crypts are 
irregular spaces which unite together to form wide inhalant canals running almost 
vertically inwards. 

The choanosomal ground-substance is compact and finely granular; the flagellate 
chambers are subspherical, close-packed, about 0°024 mm. in diameter. 

It will be seen that our sponge agrees very closely with Sollas’s re-description of the 
type [1888]. The chief distinguishing feature of the species is evidently the large 
oxyaster. 

Previously known Distribution. Fremantle, Western Australia (Carter). 

Register No., Locality, &c. cu. 2, Amirante, 18.10.05, E. 25, 44—20 fathoms. 


17. Hrylus proximus n. sp. 
(Plate 47, fig. 5.) 


The single specimen consists of an irregular mass of Siliquaria shells and sponge 
inextricably mixed together; the whole measuring about 65 by 36 by 25mm. ‘The entire 
mass is somewhat flattened dorsoventrally, but the surfaces are very uneven and irregular, 
with portions of the Siliquaria projecting freely at frequent intervals. What was pre- 
sumably the lower surface is penetrated by numerous minute, singly scattered, dermal 
pores, widely open, commonly with distinct whitish margins; numerous minute whitish 
specks scattered singly over the upper surface represent similar pores in a closed condition. 
A few open vents, about 1°5 mm. in diameter, occur singly on prominent parts of the 
upper surface. The colour of the upper surface (in spirit) is purplish or brownish grey ; 
of the lower surface greyish yellow. 

The main skeleton is a confused interlacement of large oxea, with a few short-shafted 
trieznes beneath the dermal crust. Some at any rate of the trieenes have their cladi 
extended paratangentially beneath the surface. There is a thin dermal crust formed of 
the thin, plate-like aspidasters, which also occur scattered abundantly in the choanosome. 

Spicules :—(1) Short-shafted trienes (Plate 47, fig. 5 a); much resembling calthrops, 
but with the cladi extended almost at right angles to the shaft; cladi may be a little 
longer or a little shorter than the shaft. Shaft and cladi sharp-pointed, stout or slender ; 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 259 


_cladi occasionally bifurcate at the apex. In a typical example the shaft measured about 
0-2 by 0°03 mm., cladi about the same. The forms with slender rays are perhaps young. 

(2) Oxea (fig. 56); fairly stout, fusiform, slightly curved and gradually and 
sharply pointed; varying occasionally to stylote (fig. 5b’) or even strongylote (fig. 5 6”) 
forms; often slightly centrotylote; typical size about 0-7 by 0:025 mm. 

(8) Aspidasters (fig. 5c—5c””); thin, plate-like, oval; usually with fairly even 
outline; surface beset with numerous, scattered, slightly stellate, short spines or tubercles ; 
size about 0°15 by 0°07 mm. The young forms are very thin, smooth, oval plates, with a 
marginal fringe of slender spines ; radially striate when very young. 

(4) Chiasters (fig. 5d); polyactinal strongylasters and tylotasters ; about 0°012 mm. 
in total diameter ; sometimes with a distinct centrum. 

(5) Microrhabds (fig. 5e); smooth, fusiform, usually slightly curved, usually sharply 
pointed, often centrotylote ; size about 0°053 by 0:004mm. Abundantly scattered in the 
choanosome and in the pore-sphincters. 

With regard to the microscopical anatomy of the soft parts in this form I have only 
to note two points. The inhalant pores are situated each in a distinct gap in the dermal 
armour of aspidasters, and each is provided with a thin sphincter membrane strengthened 
by numerous microrhabds. The interior of the sponge is charged with numerous small, 
brown, granular pigment-cells, especially abundant near the surface. 

This species comes very near to Hrylus carteri Sollas [1888], originally described by 
Carter from the Gulf of Manaar under the name Stelletta ewastrum [1880 B, p. 135], which 
was also associated with Siliquaria. The most important difference lies in the fact that 
the microrhabds (microxea) are smooth, as usual in the genus, instead of minutely spined 
as described and figured by Carter for the type of the species. 

To judge from the emended description given by Sollas [1888], Ridley’s [1884 c] 
Erylus cylindrigerus from the Mascarene Islands is perhaps even more closely related to 
the present species, differing, however, in the lozenge-shaped form of the aspidaster, the 
form and size of the small asters, and the (constant ?) strongylote form of the main skeleton 
spicule. It appears to me highly probable that all three species will have to be united in 
the future. 

Register No., Locality, dc. vit. 7, Cargados, 30 fathoms. 


Family Donatiidee 
=Tethyade or Tethyide auctorum. 


_ Astrotetraxonida without tetractinellid megascleres. With a strongly developed 
fibrous cortex. Main skeleton composed of radially arranged styli (or oxea?). Microscleres 
euasters of various forms, including large spherasters, to which microrhabds may be added. 

This family appears to have originated from some corticate stellettid ancestor of 
the Aurora type. Indeed, if we admit such “epipolasid” forms as Aurora distinctus, 
A. cribriporosa and A. sterrastrea amongst the Stellettidee, it is not easy to frame a 
diagnosis by which the Donatiide can be logically excluded from the same family. 

33—2 


260 PERCY SLADEN TRUST EXPEDITION 


It appears, however, that there is a small group of genera, including Donatia (Tethya), 
Tethyorhaphis and Xenospongia, of monophyletic origin and closely related to one another, 
in which the loss of the tetractinellid megascleres is absolutely constant and forms the 
principal distinguishing feature, instead of occurring sporadically as amongst the Auroras. 
This character, taken in connection with the characteristic spherasters* and the very. 
strongly developed cortex, seems to form a sufficient justification for retaining the family. 

Hitherto I have employed the name “ Tethyide” for this family, on the supposition 
that the proper name of the typical genus was Tethya and not Donatia. It will be 
remembered that Sollas expressed himself very emphatically on this point. He says 
“ Tethya lyncurvum is a combination that by the accepted laws of nomenclature cannot - 
possibly be disturbed, since the species is the type of the genus so named by Lamarck, 
and accepted and redefined by O. Schmidt in 1862” [1888, p. cxx1]. Until recently the 
name Tethya has been accepted in this sense by nearly every writer, including Vosmaer, 
Lendenfeld, Thiele, Topsent and Lindgren, | 

In 1903, however, Thiele, after consistently using the name Tethya for the genus in 
question throughout the body of his paper on the “Kieselschwimme von Ternate,” 
remarked in a footnote that by strict application of the laws of nomenclature Tethya must 
be substituted for Craniella and Donatia for Tethya. In 1905 Baer followed this up 
by proposing the family name Donatiidee, and Lendenfeld [1903], Topsent [1906 B] and 
Hentschel [1909] have all fallen into line. 

It must be admitted that there is nothing in Lamarck’s original paper [1815] to 
justify Sollas’s confident assertion. The genus Tethya is. there quite unrecognisably 
defined and Tethya lyncurium is the fifth species to be described. The first is Tethya 
asbestella, which, whatever it may be, is certainly not congeneric with 7. lyncurvum. It 
is true that 7. lyncurvwm comes before T. craniwm, which Lamarck places sixth, but that 
hardly justifies us in accepting it as the type species of the genus. According to Vosmaer 
[1887], followed by Lendenfeld [1903], the fourth of Lamarck’s Tethyas, 7. lacunata, is 
a Geodia! As the generic name Geodia was only proposed by Lamarck himself on a later 
page of the same volume as Tethya, it certainly looks as if neither T. lyncurium nor 
T. cranum had a right to the name Tethya on the ground of priority of mention. But 
then there are three other species before T. acunata. What these may be I do not know, 
and I doubt if anyone else does, but if the types are still in existence further researches 
might upset any decision as to priority which might now be made. 

On the whole the wisest course would seem to be to abandon the generic name Tethya 
altogether and to use Nardo’s name Donatia, proposed in 1833, for D. lyncurvum and its 
congeners, and Baer’s name Donatiide for the family. 


Genus Donatta Nardo [1833] 
= Tethya auctorum. 


Donatiidee of usually more or less spherical form; megascleres styli; microscleres 
large spherasters, with smaller chiasters or oxyasters or both ; without microrhabds. 
Many species of Donatia have been described by various authors, but the question 


* In the aberrant genus Tuberella, however, the microscleres have also completely disappeared. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 261 


of their mutual relationships is no less difficult than that of the name of the genus. 
There is no doubt that members of the genus exhibit very great variability in all those 
characters that might be used for the purpose of specific distinction ; in the character 
of the surface, the form and size of the microscleres, the arrangement of the microscleres, 
the thickness of the cortex, &c. 

Lindgren [1898] proposed to arrange the described forms in three groups, each 
of which might be regarded as a species. The names of the three species would then 
be lyncurium, ingalli and japonica respectively. This arrangement is based on the 
characters of the smaller asters. In Donatia lyncurium the smaller asters are supposed 
to be oxyasters (0°012—0°02 mm. in diameter) only, variable in form and size but never 
with tylote rays. In D. engalli the smaller asters are tylote chiasters (0°006—0:016 mm.) 
and oxyasters (0°02—0°052 mm.). In D. japonica the smaller asters are all tylote 
chiasters (0:008—0°016 mm.). D. lyncurium would appear to be characteristic of the 
North Atlantic and Mediterranean; D. ingalli of the Red Sea, Indian Ocean, Malay 
Archipelago, Australia and the Pacific; D. japonica of the Philippines, Java and 
Australia. I do not think that this arrangement can be accepted, the eee of 
species appears to be too comprehensive. 

In my report on the sponges of Ceylon [1905] I proposed to regard all the specimens 
of Donatia in Professor Herdman’s collection as belonging to three. varieties of Donatia 
(Tethya) lyneurium, which I distinguished as a, b and ¢ respectively. The undoubted 
existence of transitional forms of the small asters at first sight seems to make it impossible 
to recognise such forms as D. lyncurium, D. ingalli, D. seychellensis and D. japonica 
as distinct species. There is no reason, however, why either specific or varietal 
distinctions should be based exclusively upon the characters of the chiasters and 

oxyasters. The form, size and arrangement of the large spherasters, and the arrange- 
ment of the megascleres may also prove useful in this respect, and by taking these 
into account I now believe it possible to arrive at fairly satisfactory specific distinctions. 

It is impossible to discuss in this place all the species and varieties that have 
been proposed. Nothing but a thorough and tedious revision of the whole genus, 
based, if possible, upon the examination of type specimens, can lead to any really 
satisfactory general conclusions. 

There are thirty-one specimens of Donatia in the “Sealark” collection, and after 
careful examination, including measurement of the microscleres of all of them, I have 
decided to arrange them in five species. None of the specimens are much more than 
an inch in diameter and many of them are much less; they attain nothing like the 
size often seen in Australian Donatias. 

It will be seen from the locality lists that the different species occur very much 
mixed together. I have found three species (lyncwriwm, japonica and seychellensis) in 
the same jar from Salomon, presumably all obtained in the same haul of the dredge, 
and a different three (japonica, seychellensis and ingalli) in the same jar from Praslin 
Reef. These facts show how necessary it is to examine every specimen microscopically 
before determining to which species it belongs. 


262 PERCY SLADEN TRUST EXPEDITION 


18. Donatia lyncurium auctorum. 


(Plate 48, fig. 1.) 
Tethya lyncuriwm auctorum. 


(For synonymy and literature of the species vide Lendenfeld [1896].) 


There is in the collection a single very small specimen, only about 8 mm. in diameter, 
which appears to me to be indistinguishable from the European form. It has a strongly 
tessellated surface and a few small, root-like attachment processes, and the colour in spirit 
is nearly white. 

The cortex is about 0°86 mm. thick and there appear to be no large intracortical 
crypts. The radiating bundles of the main skeleton pierce the cortex and spread out into 
wide brushes, the projecting ends of the component spicules of which render the surface 
slightly hispid. Beneath the cortex the intervals between the well-defined skeletal 
bundles are occupied by numerous loose styli or subtylostyli radially arranged. 

The large spherasters (Plate 48, figs. 1a, 16) are scattered irregularly and rather 
sparsely in the cortex. They are abundant in the outer part of the choanosome mixed 
with smaller forms, possibly developmental stages. When fully grown they measure 
about 0°07 mm. in diameter. Their sharp conical rays are about half as long as the 
diameter of the centrum. 

The cortical chiasters (fig. 1c) are strongylasters or oxyasters, rarely faintly tylote, 
with usually more than six very slender rays and no conspicuous centrum ; total diameter 
about 0°012 mm. Curiously enough these spicules appear to be absent from the surface, 
where, it will be remembered, the cortical chiasters are usually most abundant. The 
choanosomal chiasters (fig. 1d) are rare and closely similar in form and size to those of 
the cortex. 

Previously known Distribution. European Seas, &c. (For further particulars wde 
Lendenfeld [1896 |.) 

Register No., Locality, de, oxx. 2c, Salomon, 10—14 fathoms. 


19. Donatia japonica (Sollas). 


(Plate 48, fig. 2.) 


1 Tethya lyncuriwm Desz6 [1878]. 

Tethya japonica Sollas [1888]. 

Tethya japonica Lindgren [1898]. 
Tethya lyncurium var. a. Dendy [1905]. 
Donatia japonica Topsent [1906 8]. 
Donatia parvistella Baer [1905]. 

Tethya lyncurium Row [1911]. 


(For other possible synonymy vide Lindgren [1898] and Hentschel [1909].) 


There are in the collection nine specimens which I think must be referred to this 
species. They are all more or less spherical in form, and the surface exhibits the usual 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 263 


range of variation from strongly tessellated in polygonal areas to conulose and proliferous. 
Although the varying character of the surface causes the specimens to differ very 
strikingly in external appearance I do not think that it implies any specific or even 
varietal distinction, but merely phases of growth, possibly associated with a periodic 
activity in throwing off buds*. None of the specimens exceed a diameter of about 
20mm. The colour in spirit is light grey or yellowish. 

The cortex is only about 1:0 mm. thick, with more or less well developed intra- 
cortical crypts from which narrow inhalant canals run into the choanosome. 

The spicule-bundles of the main skeleton penetrate the cortex and either spread out 
in brushes at the surface (in the tesserze) or are continued into well-marked conuli. 
In the outer part of the choanosome, between the spicule-bundles, numerous loose 
megascleres (styli or subtylostyli) are radially arranged. These cease abruptly just 
beneath the cortex. 

The large spherasters are rather sparsely scattered in the cortex. In the choanosome 
their numbers vary greatly and they are apt to be mixed with small forms that probably 
represent, in part at any rate, developmental stages. 

The small tylasters form a distinct dermal layer, as well as occurring scattered in cortex 
and choanosome. 

The spherasters (Plate 48, figs. 2 a, 2b), when fully grown, range from about 0:04 to 
about 0°07 mm. in total diameter. They have sharp conical rays whose length is about 
half the diameter of the large centrum. 

The cortical and choanosomal tylasters (fig. 2c, 2d) closely resemble one another 
both in form and size and range from about 0°0082 to 0°0164 mm. in total diameter. 
They usually have more than six rays, but in R.N. xiv. 2 E they are usually six-rayed. 

The chief distinguishing features of this species are the tylote character of the cortical 
and choanosomal chiasters, the absence of oxyasters and the comparative scarcity of 
spherasters in the cortex. The choanosomal chiasters may, however, occasionally lose the 
heads of the rays and exhibit a transition to those of D. ingalli (fig. 2 e). 

I pointed out in my Report on the Ceylon sponges that the tylote character of the 
chiaster may occasionally occur even in a British Donatia, and I must now add that Deszé, 
as far back as 1878, figured typical tylasters for a specimen of Donatia from Naples, 
identified by him as Tethya lyncurvwm. Perhaps he really had D. japonica before him. 

Previously known Distribution. Manila (Sollas); Java Seas (Lindgren) ; Ceylon 
(Dendy); Zanzibar (Baer); Red Sea (Topsent, Row); ?Naples (Deszé). (For other 
possible localities vide Lindgren [1898] and Hentschel [1909 ].) 

Register Nos., Localities, dc. XLVI. 2 &, XLVI. 3, XLIx., all from Praslin Reef; 
LIV. 2 B, Coetivy ; LViI. 10 a, 8B, Coin, Peros ; uxir., Lagoon, Diego Garcia, 8.7.05; cvIt. 
2, Amirante, 14.10.05, EK. 17, 12—18 fathoms; cxx. 2 B, Salomon, 10—14 fathoms. 


* Compare Sollas’s remarks on this subject in the case of Donatia seychellensis [1888]. 


264 PERCY SLADEN TRUST EXPEDITION 


20. Donatia imgallr (Bowerbank). 


(Plate 48, fig. 3.) 


Tethea ingalli Bowerbank [1872 4). 

Tethya ingalli (pars) Sollas [1888]. 

Tethya ingallt (pars) Lindgren [1898]. 

Tethyu lyncurium var. b. Dendy [1905]. 
Donatia Ingalli (pars) Hentschel [1909, 1912]. 


This is much the commonest form of Donatia in the collection, being represented by 
seventeen specimens. These range in form from a flattened crust, with wide-spreading 
base and strongly convex upper surface (R.N. cx. 7), to the usual subspherical form with 
root-like processes of attachment spreading broadly over the substratum. The surface 
is usually more or less strongly tessellated in polygonal areas. The colour in spirit is 
yellowish grey. The subspherical specimens attain a diameter of about 25 mm. 

The cortex is some three or four millimetres in thickness and densely charged 
throughout with large spherasters forming an almost solid mass. 

The spicule-bundles of the main skeleton penetrate the cortex and spread out into 
brushes beneath the surface as usual, but there are, usually at any rate, no loose, radially 
arranged megascleres beneath the cortex and between the bundles. 

The large spherasters (Plate 48, figs. 3a, 3b), when fully grown, range from about 
0-08 to 0°14 mm. in diameter, with rays about half as long as the diameter of the large 
centrum. Though most abundant in the cortex they are often common also in the 
choanosome, where they may be associated with numerous much smaller forms*. Some- 
times comparatively small forms, with very short rays, occur in the cortex just beneath 
the surface (R.N. oxi. 18, fig. 3a). In the larger forms the rays may occasionally _ 
branch (R.N. cxrx. 10 B). 

The cortical chiaster is a typical tylaster (fig. 3c) with usually more than six rays, 
from about 0°012 to 0°016 mm. in total diameter. 

The choanosomal chiaster ranges from a typical tylaster (fig. 3d) to an oxyaster or 
strongylaster (fig. 3d’) with many short rays, which may be sen roughened, and 
a total diameter of about 0°012 to 0°02 mm. 

The chief distinguishing features of these specimens are the great thickness of the 
cortex, the close-packed arrangement of the spherasters in the cortex, and the strong 
tendency of the choanosomal chiasters to lose the heads of the rays and assume the form 
of many-rayed oxyasters. 

I base my conception of this species upon Bowerbank’s original description, the 
emended description by Sollas, and my own study of one of Bowerbank’s specimens. 
I conclude that the “Sealark” and Ceylon specimens differ from the Australian types in 
the shorter rays and less strongly developed spination of the choanosomal oxyasters. As 
regards these spicules the typical form seems to approach D. seychellensis, while differing 


* T have in all the species assumed that these small choanosomal forms are immature and have not 
included them in the range of size. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 265 


from that species in the structure of the cortex. I refrain at present from expressing any 
opinion as to the supposed identity of D. robusta and D. cliftoni [Bowerbank 1873 4] 
with this species. ; 

Previously known Distribution. §.W. Australia (Bowerbank, Hentschel); Bass Str., 
Port Jackson (Sollas) ; Queensland (Berlin Museum ; a section in my possession labelled 
Tethya fissurata Ldf. and numbered 1120); ?Java Sea and Gaspar Strait (Lindgren) 
Ceylon (Dendy). 

Register Nos., Localities, dc. XLVI. 2 B, C, D, F, Praslin Reef; Lit. 8 A, B, LIV. 1, 
2 A, C, D, all from Coetivy; ex. 7, cxr 4, 5, cxu. 1 A, B, all from Egmont Reef; cxrx. 
10 A, B, Salomon. 


) 


21. Donatia seychellensis (Wright). 


(Plate 48, fig. 4.) 


Alemo seychellensis Wright [1881]. 
Tethya seychellensis Sollas [1888]. 

Tethya seychellensis Keller [1891]. 
Tethya seychellensis Topsent [1893 &]. 
Tethya ingalli (pars) Lindgren [1898]. 
Tethya seychellensis Kirkpatrick [1900 a]. 
Tethya lyncurium var. c. Dendy [1905]. 
Donatia Ingalli Topsent [1906 8]. 

2 Donatia Ingalli (pars) Hentschel [1909]. 
Tethya seychellensis Row [1911.] 
Donatia seychellensis Dendy [1915]. 


This species is characterised above all by the large choanosomal oxyasters (Plate 48, 
fig. 4d), usually with six, often branched rays. The hexradiate character of these spicules 
has already been pointed out by more than one writer and seems to be fairly constant. 
In the “Sealark” specimens the cortical and choanosomal tylasters (fig. 4c, d’) are also 
typically hexradiate. The measurements of the three kinds of aster in the ‘“‘Sealark” 
specimens are as follows:—Large spheraster (fig. 4a), when fully grown, 0:07 mm. ; 
cortical tylaster 0°012 mm. ; choanosomal oxyaster (total diameter) 0°04 mm. 

The three specimens in the collection have the usual subspherical form and more 
or less strongly tessellated surface, and all of them, in parts, show remarkably wide 
gaps (pore-grooves) between the tessere. The largest is about 25 mm. in diameter. 
The colour in spirit is grey or yellowish. One specimen only shows a few elongated, 
slender conuli (? budding). 

In all three the cortex is very lacunar and does not contain nearly so many 
spherasters as in the specimens of D. imgalli, while beneath the cortex and between the 
radiating spicule bundles (which pierce the cortex and spread out as usual) are found 
numerous loose, radially arranged styli, as in D. japonica. 

The large red Donatia so common in the neighbourhood of Port Phillip, Victoria, 
possesses large oxyasters very similar to those of D. seychellensis, but I am inclined to 
think, on account of other characters, that it is specifically distinct both from D. seychel- 
lensis and D. ingall. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 34 


266 PERCY SLADEN TRUST EXPEDITION 


Previously known Distribution. Seychelles (Wright) ; Samboangan (Sollas) ; Flinders 
Passage, Torres Straits (Sollas); Red Sea (Keller, Row, Topsent); Okhamandal and Gulf 
of Manaar (Dendy); ?South West Australia (Hentschel). 

Register Nos., Localities, de. xuvit. 2 A, Praslin Reef; cxiir. 1 c, Egmont Reef; 
oOxx. 2 A, Salomon, 10—14 fathoms. 


22. Donatia stella-grandis n. sp. 
(Plate 44, fic. 8; Plate 48, fig. 5.) 


The single specimen (Plate 44, fig. 8) is irregularly spherical, about 25 mm. in 
diameter, attached at one (evidently the lower) side to a small mass of nullipore. Part 
of the surface is very distinctly tessellated, the flat polygonal tesseree beg separated by 
wide pore-grooves ; elsewhere the tesseree are obsolete and the pore-grooves hardly 
discernible. There are two slight mammiform elevations on the upper surface, each of 
which probably bears a small vent, now closed. The colour in spirit is dull greyish 
yellow and the texture incompressible and of almost stony hardness. 

There is nothing peculiar to notice about the arrangement of the megascleres. They 
radiate in stout, widely separated bundles from the interior of the sponge and penetrate 
the cortex. On approaching the surface these bundles spread out into brushes of spicules 
whose ends may project slightly beyond the surface. 

The cortex is only about 0°7 mm. thick, and cortex and choanosome alike are densely 
packed with an almost solid mass of spherasters of various sizes. 

Spicules :—(1) Styli (Plate 48, fig. 5a), tylostyli (fig. 5a’) or strongyla (fig. 5 a”) ; 
measuring up to about 1°9 by 0°038 mm. These spicules are typically fusiform, but with 
the centre of the spindle much nearer to the broad end. The broad end is never much 
narrowed as compared with the middle of the shaft and is often distinctly tylote; the 
narrow end (centrifugal) may be sharply pointed or truncated. 

(2) Spherasters (fig. 56); rays elongated, conical, sharp, often shghtly bent, 
sometimes slightly and irregularly branched, about as long as the diameter of the 
centrum; total diameter very variable, up to about 0°25 mm. The branching of the 
rays seems to be confined to the larger forms, which are very numerous. 

(3) Small chiasters (figs. 5¢, 5d); with about 10 fairly stout, cylindrical rays, 
usually slightly roughened towards the ends, which may appear slightly tylote or simply 
strongylote ; total diameter up to about 0:016 mm. Abundant in cortex (especially at 
the surface) and in choanosome. 

(4) Chiasters (fig. 5c’) of about the same size but with slender oxeote rays 
(?roughened) ; these seem to occur in both choanosome and cortex but are not nearly 
so common as the other chiasters, from which, however, they cannot be sharply separated. 

This species is distinguished from all the other species of Donatia known to me, with 
the exception of Hentschel’s Donatia tylota [1912], by the enormous size frequently 
attained by the spherasters. The largest ones are about twice the diameter of those of 
any other specimen in the collection, and six times the diameter of those of a specimen of 
D. japonica from Praslin (R.N. xtrx.). Bowerbank speaks of the corresponding spicules 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 267 


in his Tethea robusta [1873 4] as being very large, but they are only one-fifth the diameter 
of those of the present species, according to his measurements and figure. In Donata 
tylota, however, they seem to be nearly as large as in D. stella-grandis. 

Register No., Locality, &c. cvt. 5, Amirante, 13.10.05, E. 16, 39 fathoms. 


Family Chondrosiide. 

Corticate Astrotetraxonida with complex canal system and small flagellate chambers. 
Without megascleres. Microscleres, when present, euasters. 

This family, as at present understood, comprises only the two genera, Chondrilla and 
Chondrosia. These may both be regarded as reduction forms derived from some stellettid 
ancestor such as Aurora. The spherasters of Chondrilla closely resemble those of Aurora, 
but the megascleres have entirely disappeared. In some specimens of Chondrilla mixta 
the asters have already become very rare, and. these seem to lead the way to the genus 
Chondrosia, in which all spicules have entirely disappeared. 


Genus CHONDRILLA Schmidt [1862]. 


Chondrosiidz with microscleres (euasters of one or more kinds). 


23. Chondrilla australiensis Carter. 
(Plate 48, fig. 6.) 

Chondrilla australiensis Carter [1873 c]. 

Chondrilla australiensis Lendenfeld [1886 a]. 

Chondrilla australiensis Lindgren [1897, 1898]. 

Chondrilla australiensis Dendy [1905, 1915}. 

Chondrilla australiensis Hentschel [1909, 1912]. 

This species is characterised by the presence of two kinds of aster, spherasters 
(Plate 48, fig. 6a) about 0°03 mm. in. diameter, and oxyasters (fig. 6b) of somewhat 
smaller size with roughened and sometimes slightly branched rays. The spherasters are 
characteristically cortical in distribution and the oxyasters characteristically choanosomal, 
but spherasters occur also in the choanosome and oxyasters in the cortex. 

It seems possible also that the light colour as compared with some other species may 
be a more or less constant character. Carter originally described the species as “of a 
dirty yellow or buff colour.” Lendenfeld copies this. Hentschel speaks of the colour of 
specimens from 8.W. Australia as mostly clear greyish-yellow, almost white in some small 
Specimens, but in places brown to black-brown. The Ceylon specimens collected by 
Prof. Herdman were of a greyish colour (in spirit). The “Sealark” specimens range from 
almost quite white all over (R.N. xuiu. 2) to light brown (in spirit); they form flattened 
or lobular crusts of the usual appearance. R.N. xxx 1 is growing upon a specimen of 
Derecitopsis minor. 

In some specimens (R.N. x1ir. 2, 4) the oxyasters are so nearly smooth as to resemble 


’ 


very closely those of C. mixta. 

Previously known Distribution. Port Jackson, E. Coast of Australia (Carter, Lenden- 
feld) ; Shark’s Bay, S.W. Australia (Hentschel); Okhamandal and Ceylon (Dendy); coast 
of Cochin China (Lindgren) ; Aru Islands (Hentschel). 

34—2 


268 PEROY SLADEN TRUST EXPEDITION 


Register Nos., Localities, dc. Xxx. 1, 3, XLII. 2, 4, all from Cargados Carajos, 
30.8.05, B. 18, 30 fathoms; Lxxxvir., Amirante, 9.10.05, E. 6, 28 fathoms; cxxxv. 2, 
Seychelles, 13.10.05, F. 9, 37 fathoms. 


24. Chondrilla mixta Schulze. 


(Plate 48, fig. 7.) 

Chondrilla mixta Schulze [1877 cl]. 

?Chondrilla mixta Ridley [1884 c]. 

Chondrilla mixta Lindgren [1897, 1898]. 

Chondrilla mixta Kirkpatrick [1900 8]. 

This species was described by Schulze from the Red Sea. Schulze had only a single 
specimen, of which he observes “Dasselbe stellte eine blassgraue, braungefleckte Kruste 
von 2—4 Mm. Dicke mit unregelmissig welliger aber glatter Oberfliiche dar.” 

The “‘Sealark” specimens are for the most part dark brown or nearly black on the 
surface, owing to the strong development of pigment-granules in and beneath the cortex. 

The spicules are by no means numerous and in some specimens almost disappear from 
the cortex, so that it is hard to find anything but the choanosomal oxyasters. 

The characteristically cortical spherasters (Plate 48, fig. 7a) resemble those of 
C. australiensis and measure about 0°03 mm. in diameter. The choanosomal oxyasters 
(fig. 7 b) are of about the same size or a little smaller. They differ from those of C. austra- 
liensis in having smooth, unbranched rays. Apart from the possible difference in colour 
this seems to be the only feature that distinguishes the two species, and it may well be 
doubted whether the two are more than varietally distinct. 

The specimen described by Ridley from the Amirante group was of a pale brown 

or buff colour and had oxyasters frequently with branched rays; it was probably 
— C. australiensis. ri 

Two specimens described by Lindgren from Java and Gaspar Straits were blue-black 
on the outside with a finely granulated surface due to the presence of very minute papillee. 

Previously known Distribution. Red Sea (Schulze); Java and Gaspar Straits 
(Lindgren) ; Funafuti Atoll (Kirkpatrick). 

Register Nos., Localities, dc. itt. 8, Coetivy ; Lvit. 1, Coin, Peros; cx. 4, Cxii1. 3, 9; 
Egmont Reef; cxir. 2, 4, Eemont Lagoon; cxx. 6, Salomon, 10—14 fathoms. 


25. Chondrilla sacciformis Carter. 


(Plate 48, fig. 8.) 

Chondrilla sacevformis Carter [1879 B]. 

Magog sacciformis Sollas [1888]. 

Chondrilla grandistellata Thiele [1900]. 

The single specimen in the collection forms an elongated, slug-shaped crust, about 
30 mm. in length, 10 mm. in greatest breadth and 6 mm. in greatest thickness. The 
margins are strongly incurved towards the base of attachment, probably owing to con- 
traction after removal from the substratum. There are five or six minute, contracted 
vents, scattered singly, each on a small mammiform projection on the upper surface. The 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 269 


texture is hard and tough. The colour of the upper surface is rather dark brown, 
internally much paler, with, to the naked eye, a sharp boundary line between cortex and 
choanosome. Under a pocket-lens the huge spherasters can be distinctly seen both on 
the surface and in the interior (when cut); they give the surface a characteristically 
harsh feel. 

The cortex is about 0°3 mm. thick and contains a great many of the large spherasters, 
for the most part arranged in a single layer at the surface. It is fibrous, and around the 
spherasters the fibres are concentrically arranged. The dark brown pigment granules are 
most abundant in the outermost part of the choanosome, just beneath the cortex, but 
occur also both in the choanosome below and in the cortex above this layer. Numerous 
narrow inhalant canals (pore-canals) penetrate the cortex vertically, each starting above 
from a single dermal pore. These canals are rendered conspicuous by the numerous 
pigment granules that accompany them. Probably they unite in groups on their way 
through the cortex as in other species of Chondrilla. 

The skeleton consists of spherasters (Plate 48, figs. 8a, 8b, 8c) only, varying much in 
size and in the shape of the rays. They are abundantly scattered in the choanosome as 
well as in the cortex. The fully grown ones, measuring about 0°14 mm. in diameter, are 
by far the most abundant. There appears to be no difference between the cortical and 
choanosomal spicules, much the same range of variations occurring in both situations. 
The small ones are probably young forms; I have measured one with a diameter of no 
more than about 0°04 mm.; they have very numerous, simple, smooth, conical rays, 
springing from an enormous centrum, the rays only about 0:°004 mm. long. The large 
ones are of two principal kinds, with intermediates :—(1) with sharp-pointed conical rays, 
somewhat inflated towards the base; often slightly roughened except for the tips of the 
rays (fig. 8a); (2) with the rays truncated and roughened at the ends (fig. 8 6), or even 
reduced to short, subcylindrical projections with roughened ends (fig. 8¢). The inter- 
mediate forms show all degrees of truncation and roughening of the rays*. 

I have already, in discussing the genus Aurorat, given my reasons for believing that 
Carter’s Chondrilla sacciformis is a true Chondrilla, and I need not repeat them here. 
The examination of a type specimen in his cabinet has convinced me that the “Sealark” 
specimen is specifically identical, and there appears little doubt that this is also the case 
with Thiele’s Chondrilla grandistellata. I hardly know what Thiele means by the 
statement that a distinctly differentiated cortex is not present in his specimens, At the 
same time it is true that the line of demarcation between cortex and choanosome is by no 
means everywhere clearly defined in mine, possibly owing to the indifferent histological 
condition of the material. 

The enormous size of the spherasters is very remarkable in comparison with such 
species as C. australiensis, C. mixta, &c. 

Previously known Distribution. Mauritius (Carter) ; Ternate (Thiele). 

Register No., Locality, éc. x1. 4, Saya de Malha, 7.9.05, C. 19, 29 fathoms. 


* The form represented in Fig. 8¢ is probably alone fully grown, all the others being developmental 
stages. 
+ Vide p. 245. 


270 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


PERCY SLADEN TRUST EXPEDITION 


DESCRIPTION OF PLATES 


PLATE 44. 
(On specially toned paper.) 


.1.  Dercitopsis minor n. sp. R.N. xu. 6. Nat. size. 

.2.  Yodomia perfecta n. sp. R.N.x.1. Nat. size. 

. 2a. Yodomia perfecta n. sp. R.N. vi. Nat. size. 

.3.  Myriastra cavernosa n. sp. R.N. vit. 5 A. Nat. size. 

. 3a. Myriastra cavernosa n. sp. R.N. vit. 5B. Cut surface of divided specimen. Nat. size. 
.4. Aurora cribriporosa n. sp. R.N. CxIx. 12. Seen from above. Nat. size. 


Aurora rowi n. sp. R.N. cxxxi. Nat. size. 
Ecionemia laviniensis Dendy. R.N. x1. 2. Nat. size. 


5 
6 

. 7.  Rhabdodragma conulosa (Kieschnick). R.N. Lxxvit. 1a. Nat. size. 
8 


Donatia stella-grandis n. sp. R.N. ovi. 5. Nat. size. 


PuatTe 45. 


.1. Dercitopsis minor n. sp. R.N. CXt. 7. 


la. Calthrops; 10. triods; 1c. oxea; all x 330. 


. 2. Pachastrella tenuwilaminaris (Sollas). R.N. Lx xu. 2. 


2a. Calthrops and short-shafted triznes, x 60; 2b. oxea, x 60; 2c. style, x 60; 2d. strongyle, 
x 60; 2e. hair-like oxea, x 60; 2ff metasters, x 550; 2g. microxeote, x 550. 


. 3. Yodomia perfecta n. sp. 


3a. R.N. x. 1, calthrops, x60; 3a’. R.N. rx. 1, calthrops, x60; 3a”. R.N. rx. 1, reduced 
calthrops, x 60; 36. R.N. x. 1, small calthrops, x60; 306’. R.N. 1x. 1, small calthrops, x 60; 
3c. R.N. x. 1, mesotriznes, x 60; 3c’. R.N. rx. 1, mesotriznes, x 60; 3c’. R.N. vi., mesotrizenes, 
x 60; 3d. R.N. x. 1, oxeote, x 60; 3e. R.N. vi, amphiasters, x 550; 3f. R.N. vr, smooth micro- 
rhabds, x 550; 3g. R.N. vi, spmed microrhabds, x 550. 


PLATE 46. 


1. Myriastra cavernosa n. sp. R.N. vit. 5. 

la. Orthotrizenes, x 60; 1. oxea, x 60; 1c. small oxeote, x 550; 1d. chiasters, x 550. 
2. Aurora providentiv n. sp. R.N. LXXXVI. 2. 

2a. Orthotriznes, x 60; 26. anatrizne, x 60; 2b’. cladome of anatriene, x 770; 26”. cladome 
end of reduced anatriene, x 770; 2c. oxea, x 60; 2c’. small oxeote, x 770; 2d. large spheraster, 
x 770; 2e. small spherasters, x 770; 2 oxyasters, x 770. 


g.3. Aurora cribriporosa n. sp. R.N. Cxrx. 12. 


3a. Oxea, x 60; 306. large spheraster, x 770; 3c. young spheraster(?), x 770; 3d. minute 
spheraster, x 770; 3e. oxyaster, x 770. 
4. Aurora rowi n. sp. R.N. CXXXI. 

4a. Orthotrizenes, x 60; 46. oxeote, x 60; 4c. sterrospheraster, x 770; 4d—4h. stages in the 
development of the sterrospheraster, x 770; 47. small, regular spherasters, x 770; 4k. oxyasters, 
x 770. 
5. Heionemia laviniensis Dendy. R.N. x1. 2. 

5a. Dichotriznes, x 60; 5a’. cladome of dichotrizne, x 60; 5b. anatriene, x 770; 5c. oxea, 
x 60; 5d. chiasters, x 770; 5e. oxyasters, x 770; 5. intermediate forms between oxyasters and 
chiasters, x 770; 5g. microstrongylote, x 770. 
6. Asteropus simplex (Carter) R.N. LXXVIII. 2. 

6a. Oxea, x 60; 6a’. style, x 60; 6b. oxyasters, x 770; 6c. sanidasters, x 770. 
7. Dragmastra lactea (Carter) var. mauritiana noy. R.N. CXXVI. 4 5. 

7 a. Dichotriznes, x 60; 7 6. oxea, x 60; 7c. small oxyspherasters, x 710; 7d. larger oxy- 
spheraster, x 770; 7 e. trichodragmata, x 770. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 
Fig. 


Fig. 


DENDY—HOMOSCLEROPHORA AND ASTROTETRAXONIDA 271 


PLATE 47. 


1. Rhabdodragma conulosa (Kieschnick). R.N. uxxviti. 1. 

la. Plagiotrienes, x60; 16. oxea, x60; 1c. chiasters, x 770; 1d. small oxeote, x 770; 

le. microrhabds, x 770; 1/. trichodragmata, x 770. 
2. Jaspis johnston (Schmidt). R.N. uxxvu. 11. 

2a. Oxea, x 60; 2a’. stylote form, x 60; 2b. microxea, x 770; 2c. oxyasters, x 770. 
3. Geodia auroristella n. sp. R.N. LXxXxvt. 1. 

3 a. Orthotriznes, x 60; 3b. anatrizne, x 60; 3c. oxeote, x60; 3c’. style, x 60; 3c”. strongy- 
lote, x 60; 3d. sterraster, x 310; 3d’—3d”. developmental stages of sterraster, x 310; 3d’”. portion 
of surface of late developmental stage of sterraster, x 310; 3e. smooth spheraster, x 770; 3, spined 
spheraster, x 770; 3g. minute spheraster or chiaster, x 770; 3h. oxyaster, x 770. 

4. Hrylus lendenfeldi Sollas. R.N. cit. 2. 

4a. Orthotrizne, x 60; 4b. oxea, x 60; 4c. aspidasters, x 310; 4c’—40’”. developmental stages 
of aspidaster, x 310; 4c’. marginal portion of surface of aspidaster, x 770; 4d. large oxyaster, 
x 770; 4e. small oxyasters, x 770; 4 f. polyactinose oxyaster, x 770; 49. microrhabds, x 770. 

5. Erylus proximus n. sp. R.N. LXXVIL. 7. 

5a. Short-shafted triznes, x 60; 56. oxea, x60; 50’. style, x60; 506”. strongylote, x 60; 
5¢. aspidasters, x 310; 5¢’—5c’”. developmental stages of aspidaster, x 310; 5¢’’””. marginal 
portion of surface of aspidaster, x 770; 5d. chiasters, x 770; 5¢. microrhabds, x 770. 


PLATE 48. 


1. Donatia lyncuriwm auctorum. R.N. cxx. 2 ¢. 

la. Spherasters from cortex, x 310; 16. spherasters from choanosome, x 310; 1c. chiasters 
from cortex, x 770; 1d. chiasters (oxyasters) from choanosome, x 770. 
2. Donatia japonica (Sollas). R.N. ivi. 10 B. 

2a. Spheraster from cortex, x 310; 2. spherasters from choanosome, x 310; 2c. tylasters from 
cortex, x 770; 2d. tylasters from choanosome, x 770; 2. strongylaster from choanosome, x 770. 
3. Donatia ingalli (Bowerbank). R.N. exit. 1 B. 

3a. Spherasters from cortex, x 310; 3b. spherasters from choanosome, x 310; 3c. tylasters from 
cortex, x 770; 3d. tylaster from choanosome, x 770; 3d’. strongylasters from choanosome, x 770. 
4, Donatia seychellensis (Wright). R.N. cxx. 2 A. 

4a. Spherasters from cortex, x 310; 4b. spherasters from choanosome, x 310; 4c. tylasters 
from cortex, x 770; 4d. oxyasters from choanosome, x 770; 4d’. tylasters from choanosome, x 770. 
5. Donatia stella-grandis n. sp. R.N. vi. 5. 

5a. Styli, x 60; 5a’. tylostyli, x 60; 5a”. strongyle, x 60; 50. spherasters, x 310; 5c. cortical 
chiasters, x 770; 5c’. slender-rayed chiaster, x 770; 5d. choanosomal chiasters, x 770. 
6. Chondrilla australiensis Carter. R.N. xx xml. 3. 

6a. Spheraster from cortex, x 550; 6b. oxyasters from choanosome, x 550. 
1. Chondrilla mixta Schulze. R.N. ox. 4. 

7a. Spheraster from cortex, x 550; 76. oxyasters from choanosome, x 550. 
8. Chondrilla sacciformis Carter. R.N. xt. 4. 

8a—8c. Spherasters, showing variation in shape of rays, x 550. (8a and 86 are probably 
developmental stages of 8c.) 


Perey SLADEN TRUST EXPEDITION. TRANS.LINN. Soc. SER.2.Z00L.VOL.XVII .Pu. 44. 
(DENnDyY) 


W. Champneys del. : Cambridge University Press. 


TETRAXONID SPONGES. 


TRANS. LINN. Soc. SER. 2.Z00L.VOL.XVII PL. 45. 


Percy SLADEN TRUST EXPEDITION. 


(DENDY ) 


Cambridge University Press 


HL Deakin de! 


SEALARK THTRAXONIDA. 


TRANS. LINN. Soc. SER. 2.Z00L. VOL.XVIL .Pr.46. 


PERCY SLADEN TRUST EXPEDITION. 
(Denby) 


Cambridge University Press. 


HL Deakin del. 


SHALARK TETRAXONIDA. 


Percy SLADEN TRUST EXPEDITION. 


TRANS. LINN. Soc. SER. 2.Z00L.VOL.XVIL. Pu 47. 
(Denby) 


SS eS SRE SDD ae ASANTE aan ete 


sel" sewn, 
EE 


zB “a e 


Ze 


HLA ESS 


HL Deakin del. Cambridge University Press 


SEALARK TETRAXONIDA. 


PERcY SLADEN TRUST EXPEDITION. TRANS.LINN. Soc. SER.2.Zo0oL VoL.XVIL.PL. 48. 
(DENDY) 


H.L.Deakin del, Cambridge University Press. 
SEALARK THTRAXONIDA. 


No. VII—RHYNCHOTA. PART Il: SUBORDER HOMOPTERA. 


By Wm. Lucas Distant. 
(CommuNICATED BY J. STANLEY GARDINER, M.A., E.R.S., F.LS.) 


(With Plates 49—51.) 
Read 1st June, 1916. 


IN my previous communication (1913) devoted to the Heteroptera, I was able 
to enumerate 139 species; in this Homopterous section 98 species are detailed, bringing 
up the number of the Seychelles Rhynchota to 237 species. These may again be 
augmented by referring to the first collection made by Prof. Gardinet and detailed 
in these Transactions (1909) in which 3 species of Heteroptera and 3 species of 
Homoptera were included which are not contained in the present collection ; and these 
bring up the total number of species brought home by the two expeditions to 
243 species. A few others (Heteroptera) have been enumerated and described by 
Reuter and Bergroth which have not been found by either Prof. Gardiner or Mr Scott; 
a reference to these papers will be found in the Bibliography further on. 

The Homoptera, especially the smaller forms, are now being assiduously worked 
by such good Homopterists as Melichar, Matsumura and Muir, but even with these 
advantages the material from many large areas is so scanty that little comparison 
in distributional aspects is possible. Of the 98 species of Homoptera contained in 
this collection no fewer than 82 have been regarded as undescribed and have com- 
pelled the erection of 26 new genera to contain them. It would consequently be 
futile to consider that all or a very considerable portion of these new species are 
peculiar to the Seychelles, and it is even more improbable that the new genera should 
be confined to these islands. 

With the Homoptera however a few distributional peculiarities cannot be ignored. 
The large family Cicadide is only represented by two species, one of which is also 
found in, and was described from, Madagascar. The Hawaiian fauna is reported as 
without a representative. The family Fulgoride is poorly represented; neither the 
subfamilies Fulgorine nor Eurybrachydinz are represented in these collections, and 
the Dictyophorinz, Tropiduchine and Achilinee are each represented by one genus 
and one species only. Not a single Membracid was found, and the Membracidze 
appear to be equally absent from the Hawaiian Archipelago. The family Cercopide 
is only represented by one species and that belonging to the subfamily Aphrophorinz. 

SECOND SERIES—ZOOLOGY, VOL. XVII. oo 


274 PERCY SLADEN TRUST EXPEDITION 


It is at present impossible to compare the homopterous relationship of Madagascar 
and the Mascarene Islands with the similar fauna of the Seychelles, for the 
first is still practically unworked and unrecorded except in the larger and more 
conspicuous features, nor judging from the conclusions of other entomologists who 
have worked out different groups of insects is such a relationship likely to be 
pronounced. After working out a very large portion of the homopterous fauna of 
British India, I strongly incline to the view that it is in that and adjoining regions 
that the Seychellian Homoptera find their nearest relationship, though it is not 
a close one, qualified but evident. The material does not at present exist for a 
detailed comparison. 


BIBLIOGRAPHY 


Waker, Fras. Descriptions of a few species scattered throughout the Catalogues of Heteroptera 
and Homoptera published by the Trustees of the British Museum, 1850—1873. The specimens 
were presented by Prof. Percival Wright, and some are labelled “Round Island.” 

KE. Bercroru and O. M. Reurer. “Mission Scientifique de M. Ch. Alland aux iles Seychelles, 
Hétéropteres.” Rev. d’Entomol. xii. pp. 197—209 (1893). 

Livett, Martin. L. “On the insects collected by Dr Abbot on the Seychelles, Aldabra, Gloriosa 
and Providence Islands.” Proc. U.S. Nat. Mus. xix. pp. 695—706 (1897). References to the 
Rhynchota very few. 

VoELTZKOW, ALFRED. “Die von Aldabra bis jetzt bekannte Flora und Fauna.” Abhandl. Senckenb. 
Nat. Ges. xxvi. pp. 541565 (1902). In this publication is (p. 561) a list of 20 species of 
Heteroptera, of which Dr Bergroth has written to inform me that he gave the list of “18,” 
probably an error for “20,” some of which in the “Gerride ” are ascribed to a genus “ Telmaliza,” 
of which I can find neither reference nor description. Dr Voeltzkow also refers to a paper by 
the same writer in the same publication (Bd. xxvii), of which I can find no trace. 

Distant, W. L. “Sealark” Rhynchota. Trans. Linn. Soc., ser. 2, Zool.,:vol. xiii. pp. 29—47, 
Pl. 4 (1909). 

—— “Rhynchota,” Part I, Subord. Heteroptera. Trans. Linn. Soc., ser. 2, Zool., vol. xvi. pp. 139—191, 
Pls. 11—13 (1913). 


Order RHYNCHOTA. 
Suborder Homoptera. 
Family Cicadidee. 


Only two species of this family have been received from the Seychelles and 
possibly they are the only representatives of the family in these islands. Martin 
L. Linell*, writing on the “ Insects from Gloriosa Island,” mentions “a large Cicada 
closely allied to the South African Platypleura limbata Fabr.” which is doubtless the 
“Yanga seychellensis Dist., enumerated in the present paper. Another species which 
he mentions} as ‘‘very much resembling Tettigia orni from Kurope, but smaller, is 


* Proc, U. S. Nat. Mus. xix. p. 702 (1897). 
iy Locwcitaspa Oo Os 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 275 


without doubt Cicada pulverulenta Dist. The latter of these species is found in, 
and was described from, Madagascar. Although Madagascar is richly represented in 
this family, we are only enabled to record two species from the Seychelles. At present 
however the Hawaiian fauna is without a representative, and this is the status of 
many other of the smaller islands in the Seychelles region. 


Subfamily Cicadinee. 
140. Yanga seychellensis. 


Yanga seychellensis Dist., Gen. Insect. Fase. 142, p. 13 (1912); Trans. Linn. Soc. 
London, Zool. xvi. Pl. 11, fig. 22 a, 6 (1918). 


Yanga andriana Dist., Trans. Linn. Soc. London, Zool. xiii. p. 41 (1909). 


Loc. Seychelles. Praslin. Mahé: from near Morne Blanc, Cascade Estate 
(E. G. B. Meade-Waldo). 


I originally recorded this species as Y. andriana from an examination of two 
or three discoloured specimens (supra, 1909), but subsequently Mr Scott supplied a 
number of fresh examples which proved the species to be a distinct one. A pupa- 
case, evidently of this species, is labelled “Silhouette.” Mr Meade-Waldo took this 
species at light. 


141. Cicada pulverulenta. 


Cicada pulverulenta Dist., Trans. Ent. Soc. London, 1905, p. 199; Trans. Linn. 
Soc. London, Zool. xiii. p. 41, Pl. 4, fig. 8a, b (1909). 


Loc. Seychelles. Mahé: from near Morne Blanc and from Cascade Estate, both 
about 800 feet, and other localities, 1905 and 1908—9: frequently attracted by light. 
Aldabra, 1908—9 (Fryer), Madagascar. 


Family Fulgoride. 


This family is not well represented in the Seychelles. Neither of the subfamilies 
Fulgorinz nor Eurybrachydine are found in this collection, and all that can be 
enumerated are moderately small and obscure species. The subfamilies Dictyophorine, 
Tropiduchinee and Achilinze are each represented by one genus and one species only. 


Subfamily Dictyophorine. 


Only one genus, here described as new, can be included in this subfamily. The 
genus Dictyophora so widely distributed is unrepresented in the collection. 


ASELGEOIDES, gen. nov. 


Head produced, elongate, sulcate, eyes elongate, compressed; face elongate, 
centrally and laterally carinate, apex narrowed, widened to the area of the eyes and 
then obliquely narrowed to base of clypeus which is centrally carinate; rostrum 
passing the posterior coxe; pronotum short, posterior margin obliquely concave ; 
mesonotum tricarinate ; posterior tibise with two spines, one before and the other 

; 35—2 


276 PERCY SLADEN TRUST EXPEDITION 


beyond middle, first joint of posterior tarsi longer than the other two joints together, 
tegmina about two and a half times as long as broad, the apical margin rounded, 
veins mostly longitudinal, transverse veins very few, costal margin slightly depressed 
at stigma; wings broader than tegmina, a few transverse veins 6n apical half. 

Allied to Aselgeca Walk. 

142. Aselgeoides insularis, sp. n. (Plate 49, figs. 1, 1 a). 

Head, thorax and abdomen ochraceous, apical area of abdomen more or less 
testaceous ; legs flavescent ; tegmina subhyaline, the veins closely spotted with fuscous, 
the stigma fuscous, anteriorly ochraceous, wings pale fuliginous, the veins darker ; 
structural characters as in generic diagnosis. 

Long. excl. teom. 6 mm. Exp. tegm. 12 mm. 


Loc. Seychelles. Silhouette: near Mont Pot-a-eau, over 1000 feet; marshy 
plateau of Mare aux Cochons, about 1000 feet, and from forest above. Mahé: from 
near Morne Blane; country above Port Glaud, about 500—1000 feet; forest above 
Cascade Estate, 1000 feet; from stunted forest vegetation on summit of Mount 
Sebert, cirea 2000 feet; high damp forest at summit of Morne Pilot, over 2000 feet. 


Subfamily Cixiine. 


Of this abundant subfamily seven genera are here enumerated, only two of which 
were previously known, and five are proposed as new. It is noteworthy that neither of 
the somewhat universally distributed genera, Oliarus and Cixius, is included in this 
collection; Brixia however, another widely spread genus, is found in the Seychelles. 


Genus CANEIRONA™. 


Caneirona Dist., Faun. Brit. Ind. Rhynchota, vol. vi. p. 38 (1916). 


Vertex about as broad as long, the lateral margins strongly carinate, the anterior 
margin with a short central spine, the anterior lateral angles also shortly spinous and 
distinctly extending beyond eyes; face longer than greatest breadth, widened behind 
eyes towards clypeus, anterior margin more or less truncate, posterior margin angularly 
concave with an ocellus on each side, the lateral margins reflexed, centrally strongly, 
longitudinally ridged; clypeus much shorter than face, centrally ridged, laterally 
reflexed; rostrum long, passing the posterior coxee; pronotum very short, posteriorly 
angularly concave; mesonotum about twice as long as vertex and pronotum together, 
tricarinate ; abdomen moderately robust; tegmina twice as long as greatest breadth, 
gradually widened towards apex, costal margin distinctly waved, apices rounded, claval 
vein not reaching apex, apical areas distinctly delineated ; wings shorter than tegmina, 
the broadest areas of each about equal in width, discally shortly transversely veined ; 
posterior tibize unarmed. 

Caneirona is allied to Commolenda Dist., at present only known from Ceylon. 

* This genus is named after Nicolas Caneirio on whose charts the Seychelles appeared in 1502. Since 


this description was written a second species, C. indica Dist., has been received from 8. India and is 
described in the Faun. Brit. Ind. Rhynchota, vol. vi. p. 39, fig. 24 (1916). 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 277 


143. Caneirona maculipennis, sp. n. (Plate 49, fig. 9 a). 


Body and legs ochraceous, paler beneath than above, abdomen sometimes distinctly 
sanguineous ; mesonotum with the disk pale ochraceous, the lateral areas much darker ; 
tegmina subhyaline, the margins very narrowly ochraceous, with numerous small black 
spots, usually arranged about four near base, two near extremity of claval area, three 
in transverse series beyond middle, and a waved series near bases of apical areas ; 
wings hyaline, the veins darker; structural characters as in generic diagnosis. 


Long. excl. team. 45 mm. Exp. tegm. 11 mm. 


Loc. Seychelles. Mahé: Cascade Estate, forest, above 1000 feet; high forest near 
Morne Blane. Silhouette: from near Mont Pot-a-eau, and from Mare aux Cochons. 
Praslin: Cotes d’Or Estate, from Coco-de-Mer forest in the Vallée de Mai. 


CLUSIVIUS, gen. nov. 


Vertex of head about as long as broad, centrally and laterally strongly carinate, 
distinctly projecting beyond eyes, the central carination slightly anteriorly acutely 
produced ; antennze slender and inconspicuous; face longer than broad, centrally and 
laterally carinate, the anterior margin undulate between the apices of the carinations, 
two small ocelli at posterior margin; clypeus short, globose, about one-third the length 
of face, faintly centrally carinate; pronotum tricarinate, a little shorter than head; 
mesonotum tricarinate, shghtly longer than head and pronotum together; posterior 
tibiee unarmed ; tegmina more than twice as long as broad, narrower at base, distinctly 
broadened towards apical area, costal margin slightly convex, apical margin strongly 
convexly rounded, claval margin a little posteriorly produced, veins longitudinal, 
a few short oblique transverse veins on apical costal area, an oblique transverse series 
of veins on subapical area, and an imperfect series of same a little beyond middle ; 
- wings long, only a little shorter than tegmina, two transverse veins beyond middle, one 
on costal area, the other on disk. 


144. Cluswius spectabils, sp. n. (Plate 49, fig. 15 a). 


Head, pronotum and mesonotum black, the latter with a pale ochraceous curved 
fascia occupying the lateral and subposterior margins ; abdomen testaceous, the segmental 
margins darker; head beneath, sternum and legs pale ochraceous; face with broad 
anterior and posterior fasciz, anterior and intermediate tibize and tarsi, and apices of 
posterior tibiz and tarsi, black; tegmina dark shining fuscous, the claval area and a 
large costal stigmatal spot, pale ochraceous ; wings fuliginous, with an oblique subapical 
greyish line beyond middle, commencing about middle of disk and terminating on 
posterior margin; structural characters as in generic diagnosis. 


Long. excl. tegm. 4mm. Exp. tegm. 7 mm. 


Loc. Seychelles. Silhouette: from about 1500 feet, forest above Mare aux Cochons. 
Mahé: from grass in cultivated country, about 1000 feet; Cascade Estate, forest, 
1000-—2000 feet. 


278 PERCY SLADEN TRUST EXPEDITION 


MatUTINUS, gen. nov. 


Vertex of head a little longer than broad, somewhat narrow, concave, a little 
broader at base than at apex, the lateral margins strongly carinate, the apex triangulate ; 
antenne with the first and second joints robust; face very much longer than broad, 
centrally and laterally strongly carinate, the central carination prominent at apex, two 
small ocelli on posterior margin; clypeus nearly half the length of face, centrally 
carinate ; pronotum a little shorter than head, tricarinate; mesonotum slightly longer 
than head and pronotum together, tricarinate, the disk longitudinally flattened ; tegmina 
more than twice as broad as long, the veins few and longitudinal, obliquely triangulate 
beyond middle, the costal membrane broad ; wings considerably broader than tegmina”™; 
posterior tibize unarmed. 


145. Matutinus opulentus, sp. n. (Plate 49, fig. 16 a). 


Body above black; vertex of head (excluding lateral margins), and a central 
fascia to pro- and mesonota, pale ochraceous; basal area of abdomen pale testaceous ; 
body beneath imperfectly seen in the unique carded type; face black; antennz 
ochraceous ; tegmina faintly ochraceous, claval area paler with a dark fuscous spot at 
apex, the apical area somewhat broadly dark fuscous, this coloration angulated on its 
inner margin, before apex a large ill-defined whitish spot on costal margin, a similar 
but smaller spot near apex of inner margin; wings very pale fuliginous, their apices 
(excluding an upper apical pale spot) fuliginous 7. 


Long. excl. tegm. 3mm. Exp. tegm. 8 mm. 


Loc. Seychelles. Mahé: marshes on coastal plain at Anse aux Pins and Anse 
Royale, 1. 1909. 


ADOLENDANA, gen. nov. 


Vertex long, produced in front of eyes, the margins very strongly ridged, the 
disk being thus longitudinally concave, apex triangulate, base strongly concave; face 
elongate, narrow from apex to region of eyes and then ampliated and again a little 
obliquely narrowed to base of clypeus, before which there are two small ocelli, the 
lateral margins strongly ridged and centrally longitudinally carinate; apex truncate ; 
clypeus about half the length of face, laterally ridged and centrally carinate; pronotum 
very short, anteriorly conically produced; mesonotum large, a little longer than head 
and pronotumn together, longitudinally tricarinate; abdomen somewhat laterally com- 
pressed ; posterior tibize without spines; tegmina somewhat narrow at base, ampliately 
widened towards apex which is rounded, the base of costal margin distinctly gibbous, 
costal membrane wider at apex than at base, apical cells elongate and distinctly 


* Since the unique type was figured and before the description was written, the wings were unfortunately 
mutilated in an attempt to reset the carded specimen. The peculiar venation of the wings therefore rests 
on the accuracy of a painstaking and accurate artist. 

+ Cf. note to generic description. 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA | 279 


deliminated, a distinct stigmatical spot; wings shorter than tegmina but broader, 
transversely veined on upper disk. 


Allied to Briaia Stal. 
146. Adolendana typica, sp. n. (Plate 49, fig. 10a). 


Body and legs fuscous-brown, lateral areas of the mesonotum very dark castaneous ; 
tegmina pale fuliginous, the veins darker, two small spots near base, a transverse 
fascia—bifurcating posteriorly—before middle, and the apical area, more or less, fuscous- 
brown, containing some small greyish-white spots, stigmatal spot dark castaneous, 
more or less margined with greyish-white; wings pale fuliginous ; structural characters 
as in generic diagnosis. 

Long. excl. tegm. 34 mm. Exp. tegm. 114 to 12 mm. 


Loc. Seychelles. Silhouette: near Mont Pot-d-eau, all over 1000 feet ; Mare aux 
Cochons, plateau and forest above. Mahé: country above Port Glaud, about 500—1000 
feet; high forest of Morne Blane and Pilot; top of Mount Sebert, nearly 2000 feet ; 
forest above Cascade Estate; Mare aux Cochons district, from forest of rather stunted 
Capucin trees (Northew) on summit of “ Montagne Anse Major,” 2000 feet or over. 


VoOLCANALIA, gen. nov. 


Vertex of head very narrow, appearing more as a longitudinal furrow between 
the eyes, the margins carinately reflexed, scarcely or distinctly projecting in front of 
eyes; antennse very slender; face about as long or a little longer than’ broad, its 
apex truncate or more or less emarginate, narrowed anteriorly, the lateral margins 
either more or less convex, or oblique, centrally strongly carinate, laterally with the 
margins more or less carinate, clypeus broad, varying in length either about as long, 
or a little longer or shorter than face, centrally carinate; tegmina considerably more 
than twice as long as broad, the veins longitudinal to apical area where they are 
above and beneath shortly oblique, and medially. and between them there are two 
or three straight longitudinal veins, the veins are also more or less setigerous; 
wings considerably broader than tegmina, the posterior margin undulate towards the 
anal area; posterior tibize without spines; two small, more or less distinct ocelli at 
basal margin of face. 


Type. V. typica Dist. 
Volcanalia may be placed near Haplaxius Fowler, from Central America. 
’ The members of the genus are all forest species, found sitting on palm leaves, 
&e. (Hugh Scott). 
147. Volcanalia typica, sp. n. (Plate 49, fig. 13 a). 


Head, pronotum, mesonotum and apical half of abdomen, black, basal half of 
abdomen brownish-ochraceous; head beneath and sternum black, legs brownish-ochraceous ; 
tegmina blackish-brown, two costal stigmatal spots, the innermost largest, and an 


280 PERCY SLADEN TRUST EXPEDITION 


opposite spot on inner margin, creamy-white; wings fuliginous with a pale longitudinal 
suffusion ; face slightly shorter than clypeus, its apical margin truncate, only slightly 
projecting before eyes; mesonotum tricarinate; tegmina with the veins distinctly 
setigerous. 


Long. excl. tegm. 3 mm. Exp. tegm. 8 mm. 


Loc. Seychelles. Mahé: country above Port Glaud, about 500—1000 feet ; forest 
above Cascade Estate; top of Mount Sebert, nearly 2000 feet. 


148. Volcanalia atrostriata, sp. n. (Plate 50, fig. 21 a). 


Head, pronotum and mesonotum ochraceous; mesonotum with a central longi- 
tudinal fascia and the lateral areas, castaneous-brown; body beneath and legs very 
pale ochraceous; lateral margins and two central fascize to face, two central fascize 
to clypeus, black, apices of tibize and tarsi more or less pale brownish; tegmina pale 
castaneous-brown, the lateral and central area largely greyish-white, apical third more 
or less ochraceous, its inner and apical margins, and a costal spot beyond middle, 
pale castaneous-brown ; vertex of head narrow between the eyes and roundly produced 
a little in front of them, its lateral margins distinctly, darkly, carinately reflexed ; 
face about as long as clypeus, anteriorly truncate, a little anteriorly centrally produced, 
the lateral margins convexly ampliated; mesonotum finely tricarinate; tegmina with 
the veins obscurely setigerous. 

Long. 44 mm. 

Loc. Seychelles. Mahé: high damp forest at summit of Pilot, over 2000 feet ; 
high forest. behind Trois Freres, 1500—2000 feet. 

I have seen a variety of this species in which the two central black fascize to 


the face are coalesced. 
149. Volcanalia atrovaria, sp. n. (Plate 50, fig. 18 a). 


Head, pronotum and mesonotum pale ochraceous, lateral areas of mesonotum 
distinctly darker; body beneath and legs pale ochraceous; sublateral margins and two 
central narrow longitudinal fascize to face, a central spot at base and the apex of 
clypeus, and the anal abdominal appendage to female, black; tegmina greyish-white, 
more or less tinted with ochraceous, inner and outer basal marginal spots, two 
similarly placed spots beyond middle, a transverse line before apical third, a strongly 
curved longitudinal line to same and the apices of the apical veins, pale castaneous- 
brown; vertex narrow between eyes and roundly continued a little in front of same, 
its lateral margins distinctly carinately reflexed ; face about as long as clypeus, anteriorly 
truncate, a little anteriorly centrally produced, the lateral margins convexly ampliated ; 
mesonotum finely tricarinate; tegmina with the veins obscurely setigerous. 


Long. 4 mm. 


Loc. Seychelles. Silhouette: near Mont Pot-a-eau, about 1500 feet; high forest 
above Mare aux Cochons. Mahé: high forest of Morne Blanc; Mare aux Cochons 
district, 1000—2000 feet; Cascade Estate, forest, 1000—2000 feet. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 281 


150. Volcanalia designata, sp. n. 


Body and legs ochraceous; eyes black; face and clypeus ochraceous, the latter 
with a spot at base and the apex, black ; tegmina hyaline 
much suffused with ochraceous, a slightly curved and 
broken transverse fascia at about two-thirds from base, 
from which, near middle, a longitudinal fascia extends 
to apex, and the apices of the apical veins, black ; vertex 
narrow between eyes and a little roundly continued 
beyond them, the lateral margins distinctly ridged; face 
distinctly produced beyond eyes, its apex subtruncate, 
distinctly ampliated and rounded before clypeus, which pig 1 
is about as long as face; mesonotum tricarinate. 


. Volcanalia designata Dist. 


Long. 44 mm. 


Loc. Seychelles. Silhouette: high country near Mont Pot-i-eau. 


151. Voleanalia cardm, sp. n. (Plate 50, fig. 20 a). 


Vertex of head sanguineous; pro- and mesonota pale ochraceous, the latter with 
the lateral areas and a central fascia pale castaneous-brown, the central fascia 
medially traversed by a sanguineous line ; body beneath and legs pale ochraceous ; 
face with an arcuated transverse fascia, extending to anterior margin, and clypeus 
with a narrow basal fascia, sanguineous; abdomen with lateral marginal black spots; 
tegmina with the basal two-thirds greyish, suffused with ochraceous and bluish-grey 
markings, apical third pale ochraceous, anteriorly margined with a curved fuscous 
transverse fascia, and inwardly and posteriorly margined with fuscous spots, a large 
white spot on its disk and a smaller white and fuscous spot on costal margin at about 
one-fourth from apex; vertex narrow between eyes and distinctly projecting beyond 
them, its lateral margins carinately reflexed ; face a little longer than clypeus, with the 
anterior margin subtruneate, centrally moderately excavate, its lateral margins moderately 
oblique, moderately produced in front of eyes; mesonotum finely tricarinate; tegmina 
with the veins obscurely setigerous. 


Long. 5 mm. 


Loc. Seychelles. Mahé: all from stunted forest vegetation at the top of Mount 
Sebert, nearly 2000 feet, i. 1909. 


152. Volcanalia picturata, sp. n. (Plate 50, fig. 19 a). 


Vertex of head, pro- and mesonota more or less fuscous-brown, with the margins 
and carinations paler; abdomen above black, the basal area and segmental margins 
testaceous; head beneath, sternum and legs pale ochraceous, abdomen beneath black, 
the segmental margins ochraceous, face with an arcuated anterior fascia, and clypeus 
with a very broad fascia, sanguineous; tegmina very pale dull ochraceous, the veins, 
three more or less oblique fasciz and some apical spots, dull dark castaneous ; wings 
pale fuliginous, with darker longitudinal shadings ; vertex very narrow between eyes, its 

SECOND SERIES—ZOOLOGY, VOL. XVII. 36 


282 PERCY SLADEN TRUST EXPEDITION 


margins strongly carinately reflexed, slightly passing anterior margins of eyes; face 
slightly longer than clypeus, with the lateral margins moderately oblique, its anterior 
margin notched and a little passing eyes; mesonotum finely tricarinate; tegmina with 
the veins distinctly setigerous. 

Long. excl. tegm. 35 mm. Exp. tegm. 10 mm. 

Loc. Seychelles. Silhouette: high forest near Mont Pot-a-eau ; Mare aux Cochons, 
and forest above. Mahé: near Morne Blanc, about 1000 feet ; high damp forest between 
Trois Fréres and Morne Seychellois, about 1500—2000 feet; Cascade Estate, 800— 
1000 feet; Mare aux Cochons district, 1000—2000 feet. 


153. Volcanalia varicolor, sp. n. (Plate 50, fig. 16 a). 

Head, pronotum and mesonotum pale dull ochraceous, posterior margin of pro- 
notum, and apex and carinations to mesonotum stramineous; face pale stramineous, an 
arcuated transverse fascia between eyes and a very large basal spot to clypeus, 
sanguineous; sternum testaceous, legs ochraceous, anterior and intermediate legs with 
annulations to tibie and the tarsi wholly black; abdomen above black, the base 
testaceous ; tegmina dull ochraceous, the veins darker, a pale stigmatal spot on each 
side of which the costal margin is dark fuscous, and a smaller pale spot opposite the 
costal one, on inner margin; wings pale fuliginous; vertex narrow, the lateral margins 
carinately reflexed, distinctly extended beyond eyes; face distinctly longer than clypeus, 
with the lateral margins oblique, the anterior margin distinctly angulate and moderately 
extending in front of eyes; mesonotum finely tricarinate; tegmina with the veins 
distinctly setigerous. 

Long. 5 mm. 

Loc. Seychelles. Silhouette: high forest above Mare aux Cochons. Mahé: 
country above Port Glaud, about 500—1000 feet; high forest behind Trois Fréres ; 
Mare aux Cochons district, 1000—2000 feet; Cascade Estate, about 1000 feet; 
forest near Mount Harrison, 1700 feet; high damp forest at summit of Pilot, over 
2000 feet. 


154. Volcanalia fumosa, sp. n. (Plate 50, fig. 17 a). 


Head, pronotum and mesonotum testaceous, the carinations paler; face, sternum 
and legs pale ochraceous, anterior area of face and basal margin of clypeus sanguineous ; 
abdomen beneath piceous or black, the segmental margins ochraceous; tegmina with 
the basal two-thirds piceous or blackish (reflecting the dark abdomen beneath), the 
veins prominent, the costal margin, gradually widening posteriorly, and the whole 
of the apical area more or less pale brownish-ochraceous with the veins darker, a 
series of prominent spots to apical margin, preceded by two larger spots on costal 
margin, pale dull castaneous; vertex narrow between eyes, and distinctly slightly 
projecting beyond them, its lateral margins carinately reflexed, mesonotum distinctly 
tricarinate; face about as long as clypeus, with the lateral margins slightly convex, 
anteriorly distinctly projecting before eyes; tegmina with the veins distinctly setigerous. 


Long. 4 mm. 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 283 


Loc. Seychelles. Silhouette: near Mont Pot-d-eau, about 1500 feet; Mare aux 
Cochons, and forest above. Mahé: from high forest of Morne Blane and Pilot, up to 
2000 feet ; country above Port Glaud, about 500—1000 feet ; high damp forest between 
Trois Fréres and Morne Seychellois, about 1500—2000 feet; forest above Cascade 
Estate; Mare aux Cochons district, 1000—2000 feet. 


155. Volcanalia modesta, sp. n. (Plate 50, fig. 15 a). 


Vertex of head sanguineous; pro- and mesonota ochraceous; body beneath and 
legs pale ochraceous; face ochraceous, the apex sanguineous; tegmina very pale 
ochraceous, towards base reflecting the dark abdomen beneath, the margins of the 
apical area spotted with fuscous; vertex very narrow between eyes and projecting a 
little beyond them; mesonotum distinctly tricarinate; face a little longer than clypeus, 
with the lateral margins moderately oblique, its apex distinctly continued before 
eyes and moderately emarginate; tegmina with the veins distinctly setigerous. 


Long. 4 mm. 


- Loc. Seychelles. Silhouette : high country near Mont Pot-d-eau. Mahé: from country 
above Port Glaud, about 500—1000 feet; high damp forest at summit of Morne Pilot, 
over 2000 feet; high damp forest between Trois Fréres and Morne Seychellois, about 
1500—2000 feet; Mare aux Cochons district, 1000—2000 feet ; Cascade Estate, forest, 
1000—2000 feet. 


156. Volcanala uniformis, sp. n. 


Body and legs ochraceous; tegmina subhyaline with the veins stramineous, but 
when. unexpanded reflecting the ochraceous body beneath; face ochraceous, more or 
less tinted with sanguineous, a waved oblique narrow sanguineous fascia commencing 
near anterior margin and continued on lateral areas of sternum; vertex narrow 
between eyes and distinctly projecting beyond them, its lateral margins distinctly 
ridged; face a little longer than clypeus, broadened and moderately convex towards 
clypeus, centrally and laterally carinate; pronotum strongly tricarinate; venation of 
the teemina somewhat coarse. 


Long. 5 mm. 


Loc. Seychelles. Silhouette: from high country. Mare aux Cochons, and forest 
near by. Mahé: country above Port Glaud, about 500—1000 feet, forest above 
Cascade Estate and forest near Mount Harrison, 1700 feet. 

Allied to the previous species V. modesta. 


157. Volcanalia capitata, sp. n. (Plate 51, fig. 2 a). 


Vertex of head black; pronotum and mesonotum pale fuscous-brown; face pale 
fuscous or blackish, the apex sanguineous; body beneath and legs dull ochraceous ; 
tegmina ochraceous, the apical area paler, base of costal margin, an angulated trans- 
verse fascia before apical area which has two spots on its outer margin and a series 
of smaller spots on its inner margin, fuscous-brown; vertex very narrow between eyes 
and very distinctly projecting beyond them; mesonotum distinctly tricarinate; face a 

36—2 


284 PERCY SLADEN TRUST EXPEDITION 


little longer than clypeus with its lateral margins moderately simuately oblique, its 
apex truncate, and continued some distance before eyes, both face and clypeus strongly 
centrally carinate. 


Long. incl. tegm. 4 mm. 


Loc. Seychelles. Silhouette: from high forest above Mare aux Cochons. 


Genus Brixta. 
Brivia Stal, Ofv. Vet.-Ak. Férh., 1856, p. 162. 


158. Briaia mahensis, sp. n. (Plate 49, fig. 14a). 


Body and legs ochraceous, lateral areas of pronotum very pale ochraceous ; in- 
terior area of face, behind eyes, a little darker; tegmina semihyaline, the veins darkly 
setigerous, two oblique transverse fuscous fascize (one basal directed outwardly, the 
other near middle directed inwardly), two blackish spots between these fasciz, an 
arcuate blackish line before apical area, a transverse fascia in apical area and part of 
the inner apical margin fuscous, the latter with small marginal whitish spots, an 
ochraceous stigmatal costal spot; wings pale fuliginous, the venation darker; vertex 
of head narrow between the eyes, the lateral margins carinately reflexed ; face centrally 
excavate, the lateral margins moderately convexly curved and very strongly carinately 
reflexed, a distinct pale ocellus at its base; mesonotum tricarinate. 

Long. excl. tegm. 4mm. Exp. tegm. 14 mm. 


Loc. Seychelles. Mahé: from grass in cultivated country, about 1000 feet. 
A single specimen. 


159. Brixia stellata, sp. n. (Plate 49, fig. 17 a). 


Body and legs ochraceous; vertex of head carinations to pro- and mesonota and 
legs, very pale ochraceous ; face with the interior area more or less castaneous ; tegmina 
subhyaline, with an oblique fasciate-line near base, two subcostal spots before middle, 
a curved line before apical area, preceded by two oblique costal limes and some other 
discal suffusions, and an inner apical marginal fascia, fuscous, a distinct black spot 
near inner side of apical margin; wings very pale fuliginous, the veins darker; vertex 
of head distinctly extending beyond eyes, the lateral margins strongly carinately 
reflexed ; face widened towards clypeus, the lateral margins strongly carinately reflexed, 
a distinct pale ocellus at its base; mesonotum tricarinate. 

Long. excl. tegm. 4mm. Exp. tegm. 13 mm. | 


Var. a. Darker in hue, the ground colour of the tegmina being subopaque and 
the markings more dark castaneous. 


Var. b.  Paler in hue than in type; all the markings more ill-defined, some 
absent. 


Loc. Seychelles. Silhouette: from Mare aux Cochons, about 1000 feet. Mahé: 
high forest of Morne Blane; country above Port Glaud, about 500—1000 feet; high 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 285 


damp forest at summit of Pilot, over 2000 feet; high damp forest between Trois 
Fréres and Morne Seychellois, about 1500—2000 feet ; Cascade Estate; top of Mount 
Sebert, nearly 2000 feet; Mare aux Cochons district, 1000—2000 feet.  Praslin : 
from Cotes d’Or Estate. 


This species is found in great numbers in undergrowth in the damp mountain- 
forests, often in the endemic bush, Senecio seychellensis (Hugh Scott)*. 


CURIATIUS, gen. nov. 


Vertex of head moderately projecting in front of eyes, the apex centrally 
emarginate, the lateral apical angles moderately prominent, and deflected, centrally 
ridged, broadened at base between the eyes; face distinctly projecting before eyes, 
widened beneath eyes, the margins moderately convex and carinately reflexed, two 
small ocelli at base; clypeus elongate, almost as long as face, centrally carinate ; 
antennze with the first and second joints globosely thickened; pronotum very much 
shorter than head, posteriorly deeply emarginate, centrally ridged, mesonotum about 
as long as head and pronotum together, tricarinate; posterior tibize unarmed ; tegmina 
more than twice as long as broad, two series of transverse veins on apical half, the 
inner series less continuous, veins mostly setigerous, especially on claval area; wings 
with a few upper discal transverse veins. 


Allied to Briaia but differing by the long clypeus, two ocelli at base of face, &c. 


160. Curiatius insignis, sp. n. (Plate 49, fig. 12 a). 


Body above brownish ochraceous, abdomen more or less suffused with white pile ; 
body beneath and legs pale ochraceous, lateral areas of sternum more or less greyish ; 
face castaneous, the lateral margins greyish; central carination to clypeus greyish ; 
tegmina pale castaneous-brown, basal area of costal margin, a large costal stigmatal 
spot, and large apical marginal spots, pale dull ochraceous, on apical area the nerval 
interspaces are more or less greyish, to a little beyond middle the veins are darkly 
setigerous and on claval area this character is particularly marked, including the 
inner margin; wings very pale fuliginous; other structural characters as in generic 


diagnosis. 
Long. excl. tegzm. 4mm. Exp. tegm. 14 mm. 


Loc. Seychelles. Mahé: Cascade Estate, 800—1000 feet. 


A single example. 


* The association of Brixia stellata, and of two species of Wisia (WV. sulphurata and fuscofasciata), with 
low-growing plants, and particularly with the endemic bush-groundsel, Senecio seychellensis, was remarked on 
many occasions in the damp mountain-forests. These small Homoptera appeared often to frequent this 
plant in great numbers. Compare the case of a new genus and species of Lamellicorn beetle, Mesohoplia 
senecionis, which was exclusively found on the Senecio.—H. Scorr. 


286 PERCY SLADEN TRUST EXPEDITION 


Subfamily Tropiduchine. 
DARADAXOIDES, gen. nov. 


Head long, strongly produced, longer than pronotum, almost as long as mesonotum, 
‘the lateral margins somewhat obliquely straight from base to anterior margins of eyes 
and then obliquely narrowed to apex which is truncately rounded, the margins strongly 
ridged, and a central longitudinal ridge; eyes longer than broad; face long, centrally 
and laterally ridged, slightly concave at region of eyes, obliquely narrowed before apex ; 
clypeus short, broad, centrally ridged; pronotum short, discally, conically produced 
between the eyes where it is centrally and laterally ridged, depressed at lateral areas, 
the lateral angles prominent behind eyes, posterior margin angularly concave; meso- 
notum strongly, discally tricarinate; posterior tibiee with three spines a little beyond 
middle; tegmina a little more than twice as broad as long, costal membrane slightly 
broader than radial area, the first and apical margin closely obliquely veined, the 
second more sparingly veined, the discal veins longitudinal for about two-thirds from 
base, the subapical area more or less reticulate ; wings shorter than tegmina, the 
posterior margin moderately waved, longitudinally veined except on apical area where 
the veins are more or less furcate. 


Allied to Daradax Walk., a genus represented in Malacca and Borneo. 


161. Daradaxoides mahensis, sp. n. (Plate 49, fig. 5 a). 

Body and legs pale virescent; eyes a little darker and more testaceous in hue; 
tegmina semiopaque, the veins pale virescent; wings hyaline. Structural characters 
as in generic diagnosis. 

Long. excel. tegm. 54 to 7 mm. Exp. tegm. 12 to 15 mm. 

Loc. Seychelles. Mahé: 5 specimens; from near Morne Blanc; Cascade Estate ; 
top of Mount Sebert, nearly 2000 feet; Mare aux Cochons district, 1000—2000 feet. 


Subfamily Achiline. 
PARAKOSALYA, gen. nov. 


Head including eyes a little narrower than pronotum, vertex longer than broad, 
distinctly projecting before eyes, its margins carinate and with a distinct central ridge ; 
face longer than broad, towards clypeus moderately ampliate, laterally and centrally 
ridged ; clypeus about half as long as face with a strong central ridge and its lateral 
margins cariate; pronotum short, centrally tricarinate, posterior margin strongly 
angularly emarginate; mesonotum, slightly longer than vertex and pronotum, tri- 
carinate ; legs long and slender, posterior tibize with one spine before middle, posterior 
tarsi with the basal joint very long; tegmina apically moderately widened, distinctly 
ampliate behind the clavus, two series of more or less complete transverse veins on 
apical half, wings considerably wider than tegmina. 

Allied to the Oriental genus Kosalya Dist., from which it differs by the longer 
and more projecting vertex of head, the different shape of the face, the shorter meso- 
notum, and only one spine to the posterior tibize. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 287 


162. Parakosalya imsularis, sp. n. 

Head, pronotum, mesonotum, base of abdomen, body beneath and legs ochraceous ; 
abdomen above, excluding base, sanguineous ; 
tegmina creamy-ochraceous, the costal mem- 
brane much paler; wings pale fuliginous ; 
vertex with two small black impressions 
between the eyes; abdomen above with a 
distinct central ridge ; other structural charac- 
ters as in generic diagnosis. 


Long. excl. tegm. 3mm. Exp. teem. 8 mm. 


Loc. Seychelles. Silhouette: forest above 
Mare aux Cochons. 


Subfamily Derbine. 


Of the six genera here enumerated four appear to be unknown and are here 
described. There can be no doubt that, as has been remarked by other workers, 
the Derbinze are closely related to the Cixiine, and in some instances it is difficult 
to draw the line between the two groups or subfamilies. Those described here are 
all small species for which we are indebted to the assiduous collecting of Mr Hugh Scott 
and will in all probability be again discovered, either generically or specifically in 
Madagascar and the adjacent islands. 


IGuvriuM, gen. nov. 


Vertex of head narrow, prominently projecting before the eyes, widened towards 
base, with eyes very much narrower than pronotum; antennee with the second joint 
of moderate length but strongly incrassate; face narrow, the margins ridged, strongly 
projecting before eyes; clypeus short and broad, shorter than face; pronotum short 
and broad, the anterior margin sinuate, the lateral angles subacute, posterior margin 
moderately concavely sinuate; mesonotum about as long as pronotum and_ head 
together, moderately narrowed posteriorly; tegmina more than twice as long as 
broad, narrow at base, inner margin gradually arched from base to about middle where 
the tegmen is roundly broadened, costal membrane with three transverse veins, 
remaining venation better seen in the enlarged figure here given; wings short and 
narrow. 

Two carded specimens afford small opportunity for detailed structural charac- 
teristics. 


The genus is allied to Pamendanga Dist. 


163. IRguvium albomaculatum sp. n. (Plate 51, fig. 5a). 

Body and legs ochraceous with darker mottlings; eyes black; pronotum with 
two central and two sublateral fuscous spots; mesonotum with similar anterior spots 
and two larger dark castaneous posterior spots; tegmina subhyaline, the venation 


288 PERCY SLADEN TRUST EXPEDITION 


greyish-white, crossed by five fuscous fascize of which the three basal are narrowest, 
and all containing more or less prominent greyish-white spots, on costal margin near 
apex are some distinct testaceous markings; wings subhyaline, other structural 
characters as in generic diagnosis. 

Long. incl. tegm. 5 mm. 

Toc. Seychelles. Silhouette: Mare aux Cochons plateau. Mahé: Cascade Estate, 
about 800 feet. 

The description of this very distinct species has been derived from two carded 
specimens, both of which have been used for that purpose. 


Genus FESCENNIA. 


Fescennia Stal, Hem. Afr. iv. p. 198 (1866). 


This genus was founded by Stal for the reception of a species described by Coquerel, 
as Phenice bivittata from Mayotta Isld. and Madagascar. Stal however renamed it as 
F. laticeps without giving any reason. Coquerel’s description 1s much more obscure than 
that of Stal’s, but it was published in 1859, and should take precedence. I have 
not seen the type of the genus, but the two species here described seem clearly 
to belong to Mescennia. 

164. Fescennia bimaculata, sp. n. (Plate 51, fig. 11 q). 

Body and legs pale ochraceous, more or less greyishly pubescent ; eyes testaceous ; 
tegmina palely flavescent, a black spot near apex and a similar spot near middle of 
inner margin; wings subhyaline, with some opalescent reflexions; head including eyes 
about as wide as pronotum, vertex broad, triangular, the apex emarginate distinctly 
projecting beyond eyes ; elypeus long, tricarinate, a little shorter than face; second 
joint of antennze very short ; mesonotum tricarinate, the sublateral carinations obscure ; 
tegmina elongate, about three times longer than broad, beyond medium a little broadened, | 
about three times longer than clavus; wings about a third part shorter than tegmina. 

Long. excl. tegm. 4 mm. Exp. tegm. 13 mm. 

Loc. Mahé: Mare aux Cochons district, 1000—2000 feet. 

A single example only contained in the collection. 

165. Fescennia aurea, sp. n. (Plate 51, fig. 10a). 

Body and legs pale ochraceous, more or less greyishly pubescent; tegmina 
flavescent with a black spot near middle of inner margin and the apical margin 
narrowly fascescent; wings subhyaline with opalescent reflexions. Allied to the 
preceding species F. bimaculata, but the face a little broadened posteriorly, clypeus 
a little narrower; some minor differences in venation of tegmina better shown than 
described in the figures of the two species. 

Long. excl. tegm. 4mm. Exp. tegm. 14 mm. 

Loc. Seychelles. Silhouette; Mare aux Cochons. Mahé: Cascade Estate, about 
800—1500 feet. A single example from each locality. 


DISTANT—RHYNOCHOTA. PART II: SUBORDER HOMOPTERA 289 


AQUALICIUM, gen. nov. 


Vertex of head narrow, projecting beyond the eyes, broadened towards base which 
is moderately angularly excavated ; eyes large, oblong, reaching base of head; antenn 
long, first and second joints strongly incrassated, basal joint short, second joint long, 
about as long as mesonotum, subdepressed ; face long, very narrow and slender projecting 
beyond the eyes; rostrum with the second joint nearly twice as long as the first, 
the apical joint minute; pronotum short, anteriorly subconically produced, posteriorly 
moderately concave, the lateral angles a little angularly produced; mesonotum a little 
longer than head and pronotum together, obscurely tricarinate, the central carination 
distinct, the sublateral carinations oblique, short, and more or less obscure; posterior 
tibie not spined, posterior tarsi with the basal joint about as long as the remaining 
jomts together; tegmina about two and a half, or three times as long as broad; costal 
membrane broad, obliquely transversely veined beyond middle, and again before apex, 
principal veins longitudinal, a submarginal series of transverse veins, and beneath the 
second longitudinal interspace, and at the apices of the third and fourth shorter inter- 
spaces are a series of short oblique veins, apical margin rounded, inner margin concavely 
sinuate before apex; wings about as broad and little shorter than tegmina, two trans- 
verse veins on upper disk. 


166. Aquelicvwm typicum, sp. n. (Plate 49, fig. 11a). 

Body above fuscous-brown; vertex of head and a central longitudinal fascia to 
pro- and mesonota, dull greyish; body beneath and legs ochraceous, apical area of 
abdomen beneath more or less castaneous; tegmina pale fuscous-brown, apices of the 
upper apical veins sanguineous, a large spot crossing costal membrane beyond middle, 
a smaller spot near apex, apices of the lower apical veins and spots on mner margin, 
greyish-white; wings pale fuliginous, the veins darker; basal incrassated joints of 
antennee dark fuscous; face prominently extending before eyes which are black; other 
structural characters as in generic diagnosis. 

Long. excl. tegm. 2 mm. Exp. tegm. 7 mm. 

Loc. Seychelles. Silhouette: Mare aux Cochons. Mahé: near Morne Blanc; 


country above Port Glaud, about 500—1000 feet; Cascade Estate, and forest above ; 
top of Mount Sebert, nearly 2000 feet. 


167. Aquelicium elegantulum, sp. n. (Plate 51, fig. 9 a). 


Body above dark fuscous-brown ; vertex of head and a central fascia to pro- and 
mesonota, ochraceous; body beneath fuscous-brown ; clypeus and legs ochraceous, the 
latter tinted with pale sanguineous; tegmina very dark fuscous-brown, the costal 
and apical margins sanguineous, a spot near apex of clavus, two transverse lines on 
inner margin and transverse lines on inner apical margin, pale ochraceous ; vertex of 
head distinctly prominent; incrassated second joint of antennze very dark fuscous and 
more slender than in the previous species A. typicwm. 

Long. 35 mm. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 37 


290 PERCY SLADEN TRUST EXPEDITION 


Loc. Seychelles. Silhouette: Mare aux Cochons. Mahé: high forest of Morne 
Blane and Pilot; forest above Cascade Estate; Mare aux Cochons district.  Félicité 
Island: from forest. 


168. Agqueliciwum brunnescens (Plate 51, fig. 7 a). 


Body above fuscous-brown ; vertex—excluding apex—ochraceous, an obscure pale 
longitudinal fascia to pro- and mesonota; body beneath fuscous, clypeus and legs 
ochraceous, tegmina pale fuscous, the veins darker; vertex of head distinctly prominent; 
incrassated second joint of antennee very dark fuscous, in shape and substance much 
as in preceding species A. elegantulum from which it differs by its different coloration 
and pattern. 


Long. 34 mm. 
Loc. Seychelles. Silhouette: high country near Mont Pot-a-eau. Mahé: near 
Morne Blanc; Cascade Estate, about 800 feet and over. 


HQUIRRIA, gen. nov. 


Allied to the previous genus Aquelicium, especially by the strongly incrassated 
first and second joints of the antennze, the second joint very long, but differing in the 
following characters— Vertex of head considerably broader, subtriangular, excavate, apex 
subtruneate; face broader, projecting similarly before eyes, but with the apex a little 
broadened, the lateral margins strongly elevately carinate; clypeus shorter, only about 
half the length of face; tegmina about two and a half times as long as broad, costal 
membrane broad, with two oblique transverse veins beyond middle, principal veins 
oblique not longitudinal with a sub-inner series of transverse veins, inner margin not 
distinctly concavely sinuate before apex; wings distinctly narrower than tegmina, two 
transverse veins on upper disk as in Agqueliciwm. 


169. Equirria phalena, sp. n. (Pl. 51, fig. 6 a). 

Body and legs pale dull ochraceous, more or less greyishly pubescent ; antennze pale 
castaneous; tegmina subhyaline, a subcostal fascia, an oblique fascia near base, a 
discal, elongate, oblique spot, a curved fascia near apical angle, and a subapical 
marginal line, either pale or dark fuscous; wings subhyaline, posterior halves of the 
veins on basal half, infusecated; vertex of head triangular moderately projecting 
beyond the anterior margins of the eyes; other structural characters as in generic 
diagnosis. 

Long. excl. tegm. 34 mm. Exp. tegm. 155 mm. 

Loc. Seychelles. Mahé: from near Morne Blane, ca. 800—1000 feet. 


ForRDICIDIA, gen. nov. 


Allied to the two preceding genera Aquelicium and Equrria by the strongly 
incrassated basal joints of the antenne, the second joint very long; it is allied to 
Aquelicium by the longitudinal veins to the tegmina and the slender narrow face, 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 29 1 


but it differs from that genus by the shorter and broader head and by the apex of 
the face being broader, less produced before eyes and emarginate; from Kqwrria it 
differs by the longitudinal direction of the principal veins to the tegmina, the narrow 
face and the long clypeus, nearly as long as the face, but is allied to that genus by 
the shorter and broader vertex of head; the tegmina are apically more narrowed than 
in either of the above genera, for the venation of which the three figures given for 
the three genera give a better guide than does a written description. 


170. Fordicidia robusta, sp. n. (Plate 51, fig. 15 a). 


Body and legs flavescent; tegmina subhyaline, much suffused with pale fuscous, 
in typical specimens as the one figured—three basal spots crossing claval area, two 
longitudinal spots above them, followed by an irregular transverse series of spots and 
subapical shadings, in some specimens these markings are much obliterated and the 
spots diminished in number; wings subhyaline, the veins darker ; vertex of head short, 
triangular, but with eyes considerably narrower than pronotum ; mesonotum somewhat 
obscurely tricarinate ; body short and robust; face narrow, a little widened towards 
clypeus, its lateral margins strongly ridged; second joint of antennze long and in- 
crassated, other structural characters as in the generic diagnosis. 


Long. excl, tegm. 2 to 25mm. Exp. tegm. 10 mm. 


Loc. Seychelles. Silhouette: forest above Mare aux Cochons. Mahé: Cascade 
Estate, 800—1000 feet and over; high forest behind Trois Fréres, 1500—2000 feet. 


Genus NIstA. 


Nisia Melich., Hom. Faun. Ceylon, p. 53 (1903); Dist., Faun. Brit. Ind. Rhynch. 
ili. p. 809 (1906). 

The position of this genus is still swb judice; Melichar placed it in the Derbine 
and I followed him, though of course its aberrant structure among the Derbids cannot 
be overlooked. Muir (Bull. H.S. P. Ent. xii. p. 29, 1913) writes of Misia and two 
other genera: “I exclude them from the family Derbide, but, inconsistently, have 
included them in this Bulletin because several of our greatest authorities on Homoptera 
have considered them as Derbids.” Muir seems inclined to the view that Nzsza belongs 
to the Cixiine. Matsumura (Ann. Mus. Nat. Hung. xii. p. 287, 1914) places the genus 
in the Achilinee. 

Under these qualifications I leave Misia among the Derbids, a position which 
perhaps does not seem congenial and from which it will probably be ultimately 


transferred. 


171. Nisia atrovenosa. 

Meenoplus atrovenosus Leth., Ann. Mus. Genov. xxvi. p. 466 (1888). 

Nisia atrovenosa Melich., Hom. Faun. Ceylon, p. 53 (1903); Dist., Faun. Brit. 
Ind. Rhynch. iii. p. 309, fig. 150 (1906); Muir, Rept. Exp. Stat. Haw. 8. Plant. Assoc. 


Bull. 12, p. 81 (1913). 
37—2 


292 PERCY SLADEN TRUST EXPEDITION 


Loc. Seychelles. Mahé: Port Victoria, from Botanic Gardens; near Morne Blane ; 
marshes on coastal plain at Anse aux Pins and Anse Royale; Cascade, marshy culti- 
vated country near sea-level. Félicité. Praslin. Silhouette: from the marsh at Mare 
aux Cochons. Found on water-plant which grows in marshes. 


This species is also recorded from Trincomalee, Ceylon, Nias Islands, Borneo, Piroe 
and Formosa. 


172. Nisia sulphurata, sp. n. 


Body and legs pale ochraceous ; tegmina sulphur-yellow ; wings fuliginous, the veins 
darker ; head including eyes narrower than pronotum, vertex and face continuous, their 
lateral margins strongly carinate and elevate, vertex of head longitudinally concave ; 
face longer than broad; clypeus longitudinally centrally ridged; pronotum centrally 
longitudinally carinate ; mesonotum obscurely tricarinate; tegmina more than twice as 
broad as long, costal margin distinctly arched on basal half, apical area with five 
longitudinal veins, the upper connected near base, as in NV. atrovenosa. 


Long. excl. tegm. 24 to 3mm. Exp. tegm. 10 to 12 mm. 


Loc. Seychelles. Silhouette: near Mont Pot-a-eau. Mahé: from near Morne 
Blane; high damp forest at summit of Morne Pilot, over 2000 feet; high damp forest 
between Trois Fréres and Morne Seychellois, about 1500—2000 feet ; forest above 
Cascade Estate; top of Mount Sebert, nearly 2000 feet. Praslin: Cétes d’Or Estate, 
from Coco-de-Mer forest in the Vallée de Mai. 


Found in great numbers in undergrowth (Senecio seychellensis, &c.) in the damp | 


mountain-forests (Hugh Scott). 


173. Misia maculosa, sp. n. (Plate 51, fig. 4 a). 


Vertex of head and pronotum greyish-ochraceous; mesonotum castaneous with a 
large black spot at each lateral angle; face, sternum and legs pale ochraceous, the 
first centrally more or less fuscous; pro- and mesosterna transversely fuscous ; abdomen 
beneath testaceous, more or less transversely fuscous; tegmina greyish-ochraceous, an 
oblong spot beyond middle of clavus, an elongate spot above clavus, and a large spot 
—apically widened on apical area, fuscous-brown; vertex and face with the lateral 
margins strongly ridged and elevated; clypeus strongly centrally ridged ; pronotum 
centrally carinate, mesonotum moderately tricarinate; tegmina with the costal margin 
distinctly arched near middle. 

Long. incl. tegm. 6 mm. 


Loc. Seychelles. Mahé: high damp forest between Trois Fréres and Morne 
Seychellois, about 1500—2000 feet. 


174. Nisia fuscofasciata, sp. n. (Plate 51, fig. 3 a). 


Vertex of head, pronotum and mesonotum ochraceous, the latter with an oblique 
black fascia on each lateral area; body beneath and legs ochraceous, abdomen and 
sometimes the sternum with fuscous suffusions; tegmina pale ochraceous, an inner broad 
longitudinal fuscous fascia continued on apex; vertex of head and face with the lateral 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 293 


margins strongly carinately elevated; clypeus ridged; tegmina moderately arched near 
middle of costal margin. 

Long. incl. tegm. 5 mm. 

Loc. Seychelles. Silhouette: high country near Mont Pot-d-eau; Mare aux Cochons, 
plateau and jungle near by, all from marshy plateau. Mahé: Cascade Estate. 

Found in great numbers in undergrowth (Senecio seychellensis, &c.) in the damp 
mountain-forests (Hugh Scott)*. 


175. Nisia thoracica, sp. n. 

Vertex of head, face, pronotum, and legs pale stramineous ; eyes black ; mesonotum 
pale tawny-brown; body beneath ochraceous; tegmina creamy-white; anterior and 
intermediate tarsi more or less infuscate; vertex of head moderately concave, with the 
anterior lateral angles strongly pronounced; face strongly concave, the lateral margins 
strongly ridged and subparallel, much longer than broad ; pronotum narrow, the posterior 
margin strongly angulate; mesonotum with the central carination strongly developed. 

Long. incl. tegm. 5 mm. 

Loc. Seychelles. Silhouette: forest above Mare aux Cochons. 

A single example of this distinct species. 

There is another species of this genus represented by a single mutilated specimen, 
not available for specific description. It is allied to N. maculosa Dist. 


Subfamily Lophopine. 
Genus Ivinea. 

Ivinga Dist., Trans. Linn. Soc. Lond. Zool. xiii. p. 42 (1909). 

176. Ivinga typica Dist., Trans. Linn. Soc. Lond. Zool. xii. p. 42, Plate 4, 
fig. 5a (1909). 

Loc. Seychelles: found not only in the endemic mountain-forests (where it was 
commonly beaten from the endemic ‘“ Bois Rouge” tree, Wormia ferruginea, and 
from other plants), but also among non-endemic vegetation near sea-level. Silhouette : 
Mahé. Anonyme Island. Dennis Island. Praslin. Félicité. Marie Anne. 

Aldabra: 1907 (Thomasset), 1908 (Fryer). 

In describing this species I gave the colour as dark ochraceous as is also the 
invariable appearance of a very large number of specimens examined since. Mr Scott 
has however added the note—‘‘all these are light green during life.” This is a very 
common discoloration after death in many Cicadidze and other Homoptera. 


Subfamily Issinee. 


Only two species of one genus were brought home by this expedition. The genus 
(Lollius) has previously only been recorded from the Malayan and Papuan regions, 
while an allied genus Vylana, represented on the Islands of Mauritius and Bourbon, 
was unrepresented in this Seychelles collection. 

* See footnote, p. 285. 


294 PERCY SLADEN TRUST EXPEDITION 


Genus LOoLutus. 


Lollius Stal, Hem. Afr. iv. p. 209 (1866); Ofv. Vet.-Ak. Forh. xxvii. p. 762 
(1870). 

Type. JL. furcifer Stal, from the Philippine Islands. 

In his “ Monographie der Issiden,” Melichar figures his L. gratiosus from N. Guinea, 


which is scarcely typical of the genus. The two species here described agree with the 
type L. furcifer Stal. 


177. Lollius atromaculatus, sp. n. (Plate 50, figs. 7 a, b). 


Brownish-ochraceous, speckled and marked with black, tegmina with a large shining 
black spot, bounded and divided by veins a little behind middle, the veins more or 
less testaceous ; anterior and intermediate legs with black annulations, posterior femora 
black, the tibiz black at base and apex; abdomen beneath with black markings. 
principally a black longitudinal fascia near base; vertex of head concave, the lateral 
margins strongly ridged, the apical margin with three short spines, one central and 
one at each apical angle; face with the anterior lateral spines more prominent, tri- 
carinate, the lateral carinations curved inwardly, anteriorly and posteriorly, the whole 
surface speckled with black; costal and apical margins to tegmina with more or less 
elongate black spots, the apical margins broadly, obliquely truncate. 

Long. incl. tegm. 84 mm. 


Loc. Seychelles. Mahé: high damp forest between Trois Freres and Morne 
Seychellois, about 1500—2000 feet; forest above Cascade Estate, 1000—2000 feet. 


178. Lollius vurescens, sp. n. (Plate 50, figs. 8 a, b). 


Head, pronotum, mesonotum, body beneath and legs more or less ochraceous, anterior 
and intermediate legs annulated with fuscous, posterior femora, and bases of the tibize 
fuscous; coxee and base of abdomen greyish-white, basal area of abdomen beneath— 
excluding basal margin—black, with greyish-white marginal spots; tegmina pale greenish- 
ochraceous, a small black spot at base, the apical margin spotted with black; clypeus 
with two black lines or elongate spots. In structural characters this species is closely 
allied to DL. atromaculatus. 

Long. incl. tegm. 83 mm. 


Loc. Seychelles. Mahé: from forest of rather stunted Capucin trees (Northea) 
on summit of “Montagne Anse Major,” 2000 feet or over; slopes of Morne Seychellois, 
about 1500—2000 feet. 


Subfamily Ricaniine. 

This subfamily of the Fulgoridz is well represented in the. Seychelles but is 
without any representative of its dominant genera such as Pochazia, Ricama and 
Ricanoptera, which being both Ethiopian and Oriental in their distribution might have 
been expected to be found in this fauna. Privesa however occurs in these islands as 
well as in the Ethiopian, Oriental and Australasian regions. Several other distinct 
genera required recognition. 


DISTANT—_RHYNCHOTA. PART II: SUBORDER HOMOPTERA 295 


DEFERUNDATA, gen. nov. 


Vertex of head broad and narrow, anterior and lateral margins ridged, centrally 
earinate, and with eyes about as broad as anterior margin of pronotum; face slightly 
broader than long, the anterior margin truncate, centrally and sublaterally carinate, 
the lateral margins narrowing near clypeus which is also centrally carinate ; pronotum 
short, centrally longitudinally ridged, its anterior margin convex, its posterior margin 
concave ; mesonotum about twice as long as vertex and pronotum together, tricarinate, 
the lateral carinations anteriorly inwardly forked; legs of moderate length, posterior 
tibize with two spines before apex; tegmina twice as long as broad, costal margin 
prominently convexly raised at base, slightly waved beyond middle, costal membrane 
narrower than radial area near base, somewhat closely transversely veined, apical 
margin obliquely convex, inner margin beyond base, straight, outer disk transversely 
reticulately veined, two transverse lines near apical margin defining numerous narrow 
cellular spaces; wings a little shorter and about as broad as tegmina, a few transverse 
veins on upper disk beyond middle; abdomen above centrally longitudinally sulcate. 


179. Deferundata aldabrana, sp. n. (Plate 50, fig. 1 a). 


Head, pronotum and mesonotum brownish-ochraceous; vertex with two central 
longitudinal black fascize, mesonotum with anterior and lateral black spots; abdomen 
pale ochraceous, the segmental margins black or fuscous; face brownish-ochraceous, 
the lateral areas finely spotted with black; sternum and legs brownish-ochraceous ; 
tegmina bronzy-ochraceous, the posterior claval margin distinctly darker; wings very 
pale fuliginous, the veins and apical marginal area a little darker; structural characters 
as in generic diagnosis. 

Long. excl. tegm. 5mm. Exp. tegm. 165 mm. 


Toc. Aldabra: fle Michel; Takamaka (Fryer). 


ARMILUSTRIUM, gen. nov. 


Vertex of head transverse, broad, excluding eyes narrower than pronotum and 
about twice as broad as long, margins ridged, anterior margin a little rounded, anterior 
lateral angles prominent, eyes backwardly directed along the lateral margins of the 
pronotum ; face considerably broader than long, anterior margin truncate, centrally and 
sublaterally carinate, the sublateral carinations convexly meeting near anterior margin ; 
elypeus with a central carination, the lateral margins obscurely ridged ; pronotum short, 
slightly longer than vertex, centrally longitudinally ridged; mesonotum large and long, 
considerably longer than head and pronotum together, tricarinate, the lateral carinations 
anteriorly, inwardly bifurcating ; posterior tibiz with two spines beyond middle; teg- 
mina about twice as long as broad, costal membrane narrow, much narrower than 
radial area, basal cell longer than broad, longitudinal veins bifurcating beyond middle, 
a subapical series of transverse veins, preceded by a straight and well-defined im- 
pression; wings shorter and broader than tegmina, two outer discal transverse veins, 


some veins bifurcating on outer margin. 


296 PERCY SLADEN TRUST EXPEDITION 


180. Armilustrium gardineri, sp. n. (Plate 50, fig. 3 a). 

Head, pronotum and mesonotum bronzy-brown; abdomen with the base pale 
sanguineous, apical half black, sometimes with the segmental margins testaceous ; body 
beneath and legs brownish-ochraceous, apical area of abdomen beneath, blackish; teg- 
mina shining pale bronzy-brown, irrorated with paler markings, of which two central 
ones are transverse and angularly directed outwards, a prominent black spot on lower 
half of apical margin; wings fuliginous, darker on apical area; vertex, pronotum and 
mesonotum, thickly, finely granulose; face with an anterior transverse ridge; other 
structural characters as in generic diagnosis. 

Long. excl. tegm. 4 to 5mm. Exp. tegm. 10 to 14 mm. 

Loc. Seychelles. Silhouette: from highest forest, over 2000 feet. Mahé: high 
forest of Morne Blane and Pilot. One imperfect specimen from each locality. 


181. Armilustrium scotti, sp. n. (Plate 50, fig. 4 a). 

Head, pronotum and mesonotum bronzy-brown ; abdomen sanguineous, with broad, 
shining-black, transverse, segmental fascixe, extreme lateral margins sanguineous; body 
beneath and legs brownish-ochraceous, abdomen as above; tegmina greyish-white 
irregularly speckled with brownish-ochraceous venation shining yellow, a prominent 
black spot on lower half of apical margin, wings mutilated in typical specimen. 

Differs from the preceding species A. gardinera by its larger size, different 
coloration of the abdomen, and by the practical absence of the anterior transverse 
ridge to the face. 

Long. excl. tegm. 6 mm. Exp. tegm. 13 mm. 

Loc. Seychelles. Mahé: Cascade Estate, about 800—1500 feet. 


One imperfect specimen. 


CARMENTALIA, gen. nov. 


Vertex of head broad, about twice as broad as long, excluding eyes a little 
narrower than pronotum, centrally and marginally carinate, anterior margin a little 
angularly convex ; face about as long as broad, anterior margin truncate, lateral margins 
obliquely widening to posterior margins of eyes and then obliquely narrowing to 
clypeus, centrally longitudinally carinate and with two curved ridges on anterior area; 
clypeus centrally ridged; pronotum about as long as head, centrally ridged, anterior 
margin broadly convex, posterior margin moderately concave; mesonotum large and 
long, considerably longer than head and pronotum together, tricarinate, the lateral 
carinations inwardly furcate near anterior margin, posterior tibise with a distinct spine 
beyond middle; tegmina twice as broad as long, costal membrane very narrow, costal 
margin very finely spinulose, reticulately veined beyond middle, and with a subapical 
series of transverse veins; wings shorter and broader than tegmina, two transverse 
veins on anterior disk and some of the veins furcate on apical and posterior margins. 


182. Carmentalia biformis, sp. n. (Plate 50, fig. 5 a). 


Vertex, pronotum and mesonotum ochraceous; abdomen above more or less san- 
guineous ; body beneath and legs very pale ochraceous; tegmina subhyaline, claval 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 297 


area and apical area of posterior margin more or less fuscous-brown, a distinct black 
spot on lower half of apical margin; wings hyaline, the venation pale ochraceous ; 
eyes black, lateral marginal areas of mesonotum, castaneous; structural characters as 
in generic diagnosis. 

Var. a. Tegmina without the fuscous-brown suffusions, these being replaced by 
pale yellowish. 


Long. excl. tegm. 5 to 6mm. Exp. tegm. 15 mm. 


Loe. Seychelles. Silhouette: Mare aux Cochons, and high forest above. Mahé: 
near Morne Blanc, about 1000 feet ; top of Mount Sebert, nearly 2000 feet; Cascade 
Estate, about 1000 feet; forest near Mount Harrison 1700 feet. 


NEOPRIVESA, gen. nov. 


Allied to Privesa but differing by the costal membrane of the tegmina being very 
narrow for about one-fourth from base and then moderately ampliated towards apex; 
in the type specimen here figured there is a transverse vein to the lower ulnar area, 
but this is not a constant character; the face is shorter and broader, and the vertex 
distinctly conically rounded. The structure of the costal membrane to the tegmina, 
and the conically rounded anterior margin of the vertex compel generic separation. 

183. Neoprivesa fuscovaria, sp. n. (Plate 49, figs. 8, 8a). 

Body above brownish-ochraceous, more or less spotted or suffused with black ; 
margins of vertex before eyes, and carinations to pro- and mesonota, pale ochraceous ; 
body beneath and legs pale ochraceous, face finely sprinkled with fuscous; tegmina 
subhyaline with fuscous-brown markings and suffusions, the most prominent of these 
are oblique, irregular fasciz on apical area; the transverse veins to costal membrane 
are prominently fuscous; wings hyaline, the veins, apical margin and a longitudinal 
streak—widened posteriorly—before abdominal area, dark fuliginous; vertex conically 
rounded in front; pronotum with the lateral areas ochraceous. 

Long. excl. tegm. 7 mm. Exp. tegm. 19 mm. 

Loc. Seychelles. Mahé: country above Port Glaud, about 500 to 1000 feet; 
Baie Lazare; Anonyme Island. Silhouette: Mare aux Cochons. 

Aldabra; 1907 (Thomasset), 1908 (Fryer). 


Genus PRIVESA. 
Privesa Stal, Rio Jan. Hem. ii. p. 70 (1858); Melich., Ann. Hofmus. Wien, xiil. 
p. 282 (1898); Dist., Faun. Brit. Ind. Rhynch. iii. p. 386 (1906). 
Dechitus Walk., Journ. Ent. i. p. 311 (1862). 


184. Privesa fryeri, sp. n. (Plate 49, fig. 7 a). 

Body and legs ochraceous; a spot on each side of vertex and a discal central 
spot to pronotum divided by a median pale carinate line, fuscous; mesonotum more 

SECOND SERIES—ZOOLOGY, VOL. XVII. 38 


298 PERCY SLADEN TRUST EXPEDITION 


or less suffused with fuscous, the carinations pale ochraceous; apical abdominal appen- 
dages fuscous; face with four dark spots near anterior margin and a sublateral series 
of small fuscous spots on each side; clypeus with oblique dark striz on each side; 
tegmina more or less fuscous much darker in some places than in others, two large 
pale spots in costal membrane, the first usually ochraceous, the second hyaline with 
the veins ochraceous, and more narrowly continued on radial area, two large oblique 
hyaline spots on disk, a similar but smaller spot at apex, the apical margin also 
suffused with hyaline; wings hyaline, the veins, margin and a longitudinal suffusion 
near abdominal area, fuliginous; anterior angles of vertex slightly prominent; face 
with the anterior margin truncate, centrally longitudinally carinate, the lateral margins 
subparallel; mesonotum with three longitudinal carinations, the lateral ones anteriorly 
inwardly forked; posterior tibize with two strong spines. 

Long. excl. tegm. 6 mm. Exp. tegm. 16 to 17 mm. 

Loc. Aldabra (Fryer). Assumption (Fryer). 

This species is allied to P. punctifrons Sign., by the sublateral punctate markings 
to the face, a species recorded from Madagascar and the Comoro Islands. It cannot 
however be that species as Signoret describes it “élytres entiérement hyalines, sans 
macule au bord supérieur.” 


185. Privesa melanaria, sp. n. (Plate 49, fig. 6 a). 


Vertex and pronotum ochraceous with fuscous spots, the two most prominent 
being on vertex; mesonotum dark ochraceous or castaneous with black macular 
suffusions; abdomen above fuscous, the segmental margins pale ochraceous; body 
beneath and legs ochraceous, face with two transverse black fasciz and a sublateral 
series of smal] transverse spots; clypeus with dark oblique strize on each side; tegmina 
brownish-ochraceous, the costal membrane and apical margin hyaline with scattered 
brownish-ochraceous spots, the disk more or less suffused with darker markings, and 
in well-preserved specimens greyishly tomentose, a grey curved line at inner margin 
of apical area; wings dark fuliginous; vertex with the anterior angles distinctly acutely 
prominent, face thus appearing slightly concave at anterior margin, its lateral margins 
obliquely rounded; pronotum strongly centrally carinate; mesonotum longitudinally 
tricarinate, the lateral carinations inwardly anteriorly forked; posterior tibize with 
two strong spines. 


Long. excl. tegm. 6 to 64 mm. Exp. tegm. 17 mm. 
Loc. Seychelles. Mahé: Long Island, vii. 1908. Silhouette: near Mont Pot-a-eau. 


The structure and markings of the face give this species a very distinct appearance. 


PRIVESANA, gen. nov. 


Allied to Privesa Stal, from which it principally differs in the width of the costal 
membrane, which in Privesa is considerably broader than the radial area; in Privesana 
it 1s scarcely more than half the breadth of the radial area. 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 299 


186. Privesana infusca. 

Privesa infusca Dist., Trans. Linn. Soc., Ser. 2, Zool. xiii. p. 42, Pl. 4, fig. 5 @ (1909). 

Loc. Seychelles. Mahé, 1905 and 1908—9; Cascade Estate, over 1000 feet ; 
near Morne Blanc. Silhouette: Mare aux Cochons, 1908. Praslin, 1905. In Scott’s 
experience this species was exclusively found among fallen dead leaves of the endemic 
screw-pine tree, Pandanus hornei, in the mountain-forests. 


Genus OSAKA. 

Osaka Dist., Trans. Linn. Soc. Lond. Zool. xiii. p. 43 (1909). 

187. Osaka hyalina. 

Osaka hyalina Dist., Trans. Linn. Soc., Ser. 2, Zool. xiii. p. 44, Pl. 4, figs. 15, 
15 a (1909). 

Loc. Seychelles. Silhouette: near Mont Pot-d-eau, and from Mare aux Cochons, 
about 1000 feet. Mahé; near Morne Blanc, Cascade Estate, 800—1000 feet. Marie 
Anne Island. 

Aldabra: Takamaka; [le Michel (Fryer). 


188. Osaka relata. 

Osaka relata Dist., Trans. Linn. Soc., Ser. 2, Zool. xii. p. 44 (1909). 

Loc. Seychelles. Mahé; from near Morne Blanc, 1908. Cousin Island, 1905. 
Praslin, 1908. 

Aldabra: Takamaka, 1908 (Fryer). 


Subfamily FLAatin«. 


None of the larger and more handsome species of this subfamily have been 
received from the Seychelles, such as the large and conspicuous Phromma rubra Sign. 
found somewhat common in Madagascar. In fact I can only now enumerate three 
species, and we may therefore conclude that the Flatinz are very poorly represented. 


Genus ULUNDIA. 

Ulundia Dist., Insect. Transvaal, 1. p. 250 (1910). 

189. Ulundia madagascariensis. 

Elidiptera madagascariensis Sign., Ann. Soc. Ent. Fr. (3), vi. p. 199 (1860). 

Ormenis madagascariensis Stal, Hem. Afric. iv. p. 243 (1866); Melich., Ann. 
Nios ilofmist) xeviiie yy 165) Unave f) 19))(11902)): 

Loc. Aldabra: Takamaka, &c., x—xiui. 1908 (Fryer). 

This species was described by Signoret from Madagascar, and is also recorded by 
Melichar as from German East Africa. Another species is known from Natal (1910). 


Genus KETUMALA. 
Ketumala Dist., Faun. Brit. Ind. Rhynch. iii. p. 446 (1906). 
190. Ketumala rubromarginata, sp. n. (Plate 50, fig. 2 a). 
Body and legs very pale greenish (in older specimens ochraceous), more or less 
thickly greyishly tomentose; tegmina very pale greenish, or ochraceous, slightly 
38—2 


300 PERCY SLADEN TRUST EXPEDITION 


ereyishly tomentose, costal margin very narrowly from beyond middle, apical margin, 
and inner margin to apex of clavus (more broadly), sanguineous; wings milky white ; 
vertex of head including eyes narrower than pronotum, the anterior lateral angles 
a little acutely prominent; face a little broader than long with a central carination 
not quite reaching clypeus; pronotum anteriorly truncately produced; mesonotum 
with two discal carinations ; tegmina twice as long as broad, costal margin not sinuate, 
apical margin subtruncate, costal membrane in some places wider than radial area; 
clavus distinctly granulose, the granules somewhat sanguineous; posterior tibize with 
two short spines beyond middle. 

Long. excl. tegm. 64 mm. Exp. tegm. 184 to 19 mm. 

Loc. Seychelles. Silhouette: Mont Pot-d-eau, high forest, over 1000 feet. 


Genus FLATOIDES. 


Flatoides Guér., Régn. Anim. Ins., p. 362 (1838); Dist., Faun. Brit. Ind. Rhynch. 
il. p. 459 (1906). 

191. Flatoides protea, sp. n. (Plate 50, figs. 9 a, 5). 

Body above very pale virescent, inclining to pale ochraceous, abdomen above with 
the segmental margins greyishly tomentose; pronotum with three small discal black 
spots; mesonotum with six black spots of which the two lateral spots are the largest ; 
body beneath and legs pale ochraceous, more or less greyishly tomentose ; tegmina very 
pale virescent with a few darker and paler suffusions; wings creamy-white, the venation 
ochraceous; head including eyes distinctly narrower than pronotum at base; vertex 
longer than broad, somewhat attenuated towards apex, the lateral margins distinctly 
reflexed and with two small black spots on each side; face considerably longer than 
broad, a little concave, the lateral margins strongly ridged and a little concave at 
region of the eyes; tegmina about twice as long as broad, the costal membrane 
nearly three times as broad as radial area, the costal membrane and apical margin 
strongly veined; posterior tibize with two spines. 

Var. Tegmina more ochraceous in hue with black suffusions as in fig. (9d). 

Long. excl. tegm. 8 mm. Exp. tegm. 18 mm. 

Loc. Seychelles. Silhouette, from marshy plateau of Mare aux Cochons, about 
1000 feet, and forest above. Mahé, from near Morne Blane. 


Subfamily Delphacine. 


This subfamily is well represented in the present collection. I was only enabled 
to enumerate one species in the result of the former expedition; fourteen species 
comprised in five genera are now recorded. 


Genus PUNDALUOYA. 
Pundaluoya Kirk., J. Bomb. Nat. Hist. Soc. xiv. p. 52 (1902); Dist., Faun. Brit. 
Ind. Rhynch. ui. p. 467 (1906). 
192. Pundaluoya simplicia. 
Pundaluoya simplicca Dist., Faun. Brit. Ind. Rhynch. ii. p. 468, fig. 255 (1906). 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 301 


Loc. Seychelles. Mahé: near Morne Blanc; Mare aux Cochons district, 1000— 
2000 feet; Cascade, sea-level. Praslin: Cotes d’Or Estate. From grass, &c., in cultivated 
places. 

This species is already proved to have a very wide distribution. I originally 
described it from Ceylon; since then it has been received from South Nigeria where 
it was found by Dr Lamborn breeding on the young shoots of Kola and Cacao. It is 
now to be also recorded from the Seychelles. The British Museum possesses a specimen 
from Hawaii named by the late Mr Kirkaldy as Peregrinus maidis Ashm., and his 
localities Fiji, New South Wales and Java probably also refer to P. simplicia. 


193. Pundaluoya pulchella. 

Pundaluoya pulchella Dist., Ann. Mag. Nat. Hist. (8), ix. p. 190 (1912); loc. cit. 
vi. p. 135, fie. 97 (1916). 

Loe. Seychelles. Mahé: Cascade Estate, about 800 feet and over. Brit. India: 
Bengal, Travancore, Bombay. Ceylon. 


OPICONSIVA, gen. nov. 


Allied to Pundaluoya Dist. from which it differs in the elongate face, twice, 
or a little more than twice, longer than broad; vertex about as long or distinctly 
longer than broad; tegmina always quite three times longer than broad; posterior 
tibie with a short spine beyond middle, in addition to the inner long apical spur, 
and the short outer apical spine. 


Type. O. fuscovaria Dist. 


194. Opiconsiva fuscovaria (Plate 50, fig. 10a). 

Head, pronotum and mesonotum castaneous, abdomen ochraceous with darker 
shadings; body beneath and legs ochraceous; tegmina pale shining, bronzy-brown, 
costal and apical margins (narrowly), and about the lower half of apical area fuscous, 
remaining inner half paler fuscous; wings hyaline, the veins fuliginous; head (including 
eyes) narrower than pronotum, antennz brownish-ochraceous; face much longer than 
broad, marginally and medially strongly carinate, the medial carination bifurcate at 
base, obliquely narrowed near clypeus; mesonotum tricarinate; tegmina about three 
times longer than broad; posterior tibiee with a long robust apical spine and a short 
spine beyond middle. 

Long. excl. teem. 3 mm. Exp. tegm. 11 to 11§ mm. 

Loc. Seychelles. Silhouette: near Mont Pot-d-eau, and from Mare aux Cochons, 
over 1000 feet. Marie Anne Island. Mahé: Cascade Estate, 800—1000 feet; high 
forest of Morne Blanc; Anonyme Island. Found in the mountain-forests at considerable 
elevations, and also in drier forest vegetation near sea-level. 


195. Opiconsiva colorata, sp. n. (Plate 50, fig. 11 a). 

Head, pronotum and mesonotum black; posterior angle of mesonotum and about 
basal half of abdomen above, ochraceous, apical half of abdomen black ; body beneath 
and legs ochraceous; head and abdomen beneath black ; tegmina subhyaline, the veins 
ochraceous, costal margin (narrowly), apical margin (slightly broader) and inner and 


302 PERCY SLADEN TRUST EXPEDITION 


claval areas, more or less, dark fuscous; wings hyaline, the veins fuliginous ; head 
(including eyes) narrower than pronotum, antennee with the basal joimt black, the 
second joint ochraceous ; face much longer than broad, marginally and medially strongly 
carinate, the medial carination bifureate at base, obliquely narrowed near clypeus ; 
mesonotum tricarinate; tegmina about three times longer than broad; posterior tibize 
with a long robust apical spine and a short spine beyond middle. 

Long. excl. tegm. 2 mm. Exp. tegm. 6 mm. 


Loc. Seychelles. Silhouette: high country near Mont Pot-d-eau, about 1500 feet ; 
Mare aux Cochons, plateau and jungle above; low coconut-planted country near the 
coast, Pomte Etienne. Mahé: from near Morne Blanc; Cascade Estate, 800—1000 feet, 
and forest above ; marshes on coastal plain at Anse aux Pins and Anse Royale. Found 
in grasses, &c., in more open places in lower parts of mountain-forests. 


196. Oprconsiva balteata, sp. n. (Plate 51, fig. 8 a). 


Vertex of head pale ochraceous, eyes black; pronotum and mesonotum pale castaneous- 
brown, both with a broad, longitudinal, pale ochraceous fascia, abdomen above black, 
its base and apex pale ochraceous; face and clypeus ochraceous, the former with the 
lateral areas black; sternum and legs ochraceous; tegmina subhyaline, the veins 
ochraceous ; inner margin, scarcely reaching claval area, dark fuscous ; wings subhyaline, 
the venation pale fuliginous; head (including eyes) narrower than pronotum; face 
much longer than broad, the posterior lateral margins distinctly rounded. 

Long. excl. tegm. 25 mm. Exp. tegm. 6 mm. 

Loc. Seychelles. Mahé: Cascade Estate, about 1000 feet; country above Port 
Glaud, 500—1000 feet. 

By the coloration of the tegmina this species is allied to the two preceding species 
O. fuscovaria and O. colorata, but differs by the broad pale fascia to the pro- and 
mesonota, which character allies it to the two following species, O. wmsularis and 
O. derelicta, but from all these it is to be distinguished by the structure of the 
face, which is distinctly rounded posteriorly. 


197. Opiconsiva gloriosa, sp. n. 

Head, pronotum and mesonotum black ; a broad, longitudinal central pale ochraceous 
fascia, occupying the whole central area of vertex and 
passing through pro- and mesonota; head beneath 
black ; body beneath more or less black (imperfectly 
seen in unique carded specimen); legs ochraceous ; 
tegmina black, the claval area paler and with two 
longitudinal black spots at apex, about two-thirds of 
posterior marginal area pale greyish subhyaline, the 
inner apical area palely blackish; the longitudinal 
pale ochraceous fascia more or less depressed, head 
including eyes narrower than pronotum; veins to 


Fig. 3. Opiconsiva gloriosa Dist. | tegmina moderately setigerous; vertex and face dis- 
tinetly projecting beyond the eyes. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 303 


Long. incl. tegm. 54 mm. 

Loc. Seychelles. Mahé: marshes on coastal plain at Anse aux Pins and Anse 
Royale. 

A single specimen only contained in the collection. 

198. Opiconsiva insularis, sp. n. (Plate 50, fig. 12 a). 

Vertex of head, pronotum and mesonotum pale shining greyish; antennee and 
lateral areas of pro- and mesonota pale castaneous, those of the mesonotum darker ; 
face, clypeus, rostrum and legs ochraceous; facial carinzee and a spot on each side 
of mesonotum, fuscous; abdomen beneath black with marginal ochraceous spots; 
tegmina pale ochraceous, the inner area pale fuscous; vertex somewhat narrow, 
distinctly extended before eyes, its apex truncate with a submarginal carination on 
each side; pronotum about as long as vertex; mesonotum longer than pronotum 
and vertex together; posterior tibize with a spine beyond middle and a long, robust 
apical spur; face a little more than twice longer than broad. 

Long. incl. tegm. 3 mm. 

Loc. Seychelles. Silhouette: low coconut-planted country near the coast, Pointe 
Etienne. Mahé: Cascade Estate, about 1000 feet. 

199. Opiconsiva derelicta, sp. n. (Plate 50, fig. 13 a). 

Allied to O. isularis but the colour and markings much paler in hue; vertex 
of head a little longer and less truncate at apex; face slightly longer, the lateral 
margins not parallel, being distinctly narrowed before eyes. 

Long. 35 mm. 

Loc. Seychelles. Mahé: from lower country. A single specimen only is contained 
in the collection. 

200. Opiconsiva modesta, sp. n. (Plate 50, fig. 14 a). 

Head, pronotum and mesonotum ochraceous; tegmina very pale ochraceous, with 
the venation darker; body beneath and legs ochraceous, abdomen beneath with basal 
and apical black spots; vertex a little longer than pronotum, projecting a little in 
front of eyes, the apex subtruncate; mesonotum prominently tricarinate; face twice 
as long as broad, a little narrowed before eyes; posterior tibize with a spine beyond 
middle and a long robust apical spur. 

Long. 3 mm. 

Loc. Seychelles. Mahé: Cascade, cultivated country near sea-level. 


Genus NILAPARVATA. 


Nilaparvata Dist., Faun. Brit. Ind. Rhynch. iii. p. 473 (1906). 

This genus was founded for the reception of a species from Ceylon. I am unable 
to generically separate the following species, though the antennze are inserted at about 
middle of eyes and not beneath them, while the venation of the wings is setigerous, 
and the vertex not longer than broad. 


304 PERCY SLADEN TRUST EXPEDITION 


201. Nilaparvata mahensis, sp. n. (Plate 51, fig. 14 a). 


Head ochraceous with three black punctate spots ; pronotum very pale stramineous, 
with two central black spots; mesonotum castaneous, the carinations much paler ; 
body beneath and legs ochraceous; tegmina subhyaline, the veins ochraceous and 
setigerous; vertex about as long as broad, lateral margins strongly carinate, their 
anterior angles moderately acutely prominent, two oblique discal carinations commencing 
at about middle of lateral margins and angularly meeting slightly in front of anterior 
margin, basal margin carinate; face elongate, very much longer than broad, centrally 
and laterally carinate, the central carination furcate at anterior margin; clypeus strongly, 
broadly, laterally and medially carinate, about half the length of face ; antennze inserted 
at about middle of eyes, second joint much longer than first, incrassate and granulose ; 
pronotum about as long as vertex, centrally, laterally and anteriorly carinate; meso- 
notum distinctly tricarinate. 


Long. 2 mm. 
Loc. Seychelles. Mahé: Port Victoria, from grass in Botanic Gardens. 


A single specimen only procured. 


CoNSUALIA, gen. nov. 


Head projecting beyond the eyes, centrally and laterally carinate, apical margin 
a little centrally produced; face elongate, slightly broadened before clypeus, centrally 
bicarinate, clypeus centrally and laterally carinate; antennse with the first and second 
joints long and moderately robust, second a little longer than first; pronotum short, 
conically produced between the eyes, posterior margin prominent; mesonotum longer 
than head and pronotum together, with five longitudinal carinations, the three central 
longest and percurrent, the two lateral ones shorter and oblique; abdomen broad, 
robust, centrally ridged, the lateral margins flattened; posterior legs long, the tibize 
with a short spine near base, a longer spine beyond middle, and with a long robust 
apical spur; tegmina about twice as long as broad, apical margin convexly rounded, 
costal membrane without transverse veins, discal veins longitudinal, a series of transverse 
veins before apex deliminating apical cells, those at apical angle short and oblique ; 
wings much shorter but almost as broad as tegmina, two transverse veins on upper disk. 


202. Consualia robusta, sp. n. (Plate 50, fig. 6 a). 


Body dark castaneous; posterior angle of mesonotum, base and central longitudinal 
ridge to abdomen, and legs, more or less testaceous ; tegmina pale brownish-ochraceous, 
the venation fuscous; wings pale fuliginous, the veims darker; structural characters 
as in generic diagnosis. 


Long. excl. tegm. 55 mm. Exp. tegm. 10 mm. 


Loc. Seychelles. Silhouette: near Mont Pot-d-eau, over 1000 feet. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 305 


Genus Ucyops. 


Ugyops Guér., Voy. Ind.-Orient. Bélanger, Zool., p. 477 (1834); Syn. Dist., Trans. 
Linn. Soc. London, Zool. xiii. p. 44 (1909). 


203. Ugyops senescens (Plate 49, fig. 3 a). 

Ugyops senescens Dist., Trans. Linn. Soc., Ser. 2, Zool. xii. p. 44, Pl 4, 
figs. 16a, b (1909). 

Loc. Amirantes: Desroches Island, 1905. Seychelles: Mahé, Praslin, Silhouette, 
from near sea-level (not from the mountain-forests). 

I have figured a variety in which the markings of the tegmina are darker and 
the basal joint of the antenne paler than in the type previously delineated. 

204. Ugyops facialis, sp. n. (Plate 49, fig. 2 a). 

Body above ochraceous; vertex of head with four black longitudinal lines; _pro- 
notum with a few dark spots on each lateral area; mesonotum with five longitudinal 
dark carinate lines which are usually black; abdomen spotted and suffused with 
castaneous, especially on disk and lateral margins; body beneath and legs ochraceous, 
face darker in hue and suffused with small paler and darker macular markings ; 
tegmina subhyaline, the venation darker, the longitudinal veins considerably ochraceously 
broken, a series of longitudinal veins crossing disk of tegmina beyond middle and 
a few transverse veins before apex; wings hyaline, the venation fuscous; antennze 
with the first and second joints ochraceous, about apical half of second, black; head 
with the central lobe distinctly prominent at apex, face long, narrow and prominent, 
extending for some distance before eyes, moderately widened from about anterior margin 
of eyes and again slightly narrowed before clypeus, central longitudinal ridge promi- 
nent, duplex for about anterior half, lateral margins distinctly reflexed; tegmina a 
little more than three times as long as broad. 

Long. excl. tegm. 7 to 7 mm. Exp. tegm. 20 mm. 

Loc. Seychelles. Silhouette: near Mont Pot-a-eau, all over 1000 feet, and from 
Mare aux Cochons and forest above. Mahé: Mare aux Cochons district, over 1000 feet. 
Aldabra: 1908 (Fryer). In contrast to the preceding species, this species was found 
in the Seychelles at considerable elevations and in endemic-forest districts. 


205. Ugyops seychellensis, sp. n. (Plate 49, fig. 4 a). 

Body and legs bright castaneous, lateral longitudinal sulcations to face, black ; 
tegmina pale bronzy-brown, the veins darker, a few linear darker markings on claval 
vein; wings subhyaline, the veins fuliginous; head projecting a little beyond anterior 
margins of eyes; face elongate ampliated beyond eyes towards clypeus, prominently 
centrally longitudinally carinate, the lateral longitudinal black sulcations, profound ; 
antennee brownish-ochraceous, first and second joints robust ; pronotum strongly centrally 
ridged ; mesonotum with five longitudinal ridges; abdomen distinctly centrally ridged ; 
tegmina with the veins more or less distinctly finely setigerous; posterior tibize with 
three distinct spines. 

SECOND SERIES—-ZOOLOGY, VOL. XVII. 39 


306 PERCY SLADEN TRUST EXPEDITION 


Long. excl. tegm. 5 to 55mm. Exp. tegm. 14 mm. 
Loc. Seychelles. Mahé: high forest of Morne Blanc and Pilot, circa 2000 feet ; 
slopes of Morne Seychellois, 1500—2000 feet. 


AMBARVALIA, gen. nov. 


Vertex of head considerably produced in front of eyes, about as far as from 
anterior margins of eyes to base of head, a little widened on apical area, centrally 
longitudinally carinate, the carination becoming bifid in front of eyes, the margins 
elevately carinate, face prominently extended in front of eyes, apex somewhat conically 
rounded, widened towards posterior margin, centrally longitudinally cariate; clypeus 
short, broad, less than half the length of face, centrally carinate; antenne with 
the first and second joints distinctly incrassated, second much longer than first ; 
pronotum only a little shorter than mesonotum, strongly tricarinate; mesonotum tri- 
carinate, the disk moderately raised and flattened; posterior tibize with a strong apical 
spur which is serrate on its inner margin; tegmina four times longer than broad, 
veins longitudinal, on inner margin of apical area are two short oblique veins. 

I have only seen one carded specimen of this very interesting genus, the details 
of the under surface being thus indescribable. 

206. Ambarvalia pyrops, sp. nu. (Plate 51, fig. 1 a). 

Black; the carinations to vertex, pro- and mesonota obscurely ochraceous, the 
posterior margins of the latter two being also more or less finely spotted with the 
same colour; tegmina finely spotted with greyish and ochraceous, the apical area 
unspotted; anterior and intermediate tibiz basally and apically annulated with ochraceous, 
apex of posterior tibiz including the apical spur ochraceous, posterior tarsi with the 
apex of first jomt and the whole of second joint, ochraceous; structural characters 
as in generic diagnosis. 

Long. 4 mm. 

Loc. Seychelles. Mahé, near Morne Blanc, about 1000 feet. 


Family Cercopide. 


Only one species belonging to this family is contained in the collection. It is 
in the Ethiopian region that the genus which contains this species is more largely 
represented and Madagascar contributes its largest and most showy species. 


Subfamily Aphrophorine. 

207. Ptyelus maher. 

Ptyelus maher Dist., Trans. Linn. Soc., Ser. 2, Zool. xiii. p. 45, Plate 4, 
fig. 14 @ (1909). 

Loc. Seychelles. Silhouette: near Mont Pot-a-eau, over 1000 feet; highest forest, 
over 2000 feet; Mare aux Cochons, about 1000 feet. Mahé: from near Morne Blane 
and high forest of same; high damp forest at summit of Pilot, over 2000 feet; 
high damp forest between Trois Fréres and Morne Seychellois, about 1500—2000 feet ; 
Cascade Estate, 800—1000 feet; forest near Mount Harrison, 1700 feet. 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 307 


In high damp forests, often sitting on palm leaves, &c. (Hugh Scott). 

A variable species in coloration; sometimes the pale central spot to tegmina, 
before the apical area (see fig.), is obsolete, or entirely absent; the colour of the 
tegmina also varies from piceous-brown to either brownish-ochraceous or blackish. 


Family Jasside. 
Subfamily Bythoscopine. 

This subfamily is represented by five* genera, three of which are found in the 
Indian fauna, and two described for the first time may also eventually prove to belong 
to that region. Nehela is represented by eight species. The Hawaiian Islands appear 
to possess only one genus, Bythoscopus. 


Genus [p1ocEeRUs. 


Idiocerus Lewis, Trans. Ent. Soc. Lond. i. p. 47 (1836); Dist., Faun. Brit. Ind. 
Rhynch. iv. p. 184 (1907). 

208. Idiocerus scottt, sp. n. 

Vertex of head pale greenish-ochraceous ; pronotum and scutellum more virescent, 
the former with a small black spot near each lateral 


margin (in some varietal forms with two central spots Soe " 
also), the latter with two small black spots on basal \ 
margin; body beneath and legs pale greenish-ochraceous; ly 


tegmina olivaceous-brown, costal area very pale greenish- 
ochraceous and containing a large prominent black spot 


> f 
\ 
s 
near middle, extreme costal margin bright olivaceous- / 
ereen; face with a slightly darker spot at anterior 
margin and a longer similar spot from eyes to base of : 
clypeus; a somewhat elongate species; vertex of head ae, A Me Rretns exch Dicks 
nearly three times as broad as long, front, including 
face, about as long as broad; apices of the joints to posterior tarsi, black. 
Long. incl. tegm. 4 to 44 mm. 
Loc. Seychelles. Silhouette: near Mont Pot-a-eau; Mare aux Cochons, plateau and 
jungle near by. Mahé: Cascade Estate, about 1000 feet. Praslin: Cétes d’Or Estate, 
especially from Coco-de-Mer forest in the Vallée de Mai. Félicité Island: from forest. 


KRONOS, gen. nov. 


Vertex of head slightly more than twice as broad as long, rounded in front, 
moderately obliquely depressed; face globosely depressed, broader than long, lateral 
margins concavely sinuate, narrow and truncate before clypeus, which is narrow and 
about two-thirds the length of face; ocelli much nearer eyes than to each other; 
pronotum obliquely, anteriorly depressed, and moderately raised on disk, about twice 
as broad at base as long; scutellum about twice as broad at base as long, broadly 


* Galboa typica Dist. (Trans. Linn. Soc. London, Zool. xiii. p. 46, Pl. 4, fig. 11 (1909)) was described 
from the first collection made by Prof. Gardiner (Mahé) but was not contained in the larger collection 
here described. 

39—2 


308 PERCY SLADEN TRUST EXPEDITION 


foveately excavate on disk; posterior tibie thickly, strongly spinulose; tegmina more 
than twice as long as broad, a few transverse veins on costal membrane beyond middle, 
three apical cells. 

Allied to Bythoscopus but differs by the globose and differently constructed face, 
foveate scutellum, &c. 


209. Kronos typicus, sp. n. 

Vertex of head, pronotum and scutellum ochraceous, the last much paler in hue 
and with a darker spot near each basal angle; body 
beneath and legs ochraceous; apices of the posterior 
tibize and the posterior tarsi—excluding base—-fuscous; 


tegmina brownish-ochraceous, with some small dark 
spots on claval margin, narrow apical margin distinctly 
darker; face and clypeus ochraceous; cheeks more or 


less spotted with black; structural characters as in 
4 


& 
\ 

Loc. Seychelles. Mahé: from near Morne Blane 
Fig. 5. Kronos typicus Dist. and high forest of Morne Blanc and Pilot. 


A probably scarce species as the collection only contained two examples. 


generic diagnosis. 


Long. incl. tegm. 44 mm. 


Genus BytHoscopus. 
Bythoscopus Germ., in Silberm., Rev. Ent. i. p. 180 (1833); Syn. Dist., Faun. Brit. 
Ind. Rhynch. iv. p. 190 (1907). 
210. Bythoscopus indicus. 
Macropsis indica Leth., Bull. Soe. Zool. Fr. 1892, p. 209. 
Pachyopsis chlorophana Melich., Hom. Faun. Ceylon, p. 153 (1908). 
Bythoscopus imdicus Dist., Faun. Brit. Ind. Rhynch. vi. p. 227 (1916). 


Loc. Seychelles. Silhouette: Mare aux Cochons. Also found in Ceylon, Brit. 
India, and Tenasserim. 


A single specimen only secured. 


Genus NEHELA. 

Nehela Buch. White, Proc. Zool. Soc. 1878, p. 473; Dist., Faun. Brit. Ind. Rhynch. 
vi. p. 231 (1916). 

Pachynus Stal, Hem. Afr. iv. p. 127 (1866), nom. preeoce. 

Igerna Kirk., Wien. ent. Zeit. xxii. p. 13 (1903), n. nom. 

Stal’s name Pachynus was preoccupied, but its synonym Mehela Buch. White is 
available, and Kirkaldy’s new name is therefore not required. The genus is very 
widely distributed, the type being from St Helena; it is also found throughout tropical 
Africa, and British India and Ceylon. Mr Hugh Scott found this genus very abundant 
in the mountain-forests ; it appears entirely confined to the endemic forest. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 309 


Melichar has recently described and figured a Javan species under the name of 
Oncopsis nugritus (Notes Leyden Mus. xxxvi. p. 121, Pl. 3, fig. 9) which should either 
be included in Nehela, or otherwise that generic name should give place to Oncopsis 
Burm. (1837). But as Burmeister gives as typical Oncopsis the lanio Linn., it cannot 
very well apply to nigritus Melich. which I regard as an undoubted Nehela. 

211. Nehela bimaculicollis (Plate 51, fig. 25 a). 

Bythoscopus (Oncopsis) bimaculicollis Stal, Ofv. Vet.-Ak. Férh. 1855, p. 100. 

Pachynus bimaculicolls Stal, Hem. Afr. iv. p. 127 (1866); Melich., Wien. ent. 
Zeit. xxiv. p. 297 (1905). 

Agallia quadrinotata Melich., Hom. Ceylon, p. 151 (1903). 

Igerna bimaculicollis Jacobi, Kilimandjaro-Meru Exped. 12 (7), p. 129, T. 11, 
fig. 28 (1910). 

Nehela bimaculicollis Dist., Faun. Brit. Ind. Rhynch. vi. p. 232 (1916). 

Loc. Seychelles. Silhouette: forest above Mare aux Cochons. Mahé: Cascade 
Estate, forest, above 1000 feet; Anonyme Island. Also found in Ceylon, Brit. India, 
and in Hast and South Africa. 

Jacobi’s figure (supra) appears, at least, to represent a variety? 

212. Nehela spectabilis, sp. n. (Plate 51, fig. 27 a). 

Ochraceous ; two spots on vertex of head, an oblique line on each side of pronotum 
from inner margins of eyes and two large spots at base of same, a spot at each basal 
angle of the scutellum, and a broad submarginal fascia to the tegmina—united to 
the margin near base—black; body beneath and legs ochraceous ; face with two spots 
on apical margin, an oblique spot behind eyes, followed by an elongate spot on each 
side before clypeus, three spots on disk, and a central fascia to clypeus, black; sternum 
and base of abdomen black; structure of face better shown by the figure here given. 

Long. incl. tegm. 5 mm. 

Loc. Seychelles. Mahé: Mare aux Cochons district, 1000—2000 feet. 

A single specimen of this handsome species was alone found in this collection. 


213. Nehela elegantula, sp. n. (Plate 51, fig. 24 a). 

Ochraceous ; two spots on vertex nearly equidistant from eyes and from each other, 
between them a central longitudinal line, a large spot on each side of pronotum 
before middle, between them a central longitudinal fascia, and a curved anterior 
fascia on each side behind eyes, a spot at each basal angle of scutellum and a transverse 
spot on disk ; tegminal veins and a very broad submarginal fascia, two anterior marginal 
spots to face and two smaller spots beneath them, a curved marginal spot on each 
side behind eyes and a spot at apex of clypeus, black; vertex of head narrow, sub- 
conically rounded anteriorly ; anterior tibie and tarsi pale fuscous. 

Long. incl. tegm. 5 mm. 

Loc. Seychelles: a fairly long series, entirely from the endemic forests. Silhouette : 
Mare aux Cochons, plateau and forest above, about 1500 feet. Mahé: country above 
Port Glaud, about 500—1000 feet ; high damp forest at summit of Pilot, over 2000 feet ; 


310 PERCY SLADEN TRUST EXPEDITION 


from near Morne Blanc; slopes of Morne Seychellois, about 1500—2000 feet. Praslin : 
Cotes d’Or Estate. 

In some specimens the thoracic markings are diminished in size, but in all examples 
the broad black submarginal fascia to the tegmina: is constant. 


214. Nehela conspicua, sp. n. (Plate 51, fig. 26 a). 

Vertex of head pale ochraceous with two large central black spots; pronotum, 
scutellum and tegmina black, a central spot and posterior lateral margins to scutellum 
and inner margins to tegmina pale ochraceous ; face black, anterior margin—including 
two black spots—a spot on each side before eyes, and the posterior lateral margins, 
pale ochraceous; legs and abdomen beneath pale ochraceous, the latter with transverse 
black fasciz, apices of anterior tibize and tarsi blackish. 

Long. incl. tegm. 44 to 5 mm. 

Loc. Seychelles. Mahé: Cascade Estate, about 1000 feet; Mare aux Cochons 
district, 1000—2000 feet. 


215. Nehela lineoligera, sp. n. 

Vertex of head dull ochraceous with two prominent black spots; pronotum 
piceous or blackish, the posterior margin and two small discal 
spots on each side ochraceous, a large darker black spot on 
each side behind eyes; scutellum blackish, the lateral margins 
pale ochraceous; body beneath and legs ochraceous, anterior 
tibiz and tarsi, apices of intermediate tibiee and tarsi, and 
apices of posterior tarsi, black; tegmina blackish, costal margin, 
a distinct central longitudinal fascia, and the veims—more or 
less—very pale ochraceous or greyish; face imperfectly seen in 
unique carded specimen. 


Long. incl. tegm. 5 mm. 
Fig. 6. Nehela lineoligera Loc. Seychelles. Silhouette: high country near Mont 
pe Pot-a-eau. 

216. Nehela scutellata, sp. n. (Plate 51, fig. 28 a). 

Black ; anterior margin of vertex (excluding centre), a spot before eyes, narrow 
posterior margin to pronotum, the scutellum, interior margins of tegmina, body beneath 
and legs, ochraceous ; face with anterior margin, a spot near inner margin of each eye, 
and a central fascia to clypeus, black. 

Long. incl. tegm. 4 mm. 

Loc. Seychelles. Mahé: slopes of Morne Seychellois, about 1500—2000 feet. 

A single specimen of this species only received. Apart from its distinct coloration 
and markings the structure of the face is very distinct, and this is better shown by 
the figure than by the less satisfactory method of a detailed description. 


217. Nehela flavolineata, sp. n. (Plate 51, fig. 23 a). 


Black ; longitudinal, fasciate spots to vertex of head, two minute discal spots 
and posterior submarginal fascia to pronotum, lateral margins of scutellum, inner and 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA dll 


lateral margins and veins to tegmina, head beneath, sternum and legs, ochraceous ; face 
with a short central longitudinal fascia, a spot on each side of anterior margin, a large 
spot on each side behind eyes and apex of clypeus, black ; anterior tibize fuscous; vertex 
of head more evenly rounded than in preceding species. 

Long. incl. tegm. 44 mm. 

Loc. Seychelles. Silhouette: near Mont Pot-a-eau, highest forest, over 2000 feet ; 
Mare aux Cochons, plateau and forest above. Mahé: top of Mount Sebert, nearly 
2000 feet. 

A superficial but constant characteristic of this species is the posterior pale sub- 
marginal fascia to the pronotum. 


218. Nehela aterruma, sp. n. (Plate 51, fig. 22 a). 

Head above, pronotum, scutellum and tegmina black, vertex with fine obscure 
yellowish streaks on each side near eyes; lateral margins of scutellum and outer 
and inner margins of tegmina pale ochraceous; body beneath and legs pale ochraceous, 
face with a transverse anterior marginal fascia (sometimes broadly centrally longi- 
tudinally continued as in figure, or with the anterior transverse fascia only), anterior 
tibiz, apices of anterior tarsi, and transverse abdominal fasciz, black. 

Allied to the preceding species N. flavolineata from which it differs by the different 
structure and markings of the face as shown in figures, smaller size, and more melanic 
coloration. 

Long. incl. tegm. 4 mm. 

Loc. Seychelles. Silhouette: near Mont Pot-a-eau, over 1000 feet, Mare aux 
Cochons, plateau and jungle above. Mahé: high forest of Morne Blanc; country 
above Port Glaud, about 500—1000 feet; high damp forest at summit of Morne Pilot, 
over 2000 feet; Mare aux Cochons district, 1000—2000 feet; Cascade Estate, forest, 
above 1000 feet, and forest near Mount Harrison, 1700 feet ; slopes of Morne Seychellois, 
about 1500—2000 feet. Félicité Island: from forest. Very numerous throughout the 
mountain-forests of Mahé and Silhouette. 


Subfamily Tettigonielline. 


This subfamily is represented by four genera, two of which are described for 
the first time, the other two being originally recorded from British India where the 
Tettigoniellinze are well represented. The Seychelles with only four genera and five 
species are poorly illustrative of the subfamily in which the Oriental region is so rich, 
and the Ethiopian region more moderately opulent. In the ‘‘ Fauna Hawatiensis,” 
Kirkaldy only enumerates one species and that one he had not seen himself. 


Genus KoLua. 


Kolla Dist., Faun. Brit. Ind. Rhynch. iv. p. 223 (1907). 
This genus has a very wide distribution and is found in the Ethiopian, Oriental, 
Australasian and Neotropical regions, especially in the last. 


312 PERCY SLADEN TRUST EXPEDITION 


219. Kolla seychellensis, sp. n. 


Head, pronotum, scutellum, body beneath and legs, flavescent; vertex with two 
prominent black spots on anterior margin ; face—exclud- 
ing a short discal central line and the lateral margins 
—pale brownish-ochraceous, with darker transverse 
striations; tegmina greyish-subhyaline, the imner area 
reflecting the dark abdomen beneath; vertex a little 


more than half as long as breadth between eyes, a 
very fine central carinate longitudinal line; ocelli near 
base, about as near to each other as to eyes; face longer 


than broad, the lateral margins a little concave near eyes. 
Long. incl. tegm. 4 mm. 


Fig. 7. Kolla seychellensis Dist. Loc. Seychelles. Mahé: Cascade Estate, 800— 

1000 ft.; Port Victoria, from grass in Botanic Gardens ; 

marshes on coastal plain at Anse aux Pins and Anse Royale; Cascade, cultivated country 
near sea-level. 


220. Kolla funeralis, sp. n. 


Head, pronotum, scutellum, body beneath and legs, flavescent, vertex with two 
prominent black spots on anterior margin; scutellum 
with a darker spot near each basal angle; tegmina 
greyish, claval area somewhat bluish-grey, followed by 
a broad black longitudinal fascia which reaches apex, 


extreme costal margin flavescent; vertex only a little 
less in length than breadth between eyes, apical margin 
distinctly conical, a central carinate line not reaching 
apex, the ocelli near base, a little nearer to eyes than to 
each other; scutellum transversely impressed near base. 


Long. incl. tegm. 3 mm. 
Fig. 8. Kolla fumeralis Dist. Loc. Seychelles. Silhouette: Mare aux Cochons, 
from marshy plateau and forest above; low coconut- 
planted country near the coast, Pointe Etienne. Mahé: country above Port Glaud, 
about 500—1000 feet, and near Morne Blanc. 


MAIESTAS, gen. nov. 


Allied to the genus Kolla Dist. in the general structure of the head, save that 
the ocelli are placed near the base and almost as near to eyes as to each other; another 
chief difference is in the venation of the tegmina which contains two transverse veins 
on disk, one near base, the other near middle, four large apical cells and three narrow 
ante-apical cells or areas. 


DISTANT—RHYNCHOTA. PART IIT: SUBORDER HOMOPTERA 313 


221. Maiestas illustris, sp. n. 

Vertex of head pale ochraceous, with two broad central longitudinal pale castaneous 
fasciz and a similarly coloured short longitudinal line 
between them; pronotum pale ochraceous with six pale 
eastaneous fasciz of which the central two are largest 
and curved ; scutellum with a small black spot near each 
basal angle, with two pale spots between and one beyond 
them ; body beneath and legs ochraceous, the basal area 
of abdomen sometimes shaded with piceous; tegmina 
ochraceous, when closed reflecting the dark dorsal surface 
of the abdomen beneath them, the veins greyish-white, 


the apical cells more or less spotted or suffused with 
black; anterior and intermediate femora and _ tibize 
annulated with pale brownish; vertex of head about Fig. 9. Muwiestas illustris Dist. 
as long as breadth between eyes, longly produced and 
narrowed before eyes, the basal margin moderately concave; face longer than broad, 
from near anterior margins of eyes obliquely narrowed both anteriorly and posteriorly, 
finely transversely striate on each lateral area; pronotum about twice as broad as long, 
truncate posteriorly, convex anteriorly ; scutellum about twice as broad at base as long. 
Long. incl. tegm. 34 mm. 
Loc, Seychelles. Silhouette: Mare aux Cochons, and forest near by and immediately 
above. Mahé: near Morne Blanc, country above Port Glaud, about 500—1000 feet ; 
Cascade Estate, 800—1000 feet. 


Genus Usna. 


Ujna Dist., Faun. Brit. India, Rhynch. iv. p. 239 (1907). 

This genus was founded for two species from British India, one from Ceylon, the 
other from Tenasserim. The present species here described is the largest representative 
of the genus I have as yet examined. 

222. Ujna flavidipes, sp. n. (Plate 51, fig. 20a). 

Head, pronotum, scutellum and tegmina black, the latter with a pale ochraceous 
costal marginal spot at about one-third before apex; body beneath and legs pale 
stramineous ; vertex of head broadly elongate, about as long as breadth between eyes, 
ocelli on disk in a line a little before the anterior margins of the eyes; face long, 
distinctly longitudinally carinate for less than half its length, very indistinctly carinate 
on the remaining area; pronotum shorter than vertex, lateral margins straight ; 
scutellum about as long or very slightly longer than pronotum, distinctly transversely 
impressed before apex. 

Long. incl. tegm. 5 mm. 

Loc. Seychelles. Mahé: from near Morne Blanc; Cascade Estate, forest, above 
1000 feet, and forest near Mount Harrison, 1700 feet; Mare aux Cochons district, 
1000—2000 feet. 

SECOND SERIES—ZOOLOGY, VOL. SOVIE 40 


314 PERCY SLADEN TRUST EXPEDITION 


PAGANALIA, gen. nov. 


Head nearly twice as broad between eyes as long, triangularly produced before 
eyes, ocelli near base, much closer to eyes than to each other; face about twice as 
long as broad, narrow, centrally compressed, about three times as long as clypeus; 
pronotum a little longer than vertex or scutellum, anteriorly narrowed and convexly 
produced, the apical and posterior margins truncate; scutellum much broader than 
long, basal margin longer than either of the lateral margins; tegmina considerably 
passing abdominal apex, veins simple and longitudinal. Allied to the Nearctic and 
Neotropical genus Helochara. 


223. Paganalia virescens, sp. n. 
Dull virescent, apical areas of the tegmina irregularly pale fuscous-brown; body 
ae -==x, beneath virescent, legs pale ochraceous, anterior and 
x intermediate tibize obscurely annulated with pale fuscous, 
6. posterior tibiee outwardly margined with small blackish 
ig spots at the bases of the spinules; eyes black, their 
inner margins beneath sanguineous ; structural charac- 
ters as in generic diagnosis. 


Long. incl. tegm. 6 mm. 


Loc. Seychelles. Mahé: Long Island. 


Fig. 10. Paganalia virescens Dist. 


Subfamily Jassine. 


This subfamily is not so largely represented in the collection as might have been 
expected. The species are contained in eight genera, six of which have a wide 
distribution, and only two have had to be considered as previously undescribed. 

Two species, Chelusa seychellensis Dist., and Athysanus imsularis Dist., previously 
brought home by Prof. Gardiner, and described in these Transactions (1909), were 
unrepresented in this much larger collection. 


Genus ACROPONA. 


Acropona Melich., Hom. Faun. Ceylon, p. 168 (1903); Dist., Faun. Brit. Ind. 
Rhynch. iv. p. 300 (1908). 

224. Acropona prasina (Plate 51, fig..17 a). 

Gypona prasina Walk., List Hom. Suppl., p. 258 (1858). 

Acropona prasina Melich., Hom. Faun. Ceylon, p. 168 (1903). 

? Walkerv Kirk., Entomologist, xxxili. p. 294 (1900). 

Eogypona walkert Kirk., l.c. xxxiv. p. 39 (1901). 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 315 


Loc. Seychelles. Silhouette: from highest forest, over 2000 feet. Mahé: Cascade 
Estate, 800—1000 feet ; top of Mount Sebert, nearly 2000 feet. Praslin: Cétes d’Or 
Estate. Originally described from Ceylon, and also recorded from the Maldives*. 


225. Acropona gardineri, sp. n. (Plate 51, fig. 21 a). 


Above bright olivaceous-green; body beneath and legs pale green; anterior margin 
of vertex of head and subanterior margin of face, continued on each side to near eyes, 
sanguineous ; differs from the preceding species, A. prasina, by the darker olivaceous- 
green coloration above, the sanguineous apical fasciee to vertex and face; vertex of head a 
little shorter ; pronotum with a more or less distinct transverse impression before middle. 

Long. incl. tegm. 10 to 12 mm. 

Loc. Seychelles. Mahé: Cascade Estate, about 1000 feet; from forest of rather 
stunted Capucin trees (Northea) on summit of ‘Montagne Anse Major,” 2000 feet 
or over. Praslin: Cétes d’Or Estate. 

226. Acropona sladen, sp. n. (Plate 51, fig. 19a). 

Body above dark sanguineous, body beneath and legs pale sanguineous ; anterior 
margin of vertex narrowly ochraceous, anterior margin of face ochraceous, subanterior 
margin dark sanguineous; apart from the different coloration this species differs from 
the two preceding by the broader, and laterally more sinuate face; the scutellum 
also possesses two distinct longitudinal ridges with a posteriorly transverse convex 
impression between them. 

Long. incl. tegm. 11 mm. 

Loc. Seychelles. Praslin: Cotes d’Or Estate. 


Genus JASSUS. 


Jassus Fabr., Syst. Rhyng. p. 85 (1803); Syn. &c., Dist., Faun. Brit. Ind. 
Rhynch. iv. p. 327 (1908). 

227. Jassus indicus (Plate 51, fig. 13 a). 

Celidia indica Walk., List Hom. iii. p. 855 (1851). 

Tettigonia jactans Walk., 1. c. Suppl., p. 357 (1858). 

Jassus deplanatus Spangb., Ofv. Vet.-Ak. Forh. 1878, no. 8, p. 23. 

Jassus indicus Dist., Faun. Brit. Ind. Rhynch. iv. p. 327; fig. 210 (1908); Trans. 
Linn. Soe. Zool. xi. p. 46 (1909). 

Loc. Seychelles: though not a native species, this was found largely in the high 
endemic mountain-forests. Silhouette; near Mont Pot-d-eau, over 1000 feet; Mare aux 
Cochons, marshy plateau and jungle above. Mahé: from near Morne Blanc; from grass 
in cultivated country, about 1000 feet ; Cascade Estate, 800—1000 feet; top of Mount 
Sebert, nearly 2000 feet; Mare aux Cochons district, 1000—2000 feet. Also found 

* T have it recorded in my journal that a number of specimens of Acropona were taken in a particular 
dry, scrubby type of forest-vegetation, which is found especially in connection with areas of bare granite 
“glacis.” The Acropona were obtained thus in Mahé (Cascade Estate) and in a valley behind Grande Anse, 


Praslin. But whether these statements refer to A. prasina and A. gardineri equally, or to one more than 


the other, is very hard to say.—H. Scorr. 
40—2 


316 PERCY SLADEN TRUST EXPEDITION 


widely distributed in Brit. India, Burma and Tenasserim. The specimen here figured 
is of a varietal character, but one which is also found in India. I have (supra, 1908) 
described the variation of the species. 


228. Jassus deternunatus, sp. n. (Plate 51, fig. 16 a). 

wee allied to J. zndicus, but with the face testaceous with a bond central 
longitudinal black fascia which neither reaches base nor apex, cheeks with a prominent 
black spot on each side; rostrum and legs ochraceous, apex of rostrum, apices of 
posterior tibize and apices of all the tarsi, black; posterior tibiz finely spotted with 
black. 

Long. incl. tegm. 7 mm. 

Loc. Seychelles. Mahé: Cascade Estate, about 1000 feet. 


A single carded specimen of this distinctly marked and coloured species. 


LIMENTINUS, gen. nov. 


Allied to Jassus, but with the vertex of head distinctly longer than broad, the 
apex considerably broader than base, rounded, and distinctly projecting beyond the 
anterior margins of the eyes, strongly centrally and laterally carinate, ocelli near apical 
margin, much farther apart from each other than from the lateral apical angles; face 
very long, about as long as vertex, pronotum and scutellum together, elongate, very 
shghtly convex, its anterior margin distinctly produced beyond the anterior margin 
of the eyes; scutellum small, centrally carinate, less than half the length of face ; 
other characters generally as in Jassus. 

229. Inmentinus aldabranus, sp. n. (Plate 51, fig. 12 a). 

Vertex of head ochraceous, black between the carinations; pronotum and scutellum 
black; body beneath and legs ochraceous; face and clypeus with broad lateral black 
fascize ; apices of the posterior tibiee black ; tegmina dull dark ochraceous, the venation 
broadly black, costal membrane black, with a longitudinal line before middle, and two 
prominent longitudinal spots beyond middle, pale ochraceous, the veins on apical area 
spotted with pale ochraceous, claval veins also finely spotted with ochraceous; pro- 
notum obscurely transversely strigose; scutellum strongly transversely impressed near 
middle; other structural characters as in generic diagnosis. 

Long. 8 mm. 


Loc. Aldabra: Esprit I. (Fryer). 


Genus Athysanus. 


Athysanus Burm., Gen. Ins. I. t. xiv. subj. 2 (1840). 


Of this widely distributed genus only one species is contained in the collection 
and is here described. Another, A. imsularis Dist., formerly brought home by Prof. 
Gardiner from the Amirantes, is not here included. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 317 


230. Athysanus frontalis, sp. n. 

Pale flavescent, the tegmina subhyaline; vertex of head with an angulate black 
linear fascia between the eyes; anterior and inter- 
mediate femora biannulated with black, extreme apices 
of posterior femora, and apices of joints of posterior 
tarsi, black or blackish; anterior third of face regularly 
blackly transversely striate with a central longitudinal 
ochraceous spot, remaining area reticulately blackly 
striate ; clypeus with the lateral margins black ; body 
beneath more or less shaded with black ; face elongate, 
about as long as breadth between eyes; vertex of head 
subconically produced before eyes ; head with the eyes 
somewhat wider than the pronotum. 


ib ong. evall. tegm. 44 to 5mm. Fig. 11. Athysanus frontalis Dist. 


Loc. Seychelles. Mahé: Port Victoria, from grass in Botanic Gardens; Cascade 
Estate, 800—1000 feet; low country—various localities. 


MATSUMURANA, gen. nov. 

Head much broader than long, convexly rounded before eyes, which posteriorly 
overlap the anterior lateral margins of the pronotum; face broad and moderately 
globose, its lateral margins convexly rounded, about as broad as long; clypeus about 
half the length of face; pronotum about twice the length of vertex, anterior margin 
convex between the eyes, posterior margin truncate; scutellum shorter than pronotum, 
moderately transversely impressed before apex; tegmina about three times as long as 
broad, apices moderately narrowed and rounded, lateral margins convex, claval area 
broad, three apical cells; posterior tibize longly spinulose outwardly, more shortly and 
finely spinulose inwardly. 

Allied to Dagama Dist., a genus at present only recorded from Natal, from which 
it differs by the more conically produced vertex in front of eyes, the broader and 
shorter face, and by having only three apical cells to tegmina; it also has affinities 
with Mimotellie Mats. 

231. Matsumurana facialis, sp. n. 

Above cinnamomeous-brown; vertex of head with three more or less distinct 
narrow pale ochraceous transverse fasciee, the extreme 
anterior margin also of that colour; pronotum with 
four transverse pale ochraceous fascie ; scutellum 
with a dark spot near each basal angle, and four 
pale spots, two near anterior margin, and one on 
each lateral margin; tegmina with paler suffusions 
and some irregular black spots, of which the principal 
are three oblique spots on each lateral margin, a spot 
at apex of clavus, the apical areas also black ; apical 
area of abdomen above black ; structural characters 


as In generic diagnosis. Fig. 12. Matswmurana facialis Dist. 


318 PERCY SLADEN TRUST EXPEDITION 


Long. 34 mm. 


Loc. Seychelles. Silhouette: forest above Mare aux Cochons. Mahé: country 
above Port Glaud, about 500—1000 feet. 


Genus BALCLUTHA. 


Gnathodus Fieb., Verh. z.-b. Ges. Wien, xvi. p. 504 (1866), nom. preeoce. 
Balclutha Kirk., Entomologist, xxxiii. p. 242 (1900), nom. n. 


An almost universally distributed genus. 


232. Balclutha chersonesia, sp. n. 


Bright golden-yellow, tegmina much paler except on claval area; scutellum with 


a dark spot near each basal angle; ocelli near the eyes distinctly visible near passage 


of vertex to front; vertex 


ay 


SRSSSASSS 
SS 


Fa 


Een Nate, SUS ela MN CNRS 


Fig. 13. Balelutha chersonesta Dist. 


with a narrow central longitudinal line; face with a 
distinct central longitudinal line and the lateral margins 
pale, the striated area more pale piceous; scutellum 
distinctly transversely impressed. 

Long. incl. tegm. 34 to 4 mm. 

Loc. Seychelles. Silhouette: Mare aux Cochons, 
and forest near by; low coconut-planted country near 
the coast, Pointe Etienne. Mahé: from near Morne 
Blane; country above Port Glaud, about 500—1000 
feet; Cascade Estate, 800—1000 feet; Anonyme Island; 
marshes on coastal plains at Anse aux Pins and Anse 


Royale. 


233. Balclutha varicolor, sp. n. 


Head and seutellum dark ochraceous; pronotum and tegmina pale greenish-ochraceous; 


Fig. 14.  Balelutha varicolor Dist. 


abdomen above black, the segmental margins and apex 
ochraceous ; body beneath and legs ochraceous ; tegminal 
veins mostly paler in hue; vertex of head with a central 
longitudinal ridge; pronotum moderately arched and cen- 
trally ridged ; scutellum distinctly transversely impressed. 

Long. incl. tegm. 3 to 35 mm. 

Loc. Seychelles. Silhouette: Mare aux Cochons ; 
low coconut-planted country near the coast, Pointe Etienne. 
Mahé: near Morne Blanc, about 1000 feet ; from grass in 
cultivated country, about 1000 feet. 


Genus SCAPHOIDEUS. 


Scaphoideus Uhler, Trans. Maryl. Ac. Sci, p. 33 (1888). 
This very widely distributed genus has three representatives in the Seychelles. 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 319 


234. Scaphoideus vagans, sp. n. 


Very pale ochraceous; vertex of head with six small brown submarginal spots 
arranged in pairs, a central arcuate pale brownish fascia 
containing two dark spots, and two pale brownish sub- 
quadrate spots on the basal area; pronotum with four 
basal spots, two small central discal spots, and an arcuate if 
broken narrow fascia near anterior margin, all pale 
brownish ; scutellum with pale brownish markings and 
a dark spot on each lateral margin; tegmina with linear 
blackish markings, more pronounced on apical area where 
they are more numerous and smaller; face with dark 
transverse fascize for about two-thirds its length, cheeks 
with two dark oblique lines on each side; abdomen 
beneath more or less shaded with black; vertex of head 
about as long as breadth between eyes, transversely depressed near middle; pronotum 
about as long, or only slightly longer than scutellum, the latter with a distinct curved 
central impression. 


Fig. 15. Scaphoideus vagans Dist. 


Long. incl. tegm. 3 mm. 


Loc. Seychelles. Silhouette: Mare aux Cochons, and forest near by. Mahé, country 
above Port Glaud, about 500—1000 feet; Port Victoria, from grass in Botanic Gardens 
and from grass in cultivated country; Cascade Estate, 800—1000 feet; Mare aux Cochons 
district, 1000—2000 feet ; Cascade, cultivated country near sea-level; forest near Mount 
Harrison. 


From grass &¢. in cultivated places. 


235. Scaphoideus tessellatus, sp. n. 


Very pale ochraceous ; vertex of head with four very pale brownish apical marginal 
spots, and a cluster of about six dark brown spots on ____ {Uae Pes 
disk ; pronotum with two dark brown spots on anterior : 
margin and blackish transverse linear markings on 
disk ; scutellum very pale, a blackish spot near each 
basal angle, and some small dark spots on discal and 
apical areas; tegmina very much tesselated with 
blackish linear markings, of which the most prominent 
are spots on costal and claval margins near bases ; head 
beneath very pale with a slight virescent tint, face with 
obscure darker transverse striations ; body beneath and 
legs very pale ochraceous; abdomen with black suftu- 
sions, especially on apical area; posterior legs with 
femoral streaks, apices of tibize and bases of spinules, black, the tibize also blackly 
annulate ; face much longer than breadth between eyes; scutellum about as long as 
pronotum; vertex of head about as long as breadth between eyes. 


Fig. 16. Scaphoideus tessellatus Dist. 


320 PERCY SLADEN TRUST EXPEDITION 


Long. incl. tegm. 5 mm. 


Loc. Seychelles. Silhouette: Mare aux Cochons, plateau and jungle near by and 
forest above. Mahé: from near Morne Blanc, and Mare aux Cochons district. 


A species taken in high forest. 
236. Scaphoideus seychellensis, sp. n. 


Body, tegmina and legs pale ochraceous; vertex of head with six small anterior 
submarginal spots and a T-shaped spot on disk, fuscous- 
brown, the surrounding areas of these spots greyish- 
white; pronotum with greyish-white suffusions; scutellum 
with the whole central area greyish-white, and with an 
ochraceous spot near each basal angle; face with a series 
of dark transverse linear spots on each lateral area, larger 
and nearly meeting between eyes; tegmina with the 
veins paler, and with testaceous spots on anterior half 


and at apical margin; vertex about as long as breadth 
between eyes; scutellum about as long as pronotum ; 
face much longer than broad, very slightly concave 
near eyes, narrowed and slightly rounded towards clypeus. 


Fig. 17. Scaphoideus seychellensis Dist. 


Long. incl. tegm. 35 mm. 
Loc. Seychelles. Silhouette: Mare aux Cochons. 


A single example. 


Genus PARALIMNUS. 


Paralimnus Matsum., Termész. Fiizetek, xxv. p. 386 (1902). 


Distributed in the Eastern Paleearctic, Oriental, Malayan and Australasian regions, 
and recorded from East Africa. Now found in the Seychelles. 


237. Paralimnus silhouettensis, sp. n. (Plate 51, fig. 18 a). 


Head, pronotum, scutellum and tegmina, glossy bronzy-brown; scutellum with two 
pale spots on each lateral margin and a larger darker or blackish spot near each 
basal angle; tegmina with the veins darkly prominent and with more or less numerous 
pale greyish spots; body beneath and legs pale ochraceous; face longer than broad, 
its margins sinuate before eyes, between lateral margins of face and eyes the colour 
is darker with two small pale spots; vertex of head somewhat angularly produced 
in front of eyes, nearly twice as broad as long, ocelli on anterior margin very near 
eyes; scutellum broad, basal margin broader than length of either lateral margin, 
transversely impressed near middle; tegmina longer than abdomen, four apical cells, 
three ante-apical cells, a transverse vein on disk. 


Long. incl. tegm. 6 to 7 mm. 


Loc. Seychelles. Silhouette: from marshy plateau of Mare aux Cochons, about 
1000 feet; forest above Mare aux Cochons. 


Fig. 1. 
Fig. 2. 
Fig. 3. 
Fig. 4. 
Fig. 5. 
Fig. 6. 
Fig. 7. 
Fig. 8. 
Fig. 9. 
Fig. 10. 
Fig. 11. 
Fig. 12. 
Fig. 13. 
Fig. 14. 
Fig. 15. 
Fig. 16. 
Fig. 17. 
Fig. 1. 
Fig. 2. 
Fig. 3. 
Fig. 4 
Fig. 5. 
Fig. 6. 
Fig. 7. 
Fig. 8. 
Fig. 9. 
Fig. 10. 
Fig. 11. 
Fig. 12. 
Fig. 13. 
Fig. 14. 
Fig. 15. 
Fig. 16. 
Fig. 17. 
Fig. 18. 
Fig. 19. 
Fig. 20. 
Fig. 21. 
Fig. 1. 
Fig. 2. 
Fig. 3. 
Fig. 4 
SECOND 


DISTANT—RHYNCHOTA. PART II: SUBORDER HOMOPTERA 


y 


EXPLANATION OF PLATES 49—5l1. 


PLATE 49. 


Aselgeoides insularis, gen. et sp. n. a, face and clypeus. 
Ugyops facialis, sp. n. a, face and clypeus. 

Ugyops senescens, var. a, face and clypeus. 

Ugyops seychellensis, sp. n. a, face and clypeus. 
Daradaxoides mahensis, gen. et sp. n. a, face and clypeus. 
Privesa melanaria, sp. n. a, face and clypeus. 

Privesa fryeri, sp. n. a, face and clypeus. 

Neoprivesa fuscovaria, gen. et sp. n. a, face and clypeus. 
Canetrona maculipennis, gen. et sp. n. a, face and elypeus. 
Adolendana typica, gen. et sp. n. a, face and clypeus. 
Aqueelicium typicum, gen. et sp. n. a, face and clypeus. 
Curiatius insignis, gen. et sp. n. a, face and clypeus. 
Volcanaha typica, gen. et sp. n. a, face and clypeus. 
Brizia mahensis, sp. n. a, face and clypeus. 

Clusivius spectabilis, gen. et sp. n. a, face and clypeus. 
Matutinus opulentus, gen. et sp. n. a, face and clypeus. 
Brimia stellata, sp. nu. a, face and clypeus. 


Puate 50. 


Deferundata aldabrana, gen. et sp. n. a, face and clypeus. 
Ketumala rubromarginata, sp. n. a, face and clypeus. 
Armilustrium gardineri, gen. et sp. n. a, face and clypeus. 
Armilustrium scotti, sp. n. a, face and clypeus. 
Carmentalia biformis, gen. et sp. n. a, face and clypeus. 
Consualia robusta, gen. et sp. n. a, face and clypeus. 


Lollius atromaculatus, sp. n. a, face and clypeus. 6, side view and legs. 


Lollius virescens, sp. n. a, face and clypeus. 06, side view and legs. 


Flatoides protea, sp. n. a, face and clypeus. 6, side view of tegmen. 


Opiconsiva fuscovaria, gen. et sp. n. a, face and clypeus. 
Opiconsiva colorata, sp. n. a, face and clypeus. 
Opiconsiva insularis, sp. n. a, face and clypeus. 
Opiconsiva derelicta, sp. n. a, face and clypeus. 
Opiconsiva modesta, sp. n. a, face and clypeus. 
Volcanalia modesta, sp. n. a, face and clypeus. 
Volcanalia varicolor, sp. n. a, face and clypeus. 
Volcanalia fumosa, sp. n. a, face and clypeus. 
Volcanalia atrovaria, sp. n. a, face and clypeus. 
Volcanalia picturata, sp. n. a, face and clypeus. 
Volcanalia cardw, sp. n. a, face and clypeus. 
Volcanalia atrostriata, sp. n. a, face and clypeus. 


Puate 51. 


Ambarvalia pyrops, gen. et sp. n. a, face and clypeus. 
Volcanalia capitata, sp. n. a, face and clypeus. 

Nisia fuscofasciata, sp. n. a, face and clypeus. 

Nisia maculosa, sp. n. a, face and clypeus. 


SERIES—ZOOLOGY, VOL. XVII. 


41 


321 


322 PERCY SLADEN TRUST EXPEDITION 


Fig. 5. Iguvium albomaculatum, gen. et sp. n. a, face and clypeus. 
Fig. 6. Hquirria phalena, gen. et sp. n. a, face and clypeus. 
Fig. 7. Aquclicium brunnescens, sp. n. a, face and clypeus. 
Fig. 8. Opiconsiva balteata, sp. n. a, face and clypeus. 

Fig. 9. <Aqueliciwm elegantulum, sp. n. a, face and clypeus. 

Fig. 10. Fescennia aurea, sp. n. a, face and clypeus. 

Fig. 11. Fescennia bimaculata, sp. n. a, face and clypeus. 

Fig. 12. Limentinus aldabranus, sp. n. a, face and clypeus. 

Fig. 13. Jassus indicus. a, face and clypeus. 

Fig. 14. Nilaparvata mahensis, sp. n. a, face and clypeus. 

Fig. 15. Fordicidia robusta, gen. et sp. n. a, face and clypeus. 

Fig. 16. Jassus determinatus, sp. n. a, face and clypeus. 

Fig. 17. Acropona prasina. a, face and clypeus. 

Fig. 18. Paralimnus silhouettensis, sp. n. a, face and clypeus. 

Fig. 19. Acropona sladeni, sp. n. a, face and clypeus. ~ 

Fig. 20. Ujna flavidipes, sp. n. a, face and clypeus. 

Fig. 21. <Acropona gardineri, sp. n. a, face and clypeus. 

Fig. 22. Nehela aterrima, sp. n. a, face and clypeus. 

Fig. 23. Nehela flavolineata, sp. n. a, face and clypeus. 

Fig. 24. Nehela elegantula, sp. n. a, face and clypeus. 

Fig. 25. Nehela bimaculicollis. a, face and clypeus. 

Fig. 26. Nehela conspicua, sp. n. a, face and clypeus. 

Fig. 27. Nehela spectabilis, sp. n. a, face and clypeus. 

Fig. 28. Mehela scutellata, sp. n. a, face and clypeus. 


TRAINS, JLIONIN. SOG. SIBIR, 2 ZOOL, WOIL, XOVIN 


Percy SnapEN Trust EXPEDITION. 


PL. 49 


(Distant) 


H. Knight del 


RISMINGTOTA INROM SieVCisvs LILIES 


Percy Siapen Trust Expepirion. 


(Distanr) TRAINS, JLIUNIN. SOG, SER. 2 ZOOL, VOI, 2 PIL, 


H. Knight del 


RHYNCHOTA FROM SEYCHELLES 


Percy Srapen Trust Expepirion. 


(DisTant) 


TRANS. LINN. 


SOC, SIR, 2 ZOOL, WOl, WU, IIL, Bil 


ii 

Hf d 
4h j 
Aix 4 
y ; 
4 

i) 

Ay 

iN 


Pe 


H. Knight del 


SS, 


RHYNCHOTA FROM SEYCHEEEES 


No. VIII—ON THE PONTONIIN A. 


By L. A. Borraparne, M.A. 


(Lecturer in Zoology in the University of Cambridge, Fellow, Dean, and Lecturer 
of Selwyn College.) 


(CommuniIcaTeD By Pror. J. Strantey Garpiner, M.A., F.R.S., F.L.S.) 


(Plates 52—57.) 


Read 2nd November, 1916. 


FOREWORD. 


Tue collections made during Professor Stanley Gardiner’s two expeditions to the 
Western Indian Ocean are particularly rich in Pontoniinz, and contain many new 
Species, some of which are of peculiar interest. In view of these facts, and of the 
large additions to our knowledge of the group that have been made since my revision 
of it in 1898, I have thought well again to revise it as a whole*. The results of my 
labours are embodied in the present paper. 

The following is a list of the species collected, with a statement of the localities 
in which they were taken: 


1. Urocaridella gracilis Borradaile, 1915. Suvadiva, Kolumadulu, and Haddu- 
mati Atolls. 

2. Urocaris sp., ? U. longicaudata Stimpson, 1860. N. Malé Atoll, Amirante. 

3. Urocaris psamathe de Man, 1902. N. Malé Atoll, Diego Garcia. 

4. Palemonella tridentata Borradaile, 1899. Goidu, Goifurfehendu Atoll, 8S. 
Nilandu Atoll, Salomon, Coin, Peros Banhos. 

5. Palemonella elegans Borradaile, 1915. Salomon Is. 

6. Palemonella longirostris Borradaile, 1915. Reef, Naifaro, Fadiffolu Atoll. 


* The present article was finished towards the end of the year 1914. Since then there have been 
described the following new species which probably or certainly belong to the Pontoniine. 

(1) Paratypton siebenrocki Balss, 1915 (Zool. Anz. xlv. p. 83), Red Sea and Samoa. A remarkable 
coral commensal, superficially resembles Conchodytes. As neither the telson nor the gill formula are 
described it is not certain that this is a member of the Pontoniine. 

(2) Urocaris indica Kemp, 1915 (Mem. Ind. Mus. v. p. 279), Chilka Lake, India. Perhaps accounts 
for Pearson’s report of U. longicaudata from Ceylon. Lives in water whose salinity varies, and which is at 
times nearly fresh. 

(8) Periclimenes demani Kemp, 1915 (loc. cit. p. 275). A member of the subgenus Falciger, From 
the same waters as U. indica. 

(4) Periclimenes (Hamiger) nove-zealandie Borradaile, 1916 (Brit. Antarct. (“Terra Nova”) Exp. Rep., 
Zool., 111, 2, p. 87, 1916), New Zealand. Representative of a new subgenus and possibly of a new genus. 
First chela has fingers fringed with long curled hairs, second pair unequal, one very large and of abnormal 
shape. Hepatic spine absent. Ischiomeropodite of third maxilliped somewhat broadened. 


41—2 


324 PERCY SLADEN TRUST EXPEDITION 


7. Periclimenes (Cristiger) frater Borradaile, 1915. Reef, Egmont, 
8. Periclimenes (Cristiger) incertus Borradaile, 1915. S. Nilandu Atoll. 
9. Periclimenes (Cristiger) brocki (de Man), 1887. Suvadiva Atoll. 

10. Periclimenes (Corniger) ceratophthalmus Borradaile, 1915. Hulule, Malé Atoll. 

11. Periclimenes (Corniger) cornutus Borradaile, 1915. Hulule, Malé Atoll. 

12. Periclimenes (Falciger) vitiensis Borradaile, 1898. Coetivy, Seychelles. 

13. Periclimenes (Falciger) affinis Borradaile, 1915. Salomon Is. 

14. Periclimenes (Falciger) dubius Borradaile, 1915. Minikoi. 

15. Periclimenes (Falciger) nilandensis Borradaile, 1915. S. Nilandu Atoll. 

16. Perichimenes (Falciger) seychellensis Borradaile, 1915. Praslin, Seychelles. 

17. Periclimenes (Falciger) brockettt Borradaile, 1915. N. Malé Atoll. 

18. Periclimenes (Falciger) suvadivensis Borradaile, 1915. Suvadiva Atoll. 

19. Periclimenes (Falciger) borradaile: Rathbun, 1904 Haddumati Atoll. 

20. Perichimenes (Falciger) kolumadulensis Borradaile, 1915. Kolumadulu Atoll. 

21. Perichimenes (Falciger) compressus Borradaile, 1915. Saya de Malha, 145 fms. 

22. Periclimenes (Falciger) spiniferus de Man, 1902. Goidu, Goifurfchendu Atoll, 
Hulule, Malé Atoll, Coetivy, Seychelles, Salomon Is., Diego Garcia. 

23. Periclimeneus fimbriatus Borradaile, 1915. Mulaku Atoll, Providence, 
39—50 fms. 

24. Periclimeneus robustus Borradaile, 1915. Amirante, 29—39 fms. 

25. Harpiliopsis beaupresi (Audoin), 1825. Goidu, Goifurfehendu Atoll, Hulule, 
Malé Atoll, Barachois, Diego Garcia. 

26. Harpiliopsis depressus (Stimpson), 1860. Goidu, Goifurfehendu Atoll, Hulule, 
Malé Atoll. Reef, Naifaro, Fadiffolu Atoll. Muinikoi. Coetivy, Seychelles. Coin, 
Peros Banhos. Salomon Is. 

27. Coralliocaris graminia (Dana), 1852. Coetivy, Seychelles. 

28. Coralliocaris nudirostris (Heller), 1862. Goifurfchendu Atoll, Coetivy, 
Seychelles. 

29. Coralliocaris macrophthalma (H. M.-Edwards), 1837. Saya de Malha, 26 fms. 

30. Coralliocaris japonica Ortmann, 1891. Hulule, Malé Atoll. Coin, Peros 
Banhos. Coetivy, Seychelles. Salomon Is. Saya de Malha, 26 fms. 

31. Anchistus muersi (de Man), 1888. Hulule, Malé Atoll. Egmont, Seychelles. 

32. Pontonides maldivensis (Borradaile), 1915. Fadiffolu Atoll. 

33. Conchodytes tridacne Peters, 1851. Hulule, Malé Atoll. Minikoi. 

34. Conchodytes meleagrine Peters, 1851. Salomon, Farquhar, N.W. Cheval. 


Rather more than half the species in the above list were new, and among these 
are representatives of three genera previously unrecognized. I have established a fourth 
genus for two of the species that were already known to science. 

It is unfortunately not at present possible to reach any faunistic conclusions con- 
cerning the Carides of the Western Indian Ocean. 

My revision is not based solely upon the specimens placed in my hands by 
Professor Gardiner. By the kindness of Mr F. A. Potts I am enabled to include in 


BORRADAILE—ON THE PONTONIIN 2 325 


it the results of the examination of a small but interesting collection made by him 
in the Torres Straits, which contained three species new to science, one of them the 
representative of a new genus. I have also re-examined the material in the Cam- 
bridge University Museum of Zoology, and am under much obligation to Dr W. T. 
Calman for enabling me to do the same with the Pontoniine of the British Museum. 

I have not restricted myself to the systematics of the subfamily. The un- 
fortunate destruction in transit of a great part of the crustaceans collected by myself 
in the Island of Minikoi has deprived me of the numbered specimens to which my 
field notes applied, and thereby rendered it impossible for me to give information upon 
the natural history of various prawns, but I have endeavoured to elucidate to some 
extent the conditions under which Pontoniine live, and the connection between their 
structure and their habits. It has seemed to me advisable also to discuss briefly 
certain questions of morphology and genetics, and impossible to omit consideration of 
the relationship of the genera of Pontoniinze with one another and with other prawns. 

Short definitions of the new species and of three of the new genera have 
already appeared in the Annals and Magazine of Natural History for February, 1915. 
Fuller diagnoses, with notes on some other species, will be found in the course of the 
present article. 


A Monograph of the Pontoniine. 


It is one of the paradoxes of Marine Natural History that the Decapod Crustacea, 
the most active and nervous of the invertebrates of the sea, are also among the most 
prone to avail themselves of opportunities of shelter and concealment, and make use 
of the most subtle adaptations to this habit. In describing the collection of crabs 
made by Professor Gardiner and myself in the Maldive Islands and Minikoi*, I found 
oceasion for some remarks upon this phenomenon with regard to the various groups 
of Brachyura. Among the prawns, two families are pre-eminent in the same respect. 
Professor Coutiére’s fine researches upon the Alpheidze have illuminated one of these 
cases. The other is that of the Pontoniine. Although the members of this group 
have not so rich an originality as have the Alpheidz in the production of bizarre 
modifications of structure, they are in one way even more interesting, for they show a 
more complete series of stages in the transformation of the primitive caridoid facies, 
ranging from wholly free-living forms such as Urocaris to commensal and very con- 
siderably degenerate genera such as Conchodytes. The success which has attended 
their strategy of retreat makes them so abundant that they bulk largely in any well- 
made collection of prawns from the Tropics, and their species are numerous and often 
closely related, though generally perfectly distinct. Moreover, the assembling of the 
species into genera and other groups is a matter of great difficulty, partly because 
they present an almost unbroken series of degrees of modification, associated, no doubt, 
with a gradually increasing dependence upon the host, and partly because among 
species of similar habits similar features reappear with bewildering frequency. They 
are therefore of as much importance to the systematist as to the naturalist. 


* Gardiner’s Fauna of the Maldives. 


326 PERCY SLADEN TRUST EXPEDITION 


The Pontoniinzee are members of the Paleemonidee, the central and typical family 
of the Carides. By the loss of exopodites and mastigobranchs from the legs, the 
reduction of the gill formula, and the position of the last joint of the second 
maxilliped at the side of the preceding joint, this family has discarded the primitive 
organization which still prevails in such groups as the Pasiphzoida and the Hoplo- 
phoroida. In the well-marked cleft of the mandible, the retention of the distal 
“lacinia” of the maxilla, the shape of the legs of the first two pairs, with their un- 
divided carpopodites and relatively unspecialized chelee, and in a certain absence of 
exaggeration in all the features of the body, they appear more conservative than the 
remaining members of the tribe, to some of which, indeed, and in particular to the 
Crangonoida, it seems possible that their near ancestors may have given rise. 


The Subfamilies of the Palemonide. 


The members of the Paleemonide fall into four groups, which we may rank as 
subfamilies, defining them briefly by means of the following key: 

I. None of the bristles at the end of the larval telson become in the adult 
transposed on to the anterior part of that organ, which is therefore unarmed on its 
back and sides. The surface of the molar process of the mandible is closely ridged. 
[There is a pleurobranch on the third mazxilliped. | 

Desmocaridine Borradaile, 1915. 

Il. Two pairs of the bristles at the end of the larval telson become in the adult 
transposed on to the back of that organ. The surface of the molar process of the 
mandible bears some half-dozen large knobs or crests. ) 

A. The end of the telson bears six spines. [There is no pleurobranch to the 


third maxilliped. | 
Pontonune Kingsley, 1878. 


B. The end of the telson bears four spines and a varying number of feathered 
bristles. ! 

1. The side of the carapace is traversed by a suture. The outer flagellum 
of the antennule is but slightly cleft. There is no pleurobranch to the 
third maxilliped. 

Typhlocaridine Annandale and Kemp, 1913. 

2. The side of the carapace has no suture. The outer flagellum of the anten- 

nule is deeply cleft. There is a pleurobranch to the third maxilliped. 
Palemonine Kingsley, 1878. 


The peculiarities set forth in this key belong to all members of the subfamilies, 
which by their means can be absolutely separated. In elucidating the relations of 
the subfamilies to one another, however, account must also be taken of certain universal 
characteristics by which the affinities of the groups are indicated. 

Of the four subfamilies, that of the Desmocaridine is the most primitive, as has 
been well shown by M. Sollaud, the describer of its only genus, Desmocaris*. Among 


* C. R. Ac. Sci. clii. p. 913 (1911). 


BORRADAILE—ON THE PONTONIINA 327 


other features, M. Sollaud finds the following to indicate its nearness to the ancestral 
form of the family: (1) the condition of the telson, (2) the retention of the larval 
supraorbital spine, (3) the armature of the molar process*, which recalls that of the 
primitive Acanthephhyra (Hoplophoridz), (4) the simple form of the epipodite of the 
first maxilliped, which in most other Palzemonide is bilobed, (5) the relation of 
the propodite of the second maxilliped to the dactylopodite, which is here largely free, 
instead of lying wholly alongside the propodite, (6) the armature of the fingers of the 
chelze, which consists of fine spines arranged comb-wise, as in many Hoplophoride and 
Peneeidze, and not of a cutting edge or coarse teeth, as in most Carides, (7) the gill-formula, 
which comprises a pleurobranch for each leg and one for the third mazxilliped, one 
arthrobranch for the latter limb, and a podobranch on the second maxilliped. At the 
same time, Desmocaris is not without features which constitute a departure from the 
primitive state. As such may be cited (8) the loss of the mandibular palp, (9) the dis- 
appearance of the cleft between the two divisions of the outer “lacinia” of the maxilla, 
and (10) great reduction of the podobranch of the second maxilla. 

The Paleemonine are less primitive than Desmocaris in respect of the characters 
numbered (1), (2), (8), (5), and (6) above, and most of them have a bilobed epipodite 
on the first maxilliped, though in regard to (9), (10), and generally also to (8), they are 
more primitive. The Typhlocaridine are an aberrant branch of the Palemonine stock. 

On the other hand, the less specialized of the Pontoniine are in many respects 
near to the ancestral type. It is true that in the condition of the telson and of the 
molar process}, and in the loss of the pleurobranch of the third maxilliped, all 
Pontoniinze are unprimitive, but (a) many keep the supraorbital spine, (6) while nearly 
all have lost the mandibular palp, Urocaridella and Palemonella keep this structure, 
though it is here of two joints, and not of three as in the Palzmonine, (c) nearly 
all have a double lacinia on the maxilla, (d) some have a simple epipodite to the first 
maxilliped, and in many cases it is but slightly cleft, (e) some (belonging it is true 
to the more modified genera Harpiliws and Coralliocaris) have a free, or nearly free, 
dactylopodite of the second maxilliped, (f) though the anterior pleurobranch is lost, 
Urocaridella, and according to Nobili some Ancyclocaris, have a podobranch on the 
second maxilliped, and (g) several species of Periclimenes have the comb arrangement 
on the fingers of the first chela, and in at least one (P. denticulata) it is found also 
on the second. Moreover (h) in the form of the outer flagellum of the antennule, 
which is generally but slightly cleft, the Pontoniinz present a primitive feature not found 
in either Desmocaris or the Paleemoninze. Since, however, this feature is in the present 
subfamily found in the genera which are otherwise not primitive, it seems likely that 
it is here not ancestral but a reversion. 

It would appear therefore that, though Desmocaris has departed least from the 
structure of the ancestral paleemonid, the Pontoniine left the main stem before the 
evolution of the first representative of the Paleemonine. 


* The molar surface is a plain, round area, about half of which is covered by toothed ridges. These 
are probably an exceedingly primitive feature, for they are found in many Branchiopoda. 
+ But see below, p. 335. 


328 PERCY SLADEN TRUST EXPEDITION 


Desmocaris and the Palzmoninz are active, free-living prawns, mostly inhabiting 
fresh or brackish water. The Pontoniinz, on the other hand, are marine, and most 
of them lead a hidden and sluggish life, depending for shelter upon other animals, 
such as corals, bivalves, echinoderms, and ascidians. Are we to consider that the 
non-primitive characters which differentiate this group have any connection with their 
mode of life? Clearly, since not all Pontoniinee lead a sluggish existence, it is not 
possible to regard the diagnostic characters of the subfamily as direct adaptations 
to such existence. In any case it would be hard to see this significance in the 
armature of the telson, or in the structure of the mandible, which is no doubt con- 
nected with some peculiarity, either in the food or mn the mode of feeding, shared 
by the Palemoninz. It is, indeed, somewhat remarkable to find that the mandible 
of the most advanced commensals of the subfamily does not differ essentially from 
those of its most primitive members. But M. Sollaud is quite possibly right in con- 
sidering* that the reduction of the gill series, though it cannot have been caused by 
the change to a less active life, has tended to bring about such a change. If, how- 
ever, .the characters of the Pontoniine as a whole cannot be regarded as direct 
adaptations to a sheltered life, there is within the group a long series of such 
modifications, exhibited by almost every organ of the body, and often traceable from 
genus to genus in a striking manner. 


The History of the Pontoniine. 


The subfamily Pontoniinz comprises fifteen genera, described at various dates 
from 1829 to the present day. The first of these to be established was Pontonia, 
founded by Latreille in 1829+, for a species, commensal with bivalves, which had been 
named by Petagna Astacus tyrrhenus (Plate 57, fig. 29). By 1837, when H. Milne- 
Edwards published his Histowre Naturelle des Crustacés, the number of species assigned 
to this genus had risen to four. Roux in 1831 gave the name Pelzas to two prawns 
of more active habit which are now known as Periclimenes scriptus and P. ame- 
‘thysteus, Pelias being preoccupied as a generic name. Periclimenes was given its 
present name in 1844, by Costa, who at the same time founded Typton for a species 
living in sponges. In 1851, Peters established Conchodytes for two very specialized 
species related to Pontonia. Hitherto isolated genera had been founded, but in 1852 
the first steps were taken towards recogniziug the unity of a group of forms which 
was to become the nucleus of the present subfamily. In that year Dana, reporting 
on the Crustacea of the United States exploring expedition, separated from Pontona 
certain coral-haunting species which he erected into the genera Oedipus and Harpilius. 
Conchodytes he did not recognize. At the same time he described as allied to these 
a genus Anchistia, which has subsequently proved to be the same as Costa's Peri- 
climenes, and indicated in a footnote that the true position of T'ypton was in this 
neighbourhood. The name Oedipus was already in use, and Stimpson accordingly 
changed it in 1860 to Coralliocaris. Palemonella, also founded by Dana, was placed 
by him between Anchistia and Palemon, and Stimpson, in founding Urocaris, placed 


ATCA Reclinsp yells: + References will be found below. 


_ 


BORRADAILE—ON THE PONTONIINA 329 


it between Anchistia and Palemonella. Both Palemonella and Urocaris consist of 
gracefully built species, and until recently it has generally been considered that their 
affinities are Paleemonine rather than Pontoniine. In 1878 Kingsley founded a sub- 
family Pontonine for the genera Pontonia, Coralliocaris, Harpilius, Anchistia, Urocaris, 
and Typton, together with the Huryrhynchus of Miers and Palemonetes of Heller 
which are certainly Paleemonine and have been treated as such by all subsequent 
authors. In 1879, however, Kingsley treated the members of the Pontonine merely 
as a section of his Palemonine. In 1888 Bate raised them to the rank of a 
family—the Pontoniide. In 1890 Ortmann made considerable contributions to our 
knowledge of this family, and in 1899 he added to it the Hymenocerine, placing 
its original members in a subfamily Pontoniine. The true affinities of Hymenocera, 
however, are not represented by this arrangement*. In 1898 I revised the Pontoniide, 
establishing a new genus’ Anchistus for some species intermediate between Pericli- 
menes and Pontonia. Since then Nobili has founded in 1902 the genus Coutierea, 
and in 1906 Stegopontonia, both related to Coralliocaris. Ancyclocaris, established 
in 1902 by Schenkel for a prawn which is now known to shelter under the pro- 
tection of a sea anemone, was connected with the Pontoniide by Nobili in 1906. 
In 1907 I reverted to the view that the Pontoniine should rank as a subfamily 
of the Palemonide. Finally, Sollaud, in an illuminating article (1910), discussed the 
characteristic features of the Pontoniide, and definitively included among them the genera 
Palemonella and Urocaris. In the present paper five new genera—Uvrocaridellat, 
Pontomopsist, Periclimeneust, Pontondes, and Harpiliopsis—are proposed, and four 
subgenera established within the genus Periclimenes?. 

The history of the Pontoniine as a unit of classification may be summed up as follows: 


Subfamily Pontoniine Kingsley, 1878. 


Pontonine Kingsley, Bull. Essex Institute, x. p. 53 (1878). 

Pontonnde Bate, “Challenger” Macrura, p. 927 (1888). Ortmann, in Spengel, Zool. 
Jahrb. Syst. v. p. 460 (1890). Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 376 (1898). 
Rathbun, Bull. U. S. Fish. Comm. xx. 1. p. 120 (1901). Sollaud, C. R. Ae. Sc.., 
Gi. OD, Wiss (LUO). 

Pontoniine Ortmann. Bronn’s Thierreich, v. 11. pp. 1124, 11380 (1899). Borradaile, 
Ann. Mag. Nat. Hist. (7), xix. p. 472 (1907); Ib. (8) xv. p. 206 (1915). 


The Organization of the Pontoniinze. 


The connection between structure and mode of life is very clearly shown by the habit 
of body of Pontoniinze. In Urocaridella (Plate 58, tig. 2) and Urocaris, whose members are 
probably all free swimmers, the body is slender and strongly compressed, with a very long 
sixth abdominal segment. In Palemonella (Plate 58, fig. 5) and Periclimenes (Plate 52, 
fir. 1), which wander, with an activity that probably varies from species to species, over 

* See pp. 405, 408, 410 in this volume. 
+ Preliminary descriptions of these genera have been published in the Annals and Magazine of Natural 


History for February, 1915. 
+ Marygrande Pesta, 1911, is not, in my opinion, distinct from Anchistus. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 42 


330 PERCY SLADEN TRUST EXPEDITION 


the surface of various objects on the sea-floor, or shelter without quiescence upon the 
bodies of echinoderms, where they have neither a stable foothold nor great protection, the 
habit of body is very much that of the Leander of the British coast—slender, but without 
the attenuation of Urocaris. Periclimenes (Plates 54,55, figs. 10—16) shows considerable 
variety in body form, and this is no doubt connected with differences in habitat and 
activity. Periclimeneus (Plate 55, figs. 19, 20) is an aberrant group of Periclimenes 
which has taken on an Alpheus-like habit of body, no doubt in accordance with some 
special mode of life. Pontonivpsis, commensal with a crinoid, is more modified in 
respect of some of its organs than is Periclimenes, but in regard to habit of body 
is in much the same condition. Ancyclocaris, living within the zone of protection of 
a large sea-anemone, whose mouth it even enters at times, has a more stoutly built 
body with, in the female, a hump on the back of the thorax which suggests some 
peculiarity in the ovary. Harpiliopsis. Harpilius, and Coralliocaris are adapted by 
their heavy, depressed bodies to a sluggish life among the branches of coral stocks, 
in that habitat which swarms with Trapezia and other small crabs, and is such a 
rich collecting ground for the marine zoologist in the tropics. Stegopontonia is an 
aberrant member of this group of genera, adapted for an existence spent among the 
spines of a sea-urchin, and Coutierea, also related here, bears, with its spiny, keeled 
abdomen, and long rostrum and supraorbital spines, the characteristic appearance of a 
deep sea crustacean. Anchistus, Pontonia and Conchodytes, which live within the shells 
and tests of bivalves and ascidians, have reached the highest degree of specialization 
in the subfamily. Their swollen, clumsy bodies, though not more depressed than those 
of Coralliocaris and Harpiliopsis, have in some cases almost lost the semblance of 
prawns, and suggest degeneration as strongly as that of the crab Hapalocarcinus, 
to which they bear no little resemblance. Typton, though heavily built, is compressed 
and prawn-like, and is no doubt adapted by its shape to life in the canal system 
of the sponges which it inhabits. In all these cases, the male, though his body tends 
to undergo the same modifications as that of his mate, has a more normal appearance 
than she, principally because his abdomen is less enlarged (Plate 57, fig. 29). 

Thoracic sterna are distinct and well formed in all genera of the subfamily, 
though narrow between the maxillipeds, and there is not in this respect the difference 
that might be expected between Periclimenes and the more stoutly built forms. In 
Urocaridella and Urocaris, the sterna are narrow, in correspondence with the com- 
pression of the body. 

The abdomen of the Pontoniine is of the caridean type, with a fairly large 
first segment, a very large second segment, and a bend between the third and fourth 
segments. This bend is most marked in Urocaridella and Urocaris, but it is distinct 
in all other genera except Pontonia, Pontonides, and Conchodytes, where it is merged 
into a general curvature of the hinder segments. In the more sedentary genera the 
pleura of the first three, segments of the female are very large and foliaceous, and 
form, with the incurved tail fan, a great pocket for the eggs. In Urocaridella and 
some species of Urocaris the hinder part of the third tergum is swollen into a hump, 
and bears a hooked process which overhangs the succeeding segment. 


BORRADAILE—ON THE PONTONIIN A 331 


Like other prawns, the Pontoniine have an armature of spines on prominent 
parts of the body. The use of this armature is realised at once when a prawn is 
held alive in the hand. With its abdomen bent under the thorax, the animal is a 
wedge-shaped object with the broad end in the direction of escape—that is back- 
wards—and all its spines projecting forwards, so that they preserve for it any ground 
which it gains in its struggles. The armature includes the rostrum and its teeth, 
spines on.the carapace, on the two pairs of antennae, and often on the uropods, and 
in a few cases a hooked projection at the back of the third abdominal segment. As 
might be expected, it is less developed in the heavily built sedentary forms than in the 
lighter species which live a more exposed life. 

The rostrum is probably seen in its earliest form in some of the species of Perv- 
climenes. Typically, in this genus it is a straight, compressed, lanceolate structure 
(Plate 54, fig. 10 a), of about the same length as the region of the carapace behind it, and 
bears above and below a fairly deep, toothed crest, the upper crest being continued upon the 
carapace by a row of two or three teeth. This form of rostrum is well seen in P. scriptus. 
In certain cases it loses some or all the teeth of the ventral crest (Plate 54, fig. 9 a) or 
even the crest itself. Sometimes, as in P. brevinaris, it becomes shorter. In other 
species, as in P. ensifrons, it is curved upward at the end, so that the upper edge becomes 
concave. Often, as in P. spiniferus (Plate 52, fig. 1 a), the curved rostrum is longer and 
slenderer than the lanceolate type, and this tendency reaches its height in P. borradailei 
and P. kolumaduluensis. Uvrocaridella (Plate 58, fig. 2) has a very long and much 
upeurved rostrum, toothed above and below, with the dorsal crest high at the base. 
Urocaris loses the ventral teeth of this rostrum, and in some species has the organ 
shortened by the loss of the slender distal part. Palemonella has a shallow rostrum, 
usually rather short and nearly straight. In Ancyclocaris it is short and straight, but 
of a good depth. The more sedentary genera show varying degrees of reduction and 
modification of this rostrum. The reduction is least marked in Harpiliopsis, Harpilius, 
and Coralliocaris. In Harpiliopsis and Harpilius the rostrum hardly differs from that 
of Periclimenes, but is rather small, and wide at the base. In Coralliocaris it is wide 
and shallow and the teeth show signs of reduction, though they are usually present, at 
least on the upper side. The free end is pointed both in dorsal and in side view, and 
is not curved downwards. In Anchistus, Pontonia (Plate 57, fig. 29), and Conchodytes, 
on the other hand, the rostrum, though it is more or less depressed at the base, is 
(except in some species of Pontonia) deep and strongly compressed in the distal part. 
The tip is bent downwards, and in side view rounded or diminished abruptly to a point. 
Teeth are generally absent, but in some species of Anchistus there are small dentations 
near the tip. In Typton the rostrum is peculiar, being small, compressed, almost or 
quite toothless, and bent upwards at the free end, which is pointed. Stegopontonia and 
Pontomopsis (Plate 57, fig. 27), Pontonides (Plate 57, fig. 28), and some species of 
Pontoma, differ from all other Pontoniine in having a broad and very shallow rostrum, 
lanceolate or triangular as seen from above. 

Of the spines of the carapace the most persistent is the antennal, which is never 
entirely lost, though in some species of Conchodytes it becomes very blunt. The 

42—2 


332 PERCY SLADEN TRUST EXPEDITION 


hepatic spine is generally present in free-living species but it may be absent even 
here. It is found in Harpiliopsis but not in most species of Harpilius and Coralliocaris, 
or in the still more sedentary genera. The supraorbital spine is kept only by some 
Periclimenes (Plate 52, fig. 1), Coralliocaris rhodope, Typton and Couterea. In the 
latter two cases it is very large. True branchiostegal spines are never found. There is 
a pterygostomian spine in Coutierea. The lower angle of the orbit may be produced, 
but is generally blunt. 

Here may be mentioned the grooves of the carapace. These are not well marked 
but very persistent. Groove b of Boas* is always present and there is usually some 
trace of e, though the latter never crosses the back. 

The telson corresponds in width with the stoutness of the body, but always 
narrows towards its hinder end, which is triangular, with sometimes, in primitive forms, 
a pointed projection in the middle (Plate 52, fig. 1p). On the dorsum of the telson are 
two pairs of stout moveable spines. On each side of its hinder end are three spines, 
of which the outer is short and stout, and the intermediate long and strong. The 
sub-median is usually shorter than the intermediate and longer than the outer. It is 
also more slender, and in primitive forms often feathered (Plate 58, fig. 2 p), showing thus 
a transition to the feathered bristles of other Paleemonide. In Conchodytes, Pontoma, 
and some Anchistus, however, the sub-median and intermediate spines are nearly equal, 
and in such cases they are usually of a clumsy, degenerate shape. The end of the 
telson also often bears two or three pairs of fine, unfeathered bristles, whose presence 
bears no relation to the degree of degeneracy of the species. 

The eyes are set widely apart on a region immediately below the rostrum. 
Between them is a low swelling on which is seen the remains of the larval median 
eye (Plate 52, fig. 1c). Hach eye-stalk starts with a narrow flexible region and suddenly 
expands into a wide region with stouter walls. This region is usually a little flattened, 
and at its end bears the cornea, which generally swells out beyond it at the sides. 
Such an eye is well seen in Periclimenes (Plate 52, fig. 1 b) and the other little-modified 
genera, and also in Harpiliopsis, Harpilius, and Coralliocaris. In Anchistus, Pontonia, 
and Conchodytes, however, the eye, as might be expected, is smaller. It is also here 
more cylindrical, and the cornea is either of equal width with the stalk or even 
narrower. The eye of Pontonides (Plate 57, fig. 28 b), in spite of the general resemblance 
of this prawn to Pontonia, is well developed and of the Periclimenes type. In Ancycio- 
carts, on the other hand, though the build of the body is Harpilius-like, the eye 
resembles that of Pontonia. Several interesting peculiarities are found. In Perieli- 
mencus (Plate 55, fig. 20 a) the moderately large cornea has a curious cup-shaped depression 
on the outer and lower side. In Pontonopsis (Plate 57, fig. 27 b) the eye is large and 
sub-globular, the cornea occupying one-half of the sphere. In the sub-genus Cornger, 
of Periclimenes, the cornea is ogival, and in P. (C.) ceratophthalmus (Plate 54, fig. 9 b) this 
peculiarity culminates in the appearance of a papilla, such as is found in Phyl- 
lognathia and in the amphibious Ocypode ceratophthalmus. It would be interesting 
to determine the physiological result of the structure which thus recurs in ‘genera 

* K. Danske Vidensk. Selsk. Skrifter (6), ii. 1. p. 25 (1880). 


BORRADAILE—ON THE PONTONIINA 333 


which are so widely separated, and differ even in respect of the medium through which 
light reaches the eye. 

Genera with depressed bodies have the carapace hollowed behind the eye to give 
room for it to be turned backwards. In Coralliocaris this depression becomes a 
sharply-marked pit. There is here a striking repetition of the condition which in the 
crabs may be traced through further stages till it ends in the formation of a closed orbit. 

In the antennules (Plate 52, fig. 1c), the basal joint bears on its outer side a thin 
flange which, generally speaking, varies in width with the body. At the base of this 
flange, on the outer side, stands the stylocerite, a forwardly-curved, strong spine, which is 
usually less than half the length of the joint, but sometimes, as in Harpiliopsis and 
Coralliocaris, becomes more important.. At or near its end the flange bears another 
spine, smaller than the stylocerite. In Urocaridella and Urocaris the flange has a 
rounded, fringed tip, projecting beyond the end of the joint, with the distal spine at 
its side, so that the whole structure recalls the antennal scale. In Palemonella and 
Periclimenes the end of the flange, slightly concave or convex, and usually fringed, 
is nearly on a level with the end of the joint, and bears its spine at its outer angle. 
In some species of Periclimenes (P. frater, etc., Plate 58, fig. 6 c) there are two spines. In 
the heavier genera there is an unfringed projection forwards which bears the spine at 
its tip. In some species of Conchodytes this projection is blunt, the spime having dis- 
appeared. A well-developed statocyst is always present, which is not the case in all 
Carides. The second and third joints of the stalk vary a good deal in form. Typically 
they are sub-cylindrical and about as long as broad. 

The inner flagellum is simple. The outer is more complex and shows variations 
of some importance. It is always thickened at the base, and bears in the thickened 
region a fringe of coarse, sensory hairs. At the end of this region, in all genera 
except Typton, a cleft separates a long, slender flagellum from a short, thick one, which 
is a continuation of the thickened region. The depth of the cleft varies. Usually it is 
less than the length of the uncleft basal region. In most Periclimenes (Plate 52, fig. 1 ¢), 
some Palemonelle, and the heavier genera, it is a good deal less. In some Periclimenes 
(as P. compressus, Plate 55, fig. 18), some Palemonelle (as P. bater), and in Urocaris, 
it is nearly or quite as long. In Urocaridella (Plate 58, fig. 2a) and Ancyclocaris it is 
longer than the basal region, and the antennule is then said to be “ triflagellate.” 
This is the condition found in other Palzemonidee. Since the genera of Pontoniinze 
which exhibit it are in other respects more primitive than the rest of the sub- 
family, it would appear to be in their case an ancestral feature, though of course the 
appearance of the third flagellum is in itself an innovation, and is one of the non- 
primitive features of the Paleemonide. The shortening of the cleft in higher Pon- 
toniine is thus a return to primitive conditions, not a retained primitive feature. It 
is comprehensible that the lessened activity of these members of the subfamily should 
remove the necessity for a very mobile sensory apparatus. 

The antenna (Plate 52, figs. 1c and ¢’) is of the normal caridean type. On the first 
joint’ (coxocerite) the tubercle for the opening of the green gland takes the form of a 
well-developed process on the inner side. The second joint (basicerite) is divided 


334 PERCY SLADEN TRUST EXPEDITION 


longitudinally into two parts*—an inner, which bears the flagellum and its stalk 
(together forming the endocerite), and an outer, which bears the exocerite in the form of 
the scale (scaphocerite). On this outer division stands also a forwardly-directed spine, 
which may, as in Harpiliopsis, be very long and strong, or may be represented merely 
by a slight projection of the jomt. As this spine is not homologous with the stylo- 
cerite of the antennule, standing indeed on a different joint, the same name should 
not be applied to it. The scale has a broad, fringed expansion and a stout, un- 
fringed outer border, which ends in a distal spine. Its most remarkable variation is 
found in Typton, where it is reduced to a vestige, which does not reach as far as the 
beginning of the flagellum, though it shows traces both of the expanded region and 
of the distal spine. In the endocerite, the flagellum stands on a two-jointed stalk. It is 
doubtful whether there is any justification for calling the joints of this stalk ‘‘ischiocerite” 
and ‘‘merocerite.” The flagellum, like those of the antennule, is shorter in the sedentary 
than in the active forms. It is shortest im Conchodytes tridacne and C. meleagrine. 
The mouth parts show, on the whole, an absence of modification, either within 
the subfamily or in comparison with those of related groups, which is rather striking 
in view of the specialized nature of the food of many of the commensal forms—such, 
for instance, as those which live with crinoids or lamellibranchs—consisting as this 
must largely of minute organisms collected by the feeding currents of the host. The most 
remarkable features exhibited by the organs in question are (1) the tendency which 
appears in various genera to a broadening of some or all the joints of the third maxilliped, 
(2) in the Pontoma group, that the inner lacinia of the maxillule is very wide and hairy. 
The connection of these features with the functions of the organs is discussed below. 
The mandible (Plate 52, fig. 1 and Plate 57, figs. 26 d—d’”) is deeply cleft into two 
diverging processes, both directed obliquely towards the median plane of the body. One of 
these—the incisor process—is thin and ribbon-like, trending at its base downwards, but 
curving inwards and at the same time twisting its outer edge forward, so that, while at 
its base it is nearly vertical, with its width transverse to the body, at its free end it is 
* (June, 1917. These parts, of which the outer stands always a little forward of the inner, and in 
some Decapoda, as Upogebia, becomes almost wholly distal to it, probably represent the two components 
(protobasipodite and metabasipodite) of the basipodite (symbasipodite). I have recently discussed the 
composition of this segment (Proc. Zool. Soc. London, 1917, p. 53). The homologies of the antenna 
present a very difficult problem. ,It would appear from the case of the Mysidacea that the segment upon 
which the green gland opens is the second of the primitive series, and in that case the first, or precoxa, 
must in other cases have disappeared, by excalation or by fusion with the head or with the coxopodite. 
This suggestion is supported by the fact that the maxillary gland of Stomatopoda also opens upon the 
second segment of the limb to which it is adjacent (Proc. Zool. Soc. Joc. cit.), and that the gonoducts of 
the Decapoda, probably homologous with the ducts of the excretory glands, open upon what development 
shows to be the second segment of the legs (except where, as in the crabs, the openings have secondarily 
shifted to the sterna). If Hansen be right in interpreting as segments the inconspicuous structures which 
undoubtedly exist in the antenna of Webalia proximal to the first apparent segment and between those 
which appear to be the second and third, then there is in that genus a very complete and instructive 
retention of the primitive segmentation of the limb. The segments will be, in succession, precoxa (pleuro- 
cerite), coxocerite, protobasicerite, metabasicerite, and the so-called ischiocerite and merocerite incompletely 
fused. If, on the other hand, Hansen’s view be not accepted, then the absence of exopodite and of visible 


opening of the green gland leaves the homologies of the segments obscure, though the flexion of the limb 
suggests that the incompletely double third joint belongs to the endocerite proper. | 


BORRADAILE—ON THE PONTONIINA 339 


nearly horizontal, with its width longitudinal in the body. The other—the molar 
process—is stout and subrectangular in section, and slants dorsally, to end obliquely 
truncated on the median plane. In Urocaridella (Plate 58, fig. 2d”) and Palemonella 
(Plate 53, fig. 5d"), a short, two-jointed palp is present, standing on the anterior side of 
the limb, at the base of, and just dorsal to, the incisor process, behind which it is partly 
hidden in ventral view. The incisor and molar processes perhaps represent the first 
two endites of the primitive crustacean appendage*. The palp represents the rest of 
the protopodite and the endopodite. Its first joint is probably the basipodite, but, in 
view of the facts of meristic variation, it is doubtful whether there would be any validity 
in such a statement as that the second is the ischiopodite. Still less can the third joint 
be regarded as the meropodite alone. 

The incisor process usually ends in three teeth, the middle one of which is shorter 
than the others, but the intermediate teeth may be more numerous and sometimes differ in 
number on the two sides. Thus Coralliocaris japonica has two on one side and three on the 
other, while in Conchodytes meleagrine (Plate 57, fig. 26 d) there are three and four, nearly 
equal to the outer teeth. The molar process ends in a roughly square, concave surface, 
surrounded by an incomplete wall made up of projecting lobes, from four to five in number. 
All but one of these have crescentic or horseshoe-shaped rims, open towards the middle. 
The remaining lobe has an unbroken rim, and a great part of its circumference is beset 
with a fur of bristles or a rough patch of tubercles. Opposite this lobe the wall is lower 
than elsewhere, and in the left mandible is also roughened. This roughening of the molar 
surface is, | think, the last remains of the clothing of bristles described for the Alpheidee 
by Coutiéret, who also figures in some cases an oval isolated area on one side of the 
process. The ridges which in Amphibeteus and some Hippolytide represent a part 
of the bristly surface may perhaps be transitional between this condition and that 
found ‘in Desmocaris (p. 327). There are only very slight traces of a roughened 
surface in Leander, so that possibly the possession of a considerable extent of such a 
surface should be added to the list of primitive characters of the Pontoniinze which 
are not found in Palemonine. The lobes differ a good deal in shape, and there 
is only a general correspondence between those of the mandibles of the two sides. So 
far as this correspondence goes, it is not the mirror-likeness usually found in paired 
structures, but the two arrangements are reversed, so that the rough lobe of each molar 
process overhangs the lowest part of the edge of the other and the surfaces interlock. 

The mandibles lie in a chamber enclosed between the lips (Plate 57, figs. 26s and ¢), 
the hood-like upper lip (labrum) standing in front of them and the large, bilobed lower lip 
(metastoma, Plate 52, fig. 1 q) behind, while the swollen bases of the mandibles themselves 
close in the chamber at the sides. There are two openings to the lip chamber—a 
narrow median slit between the lobes (paragnatha) of the lower lip, and a wider 
transverse gap between the upper and lower lips. The incisor processes close the 

* On the other hand, the incisor process may be merely an outgrowth from the molar process 
(gnathobase). An interesting analogue to it is seen in the flange at the end of the grinding surface of 
the mandible of Apus. As it stands, this flange is on the hinder side of the limb, but the shape of the 


mandible gives reason to suppose that it has been rotated backwards. 
7 Ann. Sci. Nat. Zool. (8), ix. pp. 152—157 (1899). 


336 PERCY SLADEN TRUST EXPEDITION 


transverse opening, meeting in the middle line. The molar processes meet deeper in 
the chamber, just under the opening of the gullet. 

In describing the maxillules and maxille, account must be taken of the fact 
that there is no agreement among authorities either as to the names to be applied 
to the leaf-like processes, usually known as lacinie, which are borne on these limbs, 
or as to the joints which these processes represent. I shall assume the correctness of 
the following view. The maxillules and maxilla of adult Decapoda can be derived 
from a type provided with five endites and a distal region which probably represents 
several further segments. Of these endites, the first stands in the region which 
represents the precoxal joint, the second proceeds from the region of the coxopodite, 
the third and fourth from that of the basipodite, and the fifth from that of the 
ischiopodite. The distal region forms the remainder of the endopodite. The lacinize 
of the maxillule represent the first and third endites. I shall call them the nner and 
outer lacone. The two cleft lobes usually known as the “lacinie” of the maxilla 
represent each two endites. I shall call them the first and second lobes. The first 
represents the endites of the precoxa and coxopodite, the second those of the basi- 
podite. Boas* and Coutiéret have shown that the endite of the ischiopodite is 
present in the larva, and in the adult is represented by a slight swelling often found 
at the base of the endopodite. The laciniz of the first maxilliped vepresent the 
coxopodite and basipodite (two fused). 

In the maaillule (e in figs.) the inner lacinia curves towards the outer. It is of varying 
width, widest in Conchodytes (Plate 57, fig. 26¢) and Pontonia, wide also in Anchastus 
(Plate 56, fig. 25e), moderate in Periclimenes (Plate 52, fig. le), narrow in Harpiliopsis 
(Plate 56, fig. 22¢) and Urocaris (Plate 58, fig. 3¢). It bears a varying number of bristles, 
of which a bunch at the end are stouter than the rest except in Conchodytes. The outer 
lacinia also varies in width. It has an edge directed towards the middle line of the body 
and bears along this edge a number of stout spines and usually also some that are more 
slender. In Anchistus, Pontonia, and Conchodytes both lacinie are very hairy. The 
endopodite is bifid at the tip, the proximal branch, which possibly represents the - 
fifth endite, being curved in a short spiral, the distal branch nearly straight. The latter 
is reduced to a faint lobe in Harpiliopsis (Plate 56, fig. 22e), Anchistus (Ib. fig. 25e), 
Pontonia, and Conchodytes (Plate 57, fig. 26e). The maxillules stand close against the 
paragnatha, the lacinize of each opposed to those of the other across the middle line, 
but not quite meeting them. The inner lacinie are behind the cleft of the meta- 
stoma, with a low median ridge between them; the outer stand beside the cleft. 
The endopodite (endognathite) is directed outwards and appears to give a purchase 
to the action of the limb by hooking its curved process around the outer edge of 
the lip, which is provided with a notch to receive it. 

In the maxilla (f im figs.) the first double lobe has disappeared, and is repre- 
sented only by a slight swelling of the edge of the limb, on which a faint notch 
sometimes still indicates its double nature. The second lobe is of a good length, 


* K. Danske Vidensk. Selsk. Skr. (6), Afd. ii. 1. pl. 2, figs. 79, 80 (1880). 
jy Ann. Sci. Nat. Zool. (8), ix. pp. 157—167 (1899). 


BORRADAILE—ON THE PONTONIIN A 337 


but tends to become simple. In this respect it is very variable, and the maxille 
of the two sides may differ in the same individual. The lobe is simple in the species of 
Pontonia and Conchodytes (Plate 57, fig. 26 f) I have been able to examine”, and also in 
the only specimen of Periclimeneus robustus I have dissected (Plate 55, fig. 20 f), in some 
Coralliocaris (as C. japonicus, Plate 56, fig. 23 f) though not in others (as C. macrophthalma, 
Plate 56, fig. 24), and, according to Sollaud, in Harpilius. In Harpiliopsis it is double, 
with obsolescent proximal lobule. From the frequent occurrence of reduction in the lobes 
of this limb in the Carides, it would seem that they are not of great physiological 
importance. The endopodite is gently swollen at the base, perhaps by the remains of the 
fifth endite, and the exopodite (scaphognathite) varies in shape and width with the gill- 
chamber, being broad in depressed forms, particularly in Harpiliopsis and Coralliocaris. 

In the first masailliped (g in figs.) two endites—the second and third—are nearly 
always recognizable, and usually separated by a very distinct notch. The edge of 
the proximal (second) endite is often indented by a shallower notch. The endopodite 
is usually simple and tapering, with a stout bristle at a short distance from the 
tip, which may perhaps represent a joint. In the Anchistus-Conchodytes group the 
endopodite is shorter and blunter, and often shows traces of a transverse joint. At 
the tip of the exopodite (exognathite) a few small joints may or may not be present, 
and at its base on the outer side is a fringed lobe (the lobe a of Boas) which generally 
varies in width with the body of the species, though it is wider than this would indi- 
cate in Urocaridella (Plate 58, fig. 2g) and some Urocaris. The epipodite (mastigobranch) 
varies greatly in size. Its outer border is usually notched, but in Periclimenes the 
notch is shallow or wanting. It is deep in Urocaridella, which is remarkable if it is to 
be regarded as an unprimitive feature. The lobes separated by the notch may be equal, 
or the distal one may be longer than the other and pointed. 

The second maailliped (h in figs.) has the typical caridean form, that is to say, 
(1) its main axis consists of six joints, owing to the fusion of the basipodite and 
ischiopodite, (2) the last two joints are bent strongly backwards, so as to lie parallel 
with the ischiobasipodite, (3) owing to the growth backward (morphologically forward) 
of a process of the propodite on the outer (morphologically mner) side of the dactylo- 
podite, the latter comes to lie not at the apex of, but along the median side of the 
propodite. Thus these two joints are compacted into a firm plate, which presents 
a long median margin, fringed with bristles, against its fellow of the opposite side. 
To the third of these characters there is an exception in the case of the genus 
Harpilius, where, according to Dana's figure for the type species (H. lutescens), there 
is a very remarkable return to the primitive condition in respect of the position, 
though not of the shape, of the dactylopodite. This arrangement is approached in 
at least one species of Coralliocaris (C. superbus). The exopodite is obscurely annulate 
in the greater part of its length, and bears at the end a comparatively small number 
of true joints. The coxopodite carries on the outer side a simple oblong epipodite, 
and on the median side a knob bearing bristles which probably represents an endite. 

* Pontonia ascidicola and Conchodytes meleagrine and tridacne. Ortmann (Zool. Jahrb. Syst. v. 
Pl. 37, figs. 9 f., 10 £.) figures a cleft in the maxilla of P. tyrrhena. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 43 


338 PERCY SLADEN TRUST EXPEDITION 


Its gill will be mentioned later. A swelling which is sometimes present at the base 
of the coxopodite may represent the precoxal endite. 

The main axis of the third maailliped (i in figs.) consists, in Urocaridella (Plate 58, 
fi. 27), of five joints, the ischiopodite being fused with the meropodite, and the propodite 
with the dactylopodite. Curiously enough, this arrangement is found also in Concho- 
dytes (Plate 57, fig. 267) at the other end of the pontoniine series. In all other genera 
the basipodite is fused with the ischiomeropodite, though the junction is still marked 
by a notch. The coxopodite bears on the outside a small, rounded epipodite, and 
often also on the inside a knob with bristles which is perhaps to be regarded as an 
endite. The exopodite is obscurely annulate and at its end there are usually a few 
longer segments, which are sometimes true joints but in other cases appear to be 
marked merely by a change in the width of the organ and the attachment of bristles. 

The long joint of the endopodite (ischiomeropodite) is in Urocaridella (Plate 58, fig. 27) 
and Urocaris (Plate 58, fig. 37) straight, with the outer side somewhat swollen at the 
base. In the other genera it is almost always more or less curved, with the concave side 
towards the middle line of the body. It is always ribbon-like, and shows throughout the 
subfamily a tendency to widen. It is narrow in Urocaridella, Urocaris, Ancyclocaris, 
Palemonella, Periclimenes, Periclimeneus, and Pontoniopsis, though in some species of 
Periclimenes (Plate 54, fig. 87) and in Periclimencus (Plate 55, figs. 19 and 2072) it isa little 
increased in width. In Harpiliopsis (Plate 55, fig. 21 and Plate 56, fig. 22 2), Coralliocaris 
(Plate 56, figs. 23 and 247), and Anchistus it is wide or narrow according to species, reaching 
in some Anchistus the greatest width it attains in the subfamily. In Harpilius it is wide, 
but narrows towards the distal end. In Pontonia and Conchodytes (Plate 57, fig. 26 2) it is 
broad. The last two joints are always a little narrower than the ischiomeropodite, but are 
of approximately the same width as it in the species in which it is narrow and in Corallio- 
caris. In Harpilius, in some species of Anchistus, and in Harpiliopsis beawpresi, they 
remain narrow though the ischiomeropodite is expanded. In Pontonia and Conchodytes 
they are wide, though the widening is not equally pronounced in all species. They 
have always a flat ventral surface but are sometimes, as in Coralliocaris, stoutly 
built. The curving of the ischiomeropodites brings the last two joints of each third 
maxilliped near to those of its fellow, so that, while the ischiomeropodites lie at the 
sides of the mouth, with a wide gap between them, in which the second maxillipeds 
are exposed, the distal parts of the limb lie side by side in front of the mouth region. 
_ A further complexity in the arrangement of the parts of the limb is brought about by 
the fact that the ischiomeropodites are twisted, so that the flat surface of the appendage, 
which in its distal part is in a horizontal plane, is in the proximal part in a plane 
between the horizontal and the vertical. This arrangement has the effect of forming 
a kind of basket below the mouth region, walled in at the sides by the ischiomeropodites, 
which are of course more efficient in that respect the wider they are, and by the long 
bristles which project downwards and inwards from the median edges of these joints. 
In front, the distal part of the limb, with its bristles, affords a surface, horizontally 
placed below the antennal region while the appendages are outstretched, which by 
bending can be brought ventrally under the mouth area to complete its enclosure. 


BORRADAILE—ON THE PONTONIINA 339 


The process of feeding has not been observed in Pontoniinz, and as yet we do 
not even know upon what they feed, though it may be inferred that the species 
which live in intimate association with sessile or subsessile animals probably share 
the food of the latter, which consists of minute organisms swept up by ciliary currents 
from the surrounding water. In regard to the use of the organs around the mouth, 
something may be gathered from a study of the prawns of the genus Leander, in which 
these organs are very similar to those of the Pontoniinze in their general structure 
and arrangement. In Leander serratus there are bristles, borne upon ridges of the 
coxopodite, basipodite, and ischiopodite of the first leg, which complete behind and 
below the basket under the jaws, but which are less well-developed or absent in 
Pontoniinee. Small particles of food may be seized and conveyed by the chelipeds 
of either pair to the region of the mouth, where they are generally received by the 
second maxillipeds, though sometimes they appear to be placed directly in charge 
of more dorsally placed structures, probably the maxillules. A large morsel occasionally 
appears to be steadied by the legs of the second pair, while those of the first pair 
tear off fragments and carry them to the jaws, but it is more often placed as a whole 
within the grasp of the second maxillipeds, which hold it in place while pieces are 
torn off it by deeper-lying organs, probably in the main by the incisor processes. In 
handling bulky masses of food the chelipeds are assisted by the third maxillipeds, 
which bend back their last two joints for the purpose. The third maxillipeds are 
also capable by the same action of scooping up food and unaided carrying it to the 
second maxillipeds, between which they sometimes thrust it with their tips. During 
these processes the basket appears to serve the purpose of keeping the food under 
control till it has been seized by the second maxillipeds. These are very important 
organs, and play an indispensable part in passing the food to the mandibles. The 
animal can still feed if the legs and third maxillipeds have been removed, but if all 
the other organs be left and the second maxillipeds cut away it is apparently incapable 
of taking food. The second maxillipeds have three principal movements. In one, 
the broad flaps in which they end open downwards like a pair of doors and with their 
stout fringes gather up the food, in another they rotate in the horizontal plane to and 
from the middle line, in the third the bent distal part of the limbs tends to straighten 
so as to brush forward any object which lies between them. Frequently these move- 
ments are combined. Once the food is past the portals formed by the second maxillipeds 
its course is hard to trace, but the following seems to be its fate. If it be small in 
quantity and finely divided, or very soft, it is abandoned to the action of the maxillules, 
by whose strong, fringed lacinis it is swept forwards and probably caused to enter 
the mouth chamber through the slit between the paragnatha™. If it be bulky or tough, 
the second maxillipeds assist the maxillules in brushing it forwards towards the incisor 
processes. The action of these latter is not so much a cutting one as a process of 
tucking the food into the mouth chamber, by first backing outwards and then moving 


* Such is the impression made. But it may be that all food is worked into the mouth chamber by 
the incisor processes, and that the function of the cleft on the metastoma is to enable the paragnatha to 
part and give room for the admission of large pieces of food. 


43—2 


340 PERCY SLADEN TRUST EXPEDITION 


inwards and rotating upwards. No doubt during this the food undergoes some tearing, 
and when the mass is large pieces have to be torn off it before they can be swallowed. 
The palp does not appear to take any mechanical part in the process. If it has a sensory 
function this is probably not of great importance, for the organ is present and absent 
in closely related genera in many cases among Carides. Finally, to enter the gullet, 
the food must pass between the molar processes, and be pounded by them as it goes. 
Their concave ends are usually found to be clogged with a pasty matter. They must 
do their work very quickly, for the movement of the mandibles, as judged by that 
of the incisor processes, ceases very soon after the food leaves the latter. How swallowing 
takes place is not clear. Parker and Mocquard suggest that the food of Decapod 
Crustaceans is caused to pass up the gullet by suction from the crop (stomach), but, 
as I have shown elsewhere*, the case of the land hermit-crabs of the genus Cenobita 
throws doubt upon this explanation. It may be that the constrictor muscles of the 
cesophagus play some part in the process. 

The first maxillipeds and the maxille probably take no very prominent part in 
manipulating the food. The feeble lobes of the maxilla are in incessant movement 
to and from the middle line as they are carried inwards and outwards by the action 
of the scaphognathite. It seems likely that their sole function is to regulate the motions 
of the latter. The large lacinia of the first maxilliped is a rather feeble structure 
with slender, silky bristles, and is not strongly moved during feeding. Probably, by 
covering the lobes of the maxilla, it prevents them from being clogged by the food. 

The part played by the paragnatha seems to be a passive one. The labrum 
undergoes active movements whose function is probably to aid in keeping the food 
under the action of the incisor processes. 

The exopodites of the maxillipeds are in constant rapid motion, setting up by 
their activity a strong current forwards from the mouth. No doubt this assists in 
carrying away the foul water from the gill chambers and the excreta of the green 
glands poured out at the base of the antenne. But it has also a significance in the 
feeding process. From time to time particles are rejected by the second maxillipeds, 
which kick them violently forwards, the distal parts of the third maxillipeds at the 
same time straightening so as to admit them to the outgoing stream, by which they 
are swept away. 

It has already been stated that the mouth-parts of the Pontoniinz, even in the 
most decidedly commensal species, are not highly modified as compared with those 
of the free-living Palzmonine. For such modifications as exist it is, however, possible 
to suggest a connection with the nature of the food. The broad third maxillipeds seem 
better adapted to shepherd a crowd of minute organisms than narrower types of those 
organs, and the large, hairy lacinie of the maxillules of the Pontonia group perhaps © 
serve the same purpose. 

The legs of the first pair are chelate, but relatively slender and never of great 
length. They are alike and equal. Their chelee are usually of simple form, but in 


* Gardiner’s Fauna of the Maldives, i. p. 79 (1901). 


BORRADAILE—ON THE PONTONIIN At 341 


several cases (Periclimenes spiniferus (Plate 52, fig. 1k), P. soror, etc.) they have a comb- 
like arrangement of fine teeth on the opposed edges of their fingers, and in others the 
tips of the fingers are double. The chela usually bears tufts of stiff hairs. The first 
three joints may show traces of the bristle-bearing ridges found in species of Leander. 

The legs of the second pair are the stoutest limbs of the body, and nearly always 
longer than the first pair. They may be equal or unequal, alike or unlike, and that 
sometimes within the limits of one genus. They are rarely of complicated shape. 
Their fingers have usually some stout teeth on the apposed edges, but sometimes 
they are blade-like. In Periclimenes denticulatus they are edged with numerous fine 
denticles. In P. petitthowarsi and P. spiniferus (Plate 52, fig. 1 /) each finger of the larger 
member of the pair bears a curious pit with raised edges, and in the great chela of 
Periclimencus (Plate 55, figs. 19, 20) a knob on one finger fits into a socket on the other, 
somewhat as in Alpheus. Curiously enough, in one of the two species as yet discovered 
the knob is on the fixed finger, while in the other it is on the “thumb” or movable 
finger (dactylopodite) as in Alpheus. 

The stoutness of the limb varies, roughly speaking, with that of the body. In 
Urocaridella (Plate 58, fig. 2a) and Urocaris it is very slender throughout. In Palemo- 
nella (Plate 58, fig. 5 a) it is stouter but still slender. In Perzclimenes it varies in stoutness 
but is never very heavy. In Ancyclocaris it is short and rather stout. In Periclimeneus 
one is massive and Alpheus-like in keeping with the general build of the body, and the 
other smaller and simpler but somewhat of the same type. In Harpilius lutescens both 
are rather slender and insignificant. In Pontoniopsis the larger is heavy, but not 
monstrous, and of fairly simple shape, while the smaller is very slender. In Harpiliopsis 
the palm is long and fairly heavy, and in Coralliocaris (Plate 56, figs. 23 /, 1’) this tendency 
is enhanced and the chela is a characteristic organ, long and heavy, swollen at the 
base, and narrowing towards the fingers, which are short. In Anchistus it is of moderate 
size, with parallel sides and fingers of a good length. In Pontoma (Plate 57, fig. 29) and 
Conchodytes it grows heavier, but the palm is still generally of fairly even width and 
the fingers generally not very short, though in some cases there is an approach to the 
condition of Coralliocaris. The same may be said of Typton. 

The thumb is on the outer side of the limb, and usually moves in a nearly horizontal 
plane, but a little upwards or downwards*. Sometimes, however, it becomes almost 
or quite vertical, and in this case it is generally above the other finger, but may be 
below, as in Coralliocaris japonica (Plate 56, fig. 23 0). 

In many cases, especially in Periclimenes (Plate 52, fig. 1 a), one or more stout spines 
are found at the end of the “wrist” (carpopodite), or “arm” (meropodite), or both. 

The mode of use of the chelipeds is as yet unknown. Presumably those of the 
first pair serve as cleaning organs, as they certainly do in Leander. They are no doubt 
also used to convey food to the mouth, as in Leander. The chelz of the larger pair 
are perhaps, as in other cases, used for seizing and tearing masses of food and as weapons 
of offence both against prey and against foes; but they seem unsuited for handling 


* This fact is exaggerated in many of the figures, by slightly twisting the hand, in order to show the 
shape of the fingers. 


342 PERCY SLADEN TRUST EXPEDITION 


the minute organisms which must form the food of some species, and it is hard to imagine 
that a Conchodytes, for instance, can have many enemies against which such weapons 
would be of any avail. They may have sexual uses, but rarely differ in the sexes. 
The peculiar features which they present in the several species are quite unexplained, 
and the study of this limb in relation to the habits of the species presents an interesting 
field of observation. 

The legs of the last three pairs are similar in all but minor details. In Urocaridella 
and some species of Urocaris they are directed forwards as in Palemon, but in Urocaris 
psamathe there appears a change in their position which increases as the build of the 
body becomes heavier, and is probably to be regarded as an adaptation to walking. 
The basipodite is here curved so that, whereas the coxo-basipodite joint is transverse 
to the main axis of the body, the basi-ischiopodite joint tends to become longitudinal, 
and the limb is thrown outwards, and thus the body has a wider area of support. A row 
of movable spines is often found on the lower side of the propodite (Plate 52, fig. 1m), 
and sometimes there are spines on other joints. The dactylopodites vary greatly. 
In the more lightly built species they are generally slender, nearly straight, and simple 
(Plate 52, fig. 1m). In Harpilius, Harpiliopsis, Coralliocaris, and Conchodytes, they are 
strongly curved. Coralliocaris (Plate 56, fig. 23 m) has at the base of the joint on the 
under side a curious prominence which is usually more massive than the terminal claw. In 
Stegopontonia this prominence is paired. Conchodytes (Plate 57, fig. 26 m) has a swelling in 
the same position. In members of various genera the end-claw may be duplicated by a 
spine underneath it, so that the limb becomes “ biunguiculate” (Plate 55, fig. 19 and 
Plate 57, fig. 26m). In some cases the end-claw is exceedingly sharp, in others it is stouter 
and blunter. No doubt each of these peculiarities is an adaptation to the substratum 
upon which the species possessing it moves. Most of them are not as yet susceptible of 
explanation, but when the claws are both sharp and curved they are often dug into the 
soft tissues of the host. 

The abdominal lambs (Plate 52, fig. 1 and Plate 53, figs. 2 o—o”) are of the ordinary 
caridean type and show few remarkable features. The appendix interna is present on the 
second to fifth pairs in both sexes, and on the second pair of the male there is an additional 
process beside the appendix. The endopodite of the first limb is always smaller than 
that of the others, and its inner edge is often crumpled over. Uvocaridella is remarkable 
for having an appendix interna on this limb. This feature occurs also in Amphipalemon 
and is one of many indications of a relationship between the Anchistioidide* and the 
Paleemonide. 

The fullest gill formula found among the Pontoniine is that of Urocaridella and 
Ancyclocaris aberrans. Here there is a pleurobranch for each leg, an arthrobranch for 
the third maxilliped, and a podobranch on the second maxilliped (Plate 53, fig. 2h), with 
a mastigobranch on each of the maxillipeds. A tiny lobe above the base of the mastigo- 
branch of the second maxilliped is perhaps the last vestige of an arthrobranch. A second 
group of genera, comprising Urocaris, Periclimenes, Harpiliopsis, Coralhocaris, Pale- 
monella, Pontonides, Periclhimeneus and Anchistus, have lost the podobranch of the 


* See p. 405 in this volume. 


BORRADAILE—ON THE PONTONIIN At 343 


second maxilliped and reduced in varying degrees the arthrobranch of the third 
maxilliped (c in figs.). In Persclimenes, however, as Sollaud has shown, there 
is often a vestige of the podobranch of the second maxilliped in the form of a finger- 
shaped process at the base of the mastigobranch (Plate 52, fic. 1h). I have not been able 
to satisfy myself of the presence of this process in some of the species I have examined, 
but it is possible that some trace of it is always present. A very minute protuberance 
found in the angle between the mastigobranch and the exopodite in Urocaris, Har- 
prliopsis, Periclimeneus and Anchistus may have the same significance. Curiously enough, 
Palemonella, in other respects the most primitive of these genera, shows a greater 
reduction of the gill of the third maxilliped than Coralliocaris or Harpiliopsis, and has 
no vestige of that on the second maxilliped. In Periclimeneus and Anchistus the 
arthrobranch of the third maxilliped is represented by a slightly folded lobe. In 
Harpilius, Pontonia, Conchodytes and Typton the gills of both maxillipeds are entirely 
lost, save for a hardly recognizable vestige at the base of the third maxilliped of 
Pontona. 

Tt will be seen that, while all Pontoniidze have undergone some reduction of the 
gill formula as compared with other Palemonide, within the subfamily a loss of gills, 
roughly speaking, accompanies the. adoption of a less active mode of life. Where there 
is less need for active respiration there is less provision of respiratory organs. In some 
cases, however (Urocaris, Palemonella, Periclimenes), the loss of gills is in advance 
of the change of habits, at least in so far as that be indicated by the build of body and 
limbs. From this fact Sollaud draws the conclusion that the reduction of the gill 
apparatus is the cause rather than the effect of the altered mode of life. In that case, 
it has presumably come about by meristic variation, more or less fortuitously, and the 
animals have survived by the adjustment of their habits to their anatomy rather than 
by the selection of variations in their anatomy which were suitable to their habits. 
The theory is attractive, and consonant with a good deal in modern biological speculation, 
but before it can be adopted more must be known about the habits of the prawns. The 
build of body is not always an indication of the mode of life, either among the Pontoniinze 
or in other prawns. Metapencus commensalis, for instance, lives in the zone of protection 
of a large sea-anemone, though its general appearance is that of allied species which are 
believed to be free-living, and various gracefully built species of Palemonella and 
Periclimenes are commensal with feather-stars. It may be after all that in the Pon- 
toniinee a change of behaviour has been the forerunner of structural change, and that 
the long series of modifications which we have traced in all parts of the body of these 
prawns, by whatever process of variation or selection it may in detail have been brought 
about, is but the result of congenital laziness in an environment which offers to the 
sluggish endless opportunities of retreat. 


The Colours of the Pontoniine. 


Most species of Pontoniinee have been described on data obtained only from spirit 
Specimens, and information concerning the colouration of the subfamily is consequently 


344 PERCY SLADEN TRUST EXPEDITION 


limited*. It is certain however that they vary greatly in this respect, the animals being 
striped, spotted, or suffused in very different ways and with very different colours, 
while the differences between species are often as great between species of the same genus 
as between those of different genera. In the cases at present known the colours are 
usually gaudy and conspicuous when the prawns are removed from their proper environ- 
ment, though, in some instances at least, their colouration harmonizes very strikingly 
with that of their natural surroundings. Nothing is known as to the colour changes 
which doubtless occur in the lifetime of the prawns, or as to the way in which there 
arises a correspondence between the colouring of the individual and that of its sur- 
roundings, or as to the value this may have to the prawn. 


On Adaptation. 


It will be clear, from various statements in the preceding pages, that our knowledge 
of the significance of peculiar features in the Pontoniine is at present very limited. 
Yet it may not be useless even now to consider the problem they present, if only because 
the alteration which is in progress in our views as to the mode of origin of such features 
may well result in an underrating of their importance in the economy of the organism. 
It has, for instance, quite lately been said that “to see fitness everywhere is mere 
eighteenth century optimism.” Now, as a repudiation of the conception of an all-but 
personified “environment,” which by “selection” forces organisms into a mould pre- 
determined by itself, this statement is undoubtedly justified. But it may also convey 
a belief that an appreciable proportion of the characteristic features of organisms is not 
correlated with peculiarities in their modes of life, and in this sense it is of much more 
doubtful validity. 

There was a time when the structural peculiarities of organisms were widely held 
to be due to the direct action of their environment in bringing them into harmony 
with itself, by means of the plasticity of the individual and the inheritance of the 
characters so “acquired.” Thus every feature was shaped to meet some demand of an 
environment which could not be escaped, and each played its part in ensuring the 
viability of the species. That conception was replaced by one in which the organism 
was allowed more initiative, in that it presented numerous small but ready-made 
modifications to its environment, by which some of them were then “ selected” to become 
the characters of an altered race. This view had in turn to be modified in the direction 
of attributing a greater importance to the organism itself, since it was discovered that 
only some of the variations presented by the organism were heritable, and that these 
were fixed and incapable of dilution or summation. Finally, it has to be recognized that 
what is selected is not this or that “character,” but the organism as a whole, with its 
powers of adapting itself to the world by various means. Thus the part held to be 
played by the environment in the production of any single character has become less and 
less, and therewith has waned the belief that every character plays some part in the 


* Dana, Nobili, Potts, Rathbun and others give information concerning the colours of various species. 


BORRADAILE—ON THE PONTONTIN & 345 


struggle for existence, since that is too often still regarded as the activity of a machine- 
like organism in the face of a fixed and unavoidable environment. 

Now it is true enough that the structural “fitness” of an organism does not consist 
in its characteristics having arisen in response to the fixed demands of an environment. 
But this does not mean that each of them is not caused to play its part in that reaction 
with the world which constitutes the life of the organism, including the habit of seeking 
an environment and maintaining itself there by the necessary measures. These habits 
are without doubt as modifiable, both in the individual and in the race, as any other 
characteristic of the organism, and one of the conditions of the persistence of any given 
structural modification is that it should not exceed the limits of the power of adjustment 
which is given by this susceptibility of modification of habit. So long as the structural 
alteration does not outrun the physiological possibilities, fitness is maintained, and it is 
incredible that this can ever be due to the alteration being indifferent to the life of the 
organism. Some disturbance it must cause, whether in the internal or in the external 
physiology, and in the long run a disturbance in internal physiology will cause a modifica- 
tion in behaviour, as in the amount of food taken, or in the sheltering of an animal which 
by any cause becomes deprived of its agility. Any belief to the contrary can only arise 
from the fact that the importance to the economy of the organism of any given structural 
modification by no means necessarily corresponds with its conspicuousness in the eyes 
of the human observer. 

It is, of course, also the case, though this is beside the present point, that alterations 
in behaviour can only take place within limits set by the structure of the organism, 
though it is not the case that every such change must involve a modification in the 
structure of the organs used. Finally, it is still true that the organism must come to 
terms with its world; but that is not to say that it is dependent upon an inevitable 
environment, or that it can maintain itself in a given set of surroundings only by a fixed 
behaviour. Adaptation, in short, is a relation between structure and habit, and only 
secondarily between structure and environment; and there is no feature of an organism 
the investigation of which can safely be regarded by the naturalist as without signi- 
ficance, and relegated by him to the student of variation. 


The Distribution of the Pontoniine. 


The majority of the Pontoniinz are members of the shallow water fauna of the 
tropical and subtropical Indopacific. Six species occur in the Mediterranean, six on the 
west coast of America, eight in the Western Atlantic, and one in South Australia. 
Three of the Mediterranean species are also recorded from British waters. Urocaris 
longicaudata and Palemonella tenwipes are stated to occur both in the West Indies and 
in the Indopacific. Pontoniine appear to be entirely absent from the colder seas. 
Most are undoubtedly benthic, but very possibly a few of the more lightly-built forms, 
such as Urocaris, will prove to belong to the plankton. One species of Urocaris is 
bathypelagic, Coutierea lives in moderately deep water, probably on the bottom, Palae- 
monella laccadivensis is dredged in 100—600 fathoms, and Periclimenes tenellus in 

SECOND SERIES—ZOOLOGY, VOL. XVII. 44 


346 PERCY SLADEN TRUST EXPEDITION 


230 fathoms. ~ Most of the subfamily are to some extent cryptozoic, and a great many 
actually commensal. A sketch of the distribution of this habit within the group has 
already been given on pp. 329, 330. Unfortunately, little is known of the behaviour of 
the Pontoniinz or of their relation to the animals which serve as their hosts. .A detailed 
study of this subject, which could only be made in the field, would undoubtedly give 
very interesting results. 


The Genera of the Pontoniine. 


The genera of the Pontoniine may be briefly characterized by means of a key, as 
follows : 

I. Body very slender and compressed. Thorax without swelling. 6th abdominal 
segment much elongate. Outer flagellum of antennule deeply cleft. A gill may be 
present,on the second maxilliped. 

A. Mandibular palp present. Gull on second maxilliped. Rostrum toothed 
below. 
Urocaridella Borradaile, 1915. 
B. No mandibular palp. No gill on second maxilliped. Rostrum without teeth 
below. 
Urocaris Stimpson, 1860. 

II. Body moderately stout, not compressed. Thorax of female swollen dorsally. 
6th abdominal segment short. Outer flagellum of antennule deeply cleft. A gill may 
be present on the second maxilliped. [No mandibular palp. | 

Ancyclocaris Schenkel, 1902. 


III. Body never very slender, or much compressed. Thorax without swelling. 
6th abdominal segment never much elongate. Outer flagellum of antennule rarely deeply 
cleft. No gill on second maxilliped. 


A. Mandibular palp present. 
Palemonella Dana, 1852. 


B. No mandibular palp. 


1. Scale of second antenna rudimentary. 
Typton Costa, 1844. 


2. Scale of second antenna well developed. 

i. Gracefully-built forms, with body little if at all depressed, rostrum well 
developed, and almost always toothed, slender legs, whose dactylo- 
podites are usually narrow and never hook-like, and antepenultimate 
joint of third maxilliped narrow (except P. brocki, where it is of 
moderate breadth). 

a. Eyes subspherical. Rostrum horizontally expanded. 
Pontonopsis Borradaile, 1915. 


BORRADAILE—ON THE PONTONIINA 347 


8. Eyes not subspherical. Rostrum vertically expanded. 
(1) Body and chelipeds Palemon-like. Dactylopodites of walking legs 


usually narrow. 
Periclimenes Costa, 1844. 


(2) Cephalothorax deep and flat-sided, recalling Alpheus. One of 
second chele very heavy, with short fingers bearing a knob and 
socket arrangement. Dactylopodites short, broad, and biungui- 
culate. 

Periclimeneus Borradaile, 1915. 


u. Clumsy-bodied forms, with body obviously depressed, at least in female, 
rostrum usually in some way reduced and often toothless, stout legs, 
ending in short, usually hooked dactylopodites, and antepenultimate 
jomt of third maxilliped moderately or very broad. 


a. Rostrum straight or upcurved, diminishing in normal fashion to a 
sharp point, usually toothed, but generally with little or no ventral 
crest. Body not very clumsy. Third maxilliped not strongly 
twisted. 


(1) Dactylopodites of walking legs without basal protuberance. Last 
two joints of third maxilliped narrow and usually contrast 
strongly in width with antepenultimate. 


(a) Third maxilliped with arthrobranch. Second maxilliped with 
last joint mediad of penultimate. 
Harpiliopsis n. gen. 
(b) Third maxilliped without arthrobranch. Second maxilliped with 


last joint posterior (distal) to penultimate. 
Harpilius Dana, 1852. 


(2) Dactylopodites of walking legs bear a basal protuberance, usually 
large. Last two joints of third maxilliped do not contrast 
strongly in width with antepenultimate, and are usually broad. 


(a) Rostrum not very broad. Basal protuberances of the dactylo- 
podites simple. 


(i) Rostrum very long. Enormous supraocular spines present. 
Abdomen carinate. 
Coutierea Nobili, 1902. 


(ii) Rostrum not very long. Supraocular spines absent or not 
very large. Abdomen not carinate. 
Coralliocaris Stimpson, 1860. 


(b) Rostrum very broad. Basal protuberances of dactyles paired. 
Stegopontonia Nobili, 1906. 
449 


348 PERCY SLADEN TRUST EXPEDITION 


8. Rostrum downcurved, toothless, usually blunt pointed, sometimes with 
a sharp point of abnormal fashion, usually with good ventral crest. 
Body usually very clumsy. Third maxilliped strongly twisted. 


(1) Last two joints of third maxilliped narrow. 
Anchistus Borradaile, 1898. 


(2) Last two joints of third maxilliped broad. 


(a) No exopodites on maxillipeds. Eyes of good size. 
Pontonides n. gen. 


(6) Exopodites on maxillipeds. Eyes small. 


(i) Dactylopodites of last three legs straight, without basal 
prominence. 
Pontonia Latreille, 1829. 


(ii) Dactylopodites of last three legs curved, with basal prominence. 
Conchodytes Peters, 1851. 


There can be no doubt that the most primitive genus of Pontoniine is Urocaridella. 
This very remarkable form has almost all the features which must have characterized 
the earliest members of the group. The complete retention of the caridoid facies, the 
forward direction and simple form of the legs, the gill formula only less by one small 
pleurobranch than that of Leander or Desmocaris, the mandibular palp, the straight, 
slender third maxilliped, the deeply-cleft outer flagellum of the antennule, all tell the 
same tale. Yet Urocaridella can certainly not be considered ancestral to the rest of the 
subfamily. It is specialized, probably for pelagic life, in its compression, in the peculiar 
form of the abdomen with the long sixth segment and projecting third segment, and 
in the long, upcurved rostrum. There can, again, be no doubt that Urocaris arose from 
Urocaridella, through such species as Urocaris psamathe, by the loss of the mandibular 
palp, of the gill on the second maxilliped, and of the lower teeth of the rostrum. The 
rest of the Pontoniinz, however, must have taken independent origin from the common 
ancestor of the subfamily, to which Urocaridella remains nearer than they, though it is 
not transitional to them. Ancyclocaris represents such a line of independent descent. 
By the retention, in one species, of the gill on the second maxilliped, by the narrow third 
maxilliped, and by the deeply-cleft outer flagellum, it is linked directly with the 
ancestral form, for the lanceolate rostrum and some features of the abdomen make 
a descent through Urocaridella impossible, but it has lost the mandibular palp, and its 
stout body and legs, with the curved dactylopodites, show an interesting parallel to 
Harpiliopsis and allied genera. Probably, however, this is due only to a somewhat 
similar mode of life. The hump on the back of the female is a peculiarity which does 
not recur in the subfamily. 

The remaining genera probably represent a third line of descent, through a form 
resembling Paleamonella and Periclimenes, from which Periclimenes has departed in 
the loss of the mandibular palp and Palemonella in the loss of the vestige of the gill 


BORRADAILE—-ON THE PONTONIIN A 349 


of the second maxilliped and the greater reduction of that of the third maxilliped. In 
other respects there is no difference between these closely related genera. While, 
however, Palemonella does not yet appear to have given rise to any daughter genera, 
Periclimenes represents a central stock from which groups of genera diverge in several 
directions. Periclimeneus is an interesting modification of this stock in which some 
of the features of Alpheus are reproduced. Pontoniopsis represents a second departure 
from the same stock. Anchistus, Pontonia and Conchodytes form a third group, charac- 
terized by degeneration in connection with the habit of living in the mantle cavity 
of bivalves and ascidians. The species described by Pesta as Marygrande mirabilis, 
which is however an Anchistus, shows the earliest stage in this degeneration. Its body 
is still fairly slender and compressed, but the downcurved, toothless rostrum, the stout 
legs, the great chela, and the broad joint of the third maxilliped betray the tendency 
which is increased in other species of Anchistus and leads eventually through Pontonia 
to Conchodytes, where it culminates in C! meleagrine. Some of the species of Anchistus 
retain, in the form of the long joint of the third maxilliped and in a minute dentation 
of the rostrum, primitive features lost by A. mirabilis. It will be recalled that Anchistus 
has a vestige of the gill of the third maxilliped which is lost in Pontonta and Concho- 
dytes, and that a peculiar breadth and hairiness of the laciniz of the maxillule are also 
less marked in the former than in the latter two of these genera. Possibly the point 
of origin of this group was somewhere in the neighbourhood of the coral-haunting 
Periclimenes aurantiacus, which has a toothless rostrum. Pontonides seems to have 
arisen near this point. A fourth group of forms probably derived from Periclumenes 
is presented by Harpiliopsis, Harpilius, Coralliocaris, Coutierea, and Stegopontonia. 
These are sluggish, generally coral-haunting forms, but not internal commensals. In 
them the legs are stout, with hooked dactylopodites, but the body, though more 
depressed than in the Pontonia group, is less swollen and degenerate. The rostrum 
loses in depth, but is straight and nearly always dentate, and the maxillules are of 
quite a different type. In Harpiliopsis and Coralliocaris the gill of the third maxilliped 
is comparatively well developed, but in Harpilius it is lost, according to M. Sollaud. 
The members of this group diverge more than those of the second group, but it is not 
necessary here to recapitulate their differences, which are given in the key above. 
Harpiliopsis is the most primitive of them, and it is quite impossible to reconcile 
the evidence either of its bodily habitus or of its maxillule with any theory of a descent 
common to it and Anchistus. Possibly it may have taken origin somewhere in the 
neighbourhood of Ancyclocaris, but it seems more likely to have arisen from Periclumenes. 
Harpilius, though outwardly it presents no very remarkable feature, is, in its loss 
of a gill and in the peculiar form of its second maxilliped, perhaps the most aberrant 
of all. Coralliocaris shows in its third maxilliped, its second cheliped, and the dactylo- 
podites of its walking legs, certain convergences with Conchodytes. 

The affinities of Typton are doubtful. Its supraorbital spines* and narrow third 

* It is possible that these spines are not present in 7’. bowvierr. Nobili states in his preliminary 


description of the species that it has ‘“‘ocular” spines, and in a later account that antennal spines are 
present, but in his figure he shows neither. 


390 PERCY SLADEN TRUST EXPEDITION 


maxilliped are primitive features. On the other hand, the uncleft condition of the 
outer flagellum of its antennule is probably an exaggeration of one of the secondary 
features of the Pontoniine. Its vestigial antennal scales and small rostrum are no 
doubt adaptations to its habitat, and in any case are without parallel im the subfamily. 
Its reduced gill formula might be expected from its habits, and need not indicate any 
affinity with Conchodytes. Its heavy, compressed body with slender walking legs ending 
in biunguiculate dactylopodites, its large chela, and, very distantly, its rostrum, rather 
suggest Periclimeneus, but in view of other differences this is probably a convergence. 
It is perhaps best regarded as independently evolved from the ancestor of Persclimenes. 


Pontonia Conchodytes Coutierea Stegopontonia 
NS : Coralliocaris 
Anchistus 
Harpilius : 
Pontonides 
Harpiliopsis 


Periclimenes 


Periclimenzeus 


Pontoniopsis 
Typton 


Palemonella 


. Urocaris 
Ancyclocaris ? 


Urocaridella 


On some difficult questions in Phylogeny. 


The making of phylogenetic trees, once a favourite pastime of zoologists, has of late 
years come under the censure of superior persons, on the ground that we know too little 
about the mode in which evolution takes place to speculate successfully as to its course. 
Yet to represent in a graphic manner the lines of resemblance and difference within 
zoological groups is a practice which dates from before the acceptance of the evolutionary 
theory, and is not without its convenience even in these days. Moreover, the objections 
to diagrammatic phylogeny apply also to any form of classification which is not purely 
empirical, and especially strongly to the attempt so to construct “keys” that they 
shall as far as possible represent real affinities. One of the difficulties which occur 
in such cases is prominent in the Pontoniine. It is presented by numerous instances 
in which classifications according to two or more different features cross one another, 
so that alternative arrangements are suggested. When one of these arrangements has 
the advantage of illustrating also the verdicts of several further characteristics the 
solution is easy, but this is often not the case. I have already described an instance 


BORRADAILE—ON THE PONTONIINA 351 


of this in another group of Crustacea* and have remarked on the appearance it presents 
of the kaleidoscopic distribution in several ways of a set of pairs of characters, and called 
attention to its suggestion of Mendelism. A simple case of this sort occurs among 
the members of the subgenus Faleager, where it is hard to decide whether the primary 
grouping of the species should be on the ground of the presence and absence of the 
supraorbital spine or on the shape of the rostrum. Many other such cases may be 
detected in the keys which are given later in this paper. Another form of the same 
difficulty arises in the sporadic distribution of a character in circumstances which make 
it impossible to trace direct connection by descent between the species in which it occurs, 
except by means of other species from which it must have been absent. ‘This is the case, 
for instance, with the supraorbital spine throughout the subfamily, and with the denti- 
culation of the fingers of the first leg in several Periclimenes. 

In earlier days these phenomena would have been even harder of explanation than 
they are at present, when convergence is receiving more attention than formerly, and 
Mendelism has risen over the zoological horizon. In many cases close examination 
of features which at first appear to be due to repetition reveals, either between the fully 
evolved organs or between stages that lead up to them, differences which show that we 
are dealing with convergence. In others, the suggestion of Mendelism is probably correct, 
the feature in question having been suppressed over a series of generations by the absence 
of some factor necessary for its development or the presence of some hostile factor. 
It may, indeed, be doubted whether much of what is known as convergence in organisms 
be not rather due in this way to Mendelism. A further question, however, arises here. 
Is the sporadic repetition of such a feature, for instance, as the supraorbital spine, 
in apparently identical reincarnation, due to the releasing of the same “factor,” which, 
once lost, cannot be regained, or is it brought about by a less organized tendency of 
the mechanism of development to fall into an identical condition from time to time ? 
Far though we be from answering them, these questions are not without interest for 
others than the systematist. But it is doubtful whether they would be asked if 
phylogenetic speculation were regarded as wholly a waste of time. 

Another problem which is continually suggesting itself concerns the mode in which 
special features arise and disappear. This has been generally assumed to take place 
by the gradual elaboration of a rudiment into the highly developed organ, and its loss, 
when it has outlasted its usefulness, by a series of vestigial stages. Now in some cases 
it is certainly possible to trace a set of incomplete forms of an organ, and it is justifiable 
to regard these as either rudimentary or vestigial, though it is often hard or impossible 
to choose one or other of those alternatives. As instances of such phenomena within 
species there may be quoted the various forms of armature of the finger of the great 
chela in Harpiliopsis depressus, where stages in the perfection of a cutting flange 
may be seen, and the varying degrees of inequality of the legs of the second pair in 
Coralliocaris graminea. It is easy to imagine that in either of these cases one form 
of the organ might come to persist to the exclusion of the others. Within the limits 
of a genus, the same thing may be seen in regard to the remarkable protuberance 

* On the Genera of the Dromiide, Ann. Mag. Nat. Hist. (7), xi. p. 303 (1903). 


392 PERCY SLADEN TRUST EXPEDITION 


on the dactylopodites of the walking legs of Coralliocaris, of which rudimentary or 
vestigial stages are shown by the subgenus Onycocaris, and in the horns of the eyes 
of Corniger, while the subfamily as a whole presents, from genus to genus, a very 
pretty example of the loss of an organ by gradual stages in the reduction of the 
arthrobranch of the third maxilliped, from the well-formed gill of Urocaridella to the 
evanescent lobe of Pontonia. But there are other cases in which an organ is always 
present in the same degree of perfection. This is perhaps not very remarkable, for 
instance, in the peculiar armature of the fingers of the chelez of certain Periclimenes, 
such as the pits of P. petitthowarsi and P. spinifera or the combs of the same and 
certain other species, where imperfect stages may well have disappeared; but it is 
most striking in the many cases of well-formed and characteristic spines on the limbs 
and body which come and go throughout the subfamily without ever a trace of 
rudimentary or vestigial stages. Of such facts as these any theory of evolution must 
take account. , 

Finally it is necessary to state that even less conception can be formed of the 
nature of a ‘‘species” in the Pontoniinze than in many other groups of animals. We 
have not the slightest knowledge as to what degree of fertility exists between the 
assemblages of individuals to which this name is given, and have not even the assurance 
that some of them are not merely allelomorphs. The attempt has been made in some 
instances to recognize certain entities as “ varieties,” but as none of these are connected 
by intermediate individuals I have felt bound, in accordance with principles which 
I have stated elsewhere*, to class them all in the same temporary category as the 
“species.” 


The Species of Pontoniine. 


In the following lists the species of the subfamily at present known are arranged 
under their genera with a key to the species of each genus. 


Genus URocARIDELLA Borradaile, 1915. 


Ann. Mag. Nat, Hist. (8), xv. p. 207. 

Definition: Body slender, much compressed ; sixth abdominal segment elongate ; 
rostrum long, slender, upcurved, with teeth above and below; outer flagellum of ~ 
antennule deeply cleft; antennal scale long, narrow; mandible with palp; second 
maxilliped with podobranch, and last joint mediad of preceding joint; third maxilliped 
slender, with arthrobranch; legs very slender, directed forwards, with simple, slender, 
nearly straight dactylopodites; first abdominal limb with appendix interna. 


Type: Urocaridella gracilis Borradaile, 1915 (Plate 58, fig. 2). 
Ann. Mag. Nat, Hist. (8), xv. p. 210. 


Definition: Rostrum nearly twice as long as carapace, much upcurved, excavate 
at base, its formula §=4°, one of the dorsal teeth standing in the middle of the 


10-12? 


* Gardiner’s Fauna of the Maldives, vol. i. 


BORRADAILE—ON THE PONTONIINA 393 


carapace, two, large, standing on a crest over the orbit, two to four, small, near 
antennule, three isolated near the tip, most of ventral teeth large and set on a crest 
between the antennules; antennal and hepatic spines present, the latter nearly in 
the branchiostegal position; antennular stalk three-quarters of length of antennal 
scale, its basal joint not much expanded, with stylocerite and distal spine small, the 
second joint shorter than the’ third, the second and third joints together shorter than 
the first ; basipodite of antenna with very small spine, scale not half length of rostrum, 
shghtly outcurved, its sides converging slightly towards the end, which is subtruncate, 
its subterminal spine not reaching the end, stalk of antenna more than half the length 
of first joint of antennule ; third maxilliped reaching or outreaching end of second joint 
of antennule; first lee outreaching antennal scale by fingers, its wrist and palm 
subequal, fingers a little longer, its arm equal to wrist and palm together; second legs 
equal and similar, outreaching antennal scale by hand and part of wrist, fingers rather 
longer than palm, wrist more than half length of palm, arm equal to wrist and palm 
together, all joints unarmed ; walking legs long, slender, with spines under the propodite, 
the longest at the end, dactylopodites long, nearly straight, simple; telson longer than 
sixth abdominal segment, shorter than uropods. 

Length of longest specimen, 40 mm. 

Maldive Is. 


Genus Urocaris Stimpson, 1860. 


Proc. Ac. Philadelphia, 1860, p. 39. Kingsley, Ib., 1879, p. 383 (1880). Rathbun, 
Bull. U.S. Fish Comm. xx. 1m. p. 126 (1901). 


Definition: Body slender, much compressed ; sixth abdominal segment elongate ; 
rostrum with a toothed crest above but toothless below; outer flagellum of antennule 
deeply cleft; antennal scale long, narrow; mandible without palp; second maxilliped 
without gill; third maxilliped slender, with arthrobranch; legs very slender, their 
dactylopodites little curved. | 


Key to the species of Urocaris : 
I. Rostrum very long and slender and much upcurved. 
U. psamathe de Man, 1902. 
II. Rostrum of moderate length, never very slender, hardly if at all upcurved. 
A. Rostrum has high, arched dorsal crest. 
U. wnfraspins Rathbun, 1902. 
B. Rostrum has low dorsal crest. 


1. Rostrum arched, with concavity below. Third abdominal segment projects 


behind. | 
a. Two teeth of dorsal crest stand behind orbit. Walking legs biunguiculate. 
U. longicaudata Stimpson, 1860. 
SECOND SERIES—ZOOLOGY, VOL. XVII. 45 


354 PERCY SLADEN TRUST EXPEDITION 


b. Five teeth of dorsal crest stand behind orbit. Walking legs have simple 
dactylopodites. 
U. esopius (Bate), 1864. 


2. Rostrum straight. Third abdominal segment does not project behind. 


a. Rostral formula ~. Second leg of great length. 
U. longipes Stimpson, 1860. 


b. Rostral formula 19541. Second leg rather short. 
U. korm (Lo Bianco), 1903. 


1. (Type.) Urocaris longicaudata Stimpson, 1860. 


Proc. Ac. Philadelphia, 1860, p. 39. Kingsley, Bull. Essex Inst. x. p. 65; Proc. 
Ac. Philadelphia, 1878, p. 330. Rathbun, Bull. U.S. Fish Comm. xx. 1. p. 126 (1901). 
? Pearson, Rep. Ceylon Pearl Fisheries, iv. p. 78, Pl. 1, fig. 5 (1905). 

Specimens in the present collection of a Urocaris resembling this species do not agree 
with the descriptions given by the authorities cited, in the following pots: (1) the 
rostrum outreaches the second joint of the antennular peduncle, (2) the flagella of the 
antennule are much longer than its stalk, (3) the first leg outreaches the antennal scale 
by the fingers, and the fingers and palm are subequal, (4) the fingers of the second 
leg are rather shorter than the palm. Further investigation of the reported occurrence 
of U. longicaudata in the Indian Ocean is desirable. 

Carolina to Brazil. Western Indian Ocean ? 


2. Urocaris longipes Stimpson, 1860. 


Proc. Ac. Philadelphia, 1860, p. 39. 
Ousima I., 20 fms. 


3. Urocaris esopius (Bate), 1864. 

Anchistia wsopia Bate, Proc. Zool. Soc. Lond., 1863, p. 502, Pl. 41, fig. 5 (1864). 
St Vincent Gulf. 

4. Urocaris infraspims Rathbun, 1902. 


Proc. U.S. Mus. xxiv. p. 903; Decap. Crust. N.W. America, p. 31 (1904). 
California to Mexico. 


5. Urocaris psamathe de Man, 1902 (Plate 58, fig. 3). 
Abh. Senckenb. Ges. xxv. p. 816, Pl. 25, fig. 51. 


Ternate. 
6. Urocaris korni (Lo Bianco), 1903. 
Anchistia Korniu, Lo Bianco, Mitt. Stat. Neapel, xvi. p. 250, Pl. 7, fig. 13 (1903), 


Periclimenes Korni? Kemp, Journ. Mar. Biol. Assoc. (N.S.) viii. p. 411 (1910). 
Mediterranean. Bay of Biscay? Bathypelagic, 400—600 fms. 


BORRADAILE—ON THE PONTONIINA 3595 


Genus ANncycLocaRIs Schenkel, 1902. 


Verh. Naturf. Basel, xii. p. 503. Nobili, Bull. Sci. Fr. Belg. xl. p. 52 (1906). 


Defimtion: Body rather stout ; cephalothorax with dorsal swelling in female ; sixth 
_ abdominal segment short; rostrum deep, of moderate length, lanceolate, with about 
half-a-dozen teeth above and one or two below; eyes small; outer flagellum of antennule 
deeply cleft; antennal scale broad; mandible without palp; second maxilliped with 
or without podobranch*, and with last jot mediad of preceding joint; third maxilliped 
slender, with arthrobranch ; legs stout, with short, simple, curved dactylopodites. 

The species of this genus are closely related, and the distinctions between them 
are hard to gather from the published descriptions. Possibly some of them will 
eventually have to be united. So far as can be learned from the diagnoses and figures 
given by their authors, they differ in the points stated in the following key: 


Key to the species of Ancyclocaris : 

I. Dorsal swelling of cephalothorax steep. Uncleft region of outer antennular 
flagellum 8-jointed. Hepatic spine farther forward than usual. [Outer edge of antennal 
scale converging towards inner. R.=%*. Fingers of second leg gaping a little. |] 

A. aberrans (Nobili), 1904. 


II. Dorsal swelling of cephalothorax gradual. Uncleft region of outer antennular 
flagellum 5—7-jointed. Hepatic spine not farther forward than usual. 
A. All rostral teeth in front of orbit. Fingers of second leg gaping a little. 
[R.=$. Outer edge of antennal scale straight. | 
A. latirostris (Lenz), 1905. 


B, One tooth of rostrum behind orbit. Fingers of second leg not gaping 
appreciably. 

1. R.=%. Cephalothorax with large swelling in middle of its length. Outer 
edge of antennal scale nearly straight t. Uncleft region of outer anten- 
nular flagellum 7-jointed. 

A. hermitensis (Rathbun), 1914. 


2. R.=8. Cephalothorax with low swelling in hinder region, Outer edge 
of antennal scale markedly convex. Uncleft region of outer antennular 
flagellum 5—6-jointed. 


A. brevicarpalis Schenkel, 1902. 


1. (Type.) Ancyclocaris brevicarpalis Schenkel, 1902. 
Verh. Naturf. Basel, xiii. p. 563, Pl. 13, fig, 21. 


Macassar. 


* Nobili (Bull. Sci. Fr. Belg. xl. p. 52) very definitely states that A. aberrans has a gill on the second 
maxilliped. I have not been able to find this in A. hermitensis. 
+ See below, p. 356. 
45—2 


356 PERCY SLADEN TRUST EXPEDITION 


2. Ancyclocaris aberrans (Nobili), 1904. 

Palemonella aberrans, Nobili, Bull. Mus. Paris, 1904, v. p. 233. 

Ancyclocaris aberrans, Nobili, Bull. Sci. Fr. Belg. xl. p. 52, Pl. 4, fig. 9 (1906) ; 
Ann. Sci. Nat. (9), iv. p. 64 (1906). 

Jibuti, under protection of Discosoma giganteum. Persian Gulf. 


3. Ancyclocaris latirostris (Lenz), 1905. 

Harpilius latirostris, Lenz, Abh. Senckenb. Ges. xxv. p. 380, Pl. 47, fig. 14 
(1905). 

Ancyclocaris (?) latirostris, Nobili, Ann. Sci. Nat. (9), iv. p. 65 (1906). 

? Eine nicht bestimmte Palemonide, Richters, Decap. Mauritius, Pl. 18, figs. 10, 
11 (1880). 

E. Africa. Mauritius ? 


4. Ancyclocaris hermitensis (Rathbun), 1914. 
Periclimenes hermitensis, Rathbun, Proc. Zool. Soc. Lond., 1914, p. 655, Pl. 1, 
figs. 1—8. 


Monte Bello Is. Torres Straits, under protection of Duscosoma, whose mouth it 
enters at times. : 


Miss Rathbun figures the antennal scale of this species without a distal spine. 
In specimens’ from Torres Straits which resemble hers in every other respect, including 
the colour pattern, the outer edge of the scale is straight from the base to a well-formed 
distal spine, and then curves abruptly inwards. 


Genus PALZMONELLA Dana, 1852. 


U.S. Explor. Exped. Rep. xiii. 1. p. 582. Kingsley, Proc. Ac. Philadelphia, 1879, 
p: 425 (1880). Bate, “Challenger” Macrura, p. 786 (1888). Ortmann, in Spengel, Zool. 
Jahrb. Syst. v. p. 513 (1890); Bronn’s Thierreich, v. 1. p. 1182 (1899). 

Definition: Body slender; sixth abdominal segment of moderate length ; rostrum 
straight or a little upcurved, shallow, with several teeth above but few below; outer 
flagellum of antennule deeply cleft or not; antennal scale of good length and moderate 
breadth; mandible with palp; second maxilliped without even a vestige of podobranch, 
and with last joint mediad of preceding joimt; third maxilliped narrow, with vestigial 


arthrobranch ; legs slender, with slender, slightly curved dactylopodites, biunguiculate 
or not in last three pairs. 


Key to the species of Palemonella : 
I. Second leg unarmed: its wrist less than half the length of its hand. 
A. No hepatic spine. 


1. Wrist of first leg longer than hand. Dactylopodites of last three legs 
simple. (R.=§.) 
P. orientalis Dana, 1852. 


BORRADAILE—ON THE PONTONIIN A 357 


2. Wrist of first leg shorter than hand. Dactylopodites of last three legs 
bear a spine in the middle of the lower edge. (R.=$.) 
P. batet n. nom. 


B. <A hepatic spine present. 
1. Last three legs biunguiculate. Second wrist more than + length of hand. 


a. Second legs unequal. R.=8. 
P. biunguiculata Nobili, 1904. 


b. Second legs subequal. R.=*%*. 
P. rathbunensis n. nom. 


2. Last three legs not biunguiculate. Second wrist less than 4+ length of 


hand. 
a. Rostrum deep, lanceolate. Palm of second leg only as long as fingers. 
(R= 59.) 


P. amboinensis Zehnter, 1894. 


b. Rostrum shallow, not lanceolate, slightly upecurved. Palm of second 
leg twice as long as fingers. 
i, R.=%, lower tooth removed from tip. 
P. affins Zehnter, 1894. 


i. R.=8 8, lower teeth near tip. 


P. laccadivensis Alcock and Anderson, 1894. 


II. Second leg bears a spine at least on wrist, which is more than half length 
of hand. . 
A. Arm of second leg bears a spine at end. Indopacific. 
1. Rostrum #, much shorter than antennal stalk. 
P. elegans Borradaile, 1915. 


2. Rostrum $=, longer than antennal stalk. 

a. Spine at end of arm of second leg subterminal. Fingers of same leg 
toothed. Teeth on lower side of rostrum well spaced. 

i. Rostrum outreaches antennular stalk by nearly half its own length, 

decidedly upeurved. Wrist of first leg half as long again as hand. 


P. longirostris Borradaile, 1915. 


ii. Rostrum but little outreaches antennular stalk, hardly upcurved. 
Wrist of first leg less than half as long again as hand. 
P. tridentata Borradaile, 1899. 


b. Spine at end of arm of second leg terminal. Fingers of same leg not 
toothed. Teeth on lower side of rostrum stand close together near tip. 
P. tenuipes Dana, 1852. 


B. Arm of second leg bears no spine. West Indian. (R.=4.) 
P. yucatanica Ives, 1891. 


358 PERCY SLADEN TRUST EXPEDITION 


1. (Type.) Palemonella tenuipes Dana, 1852. 

U.S. Explor. Exped. Rep. xii. 1. p. 582; Atlas, Pl. 38, fig. 3(1855). Stimpson, Proc. 
Ac. Philadelphia, 1860, p. 40. de Man, Arch. Naturg. liii. 1 p. 551, Pl. 22a, fig. 4 (1887). 
Heilprin, Proc. Ac. Philadelphia, 1888, p. 322 (?). Ortmann, Speng. Zool. Jahrb. Syst. v. 
p- 527 (1890). Zehnter, Rev. Suisse Zool. ii. p. 208 (1894). Nobili, Ann. Mus. Napoli, 
i. p. 6 (1901). Rathbun, Bull. U.S. Fish Comm. xxiii. p. 925 (1906), 

? Palemonella tenuipes var., Nobili, Ann, Sci. Nat. (9), iv. p. 70 (1906). 

Indopacific. Bermuda ? 

2. Palemonella orientalis Dana, 1852. 

U.S. Explor. Exped. Rep. xiii. 1 p. 583; Atlas, Pl. 38, fig. 4 (1855). de Man, 
Arch. Naturg. li. 1. p. 552 (1887). 

E. Indies, on comatulids. 

3. Palemonella yucatanica Ives, 1891. 

Proc. Ac. Philadelphia, 1891, p. 183, Pl. 5, fig. 8. 

Yucatan. 

4, Palemonella amboinensis Zehnter, 1894. 

Rev. Suisse Zool. ui. p. 200, Pl. 9, fig. 27. de Man, Abh. Senckenb. Ges. xxv. 
p. 811 (1902). 

E. Indies. 

5. Palemonella affinis Zebnter, 1894. 

Rev. Suisse Zool. u. p. 208. 

Amboina, on Actinometra. 

6. Palemonella laccadivensis Aleock and Anderson, 1894. 

Journ. As. Soc. Bengal, lxiii. p. 157 (1894). Tllust. Zool, “ Investigator,” Crust. iv. 
Pl. 26, fig. 4 (1896). Ann. Mag. Nat. Hist. (7), iii, p. 4 (1889). Rathbun, Bull. U.S. 
Fish Comm. xxui. p. 925, Pl. 22, fio. 2 (1906). 

Laccadive Is. Hawaiian Is. 100—600 fms. 

7. Palemonella tridentata Borradaile, 1899. 

Proce. Zool. Soc. Lond., 1898, p. 1007, Pl. 64, fig. 8. 

Funafuti. Western Indian Ocean. 

8. Palemonella biunguiculata Nobili, 1904. 

Bull. Mus. Paris, 1904, v. p. 233. Ann. Sci. Nat. (9), iv. p. 71, Pl. 3, fig. 6 (1906). 

Jibuti. 

9. Palemonella bate: n, nom. 

Palemonella orientalis, Bate, “ Challenger” Macrura, p. 787, Pl. 128, fig. 4 (1888). 

Philippine Is. 

10. Palemonella rathbunensis n. nom. 

Palemonella orientalis, Rathbun, Bull. U.S. Fish Comm. xxii. p. 925 (1906). 

Hawaiian Is. 


oS 


ee — eee 


BORRADAILE—ON THE PONTONIIN A 359 


11. Palemonella elegans Borradaile, 1915 (Plate 58, fig. 4). 
Ann. Mag. Nat. Hist. (8), xv. p. 210. 


Definition: Closely related to P. tridentata but with rostrum quite different, 
lanceolate, not reaching end of first jot of antennule, quite straight, of the formula 3, 
two teeth standing behind the orbit. 

Length of the single specimen, 17 mm. 

Salomon. 

It is possible that this is an abnormal specimen of P. tridentata. 


12. Palemonella longirostris Borradaile, 1915 (Plate 58, fig. 5). 
Ann. Mag. Nat. Hist. (8), xv. p. 210. 


Definition: Related to P. tridentata but differs in that: (1) the rostrum is much 
longer, outreaching the antennular stalk by not much less than half its own length, very 
decidedly upcurved, and of the formula 8, owing to the addition of a small tooth near 
the tip; (2) the third maxilliped is rather shorter, only reaching the end of the first 
joint of the antennule; (3) the wrist of the first leg is half as long again as the hand ; 
(4) the second leg is longer and more slender, and has the arm of even width, not wider 
in the middle as in P. tridentata, the spine at the end of the arm smaller, and smaller 
teeth on the fingers. 

Length of longest specimen, 17 mm. 

Fadiffolu Atoll, Maldive Is. 


Palemonella gracilis Paulson, 1875 (Red Sea Crustacea, p. 117, Pl. 17, fig. 6) does 
not belong to this genus, and appears, from its telson, not to be a member of the 
Pontonine. 


Genus PERICLIMENES Costa, 1844. 


Pelias, Roux, Mém. s.1. Salicoques, p. 25 (1831). H. M.-Edwards, Hist. Nat. Crust. 
ii. p. 881 (1837). Heller, Sitz. k. Ak. Wiss. Wien, xlv. p. 406 (1862); Crust. siidl. 
Kur. p. 254 (1863). 

Periclimenes, Costa, Ann. Ac. Aspir. Nat. a Raa ii. (1844); Faun. Reg. Napoli, 11. 1. 
(1846). Borradaile, Ann. Mag. Nat. Hist. (7), 1. p. 380 (1898). Rathbun, Bull. U.S. 
Fish Comm. xx. I p. 121 (1901). 

Anchistia, Dana, U.S. Explor. Exped. Rep. xiii. 1. p. 577 (1852). Kingsley, Proc. Ac. 
Philadelphia, 1879, p. 423 (1880). Carus, Prodr. Faun. Medit. i. p. 474 (1885). Ortmann, 
Bronn’s Thierreich, v. 1. p. 1131 (1899). 

Dennisia, Norman, Ann. Mag. Nat. Hist. (3), viii. p. 278 (1861). 


Definition: Body slender, usually somewhat compressed ; sixth abdominal segment 
of moderate length; rostrum of varying shape; outer flagellum of antennule usually 
not deeply cleft; antennal scale of moderate breadth; mandible without palp; second 
maxilliped without podobranch, though often with a vestige of it, last joint mediad of 
preceding joint; third maxilliped slender, with small arthrobranch ; legs slender, except 


360 PERCY SLADEN TRUST EXPEDITION 


second pair which may be fairly heavy ; dactylopodites of last three pairs usually simple 
and slightly curved. 

Four distinct sections may be observed in Periclimenes. (1) P. aurantiaca (Dana) 
differs sharply from all the rest in the lack of teeth on its rostrum. Quite possibly 
further information will show that it should become the type of a new genus: for the 
present it must have subgeneric rank. The remaining species fall into three groups. 
(2) In one of these the rostrum is rather short and has a convex upper edge, while the 
second leg is unarmed and its wrist short. In this group the supraorbital spine is rare. 
(3) In another group, the dorsal edge of the rostrum is convex, and the second leg of the 
only species in which this limb is known is short-wristed and unarmed, but the cornea, 
which in all other Periclimenes is subhemispherical, is here ogival, and a strong supra- 
orbital spine is present. (4) The remainder of the species, including more than half the 
genus, form a group which is characterized by a rostrum with straight or concave upper 
edge and tip nearly always upturned. The second leg generally bears a spine on arm 
or wrist or both, and almost always has the wrist of a good length. The supraorbital 
spine is common in this group. The existence of certain intermediate species, such as 
P. commensalis Borradaile and P. amethysteus (Risso), makes it undesirable that these 
sections should rank as genera. I have therefore established for them four subgenera, 
named respectively Ensiger, Cristiger, Corniger, and Falciger*. The type species of the 
genus belongs to Cristiger. 


Key to the subgenera of Periclimenes : 


I. Rostrum toothless. No spines on trunk or legs. 
Ensiger Borradaile, 1915. 


II. Rostrum toothed. Spines at certain points on trunk or legs. 
A. Cornea ogival. [Upper edge of rostrum convex. Strong supraorbital spine. ] 
Cornger Borradaile, 1915. 
B. Cornea not ogival. 

1. Upper edge of rostrum convex. Second leg with short wrist, and unarmed, 
save that angles of wrist and arm are sharp in P. graclis. Supraorbital 
spine in one species only. 

Cristuger Borradaile, 1915. 

2. Upper edge of rostrum straight or concave. Second leg rarely with short 

wrist, generally with spine on wrist or arm or both. Supraorbital 


spine common. 
Falcger Borradaile, 1915. 


Subgenus CristIcER Borradaile, 1915. 
Ann. Mag. Nat. Hist. (8), xv. p. 207. 
Definition: Rostrum toothed, with convex upper edge, short or of medium length. 
Antennal and usually hepatic spines present. Supraorbital spine very rare. Second leg 


* Since these subgenera were established, in an article in the Annals and Magazine of Natural History 
for February, 1915, I have learned that the names of two of them (Corniger and Falciger) are preoccupied 
as designations of genera. They must, however, continue to stand as subgeneric names. 


BORRADAILE—ON THE PONTONIIN AD 361 


with short wrist, and unarmed, save that in P. gracilis the angles of the wrist and arm 
are sharp. 


Key to the species of Periclimenes (Cristiger) : 
I. Rostrum much shorter than antennular stalk. Second leg shorter than first. 
[R.=8, lanceolate, two teeth behind orbit. ] 
~P. (C.) brevinaris Nobili, 1905. 


Il Rostrum little if at all shorter than antennular stalk. Second leg longer 
than first. 


A. Three or four rostral teeth behind orbit. Three teeth below rostrum. 
Atlantic and Mediterranean. 


1. Rostrum deep, horizontal. Second leg outreaches antennal scale by hand 
only. European. [R.=§&,°.] 
P. (C.) scriptus (Risso), 1826. 


2. Rostrum shallow, bent downwards. Second leg outreaches antennal scale 


by wrist and hand. W. Atlantic. [R.=%.] 
P. (C.) tenellus (Smith), 1882. 


B. No rostral tooth behind orbit. Very rarely more than one tooth below 
rostrum*. Indopacific. ; 
1. Rostrum lanceolate. 
a. Dorsal edge of rostrum bears nine or more teeth. 


i. Antepenultimate joint of third maxilliped broad. R. = B=, Fingers 
of first leg not denticulate. 
P. (C.) brocki (de Man), 1887.. 


ii. Antepenultimate joint of third maxilliped narrow. R.=14>51%. 
Fingers of first leg denticulate. 


a. First jomt of antennule bears two distal spines. No accessory 
' spinule on dactylopodites of last three legs. 
P. (C.) frater Borradaile, 1915. 


B. First joint of antennule bears one distal spine. An accessory 
spinule on dactylopodites of last three legs. 
P. (C.) soror Nobili, 1904. 
b. Dorsal edge of rostrum bears seven or fewer teeth. 
i. Ventral edge of rostrum bears more than one tooth. 
a. A supraorbital spine. [R.=$. 
P. (C.) commensalis Borradaile, 1915. 


* Only P. notatus has three. 
SECOND SERIES—ZOOLOGY, VOL. XVII. 46 


362 ‘PERCY SLADEN TRUST EXPEDITION 


8B. No supraorbital spine. 


1) R.=2. Hepatic spine present. Antennal scale bears distal spine 
3 Pp pine p 19 
at end. 

P. (C.) notatus (Heller), 1865. 


(2) R.=§. Hepatic spine absent. Antennal scale bears distal 
spine at a little distance from end. 


P. (C.) pusillus Rathbun, 1903. 


ii. Ventral edge of rostrum bears one tooth. 


a. Rostrum 5;&; longer than antennular stalk; its tip not upturned. 
Angles of arm and wrist sharp. 


P. (C.) gracilis (Dana), 1852. 


8B. Rostrum #; shorter than antennular stalk; its tip a little up- 
turned. Angles of arm and wrist not sharp. 


P. (C.) potina Nobili, 1905. 


2. Rostrum not lanceolate, with decided reversal of curve at tip. 


a. Rostrum $; outreaching antennular stalk. Hepatic spine present. 
Hand of second leg elongate. 


i. A denticle on carapace behind rostral crest. Last joint of third 
maxilliped slightly shorter than penultimate. 
P. (C.) mcertus Borradaile, 1915. 


ii. No denticle on carapace behind rostral crest. Last joint of third 
maxilliped slightly longer than penultimate. 
P. (C.) parvus Borradaile, 1898. 


b. Rostrum %; outreached by antennular stalk. No hepatic spine. Hand 
of second leg short. 
P. (C.) parasiticus Borradaile, 1898. 


1. (Type.) Perschmenes (Cristiger) scriptus (Risso), 1826. 


Alpheus scriptus, Risso, Hist. Nat. Hur. Mér. v. p. 78 (1826); Acad. Leop., 1826, 
p. 821. ; 

Pelias scriptus, Roux, Mém. s. |. Salicoques, p. 25 (1831). Heller, Sitz. k. Ak. Wiss. 
Wien, Math.-Nat. Cl. xlv. p. 406, Pl. 2, fig. 34 (1862). 

Periclimenes insignis, Costa, Ann. Ac. Aspir. Nat. Napoli, ii. (1844); Faun. Regn. 
Napoli, ii. 1. Pl. 6, figs. 1—6 (1846). 

Dennsia sagittifera, Norman, Ann. Mag. Nat. Hist. (3), vil. p. 278, Pl. 13, figs. 8—13 
(1861). | 

Anchistia scripta, Heller, Crust. stidl. Eur. p. 256, Pl. 8, figs. 18, 19 (1863). Carus, 
Prodr. Faun. Medit. i. p. 476 (1885). 

Periclimenes scriptus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 381 (1898). 

Mediterranean, Guernsey. To 30 fms. 


BORRADAILE—ON THE PONTONIIN 363 


2. Periclimenes (Cristiger) gracilis (Dana), 1852. 

Anchistia gracilis, Dana, U.S. Explor. Exped. xii. 1. p. 578 (1852); Atlas, Pl. 37, 
fic. 5 (1855), 

Periclimenes gracilis, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 381 (1898). 

Sulu Sea. 

3. Periclimenes (Cristiger) notatus (Heller), 1865. 

Anchistia notata, Heller, “ Novara” Rep., Zool. 1. iii. p. 109, Pl. 10, fig. 3 (1865). 

Periclimenes notatus, Borradaile, Ann. Mag. Nat. Hist. (7), iii. p. 382 (1898). 

Nicobar Is. 


4. Periclimenes (Cristiger) tenellus (S. J. Smith), 1882. 

Anchistia tenella, 8. J. Smith, Bull. Mus, Harvard, x. p. 55, Pl. 9, fig. 1 (1882). 
Periclimenes tenellus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p: 383 (1898). 

E. coast of N. America, 229 fms. 


5.  Periclimenes (Cristiger) brocki (de Man), 1887 (Plates 58, 54, figs. 8/—). 
Anchistia Brockit, de Man, Arch. Naturg. li. 1. p. 548, Pl. 22a, fig. 3 (1887). 
Periclimenes Brocki, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 383 (1898). | 
Amboina. Maldive Is., to 43 fms., on sea urchin. 


6. Periclimenes (Cristiger) parvus Borradaile, 1898. 

Periclimenes parvus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 384 (1898); 
Willey’s Zool. Results, iv. p. 407, Pl. 1, fig. 3 (1899). 

New Britain. 


7. Periclimenes (Cristiger) parasiticus Borradaile, 1898. 

Periclimenes parasiticus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 384 (1898) ; 
Willey’s Zool. Results, iv. p. 407, Pl. 1, fig. 4 (1899). 

New Britain, on Linckia. 


8. Periclimenes (Cristiger) pusillus Rathbun, 1903. 
Periclimenes pusillus, Rathbun, Bull. U.S. Fish Comm. xxii. m1. p. 921, fig. 71, 
Pl. 24. fig. 7 (1903). 


Hawaiian Is. 


9. Periclimenes (Cristiger) soror Nobili, 1904. 

Periclimenes soror, Nobili, Bull. Mus. Paris, 1904, v. p. 231; Ann, Sci. Nat. (9), iv. 
p. 50, Pl. 2, fig. 6 (1906). 

Jibuti. 

10. Periclimenes potina Nobili, 1905. 

Periclimenes potina, Nobili, Bull. Mus. Paris, 1905, p, 159 ; Bull. Sci. Tr. Belg. xl. 
io, Aa JPL Bh, sore, 3 (USO) 


Persian Gulf, on floating brown seaweed. 
46—2 


364 PERCY SLADEN TRUST EXPEDITION 


11. Periclimenes (Cristiger) brevinaris Nobili, 1905. 
Periclimenes Borradailet, Nobili, Bull. Mus. Paris, 1905, ii. p. 159. 


Periclimenes brevinaris, Nobili, Bull. Sci. Tr. Belg. xl. p. 42, Pl. 3, fig. 7 (1906). 
Persian Gulf. 


12. Periclimenes (Cristiger) frater Borradaile, 1915 (Plate 53, fig. 6). 
Ann. Mag. Nat. Hist. (8), xv. p. 210. 


Definition: Closely related to P. soror, but differing from it in that (1) the teeth on 
the upper edge of the rostrum are closer set towards the tip than near the base, (2) there 
are two distal spines on the first joint of the antennule, (3) the antennal scale decidedly 
outreaches the first leg, (4) there is no accessory denticle on the dactylopodites of the 
last three legs. o, 

Length of the longer of the two specimens, 13 mm. 

Seychelles, on reef. 


13. Periclimenes (Cristiger) incertus Borradaile, 1915 (Plate 53, fig. 7). 
Ann. Mag. Nat. Hist. (8), xv. p. 210. 


Definition: Closely related to P. parvus, but differing from it in that (1) the body_ 
is more slender, (2) the rostrum is shallower, (3) there is a denticle on the carapace 
behind the beginning of the rostral crest, (4) the third maxilliped is rather longer, owing 
to the greater length of the penultimate joint, which is considerably longer than the 
end joint. , 

Length of the single specimen, 11 mm. 

Maldive Is. 


The specific distinctness of this form from P. parvus is somewhat doubtful. 


14. Periclhimenes (Cristiger) commensalis Borradaile, 1915. 

Ann.’ Mag. Nat. Hist. (8), xv. p. 211. Potts, Pap. Dep. Mar. Biol. Carnegie Inst. 
Washington, viii. p. 82 (1915). 

Definition: Body rather stout, not compressed ; rostrum reaching end of antennular 
stalk, outreached by antennal scale, lanceolate, directed slightly downwards, its 
formula $; supraorbital, hepatic, and antennal spines present; cornea subhemispherical ; 
first jomt of antennular stalk broad, with moderate stylocerite and two distal spines, 
second and third joints subequal, together shorter than first joint, flagella of a fair length, 
the outer rather deeply cleft ; antenna with small basal spine, stalk nearly - reaching 
end of first joint of antennule, scale wide, with straight outer and convex inner side, 
and distal spine of good size but not reaching end; third maxilliped slender, reaching 
end of antennal stalk, its long joint a good deal curved ; first leg reaching end of thicker 
division of outer antennular flagellum, arm and wrist equal, hand shorter, fingers a little 
longer than palm; second leg short, unarmed, its wrist subconical, hand nearly equal 
to three preceding joints together, fingers equal to palm, not gaping, each with three 
small teeth near base; last three legs short, hairy, with a spine above and one below 
end claw of short, rather stout dactylopodite; telson shorter than uropods. 

Torres Straits, on Comanthus annulatus. 


BORRADAILE—ON THE PONTONIIN/®A 365 


Subgenus CornicER Borradaile, 1915. 


Ann. Mag. Nat. Hist. (8), xv. p. 207. 


Definition: Rostrum toothed, but without teeth behind orbit; its upper edge 
convex, but sometimes only slightly so. Antennal, hepatic, and strong supraorbital spines 
present. Eye ogival. Second leg at present only known in one species, where it is 
unarmed and has short wrist. 


Key to the species of Periclimenes (Corniger) : 
I. Rostrum lanceolate, outreached by antennular stalk. 


A. Rostrum deep, +. Eye without papilla. 
P. (C.) cornutus Borradaile, 1915. 


B. Rostrum shallow, 4. Eye with papilla at apex. 
P. (C.) ceratophthalmus Borradaile, 1915. 


II. Rostrum with upper edge but slightly convex and lower edge straight till 
within short distance of tip, which is slightly upturned. Antennular stalk outreached 
by rostrum. [R.=£.] 

P. (C.) amboinensis (de Man), 1887. 


1. (Type.) Perichimenes (Corniger) ceratophthalmus Borradaile, 1915 (Plate 54, 
fig. 9). | 
Ann. Mag. Nat. Hist. (8), xv. p. 211. 


Definition: Body not much compressed ; rostrum outreached barely by antennular 
stalk but distinctly by antennal scale, straight, shallow, lanceolate, bearing above four 
teeth, all in its distal half, and no teeth below; eye ogival, ending in a papilla; supra- 
orbital, antennal and hepatic spines present; first joint of antennular stalk not greatly 
expanded, with moderate stylocerite, second and third joints stout and subequal, flagella 
short; antennal stalk about 2 length of first joint of antennule, scale broad, with small 
distal tooth set well back from end; third maxilliped short, rather stout; first leg 
outreaching antennal scale by its fingers, its arm and wrist subequal, hand about half 
length of wrist; second leg outreaching antennal scale by nearly the whole of its hand, 
stout, with short wrist and no spines on any joint, hand flattened, parallel-sided, with 
fingers simple and about % length of hand; telson considerably shorter than uropods. 

Length of the single specimen, 8 mm. ; 

Malé Atoll, Maldive Is., on ecrinoid, with polycheete. 


2. Periclimenes (Corniger) cornutus Borradaile, 1915 (Plate 54, fig. 10). 
Ann. Mag. Nat. Hist. (8), xv. p. 211. 


Definition: Body moderately compressed ; rostrum outreached slightly by antennular 
stalk and rather more by antennal scale, lanceolate, deep, bearing above seven teeth, 
none behind orbit, and one tooth below; antennal, hepatic close behind antennal, and 
strong, compressed supraorbital spines present; cornea ogival; first joint of antennule 


3866 PERCY SLADEN TRUST EXPEDITION 


well expanded, with moderate stylocerite and distal spine, second and third joints stout 
and subequal; antennal scale of moderate breadth, almost straight-sided, its end truncate 
and not reached by its distal spine; third maxilliped slightly outreaching antennal stalk, 
but not reaching end of first joint of antennular stalk; first leg outreaching antennal 
scale by hand, its wrist and arm equal, hand shorter than wrist, palm and fingers sub- 
equal; last three legs stout, their propodites hairy and armed with spines, their dactylo- 
podites gently curved and ending in a sharp claw; telson shorter than uropods, 
diminishing rather suddenly near the end, which is rounded and has all the spines 
_ short. 

Length of the single specimen 12 mm. 

Malé Atoll, Maldive Is., on red and brown crinoid. 


3. Periclimenes (Corniger) amboimensis (de Man), 1887. 


Anchistia amboinensis, de Man, Arch. Naturg. lili. 1. p. 546, Pl. 22 a, fig. 2 (1887). 
Periclimenes amboinensis, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 383 (1898). 
Amboina. 7 


Subgenus FaucrcEer Borradaile, 1915. 


Ann. Mag. Nat. Hist. (8), xv. p. 207. 

Definition: Rostrum toothed, with straight or concave upper edge, nearly always 
upeurved at tip, long or of medium length. Antennal, usually hepatic, and often supra- 
orbital spines present. Second leg usually with wrist long or of moderate length, usually 
with spine at end of wrist or arm or both. 

Key to the species of Periclimenes (Falciger). 

I. Fingers of great chela each with an oval pit; those of first legs with comb-like 
arrangement of fine teeth. [R.=;5.] 

A. Supraorbital spine present. 
P. (F.) spiniferus de Man, 1902. 


B. Supraorbital spine absent. 
P. (F.) petitthouars: (Audouin), 1825, 
II, Fingers of great chela, and usually also those of first legs, not as in L 
A. Supraorbital spine present. 
1. Rostrum without teeth below. [R.=$. ] 
P. (F.) lifuensis Borradaile, 1898. 
2. Rostrum with teeth below. 
a. Second leg unarmed. Antennal scale broad; with distal spine not 
projecting beyond end. 


i. Antennular stalk reaches end of rostrum. Teeth above rostrum show 
a gap in middle of series. [R=4%. 
P. (F.) edwards: (Paulson), 1875. 


BORRADAILE—ON THE PONTONIIN 367 


i. Antennular stalk does not reach end of rostrum. Teeth above rostrum 
show no gap in middle of series. [R.=2.] 
P. (F.) mlandensis Borradaile, 1915. 


b. Second leg bears one or more spines. Antennal scale narrow; with 
distal spine projecting beyond end. 


1. Upper edge of rostrum straight at first, upcurved at end. 


a. Rostrum shallow, little excavated at base, not reaching end of 
antennular stalk. Second wrist bears one spine. 


(1) Antennal scale broad. Second wrist more than half length of 
hand. [R.=;5.] 
P. (F.) dane (Stimpson), 1860. 


(2) Antennal scale narrow. Second wrist about one-third length 
of hand. [R=#.] 
P. (F.) grandis (Stimpson), 1860. 


8. Rostrum deep, much excavate at base. Second wrist bears two or 
three spines. 


(1) Outer edge of antepenultimate joint of third maxilliped bears 
one spine. Wrist of second leg has no spine on the inner 
side. Fingers of same leg toothless. [R. =2. 

P. (F.) elegans (Paulson), 1875. 


(2) Outer edge of antepenultimate joint of third maxilliped bears 
several spines. Wrist of second leg has strong spine on inner 
side. Fingers of same leg toothed. 

(a) R.=4; its tip simple. In second leg fingers about one-third 
length of palm, and wrist nearly twice length of fingers. 
P. (F.) affinis Borradaile, 1915. 


(b) R.=§; its tip bifid. In second leg fingers. rather more than 
half length of palm, and wrist equal in length to fingers. 
P. (F.) dubius Borradaile, 1915. 
i. Upper edge of rostrum concave from base. 
a. Rostrum deep. Second wrist about as long as arm. 


(1) Antennule has flagella shorter than stalk and third joint longer 
than second. Not more than one spine at end of second wrist. 


(a) Second wrist without spine; a little longer than arm. [R=‘%=2.] 
P. (F-) ensifrons (Dana), 1852. 


(8) Second wrist bears a spine; a little shorter than arm. [R= %=.] 
P. (F.) vitiensis Borradaile, 1898. 


rs 


368 PERCY SLADEN TRUST EXPEDITION 


(2) Antennule has flagella longer than stalk and third joint sub- 
equal to second. Two spines at end of second wrist. 


P. (F.) holmesi Nobili, 1907. 


8. Rostrum shallow. Second wrist not nearly as long as arm. [R. =8.] 
P. (F.) amymone de Man, 1902. 
B. No supraorbital spine. 
1. Upper edge of rostrum straight at first, though its tip may be upturned. 
a. Rostrum without teeth below. Last two joints of antennular stalk very 
long. [R=$.] 
P. (F.) longimanus (Dana), 1852. : 
b. Rostrum with one or more teeth below. Last two joints of antennular 
stalk usually short. 
i. Second leg unarmed. 
a. Rostrum of good depth. Second wrist long. [R.==2.] 
P. (F.) americanus (Kingsley), 1878. 
8. Rostrum rather shallow. Second wrist short. 
(1) Rostrum has three teeth below, and two behind orbit, and is 
directed straight forwards. 
(a) R.=% with additional denticle at tip. First and second legs 
outreach antennal scale. 


P. (F.) compressus Borradaile, 1915. 


(b) R.=8. First and second legs do not outreach antennal scale. 
_P. (F.) amethysteus (Risso), 1826. 


(2) Rostrum has one tooth below, and one behind orbit, and is 


directed somewhat downwards. [R.=$.] 
P. (F£.) brockettt Borradaile, 1915. 


ii. Second leg bears one or more spines. 


a. Second wrist bears three spines. One rostral tooth behind orbit. 


(1) Second arm bears four spines. Antennal scale of good breadth. 
[R=4.] 
P. (F.) denticulatus Nobili, 1907. 


(2) Second arm bears one spine. Antennal scale narrow. 
P. (F.) brachiatus (Stimpson), 1860. 


8. Second wrist bears one spine. Two rostral teeth behind orbit. 


(1) Antennal scale broad, not outcurved, its distal spine not pro- 
jecting beyond its end. Rostrum not outreaching antennular 
stalk. [R.=$§.] 

P. (F.) rotumanus Borradaile, 1898. 


BORRADAILE—ON THE PONTONIIN A 369 


(2) Antennal scale narrow, outcurved, its distal spine projecting 
beyond its end. Rostrum outreaching antennular stalk. 


(a) Antennal scale shorter than carapace. Last two joints of 
antennular stalk slender. [R. = 5%. | 
P. (F.) suvadiwensis Borradaile, 1915. 


(b) Antennal scale longer than carapace. Last two joints of 
antennular stalk stout. [R.=4.] 
P. (F.) pottsc Borradaile, 1915. 


2. Upper edge of rostrum decidedly concave from base. 


a. Rostrum deep, +38. Second leg unarmed. 
P. (F.) seychellensis Borradaile, 1915. 


b. Rostrum slender, 42. Second leg bears spines on arm and wrist. 


i. Rostrum not more than 1? times length of carapace. Second legs 
not more than five times length of carapace. 


P. (F-) borradailec Rathbun, 1904. 


ii. Rostrum 24 times length of carapace. Second legs seven times length 


of carapace. 
P. (F.) kolumadulensis Borradaile, 1915. 


1. (Type). Periclimenes (Falciger) spiniferus de Man, 1902 (Plate 52, fig. 1). 


Anchistia inequimana, Heller, ‘“‘Novara” Rep., Zool. i. 11 p. 109 (1865). 

Anchistia Petitthouarsii, de Man, Arch. Naturg. lili. 1 p. 54 (1887). 

Periclimenes petitthouarsi var. spinifera, de Man, Abh. Senckenb. Ges. xxv. UI. 
p. 824 (1902). 

Periclhimenes Petitthouarsvi var. spinigera, Nobili, Ann. Sci. Nat. Zool. (9), iv. p. 49 
(1906). 

Periclimenes Petitthouars: var. spinifera, Lenz, Voeltzkow’s Reise in Ostafrika, 11. 
p- 567 (1910). 

Tahiti. E. Indies. Indian Ocean. 


2. Periclimenes (Falciger) petitthouars: (Audouin), 1825. 


Palemon Petitthouarsu, Audouin, Descr. Egypte, Hist. Nat. 1. Iv. p. 91 (1825). 
Savigny, Atlas, Crust. pl. 10, fig. 3. 

Anchistia inequimana, Heller, Sitz. k. Ak. Wiss. Wien, xliv. 1. p. 283 (1861). 

Anchistia Petitthouarsw, Kossmann, Ergebn. Reise Rot. Meeres, p. 83 (1880). 

Periclimenes Petitthouarsi, Borradaile, Ann. Mag. Nat. Hist. (7), u. p. 381 (1898). 
Nobili, Ann. Mus. Univ. Napoli, i. mm. p. 6 (1901); Bull. Sci. Fr. Belg. xl. p. 41 
(1906). Lenz, Ark. Zool. vii. xxix. p. 2 (1912). 

Periclimenes petitthouarsu, typischen art, de Man, Abh. Senckenb. Ges. xxv. 111. 
p. 824 (1902). 

SECOND SERIES—ZOOLOGY, VOL. XVII. 47 


370 PERCY SLADEN TRUST EXPEDITION 


Periclimenes Petitthouarsw, forme typique, Nobili, Ann. Sci. Nat., Zool. (9), iv. 
p. 49 (1906). 

Red Sea. Persian Gulf. 

3. Periclimenes (Falciger) amethysteus (Risso), 1826. 

Alpheus amethysteus, Risso, Hist. Nat. Eur. Mér. v. p. 77, Pl. 4, fig. 16 (1826); 
Acad. Leop., 1826, p. 821. 

Pelias amethysteus, Roux, Mém. s. |. Salicoques, p. 25 (1831). Heller, Sitz. k. 
Ak. Wiss. Wien, Math.-Nat. Cl. xlv. 1 p. 408 (1863). 

Anchistia amethystea, Heller, Crust. siidl. Eur. pp. 256, 258 (1863). Carus, 
Prodr. Faun. Medit. i. p. 476 (1885). 

Periclimenes amethysteus, Borradaile, Ann. Mag. Nat. Hist. (7), u. p. 381 (1895). 

Mediterranean. 


4. Periclimenes (Falciger) longimanus (Dana), 1852. 

Anchistia longimana, Dana, U.S. Explor. Exped. Rep. xiii. 1. p. 579 (1852); Atlas, 
Pl. 37, fig. 6 (1855). 

Periclimenes longimanus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 381 (1898). 

Loe. ? 

5. Periclimenes (Falciger) ensifrons (Dana), 1852. 

Anchistia ensifrons, Dana, U.S. Explor. Exped. Rep. xii. 1. p. 580 (1852); Atlas, 
Pl. 38, fig. 1 (1855). Miller, Verh. nat. Ges. Basel, vill. 1. p. 471 (1887). Ortmann, 
Jena. Denkschr. vill. p. 16 (1894). 

Periclimenes ensifrons, Borradaile, Ann. Mag. Nat. Hist. (7), u. p. 382 (1898). 
Nobili, Mem. Ac. Torino (2), lvii. p. 350 (1907). 

Indopacific. 

6. Periclimenes (Falciger) grandis (Stimpson), 1860. 

Anchistia grandis, Stimpson, Proc. Ac. Philadelphia, 1860, p. 39. 

Periclimenes grandis, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 382 (1898). 
Balss, Abh. k. Bayer. Ak. Wiss., Math.-Phys. Kl. i. Suppl. Bd. x. p. 49 (1914). 

1 Anchistia Petitthouarsi, Miers, “‘ Alert” Report, Crust. p. 293 (1884). 

Ousima I. Port Molle? 


7. Periclimenes (Falciger) brachiatus (Stimpson), 1860. 

Anchistia brachiata, Stimpson, Proce. Ac. Philadelphia, 1860, p. 39. 

Periclhimenes brachiatus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 382 (1898). 
Balss, Abh. k. Bayer. Ak. Wiss., Math.-Phys. Kl]. i. Suppl. Bd. x. p. 49 (1914). 

Bonin Is. 


8. Periclimenes (Falciger) dane (Stimpson), 1860. 

Anchistia Dane, Stimpson, Proc. Ac. Philadelphia, 1880, p. 39. 

Periclimenes Dane, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 382 (1898). 

Periclimenes dane (?), Borradaile, Proc. Zool. Soc. Lond. 1898, p. 1004, Pl. 63, fig. 4 
(1899). 

Tahiti. Ellice Is., on coral reef. 


BORRADAILE—ON THE PONTONIIN A 371 


9. Periclimenes (Falciger) edwardsi (Paulson), 1875. 

Anchistia Edwardsi, Paulson, Crust. Red Sea, i. p. 114, Pl. 17, fig. 2 (1875). 
Nobili, Ann. Sci. Nat., Zool. (7) iv. p. 53 (1906). 

Red Sea. 


10. Periclimenes (Falciger) elegans (Paulson), 1875. 

Anchistia elegans, Paulson, Crust. Red Sea, 1. p. 118, Pl. 17, fig. 1 (1875). Nobili, 
Ann. Sci. Nat., Zool. (9), iv. p. 52 (1906). 

Red Sea. 

11. Periclimenes (Falciger) americanus (Kingsley), 1878. 

Anchistia americana, Kingsley, Proc. Ac. Philadelphia, 1878, p. 96; Bull. Essex 
Iineis, sik, jo, WO@, Jel, 2, ine 110) (E34), 

Periclimenes americanus, Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 383 (1898). 
Revdaloove, Idyll Wis, Inela Crorraim, sec, a jo PAN (COI) 

Mid-west Atlantic. To 34 fms. 


12. Periclimenes (Falciger) rotumanus Borradaile, 1898. 

Periclimenes rotumanus, Borradaile, Ann. Mag. Nat. Hist. (7), il. p. 383 (1898) ; 
Proe. Zool. Soc. Lond., 1898, p. 1005, Pl. 638, fig. 5 (1899). 

Rotuma. 


13. Periclimenes (Falciger) vitiensis Borradaile, (1898). 

Anchistia ensifrons, de Man, Arch. Naturg. li. 1. p. 545 (1887). Ortmann, Jena. 
Denkschr. viii. p. 16 (1894). 

Periclimenes vitiensis, Borradaile, Ann. Mag. Nat. Hist. (7), u. p. 383 (1898). Proe. 
Zool. Soc. Lond., 1898, p. 1005, Pl. 64, fig. 6 (1899). Pearson, Rep. Ceylon Pearl Fish. iv. 
p. 78 (1905). 

Periclimenes ensifrons, de Man, Abh. Senckenb. Ges. xxv. 11. p. 126 (1902). Lenz, 
Abh. Senckenb. Ges. xxvii. Iv. p. 380 (1905). 

Indopacitic. 

The supposed spine at the end of the wrist of the first leg in this species is an 
appearance due to a bundle of long hairs. The spine at the end of the second wrist 
is found in both sexes and serves, together with the comparative shortness of this 
wrist and the greater average number of teeth on the rostrum, to distinguish P. vitvensis 
from P. ensifrons. The collection described by Ortmann contained members of both 
species. 

14. Periclimenes (Falciger) lifuensis Borradaile, 1898. 

Periclimenes lifuensis, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 384 (1898); 
Willey’s Zool. Results, iv. p. 405, Pl. 36, fig. 1 (1899). 

Loyalty Is. 

15. Periclimenes (Falciger) amymone de Man, 1902. 

Periclimenes amymone, de Man, Abh. Senckenb. Ges. xxv. lL p. 829, Pl. 25, fig. 53 
(1902). 

Ternate. 

; 47—2 


372 PERCY SLADEN TRUST EXPEDITION 


16. Periclimenes (Falciger) borradailei Rathbun, 1904. 

Periclimenes tenuipes, Borradaile, Ann. Mag. Nat. Hist. (7), il. p. 384 (1898); Willey’s 
Zool. Results, iv. p. 406, Pl. 36, fig. 2 (1899). 

Perichimenes borradaile:, Rathbun, Dec. Crust. N.W. America, p. 34 (1904). Nobili, 
Ann. Mus. Univ. Napoli, ii. xxi. p. 5 (1907). 

New Britain, on coral reef. 


17. Periclimenes (Falciger) holmesi Nobili, 1907. 


Anchastia tenuipes, Holmes, Occ. Pap. Calif. Ac. Sci. vil. p. 216 (1900). 
Periclimenes holmesi, Nobili, Ann. Mus. Univ. Napoli, ii. xxi. p. 5 (1907). 
Santa Catalina I. 


18. Periclimenes (Falciger) denticulatus Nobili, 1907. 


. Periclimenes Petitthowarsii var. denticulata, Nobili, Mem. Ac. Torino, (2), lvii. p. 359 
(1907). 
Polynesia. 


19. Periclimenes (Falciger) nilandensis Borradaile, 1915 (Plate 54, fig. 13). 
Ann. Mag. Nat. Hist. (8), xv. p. 211. 


Definition: Body moderately slender and compressed: rostrum distinctly outreaching 
antennular stalk but barely antennal scale, its upper edge gently concave from base, its 
formula 2; supraorbital, antennal and hepatic spines present; antennular stalk slender, 
its basal joint not much expanded, second joint longer than third, flagella nearly twice as 
long as stalk, delicate; antennal scale moderately broad, with rounded end, its distal 
spine not reaching the end, its fringe very long; first leg with wrist and arm equal, 
hand slightly shorter than either, fingers equal to palm; second legs unequal, the 
longer outreaching antennal scale by wrist and hand, arm and palm subequal, wrist 
a little shorter than either, fingers about two-thirds length of palm, simple, all joints 
unarmed; last three legs slender, with simple, nearly straight dactylopodites, and a 
row of six spines under propodite. 

Length of the single specimen, 16 mm. 

S. Nilandu Atoll, Maldive Is. 


20. Periclimenes (Falciger) affinis Borradaile, 1915 (Plate 54, fig. 11). 
Ann. Mag. Nat. Hist. (8), xv. p. 211. 


Defintion: Body graceful; rostrum of medium depth, straight for the first two-thirds 
of its length, gently upeurved at the tip, its formula %, the first two teeth standing 
behind the orbit, the last at a little distance from the tip, the first and last somewhat 
apart from the rest; antennular stalk ending between sixth and seventh rostral teeth, 
its last joint a little longer than its second joint; antennal scale slender, tapering, 
slightly outcurved, reaching end of rostrum; third maxilliped not quite reaching end of 
second joint of antennule, bearing several spines on outer edge of long joint; first leg 


just reaching end of wrist of second, with wrist about 14 times length of hand; second 


BORRADAILE—ON THE PONTONIIN 373 


leg outreaching rostrum by wrist and hand, its arm and wrist subequal, fingers about 
two-thirds length of wrist, palm a little longer than wrist, all joimts smooth; arm 
bearing at end a spine below, and wrist a spine above, a similar spine inside, and a blunt 
tooth below, fingers straight, bent slightly inwards from axis of palm, and armed with 
two or three interlocking teeth ; last three legs slender, with a few spinules on underside 
of propodites, and dactylopodites long and nearly straight; telson pointed, with naked 
submedian spines. 

Length of longer of the two specimens, 11 mm. 

Salomon I., Western Indian Ocean. 


21. Periclimenes (Falciger) dubius Borradaile, 1915 (Plate 54, fig. 12). 
Ann. Mag. Nat. Hist. (8) xv. p. 211. 


Definition : Closely related to P. affinis, but differing in that (1) the rostral formula 
is 8, (2) the tip of the rostrum is bifid, (3) the stalk of the antennule is shorter and 
stouter, (4) in the second leg the arm and wrist are stouter, while the fingers are rather 
more than half the length of the palm and about equal in length to the wrist, (5) the 
end of the telson bears a few bristles besides the ordinary spines. 

Length of the single specimen, 22 mm. 

~ Minikoi. 

P. elegans, P. affinis, and P. dubius are very closely related species and may 

eventually prove not to be distinct. 


22. Periclumenes (Faleiger) compressus Borradaile, 1915 (Plate 55, fig. 18). 
Ann. Mag. Nat. Hist. (8), xv. p. 212. 


Definition: Body a good deal compressed ; rostrum outreaching antennal scale very 
little, and antennular stalk rather more, deepest at its base, diminishing evenly towards 
tip which is slightly upturned, bearing above five teeth, of which two stand behind the 
orbit, and a minute vestigial tooth near the tip, and below three teeth; antennal and 
hepatic spines present ; first joint of antennular stalk well expanded, with strong distal 
spines, second and third joints subequal, subcylindrical, together shorter than first joint ; 
antennal scale narrow, with sides parallel, tip rounded, and distal spine not reaching 
tip, antennal stalk not reaching end of first joint of antennule; third maxilliped not 
reaching end of antennal stalk; first leg outreaching antennal scale by more than half 
the hand, its wrist, palm, and fingers subequal; second legs equal and similar, out- 
reaching antennal scale by hand, wrists short and subconical, with a notch on the inner 
side, hands long, parallel-sided, somewhat compressed, fingers less than half length 
of palm, a small tooth near the base of each moveable finger, and two or three vestigial 
teeth on the fixed finger ; last three legs long, slender, without spines, and with long, 
sharp, simple dactylopodites ; telson little shorter than uropods, long, and truncate. 

Length of single specimen, 24 mm. 


Saya de Malha. 


374 PERCY SLADEN TRUST EXPEDITION 


23. Periclimenes (Falciger) brocketti Borradaile, 1915 (Plate 55, fig. 15). 
Ann. Mag. Nat. Hist. (8), xv. p. 212. 


Definition: Body moderately compressed; rostrum not quite reaching end of 
antennal scale, but outreaching antennular stalk, shallow, directed slightly downwards, 
straight, save for a very slight upcurving of the tip, bearing above six teeth, of which 
one stands immediately behind the orbit, and below one tooth; antennal and hepatic 
spines present; first joint of antennular stalk moderately expanded, with stylocerite 
half as long as the joint, and two small distal spines, second joint wide, shorter than third 
joint ; stalk of antenna a good deal shorter than first joint of antennular stalk, scale of 
antenna rather broad, with pointed end and small distal spine which does not nearly 
reach the end; third maxilliped not reaching end of antennal stalk, its long joint rather 
broad ; first leg slightly outreaching antennal scale, with wrist a little shorter than arm, 
hand shorter than wrist, fingers longer than palm, hairy; second leg outreaching antennal 
scale by its hand and greater part of wrist, which is comparatively short, hand slender, 
fingers simple, about as long as palm; last three legs with propodites unarmed and 
dactylopodites small, slender, and set in a tuft of hairs; telson narrow, a good deal 
shorter than uropods. 

Length of the longest of the three specimens, 16 mm. 

Malé Atoll, Maldive Is., on brown crinoid. 


T have called this species after Mr W. Brockett, Head Attendant in the Cambridge 
University Laboratory of Zoology, to whose skilled and ready assistance all Cambridge 
zoologists are greatly indebted. 


24. Periclimenes (Falciger) pottsc Borradaile, 1915. 


Ann. Mag. Nat. Hist. (8) xv. p. 213. Potts, Pap. Dep. Mar. Biol. Carnegie Inst. 
Washington, vill. p. 82 (1918). 


Definition: Body moderately stout, somewhat compressed; rostrum reaching end 
of antennal scale, outreaching antennular stalk, its upper edge curving very slightly 
downward from the base and more strongly upward near the tip, its formula $, two 
teeth behind the orbit; hepatic and antennal spines present; basal joint of antennule 
elongate, with small distal spine, second and third joints stout, subequal, flagella about 
twice length of stalk, outer not deeply cleft; antennal scale shorter than carapace, rather 
narrow, outcurved, its distal spine projecting beyond its end; antennal stalk falling 
considerably short of end of first joint of antennular; third maxilliped outreaching by 
its last joint antennal stalk, slender, nearly straight, its last two joints together longer 
than preceding joint; first leg outreaching antennal scale by its hand, its arm and 
wrist subequal, hand nearly as long, and fingers slightly longer than palm; second leg 
outreaching antennal scale by hand and part of wrist, with arm a little longer than wrist, 
palm than arm, fingers and wrist subequal, a spine at end of arm and a small spine 
above at end of wrist; last three legs slender, sparsely hairy, with small spinules under 
propodites, and rather slender, slightly curved dactylopodites. 


BORRADAILE—ON THE PONTONIINA 375 


Colour, purple, brownish in parts and varying in depth with the region of the body. 
Length of longest specimen 13 mm. 
Torres Straits, on Comanthus. 


25. Periclimenes (Faleiger) suvadivensis Borradaile, 1915 (Plate 55, fig. 16). 

Ann. Mag. Nat. Hist. (8), xv. p. 212. 

Definition: Body rather slender and compressed; rostrum outreaching antennular 
stalk, outreached by antennal scale, straight, except at the end, which is gently upeurved, 
diminishing evenly to its end, with formula >", two of the teeth standing behind 
the orbit; hepatic, antennal, and suborbital spines present; antennule slender through- 
out; its second and third joints equal, its first joint equal to the second and third 
together, its outer flagellum long and only cleft near the tip; antennal scale longer than 
carapace, narrow, outcurved, its distal spine projecting at the end; third maxilliped 
reaching end of first joint of antennule; first leg outreaching antennular stalk by wrist 
and hand, arm shorter than wrist. hand about half as long as wrist, fingers slightly 
Jonger than palm; second leg outreaching antennal scale by hand, wrist; and small 
part of arm, its wrist bearing at the end a short stout spine above, its arm a small spine 
below, wrist and palm subequal, fingers rather more than half the length of palm, 
simply toothed; last three legs very long and slender, outreaching antennal scale, 
bearing a few stout hairs, their dactylopodites simple, slender, and nearly straight; 
uropods considerably outreaching telson. 

Length of the longest ofthe two specimens, 14 mm. 

Suvadiva Atoll, Maldive Is. 


26. Periclimenes (Kalciger) seychellensis Borradaile, 1915 (Plates 54, 565, 
figs. 14 a—1). 

Ann. Mag. Nat. Hist. (8), xv. p. 212. 

Definition: Body moderately stout, not much compressed; rostrum outreaching 
antennular stalk, and antennal scale, deep, its ventral edge excavate at base, its dorsal 
edge concave throughout, its formula ‘*, with two teeth behind the orbit; antennal and 
hepatic spines present ; first joint of antennular stalk moderately expanded, stylocerite 
nearly half the length of this joint, distal spine small, second and third joints subequal, 
slender, flagella longer than stalk, the outer cleft for about 4 of length of thicker 
part; antennal scale longer than antennular stalk, rather broad, subtruncate, with 
distal spine projecting beyond end; third maxilliped reaching end of first joint of 


antennule ; first leg reaching end of antennal scale, its arm, wrist, and hand subequal, 
its fingers rather longer than its palm; second legs equal, each outreaching antennal 
scale by fingers and half palm, its fingers and all joimts unarmed, wrist and arm 
subequal, hand nearly half as long again as wrist, palm and fingers subequal; last 
three legs slender, sparsely hairy, with a long spine at end of propodite, dactylopodites 
long, slender, simple, nearly straight. 

Length of the longer of the two specimens, 17 mm. 

Praslin, Seychelles. 


376 PERCY SLADEN TRUST EXPEDITION 


27. Periclimenes (Falciger) kolumadulensis Borradaile, 1915 (Plate 55, ines, 117/)). 

Ann. Mag. Nat. Hist. (8), xv. p. 213. 

Definition: Closely related to P. borradaile: Rathbun, but differing in that (1) the 
rostrum is 24 times the length of the carapace behind it (not more than 12 times in 
P. borradailet), (2) the second leg is seven times the length of the carapace (five times in 
P. borradanler), (3) the last three legs are about + longer than in P. borradailer of the 
same size, (4) the shape of the great chele is very different, the fingers of that on 
one side meeting and bearing a row of about a dozen small teeth, while those of the 
other gape widely and bear each two basal teeth and a distal cutting flange. 

Length of the single specimen, 29 mm. 

Kolumadulu Atoll, Maldive Is. 


Subgenus EnsicEer Borradaile, 1915. 


Ann. Mag. Nat. Hist. (8), xv. p. 207. 

Definition: Rostrum toothless, straight, of medium length. An absence of spines 
from trunk and limbs. Second wrist of medium length. . 

Type: Periclimenes (Ensiger) aurantiacus (Dorna), 1852. 

Anchistia aurantiaca, Dana, U.S. Explor. Exped. xiii. p. 581 (1852); Atlas, 
Pl. 28, fig. 2 (1855). 

Periclimenes aurantiacus, Borradaile, Ann. Mag. Nat. -Hist. (7), i. p. 382 (1898). 

Fiji, in coral. 


Doubtful and wrongly placed species attributed to Perzclimenes. 

Periclimenes tenuipes (Leach). 

Nobili, Ann. Mus. Univ. Napoli, 1. xxi. p. 5 (1907), alludes to this species, which 
he states to be of Mediterranean habitat. I have not been able to find the original 
description of it. 

Perichimenes hertwigi Balss, 1914, and 

Periclhimenes gorgondarum Balss, 1914. 

Abh. k. Bayer. Ak. Wiss., Math.-Phys. K1. ii. Suppl. Bd. x. p. 49 (1914). 

These species do not belong to Periclimenes, and indeed appear from statements 
regarding their telsons not to be Pontoniine. Further information regarding the mouth- 
parts and gills of these very interesting forms is desirable. 

Periclimenes hermitensis, Rathbun, 1914. 

Proc. Zool. Soc. Lond., 1914, p. 655, Pl. 1, figs. 1—3. 

The correct position of this species is probably in the genus Ancyclocaris (see above, 
p- 356). 

Periclimenes sp. de Man, 1902. 

Abh. Senckenb. Ges. xxv. I. p. 833. 

It is not possible to assign this species to any genus until further information 
concerning it be available. 


BORRADAILE—ON THE PONTONIIN A 377. 


Genus Pontoniopsis Borradaile, 1915. 


Ann. Mag. Nat. Hist. (8), xv. p. 207. Potts, Proc. Camb. Philos. Soc. xviii. 
p. 59 (1915). 

Definition: Body rather slender, compressed but with the cephalothorax somewhat 
flattened dorsally; rostrum short, shallow, lanceolate in dorsal view, toothless; a sharp 
antennal spine, but no hepatic or supraorbital spines; eye subspherical, with large 
cornea ; outer flagellum of antennule not deeply cleft; antennal scale broad; mandible 
without palp; second maxilliped without podobranch, its last joint mediad of preceding 
jot; third maxilliped with vestigial arthrobranch, narrow in all joints; legs short, 
directed outwards, rather stout, second pair very unequal, the smaller of the type of the 
first pair, the larger with long, stout, parallel-sided palm and short fingers, dactylo- 
podites of last three pairs stout, curving at tip to simple end-claw. 

Type: Pontoniopsis comanthi Borradaile, 1915 (Plate 57, fig. 27). 

Ann. Mag. Nat. Hist. (8), xv. p. 213. Potts, Pap. Dep. Mar. Biol. Carnegie Inst. 
Washington, vill. p. 81, Pl.-1, fig. 3 (1915). 

Pontomopsis sp. Potts, Proc. Camb. Philos. Soc. xviii. pp. 59, 62 (1915). 


Definition: Rostrum reaching end of second joint of antennular stalk, with slight 
keel above and deeper keel below, straight, its breadth much greater than its depth, 
about equal to greatest width of eye; basal joint of antennule broad, its expansion 
consisting of two broad spines, of which the foremost ends in the distal spinulé and 
the hinder is the stylocerite, second and third joints broad and flattened, second longer 
than third, its distal angles sharp, flagella short, outer cleft to moderate depth; 
antennal scale outreaching antennular stalk, broad, with straight outer and convex inner 
sides, and subrectangular end, which is not nearly reached by the small distal spinule; 
antennal stalk outreaching basal joint of antennule, with basal spine well developed ; 
third maxilliped not reaching end of antennal stalk, its last two joints subequal, 
together shorter than preceding joint; first leg outreaching antennular stalk by hand 
and wrist, arm and wrist subequal, hand rather shorter, fingers shorter than palm, 
hairy; longer leg of second pair outreaching antennular stalk by hand, wrist very short 
and wide, with a sharp process below at the end, hand about equal to preceding three 
joints, fingers much shorter than palm, bearing interlocking teeth ; last three legs hairy 
at end, with stout dactylopodites curving at the top rather strongly to form a sharp 
end-claw; telson rather narrow, blunt ended, shorter than uropods. 

Length of longest specimen, 8 mm. 

Torres Straits, on Comanthus. 


Genus PericLimENaus Borradaile, 1915. 


Ann. Mag. Nat. Hist. (8), xv. p. 207. 

Definition: Body rather stout, decidedly compressed, without sudden change in 
curvature of abdomen, suggestive of Alpheus; rostrum rather short, straight, with 

SECOND SERIES—ZOOLOGY, VOL. XVII. 48 


378 PERCY SLADEN TRUST EXPEDITION 


convex upper and almost straight lower edge, bearing above a row of long, close-set, 
subequal teeth, but toothless below; a strong antennal, but no supraorbital or hepatic 
spine; eye subcylindrical with rather small cornea, which has a cup-shaped depression 
on the outer side; outer flagellum of antennule not deeply cleft; antennal scale of 
moderate breadth ; mandible without palp; second maxilliped without podobranch, its 
last jomt mediad of preceding joint ; third maxilliped with vestige of arthrobranch, and 
with antepenultimate jot of medium breadth; legs rather stout, those of the second 
pair granulate, unequal, and in one of them the chela very heavy but not very long, with 
short, stout fingers, of which one bears a knob and the other a corresponding socket, those 
of last three pairs with short, stout, biunguiculate dactylopodites ; submedian and inter- 


mediate, spines of telson subequal. 


Key to the species of Periclimeneus : 


I. Two teeth of rostrum stand behind orbit. Moveable finger of second leg bears 
knob and fixed finger socket. Fringes on limbs not remarkably long. 
P. robustus Borradaile, 1915. 


II. None of rostral teeth stands behind orbit. Fixed finger of second leg bears 


knob and moveable finger socket. Fringes of limbs very long. 
P. fimbriatus Borradaile, 1915. 


1. (Type.) Periclimeneus robustus Borradaile, 1915 (Plate 55, fig. 20). 
Ann. Mag. Nat. Hist. (8), xv. p. 213. 


Definition: Body rather strongly compressed; rostrum slightly outreached by 
antennal scale and rather more by antennular stalk, directed somewhat downwards, 
sublanceolate, its formula 8; a strong antennal spine ; first joint of antennular stalk long, 
but little expanded, second and third joints subequal; antennal stalk reaching end 
of second joint of antennule, scale subovate ; antepenultimate joimt of third maxilliped 
short, rather wide at base but narrowing towards penultimate joint, which nearly equals 
it in length, last joint shorter than penultimate ; first leg outreaching antennular stalk 
by wrist and hand, wrist longer than arm, hand stout, the fingers a little shorter than 
the palm, a tuft of hairs on the moveable finger; second legs unequal, the larger without 
spines on any joint, its hand very large, stout, granulate and in parts spinulose, the | 
fingers short, abruptly hooked at the tip, the moveable one very wide, with a blunt 
process which fits into a hollow on the fixed finger and a notch which receives a sharp 
tooth of the fixed finger, the smaller hand with the same general characters as the 
larger but shorter, more swollen, and with simple edges to the fingers; the walking legs 
moderately stout, with five spines on the propodites, and the dactylopodites short, stout, 
bearing a sharp end-claw and a smaller claw below it; telson as long as uropods, of 
moderate width, narrowing somewhat towards end, which is truncate. 

In young specimens three or four of the teeth at the base of the rostrum are wanting. 

Length of the longest specimen, 14 mm. 

Amirante I., 20—39 fms. 


BORRADAILE—ON THE PONTONIIN & 379 


2. Periclimeneus fimbriatus Borradaile, 1915 (Plate 55, fig. 19). 
Ann. Mag. Nat. Hist. (8), xv. p. 213. 


Definition: Body rather stout, compressed; rostrum outreached by antennular 
stalk, and rather more by antennal scale, straight, with convex upper and almost straight 
lower edge, bearing above 4—7 subequal teeth, of which the first lies in front of the 
eyes, and without teeth below; strong antennal spine; first joint of antennular stalk 
moderately expanded, second shorter than third, outer flagellum fringed with long hairs ; 
antennal scale of moderate length, with straight outer and convex inner sides, the latter 
fringed with very long hairs, distal spine small; third maxilliped with penultimate joint 
considerably longer than last joint, last two joints together equal in length to rest of 
limb, exopodite rather short, endopodite bearing a fringe of unusually long hairs; first 
leg outreaching antennal scale by hand and wrist, its wrist equal to arm, fingers equal 
to palm, hand stout and hairy; legs of second pair unequal, the larger, which may be 
on either side, about as long as body, no spine on any joint, hand very large, granulated, 
_thick on the outer side and sharp edged on the inner, its fingers short, with a knob 
on the fixed one fitting into a socket in the moveable one, the smaller hand much lke 
the larger but with simple fingers; last three legs moderately stout, with a group of 
long spines at the end of the propodites, and dactylopodites very short and stout and 
bearing two long, hooked claws; telson shorter than uropods, narrow, tapering, with 
rounded end; uropods fringed with very long hairs. 

Length of the longest specimen, 9 mm. 

Mulaku Atoll, Maldive Is. Providence I., to 50 fms. 


Genus HARPILIOPSIS n. gen. 


Definition: Body stout, depressed ; sixth abdominal segment not elongate ; rostrum 
of moderate length, lanceolate, toothed above and below; outer flagellum of antennule 
not deeply cleft; antennal scale broad; mandible without palp; second maxilliped 
without podobranch, with last joint mediad of preceding; third maxilliped with arthro- 
branch, and with last two joints narrow, antepenultimate joint broad or narrow; legs 
stout, directed outwards, the last three with hooked dactylopodites. 


Key to the species of Harpiliopsis : 
I. Antepenultimate joint of third maxilliped narrow. 
H. depressus (Stimpson), 1860. 


Il. Antepenultimate joint of third maxilliped broad. 
H, beaupresi (Audouin), 1825. 


1. (Type.) Harpiliopsis beawpresi (Audouin), 1825 (Plate 55, fig. 21). 
Palemon Beaupresii, Audouin, Descr. Egypte, Crust. p. 91 (1825). Savigny, Atlas, 


Crust. Pl. 10, fig. 4. 
48 —2 


380 PERCY SLADEN TRUST EXPEDITION 


Harpilius Beaupresvi, Heller, Sitz. k. Ak. Wiss. Wien, xliv. L p. 280 (1861). 
Paulson, Crust. Red Sea, p. 113 (1875). de Man, Arch. Naturg. li. 1. p. 539 (1887). 
Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 386 (1898). Nobili, Ann. Mus. Univ. Napoli, 
i. mr. p. 8 (1904); Ann. Sci. Nat. (9), iv. p. 63 (1906). 

¢ Pontonia (Harpilius) dentata, Richters, Decap. Mauritius, p. 165, Pl. 17, figs. 36 
—38 (1880). 

Red Sea. EH. Indies. Indian Ocean. 


2. Harpiliopsis depressus (Stimpson), 1860 (Plate 56, fig. 22). 

Harpilius depressus, Stimpson, Proc. Ac. Philadelphia, 1860, p. 38. Rathbun, Bull. 
U.S. Fish Comm. xxii. 1. p. 920, fig. 68 (1903). 

Anchistia spinigera, Ortmann, Speng. Zool. Jahrb. Syst. v. p. 511, Pl. 36, fig. 23 
(1890). Lenz, Wo, sri, jos GSE (ILQOIL)). 

Periclimenes spinigerus, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 383 (1898); 
Willey’s Zool. Results, iv. p. 405 (1899). 

The fixed finger of the great chela in this species varies in shape. Sometimes it has 
three teeth, of which the most distal is broader than the others. Sometimes this tooth 
loses its point and becomes a blade-like flange. 

Indopacific, in corals. 


Genus Harpintus Dana, 1852. 


U.S. Explor. Exped. Crust. 1. p. 575. Kingsley, Proc. Ac. Philadelphia, 1879, p. 423 
(1880). Borradaile, Ann. Mag. Nat. Hist. (7), ul. p. 386 (1898). 

Definition: Body stout, depressed; sixth abdominal segment not elongate; rostrum 
of good length, toothed above and below, sharp ended : outer flagellum of antennule not 
deeply cleft ; antennal scale of good breadth; mandible without palp ; second maxilliped 
without podobranch, with last joint posterior to preceding joint ; third maxilliped without 
arthrobranch, and with last two joimts narrow, together longer than preceding joint, 
which is broad; legs stout, the last three with hooked dactylopodites. 


Key to the species of Harpolius* : 


I. No hepatic spine. Second leg unarmed. 
H. gerlachet Nobili, 1905. 


Il. A hepatic spine. Second leg bears one or more spines. 


A. Fingers of second leg curved inwards. Inner edge of hand concave. 
HZ. consobrinus de Man, 1902. 


B. Fingers of second leg not curved inwards. Inner edge of hand almost straight. 
FH. lutescens Dana, 1852. 


* The limits of this genus are doubtful. In most respects its characters are similar to those of the 
species on which I have founded Harpiliopsis, but the second maxilliped of the type of Harpiliws is so 
remarkable that no species which does not share this peculiarity can be retained in the genus. The other 
species here included in Harpilius are so placed provisionally, until further information as to their second 
maxilliped is available. Sollaud states that the third maxilliped in this genus has no arthrobranch, but 
does not state what species he has seen. 


BORRADAILE—ON THE PONTONIIN A 381 


1. (Type.) Harpilius lutescens Dana, 1852. 


U.S. Explor. Exped. Rep. xiii. 1 p. 576; Atlas, Pl. 38, fig. 4 (1855). Borradaile, 
Ann. Mag. Nat. Hist. (7), ii. p. 386 (1898). Nobili, Ann. Mus. Napoli, i. m1. p. 3 (1901) ; 
Ann. Sci. Nat. Zool. (9), iv. p. 63 (1906). 

Tongatabu, Red Sea. 


2. Harpilius consobrinus de Man, 1902. 


Harpilius lutescens, de Man, Arch. Naturg. lili. 1. p. 536, Pl. 22a, fig. 1 (1887). 

Harpilius consobrinus, de Man, Abh. Senckenb. Ges. xxv. 11. p. 836, Pl. 26, fig. 54 
(1902). 

E. Indies. 


3. Harpilius gerlacher Nobili, 1905. 


Bull. Mus. Paris, 1905, u1. p. 160; Bull. Sci. Fr. Belg. xl. p. 45, Pl. 4, fig. 10 (1906). 
Persian Gulf, “parmi les polypiers.” 


Genus CoRALLIOCARIS Stimpson, 1860. 


Gdipus, Dana, U.S. Explor. Exped. Rep. xii. 1 p. 572 (1852). 

Coralliocaris, Stimpson, Proc. Ac. Philadelphia, 1860, p. 38. Kingsley, Ib. 1879, 
p- 423 (1880). Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 384 (1898). Ortmann, 
Bronn’s Thierreich, Arthropoden, v. 1. p. 1131. Rathbun, Bull. U.S. Fish Comm. xx. 11. 
p. 122 (1901). 


Definition: Body stout, depressed ; sixth abdominal segment not elongate ; rostrum 
of moderate or good length, with or without teeth, usually somewhat depressed, shallow, 
not curved downwards; outer flagellum of antennule not deeply cleft ; antennal scale 
broad; mandible without palp; second maxilliped without podobranch, with last joint 

mediad of preceding joint, but often projecting far behind it; third maxilliped with 
_ arthrobranch and with last two joimts usually broad, antepenultimate joint more or less 
broadened, not contrasting sharply in width with last two joints; legs stout, directed 
outwards, one or both of second pair with large chela, swollen at base and very short- 
fingered, the last three pairs with hooked dactylopodites and a basal protuberance, which 
is usually large. 


Key to the subgenera of Coralliocaris : 


I. Dactylopodites of walking legs with an accessory spine and small basal 
protuberance. 
Onycocaris Nobili, 1904. 


II. Dactylopodites of walking legs without accessory spine, and with basal protu- 
berance of a good size. 
Coralliocaris s. str. 


382 PERCY SLADEN TRUST EXPEDITION 


Subgenus CoRALLIOCARIS s. str. 


Definition: Dactylopodites with well-developed basal protuberance, without acces- 


sory spinules, not biunguiculate; great chela generally opens horizontally, or has the 
thumb below. 


Key to the species of Coralliocaris s. str. 
I. Rostrum toothless. 


A. Rostrum reaches only to middle of first joint of antennule. Antennular 
stalk reaches middle of antennal scale. 
C. brevirostris Borradaile, 1898. 


B. Rostrum outreaches first joint of antennule. Antennular stalk nearly reaches 
end of antennal scale. 


C. nudirostris (Heller), 1862. 
II. Rostrum toothed. 
A. Rostrum bears no teeth below. 
1. Rostrum bears only one tooth above. 
C. macrophthalma (H. M.-Edwards), 1837. 
2. Rostrum bears 4—5 teeth above. 
a. Rostrum curved upwards. Hyes subspherical. 
C. atlantica Rathbun, 1901. 
b. Rostrum not curved upwards. Eyes subcylindrical. 


i. Rostrum straight, reaching middle of second joint of antennule. Great 
chela opens vertically. 
C. hecate Nobili, 1904. 


ii. Rostrum bent downwards, reaching end of second joint of antennule. 
Great chela opens horizontally. 
C. quadridentata Rathbun, 1906. 
B. Rostrum bears one or more teeth below. 
1. Rostrum bears only one tooth above. 
C. camerant Nobili, 1902. 
2. Rostrum bears 4—6 teeth above. 
a. Second legs subsimilar (often unequal). 
i. Moveable finger not of great width or very much curved. 


a. Rostrum rather deep. Great chela not long or much swollen at 


base. 
C. truncata Rathbun, 1906. 


BORRADAILE—ON THE PONTONIINA 383 


8. Rostrum shallow. Great chela elongate and swollen at base. 


(1) Third maxilliped broad. Rostrum not as long as antennular 


stalk. 
C. superba (Dana), 1852. 


(2) Third maxilliped relatively narrow. Rostrum as long as anten- 
nular stalk. 
C. japomca Ortmann, 1891. 


ii. Moveable finger very wide and much curved. 
C. graminea (Dana), 1852. 


b. Second legs quite unlike. 
C. lucina Nobili, 1901. 


1. (Type.) Corallocaris superba (Dana), 1852. 


(Hdipus superbus, Dana, U.S. Explor. Exped. Rep. xii. 1 p. 573 (1852); Atlas, 
Pl. 37, fig. 2 (1855). 

Coralliocaris superba, Stimpson, Proc. Ac. Philadelphia, 1860, p. 38. de Man, Arch. 
Naturg. Ini. 1. p. 536 (1887). Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 385 (1898). 
Nobili, Ann. Mus. Univ. Napoli, i. mr. p. 3 (1901). Ann. Sci. Nat. (9), iv. p. 55 (1906). 
Balss, Abh. k. Bayer. Ak. Wiss., Math.-Phys. K1. 11. Suppl. Bd. x. p. 53 (1914). 

(Hdipus dentirostris, Paulson, Crust. Red Sea, i. p. 112, Pl. 14, fig. 7 (1875). 

Indopacific, in corals. 


2. Coralliocaris macrophthalma (H. M.-Edwards), 1837 (Plate 56, fig. 24). 


Pontonia macrophthalma, H. M.-Edwards, Hist. Nat. Crust. i. p. 359 (1837) 
Cuvier’s R. An., Crust., Atlas, Pl. 52, fig. 3 (1849). 

(Edipus macrophthalmus, Dana, U.S. Explor. Exped. Rep. xiii. 1. p. 573 (1852). 

Coralliocaris macrophthalma, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 385 (1898). 
Nobili, Ann. Mus. Univ. Napoli, i. mr. p. 3 (1901). 

“Mers d’Asie,” Red Sea. Saya de Malha, to 26 fms. 


a 


3. Coralliocaris graminea (Dana), 1852. 


Hdipus gramineus, Dana, U.S. Explor. Exped. Rep. xi. 1 p. 574 (1852); Atlas, 
Pl. 37, fig. 3 (1855). 

Coralliocaris graminea, Stimpson, Proc. Ac. Philadelphia, 1860, p. 38. Miers, 
“Alert” Crust. p. 563 (1884). de Man, Arch. Naturg. li. © p. 536 (1887); Abh. 
Senckenb. Ges. xxv. 11. p. 840 (1902). Ortmann, Semon’s Forschungsreisen in Austral. 
v. I. p. 16 (1894). Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 385 (1898). Lenz, Abh. 
Senckenb. Ges. xxvi. Iv. p. 381 (1905). 

Coralliocaris inequalis, Ortmann, Speng. Zool. Jahrb. Syst. v. p. 510, Pl. 26, fig. 21 
(1890). Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 386 (1898) ; Willey’s Zool. Results, iv. 
p. 408 (1899). Balss, Abh. k. Bayer. Ak. Wiss., Math.-Phys. Kl. ii Suppl. Bd. x. p. 33 (1914). 


Indopacific, in corals. 


384 PERCY SLADEN TRUST EXPEDITION 


4. Coralliocaris lamellirostris Stimpson, 1860. 


Proc. Ac. Philadelphia, 1860, p. 38. Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 385 
(1898). Nobili, Ann. Mus. Univ. Napoli, i, m1. p. 5 (1901). Balss, Abh. k. Bayer. Ak. 
Wiss., Math.-Phys. K1. 1. Suppl. Bd. x. p. 53 (1914). 


Loo-Choo I., in corals. 


5. Coralliocaris nudirostris (Heller), 1862. 

Gidypus nudirostris, Heller, Sitz. k. Ak. Wiss. Wien, xliv. 1. p. 279, Pl. 3, fig. 25 
(1862). 

Coralliocaris nudirostris, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 385 (1898). 

Red Sea. 


6. Coralliocaris yaponica Ortmann, 1891 (Plate 56, fig. 23). 


Coralliocaris superba var. japonica, Ortmann, Speng. Zool. Jahrb. Syst. v. p. 509 
(1890). Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 385 (1898). 

Professor Gardiner’s collection contains a number of specimens of a Coralliocaris 
which is related to C. swperba but differs from it in the following points : 

(i) The rostrum is much narrower than in C. superba, and is at least as long 
as the antennular stalk, generally considerably longer. 

(ui) The rostral formula, is 375, generally ¢. 

(ii) The third maxilliped, though it is broader than in Periclimenes, is considerably 
narrower than in C. superba. | 

(iv) The hand of the first leg is longer, slender, and hairless, and its fingers are 
not more than half the length of the palm. 

(v) The legs of the second pair are unequal, and the arm bears below at the end 
a flange with a sharp point. The wrist bears a tooth on the inner side, but the little 
teeth found on the upper side in C. superba are wanting. The smaller chela is of simple 
form, with the fingers nearly as long as the palm. 

This species is probably the C. superba var. japonica of Ortmann. 

Japan. Indian Ocean, to 26 fms. 


7. Coralliocaris brevirostris Borradaile, 1898. 


Ann. Mag. Nat. Hist. (7), i. p. 386; Proc. Zool. Soc. Lond., 1898, p. 1006, Pl. 64 
fig. 7 (1899). 
Ellice Is. 


8. Coralliocaris lucina Nobili, 1901. 

Ann. Mus. Univ. Napoli, i. ut p. 5; Ann. Sci. Nat. (9), iv. p. 57 (1906). 

Coralliocaris lamellirostris, de Man, Abh. Senckenb. Ges. xxv. I. p. 842, Pl. 26, 
fir. 5 (1902). | 

Red Sea. Ternate. In corals. 


BORRADAILE—ON THE PONTONIIN Ai 389 


9. Coralliocaris atlantica Rathbun, 1901. 


Bull. U.S. Fish Comm. xx. 1. p. 122, fig. 26. 
St Thomas, 20 to 23 fms. 


10. Coralliocaris camerani Nobili, 1902. 


Boll. Mus. Torino, xvi. xxtml. p. 3. 
Flamenco I. 


11. Coralliocaris hecate Nobili, 1904. 

Bull. Mus. Paris, 1904, p. 232; Ann. Sci. Nat. (OP tive jo Oe}; Ib B, wares (UIGOS)), 

Jibuti. 

12. Coralliocaris rathbunt n. nom. 

Coralhiocaris tridentata, Rathbun, Bull. U.S. Fish Comm. xxi. m1. p. 920, fig. 69, 
E24) ties 1 (1906): 

Hawaiian Is., 28 to 43 fms. 


13. Coralliocaris truncata Rathbun, 1906. 


Bull. U.S. Fish Comm. xxiii. m1. p. 920, fig. 70, Pl. 24, fig. 2. 
Hawaiian Is., 24 fms. 


Subgenus OnycocaRis Nobili, 1904. 


Bull. Mus. Paris, 1904, p. 232; Ann. Sci. Nat. (9), iv. p. 60 (1906). 


Definition: Dactylopodites with small basal protuberance, with accessory spinules, 
biunguiculate; great chela opens vertically, with the thumb above. 


Key to the species of Coralliocaris (Onycocaris) : 


I. Rostrum toothless. 
C. (O.) aualitica Nobili, 1904. 


II. Rostrum toothed. 
A. R.=2. No supraorbital spine. 
C. (O.) tridentata Miers, 1884. 


B. R.==*. A supraorbital spine. 
C. (O.) rhodope Nobili, 1904. 


1. Coralliocaris (Onycocaris) aualitica Nobili, 1904. 

Bull. Mus. Paris, 1904, p. 232; Ann. Sci. Nat. (9), iv. Pl. 3, fig. 3 (1906). 

Jibuti. 

2. Coralliocaris (Onycocaris) rhodope Nobili, 1904. 

Bull. Mus. Paris, 1904, p. 232; Bull. Sci. Fr. Belg. xl. p. 49 (1906); Ann. Sci. Nat. 
(9), iv. p. 61, Pl. 2, fig. 8 (1906). 

Jibuti. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 3 49 


386 PERCY SLADEN TRUST EXPEDITION 


3. Coralliocaris (Onycocaris) tridentata (Miers), 1884. 


Coralliocaris (?) tridentata, Miers, ‘‘ Alert” Orust. p. 294. Pl. 32, fig. c. (1884). 
Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 386 (1898). 

Coralliocaris (Onycocaris) ? tridentata, Nobili, Ann. Sci. Nat. (9), iv. p. 60 (1906). 

Thursday I. 


Genus Courrerra Nobili, 1902. 


Bull. Mus. Torino, xvi. 415, p. 4. 


Definition: Body fairly slender, somewhat depressed, with regions of carapace well 
marked ; abdominal pleura ridged and armed each with a spine, and sixth abdominal 
segment rather longer than the rest; antennal and supraorbital spines very large, hepatic 
and pterygostomian spines well developed; rostrum long and narrow, but shallow and 
simple; outer antennular flagellum not deeply cleft; antennal scale of a good breadth ; 
mandible without palp (2); second maxilliped with last joint mediad of preceding joint 
and projecting but little behind it; third maxilliped narrower than in most Coralliocaris ; 
legs rather stout, those of second pair unequal but similar, with regularly ovoid palm 
and long fingers, those of last three pairs with simple, moderately curved dactyles, 
bearing basal protuberance. 


Type: Coutierea agassiza (Coutiére), 1901. 


Coralliocaris agassizi, Coutiére, Bull. Mus. Paris, vi. p. 115 (1901). 
Coutierea agassizt, Nobili, Boll. Mus. Torino, xvi. 415, p. 4 (1902). 
Barbados, deep water. 


Genus Stecorontonta Nobili, 1906. 


Bull. Mus. Paris, xi. p. 258; Mem. Ac. Sci. Torino (2), lvii. p. 360 (1907). 


Definition: Body fairly slender, a little depressed, with short carapace; which bears 
only a small antennal spine; rostrum toothless, broad, depressed, lanceolate in dorsal 
view ; outer antennular flagellum not deeply cleft; antennal scale rather narrow, with 
long distal spine; mandible without palp; second maxilliped with last joint mediad 
of preceding joint; third maxilliped broad (?); legs rather stout, those of second pair 
unequal and unlike, the smaller of Paleemonid form, the larger with elongate, swollen 
palm and short fingers, those of last three pairs with short, deep, curved dactylopodites, 
bearing a pair of basal prominences. 


Type: Stegopontonia commensalis Nobili, 1906. 

Bull. Mus. Paris, xu. p. 258; Mem. Ac. Sci. Torino (2), lvu. p. 360, Pl. 1, fig. 2 
(1907). 

Hao I., Polynesia, on Echinothria turcarum. 


BORRADAILE—ON THE PONTONTIN © 387 


Genus PoNTONIDES n. gen. 


Definition: Body stout, depressed ; sixth abdominal segment not elongate ; rostrum 
short, flat horizontally, subtriangular, curved downwards, not dentate, with slight keels 
above and below; eye well developed; outer flagellum of antennule not deeply cleft ; 
antennal scale broad; mandible without palp; maxillipeds without exopodites, second 
maxilliped without podobranch, with last joint mediad of preceding joint; third 
maxilliped with vestigial arthrobranch and with all joints broad; legs moderately stout, 
last three ending in simple, nearly straight dactylopodites without basal prominence. 


Type: Pontonides maldivensis (Borradaile), 1915 (Plate 57, fig. 28). 
Pontonia maldivensis, Borradaile, Ann. Mag. Nat. Hist. (8), xv. p. 213 (1915). 


Definition: Body much depressed; carapace broader than long, with small spine 
behind antenna and a blunt process below it; rostrum not reaching end of first joint of 
antennule, sharp pointed both in horizontal and in vertical view; first joint of antennular 
stalk broad, its spines sharp but not very long, third joint a little longer than second, 
flagella of a fair length; antennal scale ovate, longer than antennular stalk, its distal 
spine very small; last two joints of third maxilliped but little twisted from horizontal 
plane ; first leg outreaching antennular stalk by hand and part of wrist, hand and wrist 
subequal, each rather shorter than arm, fingers equal to palm; second legs unequal, the 
larger about twice as long as carapace but shorter than body, its hand long, parallel- 
sided, with short, simple fingers, smaller hand with short swollen palm and long, slender, 
simple fingers; last three legs with long, slender dactylopodites; telson shorter than 
uropods. 

Length of longest specimen, 7 mm. 


Fadiffolu Atoll, Maldive Is. 


Genus ANcuHIsTUS Borradaile, 1898. 


Ann. Mag. Nat. Hist. (7), ii. p. 387. Nobili, Bull. Sci. Fr. Belg. xl. p. 48 (1906). 
Tridacnocaris, Nobili, Ann. Mus. Genova, xl. p. 235 (1899). 
Marygrande, Pesta, Zool. Anz. xxxviii, p. 571 (1911). 


Definition: Body stout, usually swollen and somewhat depressed ; sixth abdominal 
segment not elongate; rostrum of fair length, compressed, bent or curved downwards, 
with broad, usually rounded end, dentate at the tip only or without teeth; outer 
flagellum of antennule not deeply cleft; antennal scale broad; mandible without palp; 
inner lacinia of maxillule broad and hairy; second maxilliped without podobranch, with 
last joint mediad of preceding joint ; third maxilliped with vestigial arthrobranch, and 
with last two joimts narrow, together longer than preceding joint, which is broad or 
narrow; legs stout, directed outwards, the last three with curved dactylopodites, with 
or without a small tooth on the lower side, without basal protuberance. 

; 49—2 


388 PERCY SLADEN TRUST EXPEDITION 


Key to the species of Anchistus : 
I. Last two joints of third maxilliped contrast strongly in width with ante- 
penultimate. 
A. Rostrum shallow, pointed at end. 
A. mirabilis (Pesta), 1911. 


B. Rostrum deep, rounded at end, 
A. imermis (Miers), 1884. 
Il. Last two joints of third maxilliped do not contrast strongly in width with 
antepenultimate. 
A. Rostrum toothed. 
A. miersi (de Man), 1888. 
B. Rostrum toothless. 
1. Rostrum rounded at end. Moveable finger of second cheliped much longer 
than fixed finger. 
A. biungumculatus Borradaile, 1898. 
2. Rostrum ends in a sharp point. Fingers of second cheliped subequal. 
A. spinuliferus (Miers), 1884. 


1. (Type.) Anchistus miersi (de Man), 1888 (Plate 56, fig. 25). 

Harpilius Miersi, de Man, Journ. Linn. Soc. Lond., Zool. xxii. p. 274, Pl. 17, 
figs. 6—10 (1888). Whitelegge, Mem. Austral. Mus. iii. p. 148 (1897). 

Anchistus mersi, Borradaile, Ann. Mag. Nat. Hist. (7), p. 387 (1898); Willey’s 
Zool. Results, iv. p. 408 (1899). Lanchester, Proc. Zool, Soc. Lond. 1901, p. 565 (1902). 
Nobili, Bull. Sci. Fr. Belg. xl. p. 48 (1906); Mem. Ac. Torino (2), lvii. p. 357 (1907). 


Indopacific, in various Lamellibranchs. 


2. Anchistus mermis (Miers), 1884. 

Harpilius inermas, Miers, “Alert” Crust. p. 291, Pl. 32, fig. B (1884). Miller, 
Verh. nat. Ges. Basel, vili. 11. p. 471 (1887). 

Anchistus inermis, Borradaile, Ann. Mag. Nat. Hist. (7) ii. p. 387 (1898). Pearson, 
Rep. Ceylon Pearl Fisheries, iv. p. 77 (1905). Rathbun, Proc. Zool. Soc. Lond. 1914, 
iii, p. 656 (1914). 

Indian Ocean, in Panna. 

3. Anchistus spinuliferus (Miers), 1884. 

Harpilius spinuliferus, Miers, “Alert” Crust. p. 292 (1884). 

Anchistus spinuliferus, Borradaile, Ann. Mag. Nat. Hist. (7) u. p. 387 (1884). 

Loc.? In Tridacna. 

4, Anchistus biunguiculatus Borradaile, 1898. 


Ann. Mag. Nat. Hist. (7) i. p. 387; Willey’s Zool. Results, iv. p. 408, Pl. 1, 
fic. 5 (1899). 
British New Guinea, in Tridacna. 


BORRADAILE—ON THE PONTONIINA® 389 


5. Anchistus mirabilis (Pesta), 1911. 


Marygrande mirabilis, Pesta, Zool. Anz. xxxvill. p. 571. 
Samoa, in Tridacna gigas. 


6. Anchistus (?) armatus (H. M.-Edwards), 1837. 


Pontona armata, H. M.-Edwards, Hist. Nat. Crust. ii. p. 359 (1837). 
Anchistus (?) armatus, Borradaile, Ann. Mag. Nat. Hist. (7) ii. p. 387 (1898). 
New Zealand. 


Genus Pontonia Latreille, 1829. 


Cuvier's R. An. 2nd ed. iv. p. 96. Roux, Mém. s. les Salicoques, p. 26 (1831). 
H. M.-Edwards, Hist. Nat. Crust. 11. p. 358 (1837). De Haan, von Siebold’s Fauna 
Japonica, Crust. p. 175 (1850). Dana, U.S. Explor. Exped. Rep. xii. 1. p. 570 (1852). 
Kingsley, Proc. Ac. Philadelphia, 1879, p. 422 (1880). Joliet, Arch. zool. exper. x. 
p: 19 (1882). Carus, Prodr. Faun. Medit. 1. p. 475 (1885). Borradaile, Ann. Mag. Nat. 
Hist. (7), i. p. 388 (1898). Ortmann, Bronn’s Thierreich, Arthropoden, v. 1. p. 1131 
(1899). Rathbun, Bull. U.S. Fish Comm. xx. 11 p. 121 (1901). 


Definition - Body stout, swollen, somewhat depressed ; sixth abdominal segment not 
elongate ; rostrum short, depressed, curved downwards, not dentate, with or without 
a keel below at the free end; eye more or less reduced, outer flagellum of antennule 
not deeply cleft ; antennal scale broad ; mandible without palp ; inner lacinia of maxillule 
very broad and hairy; second maxilliped without podobranch, with last joint mediad 
of preceding joint; third maxilliped without arthrobranch and with all joints broad, 
but the widest surface of the last two in a different plane from that of the ante- 
penultimate, so that they are apt to appear narrow; legs stout, directed outwards, the 
second pair with heavy chela, the last three with dactylopodite nearly straight, simple, 
and without basal protuberance. 


Key to the species of Pontonia: 
I. Rostrum reaches at least to middle of second joint of antennule. 
A. last three legs end in simple claw. Southern species. 


1. Rostrum outreaches antennular stalk. Dactylopodites of last three legs 
strong. Second legs subequal and similar. 
P. minuta Baker, 1907. 
2. Rostrum ends at middle of second joint of antennular stalk. Dactylopo- 


dites of last three legs weak. Second legs unequal and unlike. 
P. pinne Ortmann, 1894. 


B. Last three legs end in double claws. Northern species. 


1. Rostrum very narrow throughout. W. American. 
P. califormensis Rathbun, 1902. 


390 PERCY SLADEN TRUST EXPEDITION 


2. Rostrum broad at base. Old World and Atlantic. 


a. Antennal stalk as long as antennular. Hyes and first jot of antennular 
stalk subequal. Japanese. 
P. nipponensis de Haan, 1850. 


b. Antennal stalk shorter than antennular. First joint of antennular stalk 
distinctly longer than eyes. Western species. 
i. Antennal scale has a spine and outreaches antennular stalk. 
Mediterranean. 
a. Third maxilliped does not reach end of antennular stalk. Body 


bears yellow spots. 
P. flavomaculata Heller, 1864. 


8. Third maxilliped reaches end of antennular stalk. Body rose-pink. 
P. tyrrhena (Petagna). 


ui. Antennal scale has no spine and does not outreach antennular stalk 
W. Indies. 
P. grayt Rathbun, 1901. 


II. Rostrum short, not reaching end of first joint of antennule. 
A. Rostrum reaches middle of first joint of antennule. 


1. Rostrum without keel below. 
P. ascidicola Borradaile, 1898. 


2. Rostrum with keel below. 
P. mexicana Guérin, 1856. 


B. Rostrum not more than 4 length of first joint of antennule. 
P. brevirostris Miers, 1884. 


1. (Type.) Pontonia tyrrhena (Petagna) (Plate 57, fig. 29). 


Astacus tyrrhenus, Petagna, Ent. Pl. 5, fig. 5 (fide Risso). 

Gnathophyllum tyrrhenus, Desmarest, Consid. sur les Crust. p. 229. 

Alpheus pinnophylax, Otto, Mém. Ac. cur. nat. Bonn, xiv. Pl. 21, figs. 1, 2. 

Pontonia tyrrhena, Latreille, Encycl. Pl. 326, fig. 10; Cuvier's R. An. 2nd ed. 
p. 96 (1829). H. M.-Edwards, Crust. ii. Pl. 360 (1837). Cuvier’s R. An. Crust. Atlas, 
Pl. 52, fig. 4 (1849). Heller, Crust. siidl. Eur. p. 251, Pl. 8, figs. 10, 11 (1868). 
Ortmann, Speng. Zool. Jahrb. Syst. v. p. 509, Pl. 27, fig. 9 (1891). Nobili, Bull. Sci. Fr. 
Belg. xl. p. 49 (1906). 

Callianassa thyrrhenus, Risso, Hist. Nat. Eur. Mér. p. 54 (1826). 

Pontonia custos, Guérin, Expéd. Morée, Zool. p. 36, Pl. 37, fig. 1. Carus, Prodr. 
Faun. Medit. p. 475 (1885). Borradaile, Ann. Mag. Nat. Hist. (7), 1. p. 388 (1898). 

2 Pontonia parasitica, Roux, Mém. sur les Salicoques, p. 26 (1831). 

Mediterranean. In lamellibranchs and sponges. 


er = 


BORRADAILE—ON THE PONTONIINA 391 


2. Pontona nypponensis De Haan, 1850. 

Von Siebold’s Fauna Japonica, Crust. p. 180. Borradaile, Ann. Mag. Nat. Hist. 
(7), ii. p. 388 (1878). Balss, Abh. k. Bayer. Ak. Wiss., Math.-Phys. Kl. ii. Suppl. Bd. 10, 
p. 53 (1914). 

Hymenocera nipponensis, De Haan, loc cit. Pl. 46, fig. 8 (1850). 

Japan. 


3. Pontona mexicana Guérin, 1856. 

De la Sagra’s Hist. Cuba. Rathbun, Bull. U.S. Fish. Comm. xx. 11. p. 122 (1901). 

¢Pontoma unidens, Kingsley, Proc. Ac. Philadelphia, 1879, p. 422, Pl. 14, fig. 9 
(1880). Borradaile, Ann. Mag. Nat. Hist. (7), 11. p. 389 (1898). 

Mid-west Atlantic. 


4. Pontonia flavomaculata Heller, 1864. 

Verh. zool.-bot. Ges. Wien, xiv. p. 51. Carus, Prodr. Faun. Medit. i. p. 475 
(1885). Ortmann, Speng. Zool. Jahrb. Syst. v. p. 509 (1890). 

Pontoma drazone, Joliet, Arch. Zool. Exper. x. p. 108 (1882). 

Pontona phallusie, Marion, Ann. Mus. Nat. Hist. Marseille, Zool. i. (1883). 
Gourret, Compt. Rend. civ. p. 187 (1887). 

Mediterranean, in ascidians. 


5. Pontonia brevirostris Miers, 1884. 

“Alert” Crust. p. 562, Pl. 51, fig. B. Borradaile, Ann. Mag. Nat. Hist. (7), 
ii, p. 389 (1898). 

Seychelles, ‘in clamp (clam ?) shells.” 

6. Pontona pinnae Ortmann, 1894. 

Jena. Denkschr. vil. p. 16, Pl. 1, fig. 3. Borradaile, Ann. Mag. Nat. Hist. 
(7), u. p. 389 (1898). Nobili, Ann. Sci. Nat. (9), iv. p. 65 (1906); Bull. Sci. Fr. Belg. 
xl. p. 46, Pl. 4, fig. 11 (1906). 

* Cancer custos Forskal, Descrip. Anim. p. 94 (1775). 

Red Sea. EH. Africa. In Pinna. 


7. Pontonia ascidicola Borradaile, 1898. 

Ann. Mag. Nat. Hist. (7), ii. p. 389; Willey’s Zool. Results, iv. p. 409, Pl. 1, 
fig. 6 (1899). 

New Britain, in ascidian. 

8. Pontoma gray: Rathbun, 1901. 

init. WI, S_ ldtsin Conan, x6, ii, jo; WP, 

Porto Rico. 

9. Pontona californiensis Rathbun, 1902. 


Proc. U.S. Mus. xxiv. p. 902; Dec. Crust. N.W. America, p. 33, fig. 34 (1904). 
California. | 


392 PERCY SLADEN TRUST EXPEDITION 


10. Pontonia minuta Baker, 1907. 


Tr. Roy. Soc. S. Australia, xxxi. p. 189, Pl. 24, figs. 9—12. 
S. Australia. 


Genus ConcHopyteEs Peters, 1851. 


Ges. naturf. Freunde Berlin, 1851 ( fide Hilgendorf); Ber. k. Ak. Wiss. Berlin, 1852, 
p- 591. Hilgendorf. Monatsber. k. Ak. Wiss. Berlin, 1875, p. 835. Borradaile, Ann. 
Mag. Nat. Hist. (7), ii. p. 389 (1898). 


Definition : Body stout, swollen, depressed ; sixth abdominal segment not elongate ; 
rostrum short, depressed, curved downwards, toothless, with a keel below; eye small, 
outer flagellum of antennule not deeply cleft; flagella of antennule always, and that of 
antenna often, very short; antennal scale broad; mandible without palp; inner lacinia 
of maxillule very broad and hairy; second maxilliped without podobranch, with last 
joint mediad of preceding joint ; third maxilliped without arthrobranch, with all joints 
broad, but with widest surface of last two in different plane from that of ante- 
penultimate ; legs stout, directed outwards, the second pair with heavy chela, the last 
three with strongly curved dactylopodites which bear basal protuberance. 


Key to the species of Conchodytes : 


I. Flagellum of antenna less than twice length of rostrum. Latter ends bluntly. 
Distal spine of antennal scale projects beyond end cf scale. Indopacific. 


A. Fingers of chelae without projection on outer side. 


1. Rostrum outreaches antennal scale. 
C. tridacne Peters, 1851. 


2. Rostrum does not outreach antennal scale. 
C. meleagring Peters, 1851. 


B. Moveable finger of one of second chelz bears square projection on outer side. 
C. biunguiculata (Paulson), 1875. 


II. Flagellum of antenna a good deal more than twice length of rostrum. Latter 
ends in sharp spine. Distal spine of antennal scale does not reach end of scale. 
American. 

A. Palm of great chela more than half as wide as long. Third maxilliped does 


not reach end of antennal stalk Panama. 
C. margarita (Verrill), 1869. 


B. Palm of great chela less than half as wide as long. Third maxilliped slightly 
outreaches antennal stalk. W. Atlantic. 
C. domestica (Gibbes), 1851. 


BORRADAILE—ON THE PONTONIINA 393 


1. (Type.) Conchodytes tridacne Peters, 1851. 


Ges. naturf. Freunde Berlin, 1851 (fide Hilgendorf); Ber. k. Ak. Wiss. Berlin, 1852, 
p- 594; Arch. Naturg. xviii. p. 288 (1852). Hilgendorf, Monatsber. k. Ak. Wiss. Berlin, 
1878, p. 835. Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 390 (1898); Proc. Zool. Soe. 
Lond. 1898, p. 1007 (1899); Willey’s Zool. Results, iv. p. 407 (1899). Nobili, Ann. Mus. 
Genova, xl. p. 235 (1899); Ann. Sci. Nat. (9), iv. p. 66 (1906). 

Pontonia tridacne, Dana, U.S. Explor. Exped. Rep. xiii. 1 p. 371 (1852); Atlas, 
Pl. 37, fig. 1 (1885). Miers, “Alert” Rep. Crust. p. 290 (1884). ?%Ortmann, Speng. 
Zool. Jahrb. Syst. v. p. 509, Pl. 37, fig. 10 (1891). 

Indopaeific, in Trzdacna. 


The separation of this species from C. meleagrine is difficult. Peters, whose state- 
ments are none too explicit, gives several points of difference, but these all vary 
independently, with the exception of that which consists in the absence of a fringe 
to the antennal scale in C. tridacne. I have not met with this feature, and believe 
it to have been due to accident in Peters’ specimens. Examination of a number of 
specimens from various localities shows the following facts: (1) specimens in which 
the rostrum outreaches the antennal scale always have the third maxilliped falling 
considerably short of the end of the rostrum, and the arm of the first leg no longer 
than the wrist, and are always found in Tridacna, (2) specimens in which the rostrum 
is outreached by the antennal scale may have the third maxilliped reaching or ex- 
ceeding the end of the rostrum, and may have the arm of the first leg longer than the 
~ wrist, and are usually, though not always, found in Meleagrina. Form (1) is presumably 
C. tridacne, and form (2) C. meleagrine. Whether they are specifically distinct is 
another question. In both forms the chelipeds of the second pair vary greatly both 
in degree of inequality and in the actual size of the greater of them. 


2. Conchodytes meleagrine Peters, 1851 (Plate 57, fig. 26). 


Ges. naturf. Freunde Berlin, 1851 (fide Hilgendorf); Ber. k. Ak. Wiss. Berlin, 1852, 
p- 594; Arch. Naturg. xviii. p. 288 (1852). Hilgendorf, Monatsber. k. Ak. Wiss. Berlin, 
1878, p. 836. Borradaile, Ann. Mag. Nat. Hist. (7), i. p. 390 (1898). Nobili, Mem. Ac. 
Torino (2), lvui, p. 59 (1907). Pearson, Rep. Ceylon Pearl Fisheries, iv. p. 77 (1905). 

Pontonia meleagrine, Bate, “ Challenger” Macrura, p. 707, Pl. 124, figs. 1, 2 (1888). 

Pontonia tridacne, Miers, “Alert” Rep. Crust. p. 290 (1884). 

¢ Pontona maculata, Stimpson, Proc. Ac. Philadelphia, 1860, p. 38. Balss, Abh. 
k, Bayer. Ak. Wiss., Math.-Phys. Kl. ii. Suppl. Bd. x. p. 53 (1914). 

? Pontome enflée, H. M.-Edwards, Hist. Nat. Crust. 1. p. 360 (1837). 

Indopacific, in Meleagrina, and occasionally in Tridacna. 


3. Conchodytes domestica (Gibbes), 1850. 


Pontonia domestica, Gibbes, Proc. Am. Assoc. il. p. 196 (1850). Kingsley, Proce. 
Ac. Philadelphia, 1878, p. 95. 

Pontoma (?) domestica, Borradaile, Ann. Mag. Nat. Hist. (7), ii. p. 389 (1898). 

SECOND SERIES—ZOOLOGY, VOL. XVII. 50 


394 PERCY SLADEN TRUST HXPEDITION 


Conchodytes domestica, Rathbun, Bull. U. 8. Fish Comm. 1900, i. p. 122 (1901). 
Bahamas, S8.E. United States, in Pinna. Porto Santo, in Pecten (Brit. Mus.). 


4. Conchodytes margarita (Verrill), 1869. 


Pontoma margarita Smith, Verrill, Amer. Nat. i. p. 245 (1869). Lockington, 
Bull. Essex Inst. xi. p. 163 (1878). 

Conchodytes margarita, Rathbun, Dec. Crust. N.W. America, p. 34 (1904). 

Panama, in Margaritophora fimbriata. 


5. Conchodytes biunguiculata (Paulson), 1875. 


Pontona biunguiculata, Paulson, Crust. Red Sea, p. 111, Pl. 15, fig. 1 (1875). 
Conchodytes biunguiculata, Nobili, Ann. Sci. Nat. (9), 1v. p. 67 (1906). 
Red Sea. 


Genus Typron Costa, 1844. 


Ann. Ac. Aspir. Nat. Napoli, u.; Faun. Regn. Napoli, i. 1 (1846). Heller, Crust. 
stidl. Hur. p. 254 (1863). Bate, Ann. Mag. Nat. Hist. (4), i. p. 119 (1868). Kingsley, 
Proc. Ac. Philadelphia, 1879, p. 422 (1880). Carus, Prodr. Faun. Medit. 1. p. 475 
(1885). Ortmann, Speng. Zool. Jahrb. Syst. v. p. 508 (1890); Bronn’s Thierreich, Arthro- 
poden, v. U. p. 1131 (1899). Borradaile, Ann. Mag. Nat. Hist. (7), u. p. 390 (1898). 

Pontonella, Heller, Verh. zool.-bot. Ver. Wien, vi. p. 629 (1856). 


Definition: Body stout, compressed ; sixth abdominal segment not elongate; rostrum 
short, compressed, bent upwards at free end, pointed, with few or no teeth; outer 
flagellum of antennule short, not cleft; antennal scale vestigial; second maxilliped 
without podobranch, with last joint mediad of preceding joint; third maxilliped without 
arthrobranch, with narrow endopodite; legs fairly stout, one of the second pair with a 
very large chela, the dactylopodites of the last three much or little curved, biunguiculate, 
without basal protuberance. 


Key to the species of Typton: 


I. Rostrum toothless. Dactylopodites of last three legs not strongly curved. 
T. spongicola Costa, 1844. 


II. Rostrum toothed. Dactylopodites of last three legs strongly curved. 
T. bouvert Nobili, 1904. 


1. (Type.) Typton spongicola Costa, 1844. 

Ann. Ac. Aspir. Nat. Napoli, i.; Faun. Reg. Napoli, i. 1. Pl. 6, figs. 1—6 (1846). 
Grube, Ausflug-nach Triest, pp. 65 and 125. Heller, Crust. siidl. Eur. p. 254, Pl. 8, 
figs. 12—17 (1863). Norman, Ann. Mag. Nat. Hist. (4), i. p. 176 (1868). Carus, 
Prodr. Faun. Medit. i. p. 475 (1885). Ortmann, Speng. Zool. Jahrb. Syst. v. p. 508, 
Pl. 37, fig. 8 (1891). Borradaile, Ann. Mag. Nat. Hist. (7), tl. p. 390 (1898). 


BORRADAILE—ON THE PONTONIINA 395 


Pontonella glabra, Heller, Ver. zool.-bot. Ver. Wien, 1856, p. 629, Pl. 9, figs. 


1—15. 


Alpheus edwardsu, Couch, Journ. Linn. Soe. Lond., Zool. v. p- 210 (1860). 
Typton spongiosus, Bate, Ann. Mag. Nat. Hist. (4), i. p. 119 (1868). 
Mediterranean. Cornwall. In sponges. 

2. Typton boumerr Nobili, 1904. 

Bull) Mus. Paris, 1904, v. p. 233); Ann. Sci. Nat. (9), iv. p. 67, Pl. 3; figy 4 


(1906). 


Jibuti. 


EXPLANATION OF PLATES 52—57. 


Fig. 1. Periclimenes spiniferus de Man, 1902 (Plate 52). a, side view; 6, dorsal view of cephalothorax ; 


c, dorsal view of antennule and antenna with adjacent structures: a=coxocerite, 8 =basicerite 
(outer division), 6=scale, e=base of eyestalk, 11=second joint of endocerite; c’, ventral view of 
base of antenna, lettermg as in c, also 8*=inner division of basicerite, y=tubercle for 
opening of green gland, i=first joint of endocerite; d, ventral view of left mandible; d’, end 
of molar process of the same; d’’, end of molar process of right mandible; d", anterior 
view of right mandible; e, maxillule; f, maxilla; g, Ist maxilliped; h, 2nd maxilliped; 
7, 8rd maxilliped, removed with arthrobranch; h, chela of 1st leg; J, chela of 2nd leg (larger of 
pair); m, walking leg; n, 1st abdominal limb of male; 0, 2nd abdominal limb of male; 0’, 2nd 
abdominal limb of female; p, dorsal view of telson; g, paragnatha. 


. 2. Urocaridella gracilis Borradaile, 1915 (Plate 53). a, e, f, g, h, i, n, 0, p, as fig. 1; 


dad’, anterior-ventral view of right mandible; 0”, 3rd abdominal limb of male. 


. 3. Urocaris psamathe de Man, 1902 (Plate 53). e—, as fig. 1. 
. 4. Palemonella elegans Borradaile, 1915 (Plate 53). Side view of forepart of body. 
. 5. Palemonella longirostris Borradaile, 1915 (Plate 53). a, as fig. 1; d’, anterior-ventral view 


of left mandible. 


. 6. Periclimenes frater Borradaile, 1915 (Plate 53). c, basal joint of right antennule in dorsal 


view; m, end of a walking leg. 


. 7. Periclimenes incertus Borradaile, 1915 (Plate 53). Side view. 

. 8. Periclimenes brocki (de Man), 1887 (Plates 53, 54). ff g, h, 7, as fig. 1. 

. 9. Perichimenes ceratophthalmus Borradaile, 1915 (Plate 54). , as fig. 1; a, side view of carapace. 
. 10. Periclimenes cornutus Borradaile, 1915 (Plate 54). a, 6, as fig. 1. 

. 11. Periclimenes afimis Borradaile, 1915 (Plate 54). a, b, e, g, h, 2, p, as fig. 1. 

. 12. Periclimenes dubius Borradaile, 1915 (Plate 54). a, b, p, as fig. 1. 

. 18. Periclimenes nilandensis Borradaile, 1915 (Plate 54). a, b, as fig. 1. 

. 14. Periclimenes seychellensis Borradaile, 1915 (Plates 54, 55). a, b, e, f, h, 7, as fig. 1. 

. 15. Periclimenes brocketti Borradaile, 1915 (Plate 55). a, 6, as fig. 1. 

. 16. Periclimenes swvadivensis Borradaile, 1915 (Plate 55). Side view. 

_ 17. Periclimenes kolumadulensis Borradaile, 1915 (Plate 55). J, 1’, chele of second pair. 

. 18. Periclimenes compressus Borradaile, 1915 (Plate 55). a, b, as fig. 1. 

Fig. 
. 20. Perichimeneus robustus Borradaile, 1915 (Plate 55). a, b, f, g, h, 4, as fig. 1. 
. 21. Harpiliopsis beauprest (Audouin), 1825 (Plate 55). Third maxilliped. 

. 22. Harpiliopsis depressus (Stimpson), 1860 (Plate 56). e—+, as fig. 1. 


19. Perrclimeneus fimbriatus Borradaile, 1915 (Plate 55). a, b, 2, l, m, p, as fig. 1. 


50—2 


396 
Fig. 


Fig. 


Fig. 
Fig. 


Fig. 
Fig. 


Fig. 


PERCY SLADEN TRUST EXPEDITION 


23. Coralliocaris sp. ?C. japonica (Ortmann), 1891 (Plate 56). f 2, J, m, as fig. 1; a’, side 


view of rostrum; 0’, dorsal view; J’, small chela of second pair. 

24. Coralliocaris macrophthalma (H. M.-Edwards), 1837 (Plate 56). Third maxilliped. 

25. Anchistus miersi (de Man), 1888 (Plate 56). e—1, as fig. 1. 

26. Conchodytes meleagrinw Peters, 1851 (Plate 57). a’, b’, views as fig. 23 of female specimen 
with tail fan turned forward; d, d’, d’”,e, f, g, h, 1, m, as fig. 1; d”, view of end of molar process 
of left mandible from dorsal side; 7, ventral view of region of mouth, semidiagrammatic; s, the 


same, after removal of maxillipeds and maxilla; ¢, the same, after removal of maxillules and 


paragnatha. 

27. Pontoniopsis comanthi Borradaile, 1915 (Plate 57). Dorsal view of rostrum and eyes. 

28. Pontonides maidivensis (Borradaile), 1915 (Plate 57). a, 2, as fig. 1; 6, dorsal view of 
rostrum and eyes. 

29. Pontonia tyrrhena (Petagna) (Plate 57). male specimen in dorsal view, after Milne- 
Edwards; @ female specimen in dorsal view. 


—— 


Percy SLADEN Trust EXPEDITION. 


GROEN ADAITE) TRANS. LINN. SOC. SER. 2, ZOOL. VOL. XVII. PL. 52 


1q 


PONTONIINZ FROM THE INDIAN OCEAN 


Percy SnADEN Trust ExPEprrion. 


(Gee TRANS. LINN. SOC. SER. 2, ZOOL. VOL. XVII. PL. 53 


aS 


<LLLG ITTTXZEETETOE. 


PONTONIINA® FROM THE INDIAN OCEAN 


Percy SuapEN Trust EXPEDITION. 
(BorRaDAILE) TNRAING, ILIONIN, SOC, SigiR. 2, ZOOL, WOIL, 2OWIN, JPL, 4! 


PONTONIINAZ? FROM THE INDIAN OCEAN 


Prrcy StapEN Trust EXxpeEDITION. 
(BorraDAlLe) 


TOWAINS, ILIONIN, SOC, SEI, 2 ZOOL, WOIL, XWill, IPL, 8S 


PONTONIIN4Z® FROM THE INDIAN OCEAN 


Percy Siuapen Trust Experpition. 
(Borraparze) TRAINS, ILIININ, SOG, SER, 2B ZOOL, WOIL, XWill, PIL, BB 


Ae NA, CA i 


if 
WS 
SS 


PONTONIINA® FROM THE INDIAN OCEAN 


Preroy SuADEN Trust EXPEDITION. 
(Borrapal.e) 


TOVAINS, ILIMNIN, SOC, SHIR. 2, ZOOIL, WOJL, SOW IIL, 


PONTONIINAZZ FROM THE INDIAN OCEAN 


57 


No. IX.—ON CARIDES FROM THE WESTERN INDIAN OCEAN. 


By L. A. Borrapatrie, M.A. 


(Lecturer in Zoology in the University of Cambridge, Fellow, Dean, and Lecturer 
of Selwyn College.) 


(CommMuNICcATED BY Pror. J. STANLEY Garpinger, M.A., F.R.S., F.L.S.) 


(Plates 58, 59.) 
Read 2nd November, 1916. 


THE prawns which are enumerated in this paper are the residue of several large 
collections, a great part of which has already been described elsewhere. They were 
gathered by Professor J. Stanley Gardiner’s two expeditions to the Islands and Banks 
of the Western Indian Ocean, and by Mr J. C. Fryer in Aldabra. I have dealt with 
the Ponzides and Stenopides of these collections in an article in the Transactions of 
this Society which, appeared in February 1910 ((2) Zool. xiii. pt. 2, p. 257), and with 
the freshwater Carides in other articles in the same publication ((2) Zool. xi. pt. 1, p. 63, 
and xiil. pt. 3, p. 405) and in Gardiner’s Fauna of the Maldives (vol. i. p. 64). The 
Alpheide have been reported upon by Professor Coutiére in Gardiner’s Fauna of the 
Maldives (vol. ii. p. 852) and in the present volume, and the Pontonide by myself, also 
in this volume (pp. 323—396). Only 26 species remain to be dealt with here, but these 
are by no means the least important part of the collections. Twelve of them were 
new to science, including one for which it has been necessary to found a new genus”, 
and there is an exceptionally large proportion of interesting forms. The common but 
remarkable species Saron marmoratus is of course included, and this allows me to 
make some remarks concerning Mr Kemp’s recent discoveries with regard to the 
seeming dimorphism of the males, to which I first called attention in the year 1898. 
There is a new species of the genus Zhor, hitherto only known to contain 7. paschalis 
(Heller). The rare Ingur uvee, hitherto only known by the specimens described by 
myself from the Loyalty Islands, has reappeared, and I am enabled to add some 
details to my original description. Lysmatella is a new genus related to Hippolysmata 
but, somewhat strangely, unprovided with mastigobranchs upon the legs. The specimens 
which I have referred to Leander debilis throw some light upon the meaning of the 
great variability of this species and upon the nature of the numerous forms related 


* Short definitions of the new species and of the new genus have already appeared in the Annals 
and Magazine of Natural History for February, 1915. 


398 PERCY SLADEN TRUST EXPEDITION 


to it, showing that we have here quite possibly a number of distinct local races— 
a rare phenomenon among Carides. Nikoides maldivensis is a second representative of 
a genus which has not been met with since it was described in Paulson’s well-known but 
for a long time inaccessible paper on the Decapoda of the Red Sea. Lastly, a study 
of the species of Amphipalemon, Hymenocera, and Gnathophyllum has elucidated the 
relationships of these very interesting and remarkable genera, and necessitated a very 
considerable rearrangement of families in the neighbourhood of the Crangenoida. There 
is, indeed, no exaggeration in saying that the result of the examination of this little 
collection will be to bring about fundamental alterations in our ideas of the affinities 
of the higher families of the Carides. 

It is unfortunately not possible at present to draw any conclusions concerning the 
geographical distribution of the Indopacific prawns. 

The following is a list of the species, with comments upon facts of interest 
concerning them. 


Superfamily PastpPHAOIDA. 
Family PastpHa@iDé. 
Genus LEPTOCHELA. 


1. Leptochela robusta Stimpson, 1860. 


Proc. Acad. Philadelphia, 1860, p. 43. de Man, Abh. Senckenb. Ges. xxv. 11. 
p. 902 (1902). 
Haddumati Atoll, Maldive Is. 


Superfamily PANDALOIDA 
Family PANDALIDA. 
Subfamily PANDALINA. 
Genus PANDALUS. 
Subgenus PLESIONIKA. 
2. Pandalus (Plesionka) gracilis Borradaile, 1915 (Plate 58, fig. 1). 
Ann. Mag. Nat. Hist. (8), xv. p. 208. 


Diagnosis: The rostrum is long (its tip appears to be damaged in the specimen) 
and of even width throughout. At its base two strong teeth stand above the eye on 
a erest which extends backwards over the first half of the carapace. The rostrum itself 
is gently upeurved from the base; about the hindermost third of its length is unarmed; 
the rest bears below a series of small sharp teeth, set wider apart towards the tip 
of the organ, and above eight minute spinules, widely set, the last two being some- 
what larger than the rest. The eye is wider than its stalk, and has a distinct ocellus. 
The stalk of the antennule is less than the length of the antennal scale. The flagella 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN 399 


of antennule and antenna are broken short in the specimen. The third mawilliped 
reaches more than half-way along the antennal scale. The first leg is simple and 
sparsely hairy and barely falls short of the end of the third maxilliped. The second 
leg reaches the end of the first; its wrist is nine-jointed, the first five joints being 
longer and less distinct than the rest. The legs of the last three pairs are missing 
from the specimen. The third abdominal tergum is convex behind, but not produced 
into a spine. The sixth abdominal segment is longer than the fourth and fifth together. 
The telson is missing from the specimen. 

Length of specimen from tip of rostrum to tip of uropods 49 mm. 

The species appears to be related to P. martius A. M.-Edwards, 1883, and P. ensis, 
(A. M.-Edwards), 1881. 

A single female specimen was taken at a depth of 200 fathoms in the Western 
Indian Ocean. 


Genus Hetrrrocarpus. 

3. Heterocarpus umcarmatus Borradaile, 1915 (Plate 58, fig. 2). 

Ann. Mag. Nat. Hist. (8), xv. p. 208. 

A specimen taken in 637—665 fathoms near Providence Island is closely related 
to H. longirostris MacGilchrist, 1905 [Ann. Mag. N. H. (8), xv. p. 237] but distinguished 
by the loss of the hinder three-quarters of the antennal carina, of which the forepart 
is present though not sharply formed. The specimen is a good deal damaged, and may 
have presented other differences from H. longirostris. It measures 92 mm. to the tip 
of the rostrum, which is broken. Provisionally, at least, it deserves specific recognition. 


4. Heterocarpus affinis Borradaile, 1915. 
Ann. Mag. Nat. Hist. (8), xv. p. 208. 


Specimens taken in 300—500 fathoms near Saya de Malha are nearly related to 
Hf, alphonsi Bate, 1888, but show the following points of difference: (1) the rostrwm 
is much more strongly upeurved, and has three teeth on the carapace behind the 
orbit, (2) there are fewer joints in the wrists of the second legs (26 and 9, as against 
40 and 11), (3) the walking legs considerably outreach the antennal scale. The telson 
bears five pairs of spines on the dorsal surface and three pairs at the end. The longest 
specimen is 12cm. in length, including the rostrum. 


Subfamily THALASSOCARIN A. 


Genus THALASSOCARIS. 
5. Thalassocaris crinitus (Dana), 1852. 
Regulus crinitus, Dana, U.S. Explor. Exped. Crust i. p- 599; Atlas, Pl. 39, fic. 6. 


The rostrum of this species may be somewhat longer than in the specimen figured 
by Dana, and the rostral formula varies within the limits g—10 

Specimens were taken in various depths down to 80 fathoms at Amirante I. 
Cargados Carajos, the Maldives, Saya de Malha, and the Seychelles. 


400 PERCY SLADEN TRUST EXPEDITION 


6. Thalassocaris affins Borradaile, 1915 (Plate 58, fig. 4). 

Ann. Mag. Nat. Hist. (8), xv. p. 208. 

Diagnosis: The species is closely related to T. lucidus (Dana), 1852, but differs 
from it in the following points: the rostrum is less strongly upcurved, and is usually 
rather shorter. Its formula is 5=?. 
thorns, stronger than in 7. /wcidus and set wider apart. The hands of the second legs 
are of the shape of 7. crinitus—rectangular, with widely gaping fingers and a strong 


The “teeth” on the antennal scale are hooked 


basal knob on the fixed finger, and a stout tooth on the moveable finger. The merus 
in the walking legs bears the stout thorns present in 7. crinitus but neither mentioned 
nor figured by Dana for 7. luezdus. 

Length of the longest specimen 25 mm. 

Many of the specimens are mature, and some of them are considerably larger than 
the smaller specimens of 7. criutus. T. affinis appears to be intermediate between 
T. lucidus and T. crinitus. It was taken at various depths down to 26 fathoms in the 
Maldives and at Saya de Malha. 


7. Thalassocaris maldiwwensis Borradaile, 1915 (Plate 58, fig. 5). 

Ann. Mag. Nat. Hist. (8), xv. p. 208. 

Diagnosis: The body is stout and compressed. The rostrum outreaches the antennal 
scale, at first descends and then is horizontal, and has the formula *, two of the dorsal 
teeth standing behind the orbit with sometimes a vestigial tooth behind the first of 
them. Suborbital and antennal, but no supraorbital or hepatic spines are present. 
The antennal stalk is slightly outreached by the antennular, and extends to about one- 
third of the length of the antennal scale, which is without teeth on the outer edge. 
The third macilliped nearly reaches the end of the antennal scale. The jirst leg ends 
at the middle of the last joint of the third maxilliped. The second leg outreaches the 
third maxilliped and has a small chela of simple form, the wrist about as long as the 
arm, and the fingers shorter than the palm. The legs of the last three pars are sparsely 
hairy and of moderate length and stoutness. The abdominal pleura are sharp pointed. 
The third abdominal somite has no spine on its hinder edge. The telson is longer 
than the uropods and has four pairs of dorsal spines and two fixed terminal spines. 

Length of the longest specimen 19 mm. 

Peculiar features of the species are the absence of the spine on the third abdominal 
segment and of the supraorbital spines, and the feebleness of the second leg. The first 
maxilliped differs greatly from that figured by Dana for J. Jucidus, but the latter was 
probably drawn from a damaged specimen. 

Specimens were taken in Suvadiva, Mulaku, Haddumati, and 8. Nilandu Atolls in 
the Maldives. 


Superfamily PaLaMonorpa. 
Family ALPHEIDA. 


The Alpheid Prawns collected by Professor Gardiner’s Western Indian Ocean 
Expedition form the subject of a separate report by Prof. Coutiére in this volume. 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN 401 


Family Hippotytipa. 
Genus SARON. 


8. Saron marmoratus (Olivier), 1811. 

Palemon marmoratus, Olivier, Encycel. vii. (fide H. M.-Edwards). 

ippolyte gibberosus and H. marmoratus, H. M.-Edwards, H. N. Crust. ii. pp. 378, 
379, Pl. 25, fig. 8 (1837). 

IMippolyte gubberosa and H. marmorata, de Man, Arch. Naturg. lili. 1. p. 533 (1888). 

: Saron marmoratus, Borradaile, Proc. Zool. Soc. Lond., 1898, p. 1009; Kemp, Ree. 
Ind. Mus. x. p. 84 (1914). 

The rest of the synonomy of S. marmoratus will be found in the last two papers 
quoted. 

Kemp (loc. cit.) has shown that the relation of the two forms of the male of this 
species is not, as I had formerly suggested, a true dimorphism, since the individuals 
do not fall into two well-defined groups but form a graded series. This does not appear 
either in the present specimens, which are all of the gibberosus type, or in those which 
served as the basis of my earlier remarks upon the subject, in which there were two 
sharply-defined groups. I have, however, no doubt that Mr Kemp’s solution of the 
question is the correct one, and that the marmoratus characters are gradually assumed 
by the males with age. 

The collection contains 28 specimens from Salomon, Amirante, the Maldives, Minikoi, 
and Coetivy, Seychelles. 


9. Saron neglectus de Man, 1902. 


Abh. Senckenb. Ges. xxv. p. 854, Pl. 26, fig. 58. Kemp, Rec. Ind. Mus. x. p. 87 (1914). 
One specimen from Egmont Reef, Seychelles. 


Genus Licur. 

10. Lngur wee (Borradaile), 1902. 

Parhippolyte uvee, Borradaile, Willey’s Zool. Results, p- 414, Pl. 38, a ll a—g. 

Ingur uvee, Kemp, Rec. Ind. Mus. x. p. 123 (1914). 

The original description of this species omits the fact that the meropodites of 
the legs of the second pair, and the propodites of those of the last three pairs, are 
multiarticulate. 

Numerous specimens from Aldabra. 


Genus THOR. 
11. Thor maldivensis Borradaile, 1915 (Plate 58, fig. 6). 
Ann. Mag. Nat. Hist. (8), xv. p. 208. 


Diagnosis: The body is moderately compressed. The carapace is short, not keeled, 
and has large supraorbital and small antennal spines. The third abdominal segment 
projects in the middle behind. The rostrum is very short, not reaching the end of the 

SECOND SERIES—ZOOLOGY, VOL. XVII. 51 


402 PERCY SLADEN TRUST EXPEDITION 


first joint of the antennular stalk, its tip is simple, and it bears one tooth above at 
the level of the eye and none below. The antennular stalk is a little shorter than the 
antennal, its last two joints are very short and broad, and it bears a strong spine on 
each joint, those on the first two joints being sharp and external and the third broad, 
dorsal, and sutured. The antennal stalk is nearly half the length of the scale. The 
latter reaches nearly as far as the stouter flagellum of the antennule, has a convex. 
inner edge, a straight outer edge, and a nc end, and bears on the outer side a 
distal spine, which does not project as far as the end. The thard maauilliped is strong, 
and outreaches the antennal scale in the male by more and in the female by less than 
the whole of the last joint. The last and antepenultimate joints are subequal, each more 
than twice as long as the penultimate joint, the last joint is spinous, and all the joints 
are hairy. The first leg in the female is stout, simple, and shorter than the third 
maxilliped. In the male, it is as long as the body, granulate, and stout, but with 
the chela no stouter than the rest of the limb, the arm and hand are subequal, and the 
fingers about one quarter the length of the palm, on which they are bent inwards 
at an obtuse angle, each bearing a low tooth. The second leg has the wrist 5-jointed, 
with the second jomt much longer than any of the others and showing an indistinct 
ring near its proximal end. The legs of the last three pairs are alike in the two 
sexes, of medium length, with a blunt spine at the end of the carpopodite, a row of 
spinules under the propodite, and the dactyle short, stout, ending in a slender claw, and 
bearing below several moveable spinules of which the last is longer than the end claw. 
The te/son is shorter than the uropods, narrow, and tapers to an obtusely triangular 
end bearing six spines, of which the intermediate pair are the longest. 

Length of longest specimen 13 mm. 

The genus Thor has hitherto contained only one known species, 7. paschalis 
Heller 1861 (7. floridanus Kingsley). The present species has all the characters of 
the genus, including those of the mandible, but the great claw of the male is a new 
feature. The principal differences from 7. paschalis are presented by the supraorbital 
spine, the rostrum, the first and third legs of the male, and the proportions of the 
joints of the second leg. 

Specimens were taken at Malé Atoll in the Maldives, in Minikoi, and at Salomon ie 


Genus TozEUMA. 


12. Tozeuma armatum Paulson, 1875. 

Red Sea Crustacea, p. 99, Pl. 15. figs. 2a—o. Kemp, Ree. Ind. Mus. x. p. 106 (1914). 

Specimens were taken in various depths in the Maldives, the Seychelles, and 
Cargados Carajos. 


Genus Lysmara. 
13. Lysmata affinis Borradaile, 1915. 
Ann. Mag. Nat. Hist. (8) xv. p. 209. 
Diagnosis: a Lysmata closely related to L. seticauda (Risso), 1816, and to L. chaltona 
Kemp, 1914, but distinguishable from them and from the other species of the genus 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN 403 


by the following combination of features: the rostrum reaches well beyond the eyes, but yar 
ends just before the middle of the second joint in the antennular stalks. Its formula eg oS 


is-5=8, the lower teeth being small, but larger than in Z. chiltoni, and the hinder of zy 
them standing below the last upper tooth. The pterygostomial angle is rectangular ep ft, 3% 9 


and usually produced into a spinule. The thick flagellum of the antennule is fused 
to the slender one for half its own length. The antennal scale curves gently outwards 
and narrows slightly towards the end, which is truncate and distinctly outreached by 
the distal spine, the first leg slightly outreaches the scale but falls considerably short 
of the end of the third maxilliped. The wrist joint is as long as, usually a trifle 
longer than, the chela. The second legs are equal or unequal. The longer of the pair 
outreaches the antennal scale by rather less than the whole wrist. The latter has 
24—25 joints. There are double tips to the fingers. The walking legs have spinules 
below the meropodites. The first outreaches the antennal scale by the dactyle and 
nearly the whole propodite, the second by the propodite only, and the third falls a 
little short of the end of the scale. 

Length of the longest specimen 31 mm. 

Specimens were taken in Minikoi, Peros Banhos, Salomon, and the Seychelles. 


Genus H1iproLysMATA. 


14. Hippolysmata vitatta Stimpson, 1860. 


Proc. Acad. Philadelphia, 1860, p. 26. de Man, Trans. Linn. Soc. London, (2) Zool. 
ix. p. 423 (1907). Kemp, Rec. Ind. Mus. x. p. 113 (1914). 

Specimens were taken at Cargados Carajos in 30 fathoms, and in the Seychelles 
in 34 fathoms. 


15. Hippolysmata kiikenthali (de Man), 1902. 


Merhippolyte orventalis, de Man (nec Bate), Weber's Zool. Ergebn. Reise Ost-Ind. 
ile JO, 207. 

ippolyte kiikenthals, de Man, Abh. Senckenb. Ges. xxv. p. 849, Pl. 26, fig. 56 (1902). 

Hippolysmata kiikenthali, de Man, Trans. Linn. Soc. London, (2) Zool. ix. p. 426 
(1907). Kemp, Rec. Ind. Mus. x. p. 115 (1914). 

Ten specimens were taken on Egmont Reef, Seychelles. Each has a single tooth on 
the underside of the rostrum, and 14 or 15 joints in the wrists of the second legs. 


Genus LYSMATELLA. 


Borradaile, Ann. Mag. Nat. Hist. (8), xv. p. 206 (1915). 


The collection contains three specimens of a new species which would, have to 
be placed in the genus Hippolysmata, were it not for the absence of epipodites from 
the legs. In view of this somewhat important difference I have thought it best to 
establish for this species a new genus Lysmatella. 

51—2 


404 PERCY SLADEN TRUST EXPEDITION 


16. Lysmatella prima Borradaile, 1915 (Plate 58, fig. 7). 
Ann. Mag. Nat. Hist. (8), xv. p. 209. 


Diagnosis: The body is compressed, the carapace of a good length, keeled in the 
forepart and provided with a strong antennal and a small pterygostomian spine. The 
rostrum is straight at first but gently upcurved towards the tip, outreaches the 
antennular stalk, and has the formula *', the first tooth standing detached on the 
carapace, the ventral teeth smaller than the dorsal, and all the teeth sloping very 


sharply forwards. The rostrum becomes relatively longer and more strongly curved as 
the individuals increase in size. The antennule has a long, slender stalk, and the 
stylocerite short, wide and curved. The antennal scale reaches the end of the penulti- 
mate joint of the antennular stalk, and is narrow, with nearly parallel sides and a broad, 
rounded end. The third mawilliped is as stout as the first leg, in which the hand and 
arm are subequal, the wrist a little shorter than either, and the fingers gape somewhat 
widely. In the second leg the wrist has 20—22 joints, of which the last is longer 
than any of the others. The dactyles of the walking legs are provided, besides the 
end-claw, with three moveable spines on the lower side, and of these the third is larger 
than the end-claw. The te/son is barely shorter than the uropods, bears two pairs of 
moveable spines above, and is fringed with long, stout hairs. 

Length of the longest specimen 19 mm. 

Specimens were taken in Haddumati, Mulaku, and South Nilandu Atolls, Maldives. 


Family PALAZMONID. 
Subfamily PALaMoniIn &. 


Genus LEANDER. 


17. Leander debilis (Dana), 1852. 


Palaemon debilis, Dana, U.S. Explor. Exped. xiii. 1. (Crust.), p. 585, Pl. 38, figs. 6—7. 
de Man, Abh. Senckenb. Ges. xxv. ur. p. 808 (1902). 

The collection contains a large number of specimens of L. debilis from Aldabra 
which show the great variability which characterizes this species. The rostrum varies 
in length, shape, and dentation, It may be as long as, shorter than, or longer than 
the antennal scale. Its curves differ considerably in the specimens. Its formula in 
=e 
always present near the tip. The size, shape, and spacing of the teeth varies. The 
wrist of the second leg varies in length, but usually falls a little short of the end 
of the antennal scale. The antennule agrees with de Man’s description. 


these examples is most commonly 3, including the small tooth which is nearly 


There are also a number of specimens from a “barachois” in Diego Garcia. 
These are less variable than the Aldabra specimens. The rostral formula is 43, 
usually €. The average size is less than that of the Aldabra specimens, the largest 
Specimen measuring 28 mm. in length. It is quite possible that we have here two 


distinct but closely allied species, such as the LZ. debilis and L. gardineri found in 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN A05 . 


Miladumadulu Atoll (Gardiner’s Fauna of the Maldives, i. p. 98). In the case of the 
Miladumadulu species the habits and appearance of the prawns in the living state 
enabled the two forms to be distinguished with certainty. Possibly similar information 
would avail in the same way here. 


Subfamily Pontonimna. 


Many species of this subfamily are represented in the collection. They are 
enumerated in a separate article in this volume (pp. 323—396). 


Superfamily CRANGONOIDA. 


The bounds of this somewhat miscellaneous group must be enlarged to admit 
the genera Anchistioides, Amphipalemon, and Hymenocera, which, in our present state 
of knowledge appear more closely related here than elsewhere. This addition involves 
two concessions in the definition of the group: (1) if Anchistioides and Amphipalemon 
are to be admitted, it can no longer be stated that the mandible is always without 
incisor process, (2) the inclusion of HMymenocera makes it necessary to allow the 
persistence of a small representative of the second lobe of the maxilla. 


Family ANCHISTIOIDIDA. 


Borradaile, Ann. Mag. Nat. Hist. (8), xv. p. 205 (1915). 


In 1899, reporting on the Macrura brought by Dr A. Willey from the South Seas, 
I established a genus Palemonopsis for a new prawn, P. willey:, taken in New Britain. 
In 1901, Nobili proposed to change the name of this genus to Amphipalemon, on the 
ground that Palemonopsis was preoccupied, having been used by Stimpson as a 
synonymn of Palemonetes. Both by Nobili and by myself the genus has been regarded 
as belonging to the Palzemonide. 

The present collection contains two specimens, each representing a new species 
of Amphipalemon. Examination of these, and re-examination of Dr Willey’s original 
specimen, convinces me (1) that the characters of the new genus are so distinctive 
that it must become the type of a new family, (2) that the affinities of this family 
are at least as much with the Crangonoida as with the Palemonoida, (3) that the 
genus Anchistroides founded by Paulson in 1875 is closely related to Amphipalemon. 


The characters of the Anchistioidide may be stated as follows; 
(1) The body retains the typical caridoid facies. - 
(2) The rostrum is well developed, compressed, and toothed. 


(3) There is no supraorbital spine, but there may be a blunt knob nearly in 
the same position. 


(4) The outer flagellum of the antennula bears at the base a short, thick, 
accessory flagellum. The stylocerite is inconspicuous or wanting. 


406 PERCY SLADEN TRUST EXPEDITION 


(5) The antenna has a broad scale, truncate at the end, and is without a spine 
on its basipodite. 

(6) The mandible is deeply cleft, the molar process has a broad end, surrounded 
with stout teeth, the incisor process is coarsely serrate, and there is no palp. 

(7) The inner lacinia of the mamillule is round-ended, and not curved towards 
the outer lacinia. 


(8) ‘The laciniae of the maailla are aborted. 


(9) The exopodite of the first mailliped has no flagellum, and its epipod 
is simple. : 

(10) The end-joint of the second mawilliped is applied to the inner edge of 
the recurved propodite. The epipodite of this limb is discoidal. 

(11) The therd mamlliped is slender, and has no exopodite, but a short broad 
epipodite. 

(12) The legs of the first two pairs are without exopodites, chelate, with simple 
wrists, and subsimilar, but the second pair is longer and stouter than the first. 

(13) The legs of the last three pairs are alike, without exopodites and adapted 
for walking. 

(14) The first five abdominal limbs have a well-developed appendix interna, and 
in the first of them, especially in that of the male, the endopodite is small, so that it 
forms with the appendix a biramous organ. 

(15) The telson bears two or three pairs of spines above, and at the end one 
strong pair of spines and several stout bristles, of which one or a pair are feathered. 

(16) The gills comprise pleurobranchs for the legs and an arthrobranch for the 
third maxilhped. There are epipodites on the maxillipeds only. 


The family appears to be transitional between the Paleemonoida and the Crangonoida. 


The genera of Anchistioididee may be distinguished as follows: 
I. Without a blunt process of the carapace behind the eye. Scaphocerite present. 
The end of the telson bears, among others, one unfeathered bristle on each side and a pair 


of small lateral spines. 
Anchistioides Paulson, 1875. 


II. <A blunt process of the carapace behind the eye. No scaphocerite. The end 
of the telson bears, among others, more than one unfeathered bristle on each side at 
the end of the telson, but no small lateral spine. 

Amphipalemon Nobili, 1901. 


Genus AMPHIPALAMON. 


The species of Amphipalemon are closely similar in most respects, but may be 
distinguished as follows: 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN 407 


I. Three pairs of spines on the dorsal side of the telson, and a median feathered 
bristle at the hinder end. Rostrum does not reach end of antennal scale [and is very 


deep and straight at base]. 
A. willeyi (Borradaile), 1899. 


II. Two pairs of spines on the dorsal side of the telson, and a pair of feathered 
bristles at the hinder end. Rostrum at least reaches end of antennal scale. 


A. Rostrum very deep, straight at base. 
A. gardineri Borradaile, 1915. 


B. Rostrum not very deep, arched at base. 
A. cooperi Borradaile, 1915. 


18. Amphipalemon gardineri Borradaile, 1915 (Plate 59, fig. 14). 
Ann. Mag. Nat. Hist. (8), xv. p. 209. 


The most important difference between this species and A. willeyi lies in the 
arrangement of the spines of the telson. In the New Britain species, the two anterior 
pairs of spines on the dorsal side of this organ lie in its front half. In A. gardineri the 
second of them lies just behind the middle of the telson. In A. willeyi (Plate 59, fig. 13), 
the two small lateral spines of Anchastioides, which are also found in so many other Carides, 
have migrated to the dorsal surface of the telson. In A. gardineri they are altogether 
wanting. In the latter species, the adjoining spines, which thus become lateral, are 
longer than in A. willeyi. The pair of feathered bristles found in Anchistioides are 
present also in A. gardinerv; in A. willey: they are absent, but there is present 
a single median feathered bristle, shorter than those of A. gardineri and widened 
at the base. 

Lastly, in A. wlley: there are a number of unfeathered bristles, of which three 
on each side are somewhat shorter than the rest. In A. gardineri only two such 
bristles are present. 

Less important differences, which may quite possibly prove not to be constant, are: 
(1) that the rostrum has the formula $ and outreaches the antennal scale, and (2) that 
the first leg does not reach the end of the antennal scale, and (3) that the second 
leg outreaches the scale by only half the length of the palm, and its meropodite 
is relatively shorter. 

The specimen, which measures 30 mm. in length, was taken in N. Malé Atoll. 


19. Amphipalemon cooperi Borradaile, 1915. 
Ann. Mag. Nat. Hist. (8), xv. p. 209. 


This species very closely resembles the preceding, but differs from it in the following 
points: (1) the rostrum has the formula &, barely outreaches the antennal scale, and 
is arched at the base and decidedly less deep, (2) the first leg reaches the end of the 
antennal scale, (3) the second leg outreaches the scale by the whole hand, (4) the 


408 PERCY SLADEN TRUST EXPEDITION 


meropodite of this leg is longer than in either of the other species, equalling £ of the 
length of the hand, (5) the hinder pair of the dorsal spines of the telson lies farther back 
than in A. gardinerz. 

Length of single specimen 15 mm. 

S. Nilandu Atoll. 


Family GNATHOPHYLLID. 


Professsor Gardiner’s collection contains specimens of members of the genera G'natho- 
phyllum and Hymenocera and also of the very interesting species described by Dr Balss 
as Hymenocera ceratophthalma. An examination of this material makes it quite clear 
that the species in question are all members of a single family. The principal characters 
of this family are as follows: 


(1) 
(2) The rostrwm is compressed and dentate. 
(3) | 
(4) The antennule has a well-developed stylocerite, and the outer flagellum thick 
at the base and cleft for a very short distance at the end of the thick part. 


The body retains the typical caridoid facies, but is rather heavily built. 


The antennal spine alone remains on the carapace. 


(5) In the antenna the scale may be broad or rather narrow, and is rounded at 
the end, and the spine of the basipodite is short or absent. 

(6) The mandible is simple, palpless, slender, and curved. 

(7) The inner lacinia of ‘the maxillule is pointed and curved towards the outer 
lacinia. , 

(8) The first lobe of the maailla is totally lost and the second lobe is either lost 
or very small but still obscurely double. 

(9) The first maxilliped has a flagellum and the outer border of its epipodite is 
notched, but not deeply. 

(10) The end-joint of the second maailliped is applied to the inner edge of the 
recurved propodite. 

(11) The third masilliped has an exopodite, a simple epipodite, and an endo- 
podite of four joints, some or all of which are greatly broadened. 

(12) The legs of the first two pairs are chelate, without exopodites and with 
simple wrists. The two pairs are more or less dissimilar, and the second pair is the 
larger. 

(13) The legs of the last three pairs are alike, without exopodites, and adapted 
for walking. 

(14) The second to fifth abdominal limbs have a well-developed appendix interna. 

(15) The telson bears two pairs of spines at the sides, and at the end an outer 
short and an inner longer pair of spines, a submedian pair of slender feathered spines, 
and a median pointed projection. 


=o 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN 409 


(16) The gills comprise pleurobranchs for the legs, an arthrobranch for the third 
maxilliped, and in Hymenocera the vestige of a pleurobranch for the latter limb. There 
are epipodites on the maxillipeds only. 


A key to the genera of Gnathophyllide: 


I. Ischium of third maxilliped narrow and moveably sutured to merus. Rostrum 


of a good length. 
A. Mandible flattened. Lobe of maxilla present. Outer flagellum of antennule 
leaf-like, 
Hymenocera Latreille, 1829. 


B. Mandible subcylindrical. Lobe of maxilla lost. Outer flagellum of antennule 
normal. 


Phyllognathia Borradaile, 1915. 


II. Ischium of third maxilliped broad and marked off from merus by a notch 
only. Rostrum short. 
[Mandible, maxilla and antennule as in Phyllognathia. | 
Gnathophyllum Latreille, 1829. 


Genus GNATHOPHYLLUM. 


20. Gnathophyllum fasciolatum Stimpson, 1860 (Plate 59, fig. 8). 

Proe. ‘Ac. Philadelphia, 1860, p. 28. de Man, Arch. Naturg. liii. 1. p. 496 (1888). 
Abh. Senckenb. Ges. xxv. 11. p. 762 (1902). 

Gnathophyllum zebra, Richters, Meeresf. Mauritius, p. 161, Pl. 17, figs. 18—20 
and 22 (1880). 

All the specimens are completely bleached. A small antennal spine which is present 
in my specimens is not mentioned in any description of the species. 

According to Nobili (Mem. Ac. Torino (2), lvii. p. 305), this species is identical with 
G. americanus, and should be known by that name. 

Specimens were taken at Minikoi, Salomon I., and Egmont Reef, Seychelles. 


Genus PHYLLOGNATHIA. 


21. Phyllognathia ceratophthalma (Balss), 1914 (Plate 59, fig. 9). 
Hymenocera ceratophthalma, Balss, Abh. k. Bayer. Ak. Wiss. Suppl. Bd. 1, 10, 
“ p. 54 (1914). 
Phyllognathia ceratophthalma, Borradaile, Ann. Mag. Nat. Hist. (8), xv. p. 206 (1915). 
Dr Balss has kindly communicated to me his opinion that this species is in reality 
a Gnathophyllum. Its transference to the Gnathophyllide will probably meet with 
approval, but in including it in Gnathophyllum some rather considerable differences 
between its mouth-parts and those of other species of the genus have to be discounted. 
In the third maxilliped of Gnathophyllum, the ischiopodite is fused with the meropodite, 
SECOND SERIES—ZOOLOGY, VOL. XVII. 52 


410 PERCY SLADEN TRUST EXPEDITION 


though, by an exception to the rule in Carides, the boundary between these two 
joimts is marked by a notch. In H. ceratophthalma the ischiopodite is distinct and 
even moveably articulated with the meropodite, a condition which I cannot remember 
to occur in any other Caridean except Hymenocera. In the second maxilliped of 
G. fasciolatum (the only species I have been able to examine), the last two joints 
form a very large scythe-shaped organ. In H. ceratophthalma they are rather smaller 
than in most Carides. Finally, in the maxillule of G. fasciolatum the outer lacinia 
is greatly enlarged, while in H. ceratophthalma it is of quite normal dimensions. In 
the characters of the second maxilla and mandible, however, H. ceratophthalma agrees 
very well with Gnathophyllum. Both the lobes of the maxilla are lost, and the mandible 
is simple, without palp, and of a slender, curved shape, with a blunt end provided 
with small teeth. I would suggest that it is advisable to establish a new genus of 
Gnathophyllidee for Dr Balss's species. Phyllognathia would be an appropriate name 
for this species, which undoubtedly links Hymenocera with Ginathophyllum. 

In Dr Balss’s specimen the smaller leg of the second pair and the tip of the rostrum 
were missing. My specimen enables me to state that the rostrum slightly outreaches 
the antennular stalks and has the formula 7, and that the smaller leg of the second 
pair has the same features as the larger but a somewhat narrower palm and less 
pronounced serration of the moveable finger. The larger leg of this pair is missing 
in my specimen; which was taken in 8. Nilandu Atoll. 


Genus HyMENOCERA. 


The mandible of Hymenocera is of the same shape as that of Gnathophyllum save 
that it is flattened and ends in a serrated cutting edge, as though it represented the 
incisor process of the complete malacostracan mandible. In point of fact, however, there 
can be little doubt that it is really the molar process. The maxilla has a small outer 
lobe, which, however, retains traces of a notch. The telson is of precisely the same type as 
that of Gnathophyllum. The telson of Crangon is of the same type but more elongate. 
That of the Processidze is more like those of the Pontoniinz. The telson of Glypho- 
crangon is a much modified structure which has lost all spines. 


22. Hymenocera elegans Heller, 1861 (Plate 59, fig. 10). 


Verh. zool. bot. Ges. Wien, xi. p. 25; Sitzber. Ak. Wiss. Wien, xliv. 1. p. 264, 
Pl. 3, figs. 9—14 (1861). Ortmann, Speng. Zool. Jahrb. Syst. v. p. 511, Pl. 37, fig. 11. 
A specimen was taken in Coetivy, Seychelles. 


Family PRocessip#. 
Genus PRocEssa. 
23. ? Processa processa (Bate), 1888. 
Nika processa, Bate, “Challenger” Macrura, p. 527, Pl. 95 (1883). 


A specimen from the Seychelles which probably belonged to this species was 
unfortunately destroyed by an accident while it was under examination. 


BORRADAILE—ON CARIDES FROM THE WESTERN INDIAN OCEAN All 


Genus NIKOIDEs. 


24. Nikoides maldivensis Borradaile, 1915 (Plate 58, fig. 11). 
Ann. Mag. Nat. Hist. (8), xv. p. 209. 


A single specimen, taken at the Amirante Is., is closely related to NV. danae Pauls., 
1875 (Res. Crust. Red Sea i. p. 98, Pl. 14, fig. 5), but differs from it in the following 
points : (1) the rostrum is of a different shape, the dorsal tooth being larger and placed 
much farther back, (2) the exopodite of the first leg is relatively shorter, reaching 
only half-way along the merus, (3) the carpopodites of the legs of the first pair are 
of equal length, (4) there are no spines on the ischium or merus in the legs of the 
last three pairs. The specimen measured 24 mm. in length. To the differences between 
Processa and Nikoides stated by Nobili, there should be added the presence in the latter 
of a single dorsal tooth on the rostrum. 


Family GLYPHOCRANGONIDS. 
Genus GLYPHOCRANGON. 
Subgenus PLASTOCRANGON. 


25. Glyphocrangon (Plastocrangon) ceca Wood-Mason, 1891. 

Ann. Mag. Nat. Hist. Nov. 1891, p. 358; IIl. Zool. “ Investigator” Crust. Pl. 3, 
ine, Il, 

A single specimen taken in 300—500 fathoms at Saya de Malha. 


Family CRANGONID. 
Genus ANGEON. 


26. Algeon rugulosus Borradaile, 1915 (Plate 59, fig. 12). 
Ann. Mag. Nat. Hist. (8) xv. p. 210. 


One damaged specimen, taken in Haddumati Atoll, Maldives, is related to #. 
medius (Alc. and And.), 1899, but differs from it in that: (1) the beading of the 
ridges of the carapace is much coarser, (2) there is no tooth on either side of the 
base of the rostrum, (3) the large spine near the pterygostomial angle of the carapace 
stands at the end of the supramarginal, not at that of the lateral ridge, (4) the 
dactyles of the first two walking legs are longer. The length of the specimen is 
16 mm. 


Fig. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


Fig. 
Fig. 
Fig. 


PERCY SLADEN TRUST EXPEDITION 


EXPLANATION OF PLATES 58, 59. 


1. Pandalus (Plesiomka) gracilis Borradaile, 1915 (Plates 58, 59). a, side view, x 24; b, end of 
second leg, x 12. 

2. Heterocarpus unicarinatus Borradaile, 1915 (Plate 58). Side view, x 14. 

Heterocarpus affinis Borradaile, 1915 (Plate 58). Side view, nat. size. 

Thalassocaris affinis Borradaile, 1915 (Plate 58). Side view, x4. 

Thalassocaris maldivensis Borradaile, 1915 (Plate 58). Side view, x 4. 

Thor maldivensis Borradaile, 1915 (Plate 58). Side view, x 7. 

Lysmatella prima Borradaile, 1915 (Plate 58). Side view, x 4. 

Gnathophyllum fasciolatum Stimpson, 1860 (Plate 59). -a, third maxilliped, x 16; 6, second 
nexellbiged x16; ¢, maxillule, x 16; d, mandible, x 25. 

9. Phyllognathia ceratophthalma (Balss), 1914 (Plate 59). a, third maxilliped, x 16; 6, second 
maxilliped, x 16; c, first maxilliped, x 16; d, maxilla, x 16. 

10. Hymenocera elegans Heller, 1861 (Plate 59). a, third maxilliped, x 4; 6, second maxilliped, x 16; 
c, first maxilliped, x16; d, maxilla, x16; e, maxillule, x 16; f, mandible, x 25. 

11. Nikoides maldivensis Borradaile, 1915 (Plate 58). Side view, x 3. 

12. Ageon rugulosus Borradaile, 1915 (Plate 59). Side view, x 4. 

13. Amphipalemon willey: Borradaile, 1899 (Plate 59). Telson, x 35. 

14. Amphipaleemon gardinerr Borradaile, 1915 (Plate 59). Telson, x 35. 


Go I oe 90 | 


Percy Suapen Trust EXPEDITION. 
(BorRADAILE) 


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CARIDES FROM THE WESTERN INDIAN OCEAN 


Percy SLADEN Trust EXPEDITION. 
(BorraDAILr) 


TRAINS. ILININ, SOC. SIE, 2, ZOOL, WOlL, XW, PIL. SY 


CARIDES FROM THE WESTERN INDIAN OCEAN 


No. X.—LES ESPECES D'ALPHEIDZ RAPPORTEES DE LOCEAN INDIEN 
PAR M. J. STANLEY GARDINER. 


Par M. te Proresseur H. CoutiiRe. 
(Communiqué PAR M. LE ProressEuR J. STANLEY GarpDINER, M.A., F.R.S., F.L.S.) 


(Planches 60—64.) 
Lu le 20 Juin, 1918. 

Ls espéces nouvelles d’Alpheidz rapportées par le Percy Sladen Trust Expédition ont 
été décrites en 1908, dans le B. de la Soc. Philomathique. Depuis, le trés important mémoire 
de de Man, consacré aux récoltes du Siboga, est venu montrer, surabondamment, quel fonds 
on peut faire sur les Alpheidee pour caractériser la faune corallienne d'une région. Et on 
peut tenir pour certain que la liste est loin d’étre close; elle s'allongera encore d’une 
centaine de formes, au moins, lorsque toute la ceinture corallienne du globe sera aussi bien 
explorée que |’ Indo-Pacifique du Seelark et du Siboga. 

Le beau travail de de Man, malheureusement non accompagné des figures si in- 
dispensables, m’a obligé 4 revoir certaines déterminations, en particulier celle du S. neomeris, 
mais, dans l’ensemble, les espéces communes aux deux régions n'ont pas augmenté de nombre, 
. ou & peu prés. J’ai donc pu me borner & reproduire, presque textuellement, mon travail de 
1908, en y intercalant les espéces deja connues, et en y ajoutant de nombreuses figures. Ce 
mémoire est, en somme, la suite de celui que j'ai publié sur les Alpheidze des Maldives et 
Laquedives, aussi ai-je pu étre trés sobre d’indications bibliographiques. Je n’ai pas non 
plus reproduit les indications des groupes d’espéces, aujourd'hui classiques. II] serait beau- 
coup plus intéressant de pouvoir dire pourquoi il faut distinguer tant de formes spécifiques 
dans cette famille & éthologie si spéciale, mais les raisons que jen apergois sont beaucoup 


trop spéculatives pour que je me risque & les donner ici. 
Genre Automate de Man. 
1. A. salomoni H. Cout., Bull. Soc. Phil. 1908, p. 2. (Plate 60, fig. 1.) 
L’unique spécimen est une ? ovée mesurant 17°5 mm. 
Chagos, Salomon Island. 
Genre Athanas Leach. 
2. A. djiboutensis H. Cout., Alph. M. et L. p. 856, fig. 129. 
Chagos, Egmont récif. 
Genre Arete Stimpson. 
3. A. indicus H. Cout., Alph. M. et L. p. 863, figs. 134—35. 
Seychelles, Praslin reef. 
4. A. equalis H. Cout., Alph. M. et L. p. 869, fig. 138. 


Coetivy, Amirante Bank, 25—80 brasses. 
SECOND SERIES—ZOOLOGY, VOL. XVII, 53 


414 PERCY SLADEN TRUST EXPEDITION 


Genre Alpheopsis H. Cout. 

5. A. fissipes H. Cout., Bull. Soc. Phil. 1908, p. 3. (Plate 60, fig. 3.) 

L’unique spécimen de VA. fissipes est une ? de 6°5 mm. de longueur, portant 7 ceufs 
seulement, de taille comparable & ceux de I’A. equalis. 

Providence, D. 4 (50—78 brasses). 

6. A. idiocarpus H. Cout., Bull. Soc. Phil. 1908, p. 6. (Plate 60, fig. 2.) 

Lunique spécimen de lA. idiocarpus est un f de 7 mm., provenant, comme le précédent, 
de Providence, 50—78 brasses. 

Ces deux espéces constituent une augmentation de grande importance pour le genre 
Alpheopsis. Jusqu’a présent, ses deux groupes, equalis & pinces lisses, et trispinosus & 
pinces sillonnées, paraissent assez faiblement reliés. Peut-étre les formes ci-dessus décrites 
apporteraient-elles précisément le trait d’union cherché, si leurs pinces de la 1° paire étaient 
connues. 

Genre Synalpheus sp. Bate. 

7. S. metaneomeris n. nom. (Plate 60, fig. 4.) 


S. neomeris H. Cout., Alpheidee des Laquedives et Maldives nec de Man. Bull. Soc. 
Phil. xi. 5, 1908. 


S. streptodactylus de Man, Alpheidz du Siboga, :p. 226. 


Dans son magnifique travail sur les Alpheidz recueillis par l’expédition du Siboga, de 


Man établit définitivement la non-identité du véritable S. neomeris de Man avec l’espéce ; 


ci-dessus, qui en est évidemment trés distincte. Je ne puis toutefois accepter le nom de 
streptodactylus donné par de Man, le nom correspondant 4 une variété que je crois toujours 
valable, et qui est basée sur l’aspect différent des dactyles. Je reproduis ici les 2 figures de 
mon précédent travail, ot l’on voit clairement, mieux que par toute déscription, la différence 
de forme et de proportion des deux griffes. Cette forme devra donc porter le nom de 
S. metaneomeris var. streptodactylus. 

Quant & S. metaneomeris, lespéce me parait assez variable. Le sp. $ que j’ai figuré 
est un de ceux qui ont été communiqués 4 de Man et lui ont servi a la description de 
S. streptodactylus. Le scaphocérite s’étend jusqu’au milieu de l'article antennulaire distal, 
et toutes les épines sont trés allongées. Cette différence n’est pas sexuelle autant qu’il en 
a paru, elle correspond 4 ce que j'ai appelé la forme “oxyceros” qui marque chez les 
Synalphées une véritable direction évolutive, si l’on en juge par sa fréquence. Les deux 
autres spécimens figurés sont des ?, choisies parmi d’autres exemplaires pour la briéveté des 
épines antennaires et antennulaires, mais qu'il est impossible par ailleurs de séparer. 

Cette espéce est la plus abondamment représentée du genre Synalpheus dans la 
collection. Elle a été recueillie dans les localités et stations suivantes : 

Amirante Bank, st. 1, 2, 3, 5, 6, 9, 11, 13 (20—80 brasses); Saya de Malha, st. 10, 
12 (47 et 90 brasses); Cargados Carajos, st. 8, 13, 15, 19, 20 (28—33 brasses); Providence, 
st. 11 (58 brasses). 


8. S. graviert H. Cout., Alph. Mald. et Lace. p. 870, pl. 70, fig. 2. 
Amirante Bank, st. 16, 18, 22 (16—280 brasses); Providence, st. 11 (58 brasses). 


gee ting 


COUTIERE—LES ESPECES D’ALPHEIDA RAPPORTEES DE LiOCEAN INDIEN 415 


9. S. merospiniger H. Cout., Bull. Soc. Phil. 1908, p. 5. (Plate 60, fig. 5.) 
Amirante E. 25 (20—44 brasses). Une ? ovée. 


10. S. paraneomeris H. Cout. 

S. paraneomeris H. Cout., Alph. Mald. et Lace. p. 872, pl. 71, fig. 7. 

Chagos: Salomon, et Coin, Peros. Coetivy. 

11. S. paraneomeris, praslini n. var. (Plate 61, fig. 6.) 

Quelques spécimens de Praslin récif se montrent nettement distincts par trois caracteéres : 
langle supéro-interne du basicérite est nettement prolongé en une épine, alors que 
labsence d’épine en ce point est une des caractéristiques principales de l’espéce para- 
neomeris. Le méropodite de la 3° paire est plus large que dans les spécimens typiques, le 
rapport de ses dimensions étant seulement 3°5 au lieu de 4°5. Enfin les ceufs sont notable- 


ment plus gros, mesurant 1°3 mm. de grand axe. 

Seychelles, Praslin reef, 1g, 1 9. 

12. S. paraneomeris, seychellensis n. var. (Plate 61, fig. 7.) 

2 spécimens ? provenant des Seychelles différent également du type par quelques 
caracteres nets: 

Le basicérite a son angle supéro-interne, sinon épineux, au moins aigu. 

Les pattes de la 3° paire ont leur méropodite 4 fois aussi long que large, et le propodite 
porte 8 épines au lieu de 5 chez les spécimens typiques. C'est & ces exemplaires que jai 
comparés le S. otvosus. Par tout ses autres caractéres, dimensions du carpocérite, forme du 
dactyle, telson, cette variété seychellensis ne s'éloigne pas de l'espéce paraneomerrs. 

Enfin il se trouve parmi des spécimens provenant de Coin, Peros Atoll, un ¢ de la 
forme “ oxyceros,” caractérisé par la longueur de toutes ses épines antennaires et anten- 
nulaires; le stylo et le basicérite atteignent sensiblement l’extrémité de l'article antennulaire 
médian, le scaphocérite, par son épine latérale, dépasse de beaucoup le carpocérite. C'est 
pour de tels exemplaires que j’ai proposé, sur l’invitation de Miss Rathbun, le nom de 
paraneomeris prolatus, mais je trouve la dénomination commune “ oxyceros” infiniment 
plus expressive, parce qu'elle indique, comme je !’ai dit plus haut, une direction évolutive 
trés répandue, pouvant servir & expliquer comment se sont constituées de nouvelles espéces. 


18. SS. otiosus H. Cout., Bull. Soe. Phil. 1908, p. 5. (Plate 61, fig. 8.) 

Liespéce se distingue du S. paraneomeris H. Cout. par les points suivants : 

Le carpocérite est seulement 3 fois aussi long que large (au lieu de 4 fois). 

Le méropodite de la grande pince est un peu plus épais dans sa moitié proximale, et 
se termine par une pointe obtuse. 

Le méropodite de la 3° paire est seulement 3°5 fois aussi long que large, au lieu de 
4°5 fois chez le S. paraneomeris, et le propodite qui porte 5 épines est également plus massif. 

Le telson est plus large & son extrémité, le rapport de ses bases étant 1°5 au lieu 
de 1°85. 

Coetivy, un bel exemplaire ¢. 


14. S. nilandensis H. Cout., Alph. Mald. et Lace. p. 871, pl. 70, fig. 4. 
lex. Seychelles, st. 5 (44 brasses). 


ae 


416 PERCY SLADEN TRUST EXPEDITION 


15. S. nilandensis oxyceros H. Cout., Alph. Mald. et Lace. p. 871, pl. 70, fig. 5. 
1ex. Chagos, Egmont, lagoon. 


16. S. fossor Paulson. Alph. Mald. et Lace. p. 872, pl. 70, fig. 6. 
1 ex. Chagos, Diego-Garcia, lagoon. 


17. S. trionyx H. Cout., Bull. Soc. Phil. 1908, p. 6. (Plate 61, fig. 9.) 


Lespéce est trés voisine du S. fossor Paulson, dont elle se distingue par les points 
suivants : 

Les intervalles sont plus larges entre le rostre et les épines latérales, celles-ci étant, 
par suite, plus divergentes. 

Lécaille du scaphocérite atteint au moins la moitié de l'article antennulaire distal, et 
Pépine latérale dépasse toujours le carpocérite, qui est 5 fois environ aussi long que large. 
Chez le S. fossor, l’écaille antennaire atteint rarement l’extrémité de l'article antennulaire 
médian, l’épine latérale est toujours plus courte que le carpocérite, et celui-ci est 6 fois plus 
long que large. 

La grande pince est épineuse chez le S. tionyx 4 son bord palmaire antérieur, et le 
bord supéro-externe du méropodite se termine également par une forte épine. L’un et 
lautre sont inermes chez le S. fossor, surtout la paume de la grande pince. 

La petite pince du S. trionyx a pour proportions: doigts 1, longueur totale 2-4, 
hauteur 0°8. Le méropodite, dont le bord supéro-externe se termine en pointe aigué, est 2°1 


4 2°6 fois plus long que large. II est toujours plus épais que la paume. Chez le S. fossor ~ 


les proportions de la petite pince sont: doigts 1, longueur totale 2°85, hauteur 1:2, le membre 
étant plus trapu. Par contre, le méropodite gréle est 2°7 fois aussi long que large, et n’a 
que 0°6 de la hauteur de la paume. 

Le premier segment du carpe est plus long que la somme des 4 autres chez le S. trionyz, 
plus court chez le S. fossor. . 

Les proportions de la 3° paire sont chez le S. trionyx: propodite 2°35, carpe 1, méro- 
podite 2°6, celui-ci 4°4 fois aussi long que large. Chez le S. fossor, ces proportions sont : 
propodite 1:9 environ, carpe 1, méropodite 2°4, celui-ci 3°55 fois aussi long que large. 

Enfin, le dactyle est de forme nettement différente. Chez le S. fossor, la griffe ventrale 
surnuméraire a ses deux cdtés presque perpendiculaires, la griffe dorsale est trés accessoire 
par rapport a la ventrale, et le groupe de ces 2 griffes est séparé du reste du dactyle par 
une faible dépression dorsale. Chez le S. trionyx, la surnuméraire ventrale est aigué, 
dirigée en avant; la dorsale est presque aussi longue que la griffe principale, et le dactyle 
ne montre aucune concavité dorsale. 

Saya de Malha, C. 10 (26 brasses), 2 ex. f et 2; C. 19 (29 brasses), 1 ex. 2. 


18. S. triunguiculatus Paulson. 

Cargados Carajos, st. 8, 13, 17 (30—82 brasses). 

19. S. charon Heller. 

Chagos: Salomon, Peros, Egmont, récif et lagon. Coetivy. 

20. S. biunguiculatus Stimpson. H.Cout., Alph. des Mald. et Lace. p. 873, pl. 71, fig. 8. - 
Amirante Bank, st. 6, 13, 21 (20—30 brasses). Chagos, Egmont. 


ee ee es ee 


—— 


ee eee 


COUTIERE—LES ESPECES D’ALPHEIDA RAPPORTEES DE LiOCHAN INDIEN 417 


21. S. pachymeris n. nom. (Plate 61, fig. 10.) 

S. brunguiculatus var. pachymeris H. Cout., Alph. Mald. et Lace. p. 873, pl. 71, fig. 9. 

Cette forme est nettement distincte du S. biungwiculatus. Elle est représentée dans la 
collection par quelques spécimens typiques, mais aussi par une forme dont je fais une 
variété cargados: et qui sen distingue par quelques caracteéres: l’angle supéro-interne du 
basicérite est & peine épineux, le méropodite de la 3° paire ne porte que 3 épines mobiles 
sur son bord inférieur, au lieu de 5, enfin le propodite n’a que 5 épines au lieu de 8, et il 
est plus court, mesurant 1°33 du carpe au lieu de 1:47 chez les spécimens typiques. 

Cargados Carajos, 1 sp. 2 (var. Cargadosi). Providence Island. 


22. 8S. lophodactylus H. Cout., Bull. Soc. Phil. 1908, p. 7. (Plate 61, fig. 11.) 

Cette nouvelle espéce se distingue du S. biwnguiculatus Stp. par les points suivants : 

Le pédoncule antennulaire est plus court, l'article basal ne dépassant guére le rostre, 
et seulement 4 fois aussi long que large. Le stylocérite ne dépasse pas I’article basal. 

L’écaille antennaire atteint l’extrémité de l'article antennulaire médian, et son épine 
latérale ne le-dépasse que de la moitié environ de l’article antennulaire distal. 

La basicérite est complétement inerme en dessus, détail qui différencie immédiate- 
“ment les deux espéces. le carpocérite dépasse notablement l’antennule, mais il est 
seulement 4 fois aussi long que large, ayant subi le méme raccourcissement que les autres 
appendices céphaliques. ' 

La grande pince est absente sur le type. 

La petite pince a pour proportions: doigts 1, longueur totale 2°73, hauteur 0°88. Son 
méropodite est 3°3 fois aussi long que large. Le doigt mobile porte une brosse de soies 
sériées sur la moitié distale. Elles sont moins serrées et moins abondantes que dans le 
groupe /evimanus, mais n’en indiquent pas moins l’étroite parenté qui relie les deux groupes 
d’especes. Chez le S. biunguiculatus, la petite pince est plus épaisse, et le doigt mobile 
na que des traces insignifiantes de la brosse de soies sériées. 

Les pattes de la 3° paire ont pour proportions: propodite 1°9, carpe 1, méropodite 2°43. 
Ce dernier, trés gréle, est 4°5 fois aussi long que large. Le dactyle est celui du S. biwngus- 
culatus. 

Lagon de Diego, 1 seul spécimen 2. 

23. S. tumido-manus, Paulson. 

S. tumido-manus Paulson. H. Cout., Alph. Mald. et Lace. p. 876, pl. 73, fig. 14. 

Cargados Carajos, st. 2 (30 brasses), 1 ex. 3. 


24, S. hastilicrassus H. Cout., Alph. Mald. et Lace. p. 875, pl. 72, fig. 12. 
Amirante Bank, st. 21 (30 brasses); Providence, st. 11 (58 brasses). 


25. S. sladeni H. Cout., Bull. Soc. Phil. 1908, p. 8. (Plate 62, fig. 12.) 

Cette espece, l’une des plus remarquables recueillies par l’expédition, est jusqu’a 
~ présent la seule du groupe Jevimanus qui soit présente dans la région indo-pacifique, & 
Yexception peut-étre du S. levimanus haddon H. Coutiére. Ce groupe est caractérisé, 
comme je l’ai montré, par Ja présence constante sur le doigt mobile de la petite pince d’une 
brosse de longues soies raides disposées en séries transversales, et occupant les 2 de 


418 PERCY SLADEN TRUST EXPEDITION 


Yarticle. Tout en présentant de fagon trés nette ce caractére, la nouvelle espéce se place 
bien & part dans le groupe levimanus. 

Le bord frontal présente en avant des yeux une avancée considérable, si bien que la 
pointe du stylocérite est au niveau de celle du rostre, et que l’épine latérale du basicérite 
n’atteint pas tout a fait les épines latérales du bord frontal. 

Le scaphocérite est sensiblement aussi long que l’antennule, et son écaille atteint 
Vextrémité de l'article antennulaire médian, caractéres insolites dans le groupe, ainsi que la 
faible épine inférieure du basicérite. 

Le carpocérite est volumineux, 4°7 fois aussi long que large, plus long que les an- 
tennules. 

Les fouets des maxillipédes externes sont trés asymétriques en longueur et surtout en 
volume. J ignore si cette disposition est fortuite ou constante. 

La grande pince a pour proportions: doigts 1, longueur totale 3°12, hauteur 0°84. Le 
doigt mobile dépasse un peu le doigt fixe. La paume est cylindrique, et sa grande longueur 
est également une caractére insolite. 

Par contre, la petite pince ressemble beaucoup & celle du S. longicarpus Herrick, par 
sa forme, ses proportions, sa brosse de soies sériées. ° 

La 2° paire est gréle, la 3° paire également. Les proportions de cette derniére sont : 
propodite 2, carpe 1, méropodite 2°15, ce dernier article 5:4 fois aussi long que large. La 
eriffe est celle de toutes les espéces levimanus. 

Le telson a pour proportions: largeur distale 1, largeur proximale 2°7, hauteur 3 en- 
viron. 

Pour l’avancée du bord frontal, la forme de la grande pince, la gracilité des pattes 
suivantes, la forme du telson, cette espéce rappelle les espéces du groupe comatularum, 
jusqu’a présent aussi exclusives 4 l’Indo-pacitique que le groupe /evimanus Vest aux cdtes 
américaines. 

Cargados Carajos, B. 2 (30 brasses), 1 seul ex. 2. 


Genre Alpheus Fabr. 
26. A. staphylonus H. Cout., Bull. Soc. Phil. 1908, p. 14. (Plate 62, fig. 13.) 


L’espéce est trés voisine de |’A. megacheles Hailstone de la Méditerranée et de l Atlan- 
tique. Le rostre est 3 fois aussi long 4 peu pres que les dents sus-orbitaires, il est plus court 
que larticle antennulaire basal. L’article médian est 1°5 fois aussi long que chacun des 
2 autres. Le pédonculaire antennulaire, la forte épine latérale du scaphocérite, le carpocérite 
sont sensiblement égaux. 

La grande pince est trés semblable 4 celle de l'A. megacheles comme forme et pro- 
portions. La petite pince est plus gréle, la paume étant 2°43 fois aussi longue que large, 
au lieu de 2 fois, le doigt mobile n’est pas élargi, bien qu'il s'agisse d’un 3, les doigts sont 
légérement plus courts que la paume. 

La 2° paire est trés allongée, la somme du carpe et de la pince mesure 0°85 du cephalo- 
thorax, et la partie proximale du membre 1°05 de cette méme longueur, ce qui distingue 
immédiatement l’espéce de 1’A. megacheles et la rapproche de |’A. hadlstone: des Maldives. 

La 3° paire mesure 1°2 du*céphalothorax, alors quelle Végale seulement chez 


COUTIERE—LES ESPECES D’ALPHEIDA) RAPPORTEES DE LIOCEAN INDIEN 419 


YA. megacheles. Le méropodite est 5 fois aussi long que large, et le propodite est 1°4 fois 
aussi long que la carpe, au lieu de 1:2 chez l’A. megacheles. 

L’unique spécimen 3, long de 12 mm., provient de Salomon Island (Chagos). 

27. A. hailstoner H. Cout., Alph. Mald. et Lacc. p. 879, pl. 74, fig. 18. 

Seychelles, st. 8, 9 (34 et 37 brasses). Maurice. Saya de Malha, st. 15 (55 brasses). 

28. A. paradentipes H. Cout., Alph. Mald. et Lacc. p. 880, pl. 74, fig. 17. 

Amirante Bank, st. 25, 26 (20—100 brasses). Providence, st. 11 (58 brasses). 

29. A. collwmianus Stimpson, Alph. Mald. et Lace. p. 881. 

Amirante Bank, st. 7, 10, 11, 13 (20—85 brasses). Providence, st. 1, 3 (29—39 
brasses). Chagos, Salomon. Cargados Carajos, st. 20 (28 brasses). 

30. A. sewrate H. Cout., Alph. Mald. et Lacc. p. 881, pl. 75, fig. 20. 

Amirante Bank, st. 13 (20—25 brasses). Coetivy. 


31. A. malhaensis H. Cout., Bull. Soc. Phil. 1908, p. 15. (Plate 62, fig. 14.) 

L'espéce se rapproche particulitrement de I’A. sewrati H. Cout. dont elle se distingue 
par les points suivants: 

Antennes et antennules un peu plus gréles et allongées, y compris |’épine latérale du 
basicérite. 

Bord inférieur de la grande pince entiérement lisse, constriction plus.profonde entre la 
paume et le doigt fixe, extrémité du doigt mobile moins massive. 

Carpe de la 2° paire avec les 2 premiers segments presque égaux, le premier légére- 
ment plus long. 

Méropodites de la 3° et 4° paires fortement épineux & |’apex inférieur distal, celui 
de la 3° paire 3°7 fois plus long que large. Proportions du membre: carpe 1, propodite 1-25, 
méropodite 1°9 (au lieu de 1, 1°45, 2 chez 1’A. sewrati, oti le méropodite est seulement 
3 fois aussi long que large, et le propodite également trés épais). 

Saya de Malha, C. 19 (29 brasses), 1 sp. 9. Amirante, E. 2 (29 brasses), 1 sp. 2. 

32. A. macrochirus Richters. H. Cout., Alph. Mald. et Lace. p. 882. 

Chagos, Salomon Islands. Coetivy. 

33. A. ventrosus H. M. Edwards, Alph. Mald. et Lace. p- 882. 

Seychelles, Praslin reef. Chagos: Diego Garcia, barachois; Salomon; Egmont, 
lagoon. Coetivy. 

34. A. gracilis Heller. 

Chagos, Salomon. Coetivy. 

35. A. gracilis var. allwaudi H. Cout., Alph. Mald. et Lace. p. 882. 

Providence Island, 1 sp. mutilé. 

36. A. paragracilis H. Cout., Alph. Mald. et Lace. p. 883, pl. 76, fig. 22. 

Coetivy. 

Alpheus sp.? (Plate 62, fig. 15.) 
Metalpheus n. gen. ? 
Il s'agit ici d’une forme trés remarquable, peut-étre identique & 1A. rostratipes Pocock, 


420 PERCY SLADEN TRUST EXPEDITION 


ce que je ne saurais décider en l’absence du type. Cette identité ou valeur spécifique est 
d’ailleurs secondaires en l’espéce, en présence des différences qui séparent ces formes du 
genre Alpheus lui-méme, et conduisent presque 4 la formation d’un nouveau genre. 

Le rostre émerge entre les échancrures obliques des votites orbitaires, qu'il égale en 
longueur. La protection des ophthalmopodes n’est pas assurée du coté antéro-interne. 

Comme chez |’A. rostratipes, les articles antennulaires sont aussi larges que longs. 
Toutefois, le 3° est 1°5 fois aussi long que large. Le stylocérite atteint le milieu de l'article 
médian. la large écaille du scaphocérite atteint le tiers proximal, et son épine la moitié 
de l'article distal antennulaire. Le basicérite est volumineux, beaucoup plus haut que long 
surtout, avec une forte épine latérale, et le carpocérite, dépassant l’antennule de la moitié 
au moins de l'article distal, n’est guére que 1°5 fois aussi long que large. 

Le volume de l’antenne contrastant avec le faible développement du scaphocérite, 
l’épaisseur des articles et des fouets antennulaires sont des caractéres insolites chez Alpheus, 
rappelant surtout les genres Arete et Alpheopsis. 

Le labre posséde également un volume excessif. I] descend verticalement entre les 
bases des antennes, et sa hauteur égale celle du céphalothorax, de fagon & constituer une 
sorte de mufle qui donne 4 l’espéce une physionomie trés spéciale. La partie inférieure en 
est enfermée, comme dans des valves, entre les psalistomes trés élargis des mandibules, dont 
le palpe est entiérement invisible du dehors. Le psalistome porte une dizaine de faibles 
dents sur une courte portion de son bord supérieur. Le processus molaire est inclus, comme 
toujours, entre le labre et les paragnathes, trés développés également. C’est la une forme 
de la mandibule unique jusqu’a présent chez les Alpheide. 

La maxillule et la maxille ont leur forme normale. Sur le 1* maxillipede, l article distal 
du sympodite est largement arrondi, l’endopodite et l’exopodite soudés sur leur tiers proximal, 
ce dernier avec un tres étroit lobe a (Boas). 

Le 2° maxillipéde a son épipodite en forme de sac membraneux. Le 3° a son article 
distal foliacé, trés élargi (un peu plus de 2 fois aussi long que large) excavé de fagon a 
recevoir la masse du labre, des mandibules et des maxillipedes précédents, comme chez 
l Alpheopsis fissipes H. Cout. L’article distal est conique, pourvu des soies sériées habi- 
tuelles, sans épines terminales. I] y a un bourgeon d’arthrobranchie a la base du membre, 
avec une trace de bifurcation. 

Par une malchance singuliére, les 2 spécimens que j'ai étudiés ne possédent qu'une 
seule des pinces de la 1° paire, de méme que les types de lA. rostratipes. Je suis porté & 
croire que les deux pinces sont semblables. Sur un spécimen sec de 1’A. rostratipes (S. Ken- 
sington Muséum) la seule pince présente est si volumineuse qu'elle représente bien plutot 
la grande. Sur un des deux spécimens de la forme ici étudiée, la pince opposée est en voie 
de régénération, elle a dépassé le stade de forme indifférente, et apparait trés semblable a 
son opposée comme aspect général et proportions. 

La 2° paire est courte et massive. Les segments du carpe décroissent du 1®& au 4° 
ce dernier plus large que long, le 5° égal au 1°. La 3° paire a le méropodite trés massif, 
inerme, 3 fois aussi long que large. Le dactyle est bifide, avec une saillie ventrale sur- 
numéraire plus marquée chez le ¢. 

La 4° paire n’a pas d’épipodite en crochet. Les rames des pléopodes, chez le g, sont 


COUTIERE—LES ESPECES D’ALPHEIDA! RAPPORTEES DE LOCEAN INDIEN 421 


beaucoup plus courtes que le sympodite, presque dépourvues de soies et de longueur égale. 
La 2° paire fait exception en ce que la rame interne est étroite et trés longue. 

Chez la ? les rames sont plus longues et aussi plus larges que le sympodite, et le 
rétinacle de la rame interne est lui-méme trés élargi, si bien que la rame parait simplement 
bifurquée sur son tiers distal. 

Volume des fouets antennaires et de l’antenne, mhcomplete protection des yeux, volume 
du labre, forme trés spéciale des mandibules, du 2° maxillipéde, du 3° maxillipéde, des 
pinces de la 1° paire (?), briéveté de la 2° paire, réduction du nombre des épipodites, forme 
trés spéciale des pléopodes, tels sont les caractéres que l’on pourrait invoquer pour la 
séparation de |'A. rostratipes et des formes affines. Le nouveau genre pourrait recevoir le 
nom de Metalpheus sil était conservé. 

D’autre part, ces espéces montrent avec |’'A. paragracilis H. Cout. une ressemblance 
trés grande, au point que tous leurs caracteéres différentiels s’y retrouvent, y compris absence 
ad épipodite sur la 4° pavre. Mais ces caractéres sont pour la plupart atténués. I] en est 
ainsi pour le volume de l’antenne et sa disproportion d’avec le scaphocérite, pour le volume 
du labre, la grandeur du psalistome des mandibules, du sympodite du 1° maxillipéde, la 
largeur de l'article basal du 3° maxillipede, le volume de la petite pince, la briéveté de la 
2° paire, pour la forme méme des pléopodes. En un mot, lA. paragracilis ne possede plus 
en propre qu'un seul caractére le séparant nettement du genre Alpheus, l'épipodite du 
4° péréiopode. Et si lon passe 4 des espéces telles que l’A. soczalis, également trés voisine, 
ce dernier détail disparait. 

On voit donc V’intérét qui s’attache & la connaissance plus compléte de l’A. rostratipes 
(et de A. sp? qui en est peut-étre distinct) surtout pour savoir si la grande pince est ou 
non celle d'un Alpheus. 

On peut noter que |’ Alpheopsis fissipes, la nouvelle espéce décrite plus avant, posséde, 
seule du genre, les maxillipédes de la 3° paire trés élargis et les dactyles bifides. Si, la 
encore, les pinces de la 1’ paire étaient connues, peut-étre posséderait-on un repére 
précieux pour fixer la validité du-genre Metalpheus. Par son rostre, lAlpheopsis fissupes 
se rattache 4 1’A. equalis, dont les pinces ne sont pas sillonnées. On congoit trés bien dés 
lors qu'une forme telle que Metalpheus, dans hypothése ott ses pinces seraient semblables 
et non sillonnées, soit sortie d’une autre telle que | Alpheopsis fissipes. 

D’autre part, le groupe megacheles, parmi les Alphées, présente comme je l’ai montré, 
les ressemblances les plus étroites avec les Alpheopsis tel que lA. trispinosus et | A. chilensis, 
& pinces sillonnées. Comme les 2 sections du genre Alpheopsis sont fort voisines, il n’est 
pas étonnant de rencontrer chez leurs dérivés Alpheus et Metalpheus des convergences 
comme celles qui rapprochent IA. rostratipes et lA, paragracilis. 


37. Alpheus amiranter H. Cout., Bull. Soc. Phil. 1908, p. 15. (Plate 63, fig. 16.) 


Cette espéce se rapproche plus de lA. paragracilis H. Cout. que d’aucune autre, mais 
elle en est facilement séparable. 

Le rostre étroit se continue par une faible créte, un peu élargie derriére les votites 
orbitaires ; il est séparé de celles-ci, en avant, par deux trés faibles sinus concaves du bord 
frontal, le reste du bord étant régulierement convexe et inerme. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 54 


422 PERCY SLADEN TRUST EXPEDITION 


Le stylocérite est plus court que l'article antennulaire basal, ou 4 peine aussi long, le 
basicérite de l’antenne est & peu prés inerme. 

Le bord inférieur de la grande pince est droit, sans trace de constriction. Le bord 
supérieur porte au contraire une constriction transverse et un sillon longitudinal du cdté 
inféro-interne. Du cdte supéro-externe est un second sillon longitudinal trés faible. Le 
méropodite porte une forte épine 4 son bord inférieur interne. 

La petite pince différe de celle de l'A. paragracilis par sa taille rebate moindre, les 
doigts égaux & la paume, le méropodite pourvu d'une forte épine. 

Sur la 2° paire, le 1° segment du carpe n’a guére que le tiers du second. 

La 3° paire et la 4° ont le méropodite fortement épineux 4 son bord inférieur distal, le 
propodite est seulement un peu plus long que le carpe, l'un et l'autre sont beaucoup plus 
faibles que le méropodite. Le dactyle est simple. C'est une forme du membre surtout 
fréquente dans le groupe erinitus. 

Amirante E. 11 (25—80 brasses), 2 spécimens @. 

38. Alpheus baculifer H. Cout., Bull. Soc. Phil. 1908, p. 16. (Plate 68, fig. 17.) 

Je place ici cette trés remarquable espece. Elle représente un cas extréme d’allonge- 
ment des pinces chez une forme qui, par la plupart de ses caractéres, rentre dans le groupe 
obeso-manus. Crest le paralléle de lA. cylindricus Kingsley pour le groupe megacheles. 

Le rostre, trés court, dépasse 4 peine les votites orbitaires trés saillantes, et se continue 
en arriére par une créte assez marquée. L’article antennulaire médian est 1:5 fois aussi 
long que chacun des deux autres, le stylocérite est plus court que Varticle médian et ne 
fait qu'une faible saillie latérale. Le scaphocérite ne posséde qu’une écaille rudimentaire, et 
son épine latérale ne dépasse pas le tiers distal de l’article antennulaire médian. Le car- 
pocérite est a peine plus long que ce méme article. 

Les pattes de la 1° paire sont extrémement caractéristiques. La grande pince a la 
paume cylindrique, 5°5 fois aussi longue que large. Elle est tronquée 4 l’extrémité ; le 
doigt mobile, en “porte a faux,” sur plus de la moitié de sa longueur, est dirigé presque 
perpendiculairement 4 la paume. II n’y a plus trace dé sillons palmaires. Le méropodite 
est également trés allongé. 

La paume de la petite pince est de méme forme, mais les doigts sont paralléles, égaux, 
et dans le prolongement du membre. Ses proportions sont : doigts 1, longueur totale 4°34, 
hauteur 0°55. 

Les pattes de la 2° paire sont trés inégales, comme dans la plupart des espéces obeso- 
manus. la plus longue mesure 1°66 fois celle de la 3° paire, la plus petite 1°33 fois. Le 
second segment du carpe est trés légérement plus long que le premier, la pince distale a 
les doigts trés courts. 

La 3° paire a pour proportions: carpe 1, propodite 1:07, méropodite 1°46. Ce dernier 
est merme, et 3°6 fois aussi long que large. 

L’unique spécimen est une ? venant de muer dont les pleurons abdominaux sont trés 
grands, comme il est de régle dans ce groupe d’espéces. Le telson et les uropodes sont 
dans le méme cas ; la rame interne de ces derniers a son bord externe triangulaire, la suture 
de la rame externe porte 2 courtes épines. 

Tle du Coin, Peros (Chagos), une $ de 25 mm. 


COUTIERE—LES ESPECES D’ALPHEIDH RAPPORTEES DE LOCEAN INDIEN 423 


39. A. ovaliceps H. Cout., Alph. Mald. et Lace. p. 888, pl. 77, fig. 27. 

Chagos : Coin, Peros. 

40. A. stanleyi H. Cout., Bull. Soc. Phil. 1908, p. 17. (Plate 68, fig. 18.) 

Cette espéce se rapproche beaucoup de 1’A. ascensionis Ortmann, de 1’A. architectus 
de Man, de !’'A. styliceps H. Coutiére. 

La forme du bord frontal est trés semblable 4 celle de |’'A. styliceps, les votites or- 
bitaires étant toutefois reportées plus en arriére encore. L’écaille du scaphocérite atteint 
le tiers distal de l'article antennulaire médian, tandis qu’en revanche |’épine latérale atteint 
& peine le milieu de l'article distal. Le carpocérite ne dépasse que trés faiblement l’antennule. 

La grande pince est celle de l’A. ovaliceps H. Coutiére, et se distingue par suite de 
celle des 3 espéces précitées. Le doigt mobile est toutefois beaucoup plus en “ porte a 
faux.” Les sillons palmaires sont & peu prés nuls. Ses proportions sont : doigt mobile 1, 
longueur totale 3:4, hauteur 1°4. 

La petite pince a pour proportions: doigts 1, longueur totale 2, hauteur 1°72. Elle 
est, par suite, de forme banale, et bien distincte de celle de A. styliceps. Les doigts sont 
béants, linférieur trés large; leur bord inféro-externe excavé porte une rangée de fortes 
soles courtes et espacées, au nombre d'une dizaine. 

Sur la 2° paire, le 2° segment du carpe est un peu plus court que le premier. 

La 3° paire a pour proportions: propodite 1:4, carpe 1, méropodite 2°25. Ce dernier, 
presque 5 fois aussi long que large, porte une forte épine 4 son angle inférieur distal. I] 
en est de méme sur la 4° paire. Le dactyle est trés court, sans trace de bifurcation. 

Le telson a pour proportions : petite base 1, grande base 2, hauteur 3°27. Le bord de — 
luropode interne porte une série de courtes épines. 

Chez lA. architectus et 1A. ascensionis, indépendamment des autres différences, les 
méropodites 3 et 4 sont inermes. 

Amirante E. 21 (30 brasses), un unique sp. f de 18 mm. 

41. A. microstylus H. Cout., Alph. Mald. et Lacc. p. 884, pl. 76, fig. 23. Coetivy. 


42. A. phrygianus H. Cout., Alph. Mald. et Lacc. p. 886, pl. 77, fig. 25. 

Chagos: Salomon Island, lagoon. Providence, st. 1 (39 brasses). Amirante, st.- 21 
(30 brasses). 

43. A. lutini H. Cout., Alph. Mald. et Lace. p. 885, pl. 76, fig. 24. Coetivy. 

44, A. bradypus H. Cout., Alph. Mald. et Lace. p. 891, pls. 78—79, fig. 30. Coetivy. 

45. A. bucephalus H. Cout., Alph. Mald. et Lace. p. 890, pl. 78, fig. 29. 

Providence, st. 4, 7 (50 et 70 brasses). Chagos: Salomon; Diego Garcia, barachois. 
Amirante Bank, st. 9, 18, 16, 29 (20—44 brasses). Coetivy. 

46. A. aculeipes H. Cout., Alph. Mald. et Lace. p. 892, pl. 79, fig. 31. 

Amirante Bank, st. 2, 9, 11, 13, 16, 21 (20—89 brasses). Seychelles, st. 1, 5, 8, 9 
(20—44 brasses). Saya de Malha, st. 10, 19 (90 et 55 brasses). Coetivy, récif. Chagos : 
Diego Garcia. 

47. A. paraculeipes H. Cout., Alph. Mald. et Lace. p. 894, pls. 79—80, fig. 32. 


Amirante Bank, st. 13, 21 (20—30 brasses). 
54—2 


A424 PERCY SLADEN TRUST EXPEDITION 


48. A. providence: H. Cout., Bull. Soc. Phil. 1908, p. 18. (Plate 68, fig. 19.) 


Le difficile groupe crinitus, déji si riche en espéces, a fourni encore ici plusieurs formes 
nouvelles, séparées des formes connues par de minimes différences. 

Par la forme du bord frontal et des appendices céphaliques, |’A. pi ouedonees rappelle 
absolument lA. aculeipes H. Coutiére. Ici, toutefois, le rostre fait & peine saillie en avant; 
sa créte médiane est au contraire plus forte, et sélargit plus rapidement derriére les votites 
orbitaires. 

Par ses autres caractéres, l'espece se rapproche davantage de lA. paraculeipes H. 
Coutieére, sans étre toutefois intermédiaire entre les deux formes précitées. 

La grande pince, dont les proportions sont: doigts 1, longueur totale 3:33, hauteur 
1°5, a une forme régulierement ovoide ; ses doigts sont fortement infléchis en dedans. Le 
méropodite épineux est 2°2 fois plus long que large. 

La petite pince (?) est trés semblable a celle de 1A. paraculempes. Les doigts sont 
seulement plus longs, ses proportions étant: doigts 1, longueur totale 2°18, hauteur 0°8. 

Les deux premiers segments du carpe sont entre eux comme | et 1°95, rapport maguire 
que dans les deux espéces précitées. 

Pour la 3° paire, les proportions sont: propodite 1:5, carpe 1, méropodite 2°3. Ce 
dernier 4 fois plus long que large. L’épine mérale est dans le prolongement du bord 
inférieur, étroite et trés aigué. Le bord inférieur interne est absolument nu, d’ou une dif- 
férence notable d’avec les A. aculeipes et paraculeipes. 

Le bord externe du carpe porte une épine et 8—10 soies, mais celles-ci sont effilées, 
molles, et bien différentes de celles de 1A. paraculeipes. Le propodite n’a pas d’épine 
distale au bord supérieur. Le dactyle porte au bord ventral une saillie surnuméraire trés 
nette. L’ischiopodite n’a pas d’épine enfoncée. Le méropodite de la 4° paire n’est pas 
épineux. 

L’espéce est encore plus voisine de |’A. alcyone de Man. Dans cette derniére forme, 
le scaphocérite a son bord externe concave, son épine latérale plus longue; le méropodite 
de la petite pince est inerme; le premier segment du carpe mesure seulement le tiers du 
second, et il est surtout plus petit que le 5°. Le méropodite de la 3° paire porte quelques 
spinules courbées en 8, le dactyle n’est pas bifide; le carpe de la 4° paire est semblable a 
celui de la 3°. Enfin, les ceufs sont rares et volumineux, donnant vraisemblablement 
naissance & des larves mysis. 

On peut considérer l’A. alcyone et lA. providencei comme deux formes peecilogoniques, 
mais, icl comme dans la grande majorité des cas connus, il s'ajoute au caractére tiré du 
volume des ceufs une série de menues différences montrant que les deux formes sont bien 
spécifiquement distinctes. 

Providence D. 4 (50—78 brasses), 1 sp. 2. 


49. A. alcyone de Man? H. Cout., Alph. Mald. et Lace. p. 896. 
Amirante Bank, st. 13 (20—25 brasses). 


50. A. paralcyone H. Cout., Alph. Mald. et Lace. p. 895, pls. 80—81, fig. 34. 
Amirante Bank, st. 11, 13, 21 (20—80 brasses). Seychelles, st. 8 (34 brasses). 


cc — 


COUTIERE—LES ESPECES D’ALPHEIDA RAPPORTEES DE OCEAN INDIEN 4295 


51. A. superciliaris H. Cout., Alph. Mald. et Lace. p. 896, pl. 81, fig. 35. 
Amirante Bank, st. 21, 25 (209—160 et 44—20 brasses). 

52. A. pachychirus Stimpson. 

Chagos: Salomon; Egmont. Coetivy. 

Cowles a minutieusement décrit, récemment, la fagon dont cette espéce se construisait 


un tube a l’aide d’ Algues vertes pour s'y loger par couples. I] s'agit plus probablement de 
LA. frontalis = Beteus utricola Richters. 


53. A. frontalis H. M. Edwards. 


. Amirante Bank, st. 9, 11, 13 (20—80 brasses). Chagos: Egmont; Salomon. Sey- 
chelles : st. 2 (31 brasses) ; Praslin, récif. Coetivy. 


54. A. clypeatus H. Cout., Alph. Mald. et Lace. p. 897, pls. 81—82, fig. 36. 
Chagos: Egmont, récif. 


55. A. adamastor H. Cout., Bull. Soc. Phil. 1908, p. 19. (Plate 64, fig. 20.) 


Cette espéce est extrémement voisine de |’A. clypeatus H. Coutiére. Elle en différe 
par les points suivants : 

Le bord frontal présente une avancée convexe en dega de chaque volte orbitaire, mais, 
au lieu de se réunir en un rostre saillant, les votes sont séparées par un sinus médian, 
aussi large que chacune d’elles. La créte mousse du rostre, qui parait s’arréter au bord de 
ce sinus, se continue en réalité un peu en avant et sur un plan inférieur, en une pointe 
étroite et trés courte. 

Le basicérite porte une épine beaucoup plus faible que chez lA. clypeatus. 

La grande pince a pour proportions: doigts 1, longueur totale 3, hauteur 1°34. Elle 
est donc légérement plus massive que chez |’. clypeatus, ot les doigts ont aussi 1/3 de la 
longueur totale, mais ott le second rapport est 1°23 seulement. 

La petite pince présente des différences bien plus notables. Elle a pour proportions : 
doigts 1, longueur totale 2°36, hauteur 0°75. Elle est donc 3 fois plus longue que haute, 
au lieu de 2°6 chez l’A. clypeatus g. Elle rappelle par suite les proportions de l’appendice 
chez la ? de lespéce précitée, mais le doigt mobile est ici notablement élargi. I] est 
vraisemblable que la différence sexuelle dans la forme de la petite pince, chez l’A. adamastor 
dont la ? est inconnue, porte sur ce détail du doigt mobile, et qu'elle est par suite assez 
faible. 

Le méropodite inerme est 4 peine plus court que la pince et presque 3 fois aussi long 
que large. Ce dernier rapport égale 2 chez 1A. clypeatus g, le méropodite étant beaucoup 
plus court que la pince. 

Sur la 2° paire, le 1° segment du carpe égale 1°6 fois le second. Ils sont sensiblement 
égaux chez l’A. clypeatus. | 

La 3° paire est trés semblable comme proportions et forme 4 celle de lA. clypeatus, 
mais le méropodite est 4 fois aussi long que large, au lieu de 3°5 fois. 

La forme du bord frontal et les proportions de la petite pince du ¢ permettent aussi 
de distinguer aisément cette espéce de |'A. pachychirus Stimpson. 

Salomon (Chagos), un unique spécimen 2. 


426 - PERCY SLADEN TRUST EXPEDITION 


56. A. longecarinatus Hilgendorf. 

Amirante Bank, E. 13 (20—25 brasses). Providence, D. 1 (89 brasses). 

57. A. imsignis Heller. 

Chagos: Egmont; Salomon. Coetivy. 

58. A. lanceloti H. Cout., Alph. Mald. et Lace. p. 901, pl. 83, fig. 39. 
Amirante Bank, E. 11 (25—80 brasses). 

59. A. splendidus H. Cout., Bull. Mus. Paris, 1897, no. 6, p. 235. 
Seychelles: Praslin, récif. 

60. A. dasycheles H. Cout., Bull. Soc. Phil. 1908, p. 21. (Plate 64, fig. 21.) 


L’espéce se rapproche de |’A. gracilipes Stimpson par la forme du rostre. C'est une 


large pointe triangulaire 4 bords légérement concaves, 1°5 fois aussi longue que large a la 
base. Il y a sur la ligne médiane une trace de l’épine gastrique. Les bords du rostre sur- 
plombent les sillons rostro-orbitaires comme chez |’A. gracilipes, et les votites orbitaires 
présentent en avant une saillie obtuse, ne dépassant pas le bord frontal sinueux. La pointe 
du rostre n’atteint pas l’extrémité de l'article antennulaire basal, que le stylocérite dépasse 
légérement. 

Lécaille du scaphocérite égale le pédoncule antennulaire, que son épine latérale dépasse 
des 3/4 environ de Varticle distal. L’épine latérale du basicérite atteint en avant aussi loin 
que le rostre, le carpocérite ne dépasse pas l’extrémité de l'article antennulaire médian. 

Les pinces de la 1° paire sont cylindriques et entiéres, sans traces de sillons et de lobes. 
Elles possédent un revétement assez dense de longues soies sur leur face inféro-externe. La 
plus grande a pour proportions: doigts 1, longueur totale 3:2, hauteur 0:9. 

La plus petite: doigts 1, longueur totale 2°38, hauteur 0°5. Sur la 2° paire, le 1° 
segment égale 1:2 fois le second. la 3° paire a pour proportions: propodite 1°64, carpe 1, 
méropodite 1°88. Ce dernier est 5 fois aussi long que large, l'ensemble du membre étant 
eréle, mais le méropodite porte une forte épine 4 son apex inférieur. Le carpe se termine 
également par une épine. Le dactyle est simple. Il y a une épine enfoncée mobile sur 
LVischiopodite. 

Seychelles, F. 9 (37 brasses), 2 sp. 3 et ?, le plus grand mesurant 22 mm. 

61. A. percyt H. Cout., Bull. Soc. Phil. 1908, p. 21. (Plate 64, fig. 22.) 

Cette espéce se rapproche également de |’A. gracilipes, plus méme que la précédente. 

Le rostre a la méme forme lancéolée, & bords surplomblant les sillons rostro-orbitaires. 
Mais les votites orbitaires sont réguliérement hémisphériques, sans lobe antérieur saillant, 
de sorte quentre elles, le rostre et le bord frontal, il n’existe pas la partie plane du sillon 
rostro-orbitaire brusquement élargi en avant que l’on remarque chez lA. gracilipes. 

Le rostre atteint, du méme que le stylocérite, la longueur de l'article antennulaire basal. 

L’écaille antennaire atteint au moins l’extrémité du pédoncule antennulaire, quelle 
dépasse méme notablement chez les grands spécimens. Son épine latérale et le carpocérite 
comme dans l’espece précédente, et comme chez |’A. gracilipes. 

Les pattes de la 1¢ paire ne different que trés peu de celles de l’A. gracilipes. La 
grande pince est entaillée d’un profond sillon transverse un peu avant !’articulation du 
doigt mobile; la petite pince, “ baleniceps” chez les g, est munie d’une forte épine sur 


: 
| 


COUTIERE—LES ESPECES D’ALPHEIDA RAPPORTEES DE LiOCEAN INDIEN 427 


chacun des condyles articulaires du doigt mobile. Les bords supéro-externe et inféro- 
externe du méropodite se terminent chacun par une forte épine, sur les 2 pinces. 

Sur la 2° paire, les 2 premiers segments du carpe, presque égaux, sont entre eux 
comme 1, 1°15. J’ai étudié des spécimens jeunes ot le rapport était sensiblement inverse. 
La 8° paire est trés semblable & celle de l’espéce précédente comme forme et proportions. 

Cargados Carajos (30 brasses), 2 sp. f et ?, le plus grand mesurant 40 mm. Amirante 
E. 11 (25—50 brasses), 1 petit sp. 2. Coetivy, 1 petit sp. ¢. 3 

62. A. coetivensis H. Cout., Bull. Soc. Phil. 1908, p. 20. (Plate 64, fig. 23.) 

L’espéce se rapproche beaucoup de |'A. paracrinitus Miers et de la forme que j’ai 
décrite comme var. bengalensis de cette derniére. 

Les votites orbitaires sont ici atténuées en forme d’épines plates mesurant & peu prés 
la moitié de l'article antennulaire médian. Le scaphocérite a son écaille plus large, son 
épine latérale moins grande que dans les deux formes précitées, et le carpocérite est plus 
volumineux, 

La grande et la petite pince sont tout a fait celles de lA. paracrinitus bengalensis 
comme forme et proportions, la petite pince étant inerme sur le condyle articulaire externe 
du doigt mobile. Les méropodites sont inermes. 

Sur la 2° paire, le premier segment du carpe est 1°7 fois aussi long que le 2°, sensible- 
ment comme chez lA. paracrinitus. 

Sur la 3° paire, trés gréle, les proportions sont: propodite 1°45, carpe 1, méropodite 
1-7, le membre étant encore plus allongé que chez lA. paracrinitus. 

Coetivy, 1 sp. g. Chagos: Salomon, 1 sp. ? avec sa petite pince seule présente. 

63. A. alpheopsides H. Cout., Alph. Mald. et Lace. p. 901, pl. 83, fig. 40. Chagos: 
Salomon. 

64. A. paralpheopsides H. Cout. Chagos: Salomon. 

65. A. parvirostris Dana. Chagos: Egmont; Salomon. Seychelles: Praslin. Coetivy. 

66. A. hippothoé de Man. Amirante Bank, E. 13 (20—25 brasses). 

67. A. bowers H. M. Edwards, Alph. Mald. et Lace. p. 907, pl. 85, fig. 44. Chagos: 
Salomon ; Coin, Peros. Cargados Carajos. Coetivy. 

68. A. bastard: H. Cout., Alph. Mald. et Lace. p. 907, pl. 87, fig. 45. Chagos: Salomon; 
Coin, Peros. Coetivy. 

69. A. pacificus Dana, Alph. Mald. et Lacc. p. 909, pls. 85—86, fig. 47. Chagos: 
Salomon ; Coin, Peros. 

70. A. audowins H. Cout., Alph. Mald. et Lace. p. 911, pl. 87, fig. 52. Chagos: 
Salomon. 

71. A. strenuus Dana, Alph. Mald. et Lacc. p. 913, pl. 87, fig. 53. Chagos: 
Salomon. a 

72. A. strenwus angulatus H. Cout., Alph. Mald. et Lace. p.914. Chagos: Salomon. 
Amirante Bank, E. 11 (25—80 brasses). 


73. A. leptochirus H. Cout., Alph. Mald. et Lace. p. 914, pl. 87, fig. 54. Chagos: 


Egmont ; Salomon. Amirante Bank, E. 11 (25—80 brasses). Cargados Carajos. 


Fig. 
Fig. 
Fig. 


Fig. 


Fig. 
Fig. 


Fig. 
Fig. 


Fig. 
Fig. 
Fig. 
Fig. 


Fig. 


PERCY SLADEN TRUST EXPEDITION 


EXPLICATION DES PLANCHES 60—64. 


1. Automate salomont H. Cout., région céphalique vue en dessus; 1”, région céphalique vue latérale- 
ment; 1a, patte; 16, telson et uropodes. 

2. Alpheopsis idiocarpus H. Cout., région céphalique vue en dessus; 2a, patte de la deuxieme paire ; 
2b, patte; 2c, telson et uropodes. 

3. Alpheopsis fissipes H. Cout., région céphalique vue en dessus; 3a, maxillipede 111; 3, 2° péréio- 
pode; 3c, patte; 3c’, dactylopodite; 3d, ceuf. 

4. Synalpheus metaneomeris n. nom., région céphalique vue en dessus (f°); 4, 4”, la méme ($¢); 
4a, détail de lantenne; 4b, 3° patte; 4c, 4° patte; 4c’, dactylopodite; 4c”, dactylopodite; 4c”, 
var. streptodactylus dactylopodite; 4d, telson; 4e, ceuf. 

5. Synalpheus merospiniger H. Cout., région céphalique vue en dessus; 5a, détails de l’antenne ; 
5b, patte; 5c, dactylopodite; 5d, telson; 5e, ceuf. 

6. Synalpheus paraneomeris praslini n. var., région céphalique vue en dessus; 6a, patte; 6b, ceuf. 
7. Synalpheus paraneomeris seychellensis n. var., région céphalique vue en dessus; 7 a, dactylopodite. 
8. Synalpheus otiosus H. Cout., région céphalique vue en dessus; 8 a, détails de lantenne; 8 6, grande 
pince; 80’, la méme, méropodite et carpe; 8¢, petite pince; 8c’, la méme, méropodite et carpe; 8d, 
patte; 8d’, dactylopodite; 8 f, telson. 

9. Synalpheus trionyx H. Cout., région céphalique vue en dessus; 9 a, détails de l’antenne; 9 b, grande 
pince; 9b’, — méropodite et carpe; 9c, petite pince; 9d, 2° péréiopode; 97, patte; 9/', dacty- 
lopodite. 

10. Synalpheus pachymeris cargadosi n. var., région céphalique vue en dessus; 10a, patte. 

11. Synalpheus lophodactylus H. Cout., région céphalique vue en dessus; 11a, détails de l’antenne ; 
11}, petite pince; 11l¢, 2° péréiopode; 11d, patte; 11d’, dactylopodite; 11f, telson. 

12. Synalpheus sladeni H. Cout., région céphalique vue en dessus; 12a, grande pince; 126, petite 
pince; 12c, 2° péréiopode; 12d, patte; 12d’, dactylopodite; 127, telson. 

13. Alpheus staphylinus H. Cout., région céphalique vue en dessus; 18’, région céphalique vue 
latéralement; 13a, grande pince; 13a’, petite pince; 13a, la méme, méropodite ; 136, 2° péréiopode; 
13, patte. 


/ 


Fig. 14. Alpheus malhaensis H. Cout., région céphalique vue en dessus ; 14a, grande pince ; 14a’, la méme, 


Fig. 


Fig. 
Fig. 


Fig. 


Fig. 


méropodite et carpe; 146, 2° péréiopode; 14c, patte. 

15. Alpheus sp., région céphalique vue en dessus; 15’, région céphalique vue latéralement; 15a, 
mandibule; 156, maxillipéde 1; 15c, maxillipede 111; 15d, grande pince; 157, 2° péréiopode ; 
15g, patte; 159’, dactylopodite; 15h, pléopodes de la femelle; 15h’, pléopodes du male. 

16. Alpheus amirantei H. Cout., région céphalique vue en dessus; 16 a, grande pince; 16 a’, la méme, 
méropodite et carpe; 166, petite pince; 16¢, 2° péréiopode; 16d, patte. 

17. Alpheus bacuifer H. Cout., région céphalique vue en dessus; 17a, grande pince; 17, petite 
pince; 17 c, 2° péréiopode; 17d, patte; 177, telson et uropodes. 

18. Alpheus stanleyi H. Cout., région céphalique vue en dessus; 18’, région céphalique vue latérale- 
ment; 18a, détail de maxillipéde 111; 18, grande pince; 180’, la méme, méropodite et carpe; 18¢, 
petite pince; 18c’, détails de la petite pmce; 18d, 2° péréiopode; 187, telson et uropodes. 

19. Alpheus providencei H. Cout., région céphalique vue en dessus; 19 a, grande pince; 19 a’, la méme, 
méropodite et carpe; 19, petite pince; 19c, 2° péréiopode; 19d, patte; 19d’, dactylopodite; 197 
détails; 19g, telson. 


Fig. 20. Alpheus adamastor H. Cout., région céphalique vue en dessus; 20 a, détail de maxillipede 111; 20 6, 


grande pince; 20c¢, petite pince; 20 d, 2° péréiopode ; 20 f, patte. 


Fig. 21. Alpheus dasycheles H. Cout., région céphalique vue en dessus; 21a, grande pince; 21 8, petite 


pince; 21 c, 2° péréiopode ; 21 d, patte; 21 f, telson et uropodes. 


Fig. 22. Alpheus percyi H. Cout., région céphalique vue en dessus; 22’, région céphalique vue latéralement, 


22a, grande pince: 226, petite pince; 22c, 2° péréiopode; 22d, patte; 227, telson et uropodes. 


Fig. 23. Alpheus coetivensis H. Cout., région céphalique vue en dessus; 23a, grande pince; 23), petite 


pince; 230’, la méme, méropodite et carpe; 23c, 2° péréiopode; 23d, patte. 


Percy SLADEN Trust ExPrpition. 
(Courts) TRANS. LINN. SOC. SER. 2, ZOOL. VOL. XVII. PL. 60 


ALPHEIDAE 


Percy SLADEN Trust EXPEDITION. 
(CouTIERE) 


TRANS. LINN. 


ALPHEIDAE 


SOG, SIEIR. B ZOOL, WO!L, XVM. PIL. 


61 


Percy SLADEN Trust ExpPEpIrion. 
(Courriee) TRANS. LINN. SOC. SER. 2, ZOOL. VOL. XVII. PIL, 62 


ALPHEIDAE 


Percy Siapen Trust Expepirion. 


(Courrire) TOWNS, IDUNIN, SOC, SIR: 2 ZOOL, WOIL, Savi, DIL 6s 
16d ee a 


ALPHEIDAE 


Percy SLADEN Trust EXPEDITION. 
TORVAINGS. JLIONIN. SOO. SIR. 2 ZOOIL, WOJL OWI. IPIL, 64 


(CouTibRE) 


ALPHEIDAE 


No. XI1L—ON THE CEPHALOPODA OBTAINED BY THE PERCY SLADEN 
TRUST EXPEDITION TO THE INDIAN OCEAN IN 1905. 


By G. C. Ropson, B.A. 


(PUBLISHED BY PERMISSION OF THE TRUSTEES OF THE British MusEvm.) 
(CommuNIcATED BY Pror. J. StantEY Garpiner, M.A., F.R.S., F.L.S.) 


(Plates 65, 66, Text-figs. 1—6.) 
Read 17th June, 1915. 


AN examination of the Cephalopoda obtained by the “Sealark” on her cruise in the 
western regions of the Indian Ocean has been delayed since the return of the expedition 
in 1905 until the beginning of the present year [1914] when Prof. Gardiner placed them 
in the author's hands. 

The collection consists of eighteen species representing ten genera and includes a 
new genus and two new species. Although no very remarkable forms were obtained, 
the collection as a whole is not without interest, while some of the anatomical features 
revealed by dissection are of importance. 

A peculiar feature of this collection is the total absence of Myopsida (Loligo, Sepia, 
ete.). This group usually being of a littoral habitat their absence in conjunction with 
the relatively large number of pelagic and abyssal forms might be taken to shew that 
the ‘““Sealark’s” work was confined to deep-sea dredging or tow-netting far from land. 
This however was not the case. As will be seen from a study of the stations from 
which the various species were obtained some of the Polypods were taken at very 
moderate depths off the islands at which the “Sealark” touched: so that the absence 
of Myopsida is difficult to account for. 

In the present state of our knowledge of the group, Cephalopoda are a rather 
unprofitable class for use in zoogeographical studies. Those that are well known (the 
Polypods, squids and cuttlefish) have usually a very extensive range (or what appears to 
be so), being for the most part powerful swimmers. For example, Polypus fontanianus 
ranges from Chili to the Indian Ocean; while Stenoteuthis bartramu is cosmopolitan. The 
delicate abyssal forms, on the other hand, are so little known at present that it would be 
useless to generalize on their distribution. 

We have also to bear in mind the fact that the diagnoses of a great many species 
_ of Cephalopoda are based upon external characters of doubtful value, and that a different 
view of their distribution may be taken when students of this group have paid more 
attention to the description of internal parts. Distributional areas at present ranging 
over many degrees of latitude and longitude may by this process become more restricted. 
The habits of the majority of these animals, however, do not leave much ground for 

SECOND SERIES—ZOOLOGY, VOL. XVII. 55 


430 PERCY SLADEN TRUST EXPEDITION 


assuming that they will ever prove as interesting to the zoogeographer as land and fresh- _ 
water mollusca have been. 

Whenever possible, attention has been paid to the anatomy of the forms included in 
this collection, and several points of interest have been revealed. The author has been 
struck, while in the course of this work, with the necessity for a more intensive study 
of these animals for the purposes of systematic zoology. Any worker familiar with the 
Cephalopod mandible and radula will recall how at first sight these structures differ to 
a remarkably slight degree in forms placed very widely apart. It seems likely, however, 
that closer study will reveal differences between the radule of such forms. The author 
has been impressed by the way in which, in apparently similar radulz of forms otherwise 
very distinct, certain elusive and subtle differences may remain constant over a series of 
examples. The shells of certain groups of land mollusca have been separated into a 
number of species which, to the ordinary observer, exhibit very little difference one 
from another. It is claimed by conchologists that the differences between them are 
constant, however subtle and minute they may be. Very much the same sort of thing may 
be found by intensive study among the Cephalopoda. 

Among the internal organs the author has found that the genitalia and heart are 
frequently strongly characterised; and it often seems that for a provisional arrangement 
these organs would supply a more useful clue to identity than the radula and mandible. 
On the other hand due care has to be exercised, especially with regard to the heart, that 
characteristics such as are probably occasioned by the temporary physiological state of 
such organs are not registered as of diagnostic value. 

The author is indebted to Dr W. KE. Hoyle for information with regard to the 
Cranchiidee, and to the Rev. Dr H. M. Gwatkin for the loan of a series of radula-prepara- 
tions which has been of great value in determining the affinities of certain forms. 

The types and a series of other specimens have been presented by Prof. Gardiner to 
the British Museum. 


The following arrangement has been adopted from Pelseneer (14), Pfeffer (16), and 
Chun (2). 
Class CEPHALOPODA. 
Order 2. DIBRANCHIA. 
Suborder 1. DECAPODA. 
Tribe I. CEGOPSIDA. 
Ggopsida libera. 
Family 2. Onychoteuthidee. 
Onychoteuthis, sp. (immature). 


Teleoteuthis, sp. (immature). 


3. Enoploteuthide. 
Abralia (Compsoteuthis), sp. (immature). 


» Sp. (immature). 


@ 
ROBSON—ON THE CEPHALOPODA 431 
Family 6. Benthoteuthide. 
Chunoteuthis minima, n. gen., n. sp. 
8. Ommatostrephide. 
Stenoteuthis bartrami (Le Sueur). 
11. Chiroteuthide. 
~ Chiroteuthis (Doratopsis) exophthalmica Chun. 
(igopsida consuta. 
Family Cranchiide. 
S.-fam. i. Cranchiine. 
Inocranchia gardineri, n. sp. 
S.-fam. u. Taoniine. 


Taonidium, sp. (immature). 


Suborder 2. OCTOPODA. 
Tribe II. TRACHYGLOSSA. 
Family 3. Polypodidee. 


Scaeurgus umcirrhus Tiberi. 

Polypus fontanianus D’Orb. 
,  arborescens Hoyle. 
»  gardinerr Hoyle. 


»  horridus D’Orb. 

»  ?venustus Rang. 

»  granulatus Lamarck. 
~» J& Isloyle, 

oy des Dy Ds (Oy 


(HGOPSIDA. Family 2. Onychoteuthide. 

1. Onychoteuthis sp. 

One example, sex indeterminate, young, from lat. 20°S. between Maldives and Chagos, 
50 fms. (15387). 

One example, sex indeterminate, young, from between Salomon and Diego Garcia, 
Chagos, surface (1540). 

These examples are very immature, but they exhibit considerable resemblance to young 
examples of this species figured by Pfeffer (16). 

2. Teleoteuthis, sp. 

Two examples, young (7—8 mm.) from Amirante Bank, surface (1539). 

Four * a ? locality (1409). 

The above enumerated examples have been referred to this genus with much hesita- 
tion. The general appearance is in fairly close conformity with young figured specimens 
referred to the genus, (10) and (13); but they are rather more elongate than is apparently 

55—2 


@ 
432 PERCY SLADEN TRUST EXPEDITION 


usual at this stage and the arms are shorter than customary. The suckers of the tentacular 
arms are arranged in a formation more closely serried than those figured by Pfeffer 


and Chun. 


Family 3. Enoploteuthide. 

Abralia (Compsoteuthis), sp. 

One example, sex indeterminate, from off Providence, between surface and 10 fms: 
(1536). | 

The specimen is in a very poor condition, the head and arms being damaged, but 
from the characters that are apparent it is probably referable to this genus. 

Another very badly damaged specimen from 250 fathoms off Desroches Atoll (1532) 
is possibly referable to Abralia also. 


Family 6. Benthoteuthide. 


Chunoteuthis minima, n. gen. n. sp. (Plate 65, fig. 2). 

One example, sex indeterminate, from between Alphonse and Providence Islands, 
900 fathoms (1528). 

This specimen has been the source of a considerable amount of trouble. In the first 
place, it arrived in a very shrivelled-up condition, apparently having suffered desiccation 
at some time, and in consequence a good many of its external features have been 
obliterated. In the second, it does not readily fall into line with any described and figured 
(Egopsid genus. Even its family relationships are very doubtful. The result is that, 
although it certainly calls for description, its exact position is extremely problematical, 
seeing that examination of the anatomy and mantle cavity is denied by the state of its 
preservation. After much hesitation it has been decided to assign it to the Benthoteuthidee 
on the strength of its general superficial appearance. 

Chunoteuthis, n. gen. 

General appearance. Very small in size* (length of mantle from apex to base of 
arms, 3 mm., length of tentacular arms, 5-5 mm. (?+1 mm.), width 1 mm.). The head is 
large and oblong in shape with very large, prominent eyes upon which no traces of 
accessory light organs are to be seen. The body is saccular but laterally compressed} with 
a well-marked dorsal carina and a very prominent nuchal protuberance on the dorsal 
mantle edge. Posteriorly two small subquadrate fins are found. 

The tentacular arms are very long and slender. The club is not very much expanded 
and the suckers upon it are irregularly disposed. The sessile arms were so tightly 
entangled that they could not be separated without damage. They are relatively long 
and apparently subequal in size, and all exhibit prominent suckers. 

The colouring has of course disappeared, but traces of dark chromatophores appear 
upon the arms. 

C. minima, n. sp., with the characters of the genus. 

Type in the Zoological Department of the British Museum. 


* The adult condition is testified by the length of the arms both sessile and tentacular. 
7 It is not impossible that the lateral compression may be an artefact character. 


—— se 


a ee ee eee eee ee eS eee ee 


° 


ROBSON—ON THE CEPHALOPODA 433 


Family 8. Ommatostrephidz. Stenoteuthis. 


S. bartramu (Le Sueur). (Plate 66, fig. 1.) Journ. Acad. Nat. Sci. Philadelphia, 
u. pt. 1821. 90 +t. 7. (For a full synonymy cf. Pfeffer (16).) 

Two examples, $ (young), “flew on board” 30 miles W. of the Centurion Bank 

(Chagos Is.), (1416). 
' A complete account of the anatomy-of this cosmopolitan form is very desirable on 
account of its peculiar “flying” habit. Unfortunately the amount of material to hand 
prohibits the preparation of such an account in the present instance; nor, in fact, does 
the condition of the animal here described from dissection, favour a satisfactory account, 
particularly with regard to the genitalia. 

The branche are slender and elongate resembling a quill pen in general appearance. 
They are relatively very long, in individuals of 80 mm. length measuring 33—34 mm. 
There are 56 (+1) lamine, the distal ones bemg very minute. The scheme of plication, 
which is tolerably simple, is shewn in fig. 1 ¢. 

The anus exhibits two lateral valves, elongate and auriform in appearance. 

Internal anatomy. ; 

The heart (fig. 1b) is fairly regular in shape and is elongate in its sagittal axis. 
The anterior and posterior aort originate at the anterior and posterior extremities. In 
the two specimens there were no traces of auricles. It is possible that the swellings at 
the base of the efferent vessels represent these. 

The branchial hearts are large, irregularly-shaped organs, each exhibiting an 
appendix and partly separated from it by a very deep groove in which the afferent vessel 
terminates. 

Posteriorly the posterior vena cava appears to arise from two very long and narrow 
blood sinuses. From one of these a main vessel was found debouching into the vena cava. 
On the other side the specimen was too badly preserved to allow dissection to trace a 
similar connection; though there is no reason to suppose an asymmetrical condition. 

Such blood sinuses are apparently rare in the Cephalopoda and may be in some way 
related to the animal’s habit of “flying” or leaping from the water. 

The renal papille are found about 2 mm. from the base of the branchiz between 
the latter and the intestine. 

The stomach appears to bear two appendices (fig. 1a), one of these a rounded sac 
lying anteriorly, the other a narrow elongate structure occupying a terminal position. 

The first appears to be plicate internally and may represent the true stomach described 
for Loligo pealew by Verrill (18), while the second may be the equivalent of his “caecal 
lobe” described for the same animal. 

The testis is a thin elongate organ closely applied to the left-hand blood sinus and ter- 
minating anteriorly in a piriform mass of coiled tubes, the external aperture being dorsal 
to the gill heart and just near its posterior extremity. From this aperture project the 
ends of two tubules, apparently the terminal portion of Needham’s sac and the prostate (?). 
It is very much to be regretted that the condition of the single animal available for dissection 
precluded a complete examination of the genitalia. 


Be wee DD) 
a LIBRAR Yi sc! 


oS 


Z\ zee i 2} 
\¥ Lr | 
Os 


se) y te 


434 PERCY SLADEN TRUST EXPEDITION 


It may be taken for granted that this animal does not “fly” in the strict sense 
of the word, i.e. it does not maintain itself or effect progress by repeated muscular 
efforts out of the water. It should more correctly be called the “leaping squid,” as it is 
pretty clear that by the action in question it merely projects itself from the water by a 
single effort. It has been suggested that the occurrence of the animal out of the 
water may possibly be merely accidental. It is a powerful swimmer and it is possible that 
in a rough sea it might accidentally shoot out of the water when swimming close to the 
surface. From D’Orbigny’s account (4), however, in which he describes the considerable 
altitude (15—20 feet) to which it leaps and the way it is preyed on by sea-birds, I am 
inclined to think it executes the characteristic leap voluntarily in order to escape from its 
enemies. D’Orbigny says the habit has been observed in some species of Sepioteuthis as well. 
He is of opinion that it is effected by the well-developed fins on the arms; but, however 
much the latter may facilitate the leap, they can scarcely initiate it. The latter function 
is probably performed by the powerful trunk muscles exercising an abnormal pressure on 
the contents of the mantle cavity. It is possible that the power is possessed by a good 
many other squids, but has only been observed in S. bartrama by reason of its plentiful 


LrocRANCHIA GARDINERI: Fig. 1, Mandibles; Fig. 2, Radula. A. Admedian tooth. (Cam. 4, nom. imm. x 6c.) 


occurrence. 


Family 11. Chiroteuthide. Chiroteuthis. Subgenus Doratopsis. 
Chiroteuthis (Doratopsis) exophthalmica Chun. Deutsche ‘Tiefsee-Expedition. 
ibe, WG, Wt, jo, AHO, WO) 


A single specimen, sex indeterminate, from between Peros and Salomon, Chagos, 
600 fms. (1535). 


Distribution previously known: Madeira and South Indian Ocean. 


The specimen is very much damaged, only the head, tentacles and a small part of the 
body remaining. The identification is provisional. 


Family Cranchiide. Liocranchia. 


Inocranchia gardineri, n. sp. (Plate 65, fig. 1 and Text-figs. 1 and 2.) 
One example, sex indeterminate, from off Desroches Atoll, 200 fms. (1530). 


It is very much to be regretted that only a single specimen of this interesting form 
was obtained, as owing to the size and structure it is impossible to dissect out any of the 
internal organs or even to open the mantle cavity without seriously damaging the specimen. 


ROBSON—ON THE CEPHALOPODA 435 


Dimensions. Max. length of body (apex—base of arms), 10 mm. 
oy WACK ne gg fama, 
Length of tentacular arms, 9 mm. 


The surface of the mantle is smooth, semi-transparent, and of a faint ochreous colour. 
A number of small dark-red chromatophores are found dorsally scattered sparsely over the 
posterior half and disappearing anteriorly and ventrally. A couple of chromatophores 
appear in a dorsal position on the head, one on each side of the median line. 

The body is saccular and rounded posteriorly, where, in a subterminal position, a pair 
of small quadrate fins appear on the back. 

The funnel is large and projects more than 2 mm. beyond the edge of the mantle. 

The sessile arms exhibit the formula 3. 4. 2. 1. 

The tentacular arms. Only one of them is complete. They are solid, truncheon- 
like structures for more than 2°5 mm. of their length, after which they rapidly taper to a 
very fine strand continued to the “club” which is 2 mm. long and well developed. 

The lateral and admedian cartilaginous rows are furnished with numerous pyramidal 
tubercles. There are no median dorsal cartilaginous tubercles as in L. reinhardt. 

The mandibles (cf. Text-fig. 1). 

The radula. The admedian lateral tooth is enlarged so as almost to match the second 
lateral in size. There would appear to be no small marginal tooth, though it is impossible 
to be certain of this point. 

Type in the Zoological Dept. British Museum. 


Cranchiide. S.-fam. ii. Taoniine. 
Taomdium, sp. 


A damaged example, sex indeterminate, juvenile, “16:5:05,” from 125 fathoms (1534). 
Possibly referable to 7. suhmz (Hoyle). 


Family Polypodide. Scaeurgus. 


Scaeurgus umcirrhus Tiberi. (Plate 66, fig. 2.) Bull. Soc. Malac. Ital. 1880, p. 5 
(after Delle Chiaje, n. n.). 

One example, 2, from 123 fms. 8. de Malha (1406). 

a & 9 1B . , (1404). 

Distribution previously known. Mediterranean region. 

The examples are unmistakably representatives of this species, which as far as the 
author has been able to satisfy himself, has never been previously recorded E. of the 
Mediterranean area. 

It is desirable to add some further notes to the already existing knowledge of the 
anatomy of this form. One or two points appear to be of considerable importance. 

To the description of the hectocotylus given by Jatta (9) we may add the fact that 
there is no communication between the main sperm-path (the marginal sulcus) and the deep 
cleft at the extremity of the hectocotylus described by Jatta (p. 237). The function of 
this cleft is therefore somewhat difficult to understand, unless we assume that the whole 
extremity is applied as a sucker for prehensile purposes. 


436 PERCY SLADEN TRUST EXPEDITION 


The male genitalia have been already described by Marchand (13) and do not appear 
to differ in any important respects from those of Polypus. 

The female genitalia in this species appear to differ from those of Polypus somewhat 
(fig. 2). The oviducts leave the ovary in the form of broad, strap-lke ducts, narrow- 
ing down as they approach the oviducal glands. The latter are relatively very large. The 
anterior portions of the oviducts do not again contract on quitting the ovidueal glands but 
continue as stout muscular tubes to the oviducal aperture which is situated upon a round 
elevated papilla of characteristic shape surmounted by a tuft of small (possibly glandular) 
lobes which surround the aperture. 

Coelom and coelomic organs. The limits of the kidney are shewn in fig. 2. It is 
a voluminous organ and possesses extensive glandular bodies. It is divided by the afferent 
and efferent branchial vessels into three portions, the long axes of which are transverse in two 
cases, antero-posterior in the third. The latter is small and contains the reno-pericardial 
orifice which is situated at a very short distance from the entrance to the renal orifice ; 
while the former are extensive and ramify among the digestive organs in such a way as to 
make it difficult to follow them. 


5 5 << 
=) pericardium 


appindx of > 
branchial heart 


Fig. 3. A. Decapop. B. Potypus. C. Scarureus. 


The so-called aquiferous duct leaves the capsule of the ovary from a very anterior 
position and running backwards comes into close contact with the pericardium in the 
external tissues of which it runs as a fine fibrous strand gradually diminishing in size until 
it entirely vanishes near the apex of the pericardium. Very careful dissection of both sides 
failed to reveal any opening of this duct into the pericardium. If this observation should 
be confirmed by future investigators, it will be permissible to suppose that morphologically 
Scaeurgus carries us a step further than Polypus in the obliteration of the coelomic cavity, 
and would represent the third stage in the annexed diagram in which the gradual reduction 
of the coelomic cavity of the Cephalopoda is illustrated (Text-fig. 3). 

This process of separation of the gonad from the remnant of the coelom would be 
finally completed by the condition observed in Argonauta and Philonexis in which the 
aquiferous canals are suppressed altogether (Pelseneer (14)). 

It is necessary to emphasize the fact that these observations are founded upon dis- 
section of a single specimen and that due caution is necessary in employing them for any 
generalization. 

The anus is in the form of a transverse slit at the lateral extremities of which are 
found pedunculate flaps of skin which apparently function as valves and are capable of being 
folded over the aperture. 

The heart (fig. 2 a) is asymmetrical, rectangular on the left side and drawn out by the 


ROBSON—ON THE CEPHALOPODA 437 


posterior aorta upon the right side. Between the right and left efferent vessels and the 
anterior aorta are found two small vessels which run to the surface of the renal capsules. 
The genital artery leaves the heart in a median position in the antero-ventral region. The 
auricles are small, simple and roughly triangular in shape. 

The branchie are shewn in fig. 2 0. 


Polypus. 
i, 1B, Jontanianus (D’Orbigny). (Plate 66, fig. 5.) Amérique Méridionale, p. 28, 
1835. 


One example, ?, Coetivy (1405). 
Distribution previously known: Peruvian and Patagonian regions. Indian Ocean. 


Originally regarded as a South American form, this species has recently been recorded 
from the Indian Ocean (Joubin (12)). The enlargement of the proximal suckers of both 
lateral arms, regarded by D’Orbigny as a character diagnostic of the species but now found 
to be a sex-limited character found in other species, has been already discussed by Hoyle (5). 
It may possibly serve a prehensile function similar to that discharged by the enlarged 
thumb of male frogs. Insemination being effected among the Cephalopoda by means of 
the hectocotylus it would appear unnecessary for the male to be supplied with prehensile 
organs. Very little, however, is known of the actual process of sperm-transference and 
coitus, and it may be possible that some form of amplexus takes place in some species of 
Polypus in which the enlarged suckers seen in P. fontanianus and P. gardineri are used. 
That amplexus may take place in some members of a genus and not in others is attested 
_ by Boulenger's description (1) of fecundation in Batrachians, according to which amplexus 
occurs in Molge aspera and M. montana but not in M. cristata and M. alpestris. 

The mandibles. The upper is erect and exhibits a well-pointed rostrum ; in the lower 
the anterior laminz are provided with a marked posterior median angulation. 

In the radula the median tooth appears to exhibit three phases, one with lateral 
cusps placed about half-way down from the apex of the median cusp, one with similar cusps 
placed lower down and one with cusps in both positions. The third lateral tooth is stout, 
recurved at the tip and its base is well developed. 

The anus is situated within a circular depression in the mantle cavity and is furnished - 
with two elongate valves. 

The heart is markedly asymmetrical. The genital artery leaves it towards the right 
side, the anterior aorta well on the left with the root of the posterior aorta occupying the 
opposite angle. Shortly after it leaves the ventricle the anterior aorta gives off a small 
vessel which apparently runs to the stomach and intestine. 

The right auricle was found to be in an abnormal condition inasmuch as it was en- 
larged to a size exceeding that of the ventricle itself. Dissection revealed the presence 
in it of a mass of coagulated blood. The left auricle being, as is usual in the Cephalopoda, 
of small size, one is forced to conclude that the distension of the right auricle is abnormal, 
due either to pathological causes or to some temporary derangement of the circulation due 
possibly to the circumstances of the animal’s death. 

SECOND SERIES—ZOOLOGY, VOL. XVII. 56 


438 PERCY SLADEN TRUST EXPEDITION 


The genitalia (?) do not call for any particular comment. The oviducal glands are 
very small, the ovary is symmetrical and the oviducal apertures are simple and placed 
very near the anus. The genital artery enters the capsule of the ovary in an anterior 
position between the two ducts which are separate at their origin. 

The branchie (tig. 5 b, c) have their laminee diagonally plicate. 


2. Polypus arborescens Hoyle. (Plate 66, fig. 3.) Rept. Pearl Oyster Fisheries of 
Ceylon, (Roy. Soc.) Pt. 1. p. 189, 1904. 


One example, ?, from Cargados, 30 fms. (1411). 

One example, ?, from same (1403) (dark variety 2). 

These are unmistakable examples of Hoyle’s species. 

Distribution previously known: Ceylon; Zanzibar; Pacific Ocean. 

The mandibles (ef. fig. 3 b). 

The radula. The first lateral tooth is rather deep, the second singularly long and 
accompanied by a rather slender third lateral. The median tooth exhibits a very deep 
basal notch. 

The internal organs. In the example No. 1411 the ovary was so much enlarged and 
reflected posteriorly that it covered the whole of the branchial complex and practically 
obscured the gills themselves from view. Apparently by reason of this development of the 
ovary (no doubt attained by numerous other species at the time of sexual maturity) the 
following somewhat anomalous condition of the branchial complex and surrounding organs 
is to be noted : 

(1) the gills lie at a very low level, 

(ii) the heart is displaced very much to the right of the median line, 

(ui) the oviducts instead of originating from the dorsal surface of the ovary and 
running downwards are found well past the middle of the mantle cavity on the ventral side 
of the ovary, running first upwards and then downwards. 

The other example, which is very much darker in colour, shewed no such development 
of the ovary, the gills were more posterior in position and the oviducts pursued a more 
normal course. The heart, however, is still placed well towards the right-hand side. Com- 
paring the two examples we may safely conclude that at sexual maturity the extensive 
development of the ovary tends to displace the organs of the branchial complex in the 
manner described above. 

In both forms the heart is peculiarly elongate and piriform, the posterior aorta 
occupying the apex and the two efferent vessels and the anterior aorta the base. 

3. Polypus gardineri Hoyle. Fauna and Geography of the Maldive and Laccadive 
Archipelago (J. 8. Gardiner). Cephalopoda. Vol. i. Supp. 1. p. 976. 

One example, ¢, from Coetivy, 32 fathoms (1419). 

It was impossible to make any dissection of this example owing to the imperfect state 
of preservation. Superficially it agrees in all respects with Hoyle’s type description, the 
lateral arms exhibiting enlarged suckers (v. supra, p. 437). 

Hoyle supposed that this form might possibly be the young of P. fontanianus; but 
comparison of the radula of these two forms does not support this possibility. 


ROBSON—ON THE CEPHALOPODA 439 


The hectocotylus. This was referred to by Hoyle as comparable to that of P. vulgaris. 
It should however be pointed out that in the present example of P. gardineri the hecto- 
cotylus differs from that of P. vulgaris in that the seminal groove is well covered over by 
its external edge, though it is possible that the loose external edge may be contracted or 
relaxed according to the state of preservation. 

The radula (Text-fig. 4). 


Babe 


Fig. 4. Potypus GaRpinErt: Radula. Fig. 5. PoLtypus Horripus: Radula and 
Cam. 6 oc. x 6 obj. Reich. Mandibles. Cam. 6 oe. x 6 obj. Reich. 


4. Polypus horridus D’Orb. Tabl. méthod. Ann. Sei. Nat. vii. p. 54, 1826. 


One example, ¢, from Amirante, 20—25 fathoms (1418). 
&; 280 4 (AOL), 
- sex indeterminate, young, Amirante, 30 fathoms (1420). 
Distribution previously known: Zanzibar; South Africa; Red Sea; Ceylon; Malé Atoll. 
Mandibles. The distal extremity of. the anterior ramus of the lower mandible is 
markedly turned back and expanded. The upper mandible is very erect and anteriorly 
presents a remarkably straight front. 
Radula. The first laterals are elongate. The second laterals exhibit a very heavy 
external inferior angle, while the third laterals are tolerably stout. (Text-fig. 5.) 


5. Polypus venustus Rang. Magasin de Zoologie, Ann. 5-8, 1835-8, fig. 93. 

One example, $, from between Peros and Salomon, Chagos, 20 fathoms (1541). 

The author has been unable to find any modern work upon this species. There is a 
brief reference to it in Tryon’s Manual of Conchology, vol. 1, where the distribution is 
given as ‘“‘Aloiers: Island of Goree.’ Tryon places it among his indeterminable species 
and considers it to be a young specimen. 

The specimen obtained from the above locality agrees with Rang’s description (and 
figure) pretty closely. In one or two respects it differs, e.g. the chromatophores are not 
aggregated into a median dorsal clump but are found clustered towards the anterior end - 
of the mantle. Nor are the ventral chromatophores disposed in the sparse even rows 
indicated by Rang’s figure but are more irregularly distributed. Otherwise the description 
given by Rang applies very adequately to this specimen. Particularly significant is the 
agreement on the following characters : “‘un peu dorée a la partie dorsale ” and “ les bras... 
avec des séries de petites taches dorées répondant aux ventouses.” 


56—2 


440 PERCY SLADEN TRUST EXPEDITION 


Tryon’s supposition that this is a young form is borne out by the delicacy and fragility 
of the tissues but by nothing else. The condition of the arms and suckers and the 
general appearance, which is peculiar and characteristic, do not suggest that it is young. 

It is very much to be regretted that only a single specimen was found, otherwise 
some interesting anatomical characters might have been revealed. The branchial cavity 
alone was opened and found to reveal at least one interesting feature. As far as the radula 
is concerned there is nothing to separate it from a normal Polypus, but the single mandible 
examined is somewhat peculiar. 

The branchie occupy a peculiar position in the mantle cavity, being inserted into the 
visceral mass at a remarkably high point, well up into the apex of the mantle cavity. In 
form they are extremely long, flat and strap-like. 

The radula (Text-fig. 6) is normal. In the median tooth there appear to be two main 
lateral cusps, the separation and approximation of which in different teeth give rise to 
two main types indicated in the figure. 


Sas 


So 


Fig. 6. Ponypus venustus: Radula. Cam. 6 oc. x 6 obj. Reich. 


The lower mandible alone was examined, the upper member of the pair being somewhat 
damaged. It is peculiar in exhibiting an excessively deep notch and a very small anterior 
lamina. 

Distribution previously known: ‘* Algiers: Goree.” 


6. Polypus granulatus Lamarck. Mém. Soc. Hist. Nat. Paris, i. p. 20, 1799. 


One example, about one-third grown, ¢ (?), from Amirante, 22—85 fathoms (1402). 
I share Dr Hoyle’s view (5, p. 80) as to the difficulty of identifying the S. rugosa of 
Bose to which this form has been referred by Férussac and D’Orbigny. 


7. Polypus L. Hoyle. Fauna and Geography Lace. and Mald. Archip. (J. 8. 
Gardiner), Vol. 11. Supp. 1. 1905. | 

One example, sex indeterminate, young, Amirante, 20—40 fathoms (1407). 

This young example agrees in all respects with the form recorded by Hoyle, except 
that there are two, not three, ventral chromatophores. 


8. Polypus P. (Plate 65, fig. 3; Plate 66, fig. 4.) 


One example, ?, from off Peros, Diamant Is., 12 fathoms (1410). 

This specimen proved to be too young to be treated as a new species. It exhibits 
affinities with P. horsti, P. fontamanus and P. gardineri, but at the same time appears 
to be specifically distinct from these forms. 

The body is posteriorly rounded and sac-like, the eyes are tolerably prominent and 
between the latter the head is slightly wider than the body. The skin is wrinkled 


ROBSON—ON THE CEPHALOPODA 44] 


dorsally, probably by the action of the preserving agent, and exhibits a sparse array of 
papillae some of which are multifid. There are a few papillze round each eye. 

The arms exhibit the formula 2=3=4.1. There is a weakly developed umbrellar 
membrane extending a few millimetres up each arm. ; 

The colouring is a brownish purple on the back, passing to a very pale yellowish 
brown ventrally, the whole covered with numerous black chromatophores. Very charac- 
teristic is an oval patch of an intense blue-black colour, about 2 mm. long, situated on 
either side somewhat posterior to the eye at the base of the third arm. Such patches are 
found in this position in P. horsti (Joubin (12)), but although this is a common character- 
istic between the two species, the radule, as well as other structures, differ too widely 
to allow of uniting the two forms (cf. Plate 66, fig. 4a and Text-fig. of Hoyle (6)). 

Radula (ef. fig. 4 a). 

Internal anatomy. The heart is relatively very large and is markedly rounded ; the 
efferent vessels enter it at equal distances from the anterior aorta. The auricles lie very 
near the surface in close proximity to the renal aperture. 

The ovary, in accordance with the immature age of the specimen, is very small, 
much smaller in fact than the capsule. The oviducal glands are also very small and the 
oviducts long and slender. These open low down in the mantle cavity towards the median 
line in the region of the kidney. 


INDETERMINATE JUVENILE EXAMPLES. 


A single very small example from “6. 10. 05,” 140 fathoms (1538). The only salient 
characters are its squat build and the plentiful scattering of brownish-black chromatophores 
all over the mantle. 

A single juvenile example from Providence, 39 fathoms (1408). Possibly young form 
of P. granulatus. ; 


442 PERCY SLADEN TRUST EXPEDITION 


fo tS 


GOR Ss Exe E> 


Fig. 
Fig. 
Fig. 


Fig. 


Fig. 
Fig. 


Fig. 
Fig. 


LIST OF WORKS REFERRED TO. 


BouLEenGceEr, G. A. Zool. Jahrb. Bd. vi. Syst. p. 447. 1892. 
Cuun, C. Deutsche Tief-See Exped. Oegopsida. 1910. 
i Zool. Anzeig. 31, p. 82, 1906. 
D°Orsieny, A. D. Voy. Amérique Mérid. vol. 5, p. 56. 1847. 
Hoyts, W. E. “Challenger” Rep. Cephalopoda. 1886. 
P P. Z. S. 1907, p. 450. 
5 Rept. Pearl Oyster Fishery, Gulf of Manaar, (Roy. Soc.) Pt. 1. 1904. 


x Fauna and Geography of the Maldive and Laccadive Archipelago (J. 8S. Gardiner). 
Cephalopoda (vol. 2, suppl. 1.). 1905. 


Jarra, G. Flora und Fauna des Golfes v. Neapel: Cephalopoda. 1896. 
JouBin, L. Rés. Camp. Scient. (d’Albert I.): 
IX. Céphalopodes de Atlant. Nord. 1895. 

3 XVII. 3 prov. des campagnes de la “Princesse Alice.” 1900. 

3 Notes: Leyden Museum, xx. pp. 21—28. 1898. 
Marcuanp, W. Zeitschr. fiir Wiss. Zool. 86. 1907. 
PELSENEER, P. Lankester’s Treatise on Zoology. Mollusca. 1906. 
PFEFFER, J. Jahrb. Hamburg. Wiss. Anstalt. xvit. 1910. 

% Plankton Exped. Oigopsiden. 1912. 
PossELT, H. J. Vidensk. Meddelels. Natur. For. 1891, p. 301. 
VeERRILL, A. E. Trans. Connecticut Acad. Sci. v. 1879. 


EXPLANATION OF PLATES. 


PLATE 65. 


1. Lnocranchia gardineri, n. sp., anterior and posterior aspects (x 5). 
2. Chunoteuthis minima, n. gen., n. sp., anterior and posterior aspects (x 17). 
3. Polypus P., anterior aspect ; 3a, funnel; 30, oral aspect shewing disposition of suckers (x 2). 


PLATE 66. 
1. Stenoteuthis bartrama Lesueur, general appearance of mantle cavity; la, stomach and caeca; 
1b, heart; 1c, lamine of gills, seen from above. 
2. Scaeurgus unicirrhus Tiberi, genitalia, etc.; 2a, heart; 26, gill. 
3. Polypus arborescens Hoyle, female in breeding season; 3a, diagrammatic representation of the 
heart and female genitalia shewing asymmetrical condition of these organs due to growth of ovary 


at sexual maturity; the dotted outline shews normal position of these organs; 36, mandibles ; 
3c, radula (4 oc. x 6 obj.). 


4. Polypus P., heart and genitalia of female; 4a, radula (4 oc. x 6 obj.). 


5. Polypus fontamanus (D’Orbigny), heart shewing enlarged right efferent artery, probably patho- 
logical; 5a, ovary, etc.; 5b, gill; 5c, a single gill lamella; 5d, mandibles; 5e, radula. 


Lerrerine: AA=anterior aorta; A, a=auricle; a.d.=aquiferous duct; 6.h.=branchial heart; c.o.=capsule of ovary ; 


e.v.=efferent branchial vessel; GA=genital artery; g.o.—genital aperture; LAV=left efferent vessel; MHS. V.=me- 
senteric vessel; od=oviduct; ov=oviducal gland; PA=posterior aorta; RHV=right efferent vessel; 7.p.a.=reno- 
pericardial aperture; 7, y=renal vessels. 


Percy Stapen Trust ExPEpDITION. 


PL, OS 


TURANN'S, JLIUNIN, SOC. SEIR. 2 ZOOIL, WOlL, XA 


(Roxson) 


net 


Eats. 


Fig. 3b. 


Fig. 3. 


CEPHALOPODA 


Fig. 3a. 


Percy SLADEN Trust ExPEDITION. 
(Rosson) 


TRANS. ILJUNIN, SOC. SER, 2 ZOOL, WOIL, XW IIL, G6 


Gti MELEE 
So 


fi 
S 


PN 
= 
=n 


MARGINAL 


CEPHALOPODA 


[Nore.—Synonyms and Native Names are printed in italics. 


Abralia, Gray, sp., 430, 432. 
Acanthastrea = seq. 
Acanthastrea, M. Edw. & Haime, 
17, 78. 
bowerbanki, M. Edw. & Haime, 
112, 113. 
brevis, M. Edw. & Haime, 101, 
102. 
dipsacea, H. M. Kdw., 102. 
grandis, M. Edw. & Haime, 
101, 102. 
hirsuta, M. Edw. & Haime, 101, 
102. 
var. megalostoma, Klunz., 
102. 
var. microstoma, Klunz., 
110-112. 
irregularis, Quelch, 101, 102. 
spinosa, M. Edw. & Haime, 101, 
102. 
Acanthephyra, A. M. Hdw., men- 
tioned, 327. 
Acervularia ananas, M. Edw. & 
Haime, mentioned, 99. 
Achilinz, 286, 287. 
Acropona gardineri, Distant*, 315; 
ftnote, 314. 
prasina, Melich., 314; men- 
tioned, ftnote, 315. 
sladeni, Distant*, 315. 
Acropora, Oken, mentioned, 33, 121. 
Actinia equina, Zinn., mentioned, 
112: 
mesembryanthemum 
Sol.), mentioned, 8. 
Adolendana, Distant*, 278. 
typica, Distant*, 279. 
Z&geon, Gosse, 411. 
medius (Ale. & And.), 411. 
rugulosus, Borrad., 411. 
Abgina citrea, Hschsch., 200. 


(Eu. & 


INDEX. 


appear to be used for the first time.] 


A star is added to names which 


/&quorea, Pérond Lesweur, 188-190. 
macrodactyla (Brandt), 188, 
189; mentioned, 170, 171. 
maldivensis, Browne, 188, 189, 
190. 
parva, Browne, mentioned, 188, 
189. 
Agallia quadrinotata, Melich., 309. 
Aglaura, Péron & Lesueur, 196,197. 
hemistoma, Péron & Lesueur, 
196; mentioned, 171, 172. 
var. laterna, Maas, 196. 
var. nausicaa, Maas, 196. 
var. octogona, Mayer, 196. 


—— var. prismatica, Mayer, 
196. 
laterna, Haeck., 196. 
nausicaa, Haeck., 196. 
octogona, Bigelow, 196. 
prismatica, Maas, 196. 
Ahasuerus advena(Wadltl.), 141,142, 
145. 
Alcyonium digitatum, Zinn., men 
tioned, ftnote, 31. 
Alemo seychellensis, Wright, 265. 
Algol, Sollas, mentioned, 235. 
Alpheide, 400, 413-428. 
Alpheopsis, 1. Cout., mentioned, 420. 
chilensis, H. Cout., mentioned, 


421. 
equalis, H. Cout., mentioned, 
414, 421. 


fissipes, H. Cout., 414, 421; 
mentioned, 420. 
idiocarpus, H. Cout., 414. 
socialis, H. Cowé., mentioned, 
421. 
trispinosus, H.Cout.,mentioned, 
414, 421. 
Alpheus, Fabr., 418; mentioned, 
341, 349. 


Alpheus aculeipes, H. Cout., 423; 
mentioned, 424. 
adamastor, H. Cout., 425. 
alcyone, de Man, 424. 
alpheopsides, H. Cout., 427. 
amethysteus, Risso, 370. 
amirantei, H. Cout., 421. 
architectus, de Man, mentioned, 
423. 
ascensionis, Orém., mentioned, 
423. 
audouini, H. Cout., 427. 
baculifer, H. Couwt., 422. 
bastardi, H. Cout., 427. 
bouvieri, 7. WM. Edw., 427. 
bradypus, H. Cout., 423. 
bucephalus, H. Cowt., 423. 
clypeatus, H. Cout., 425. 
coetivensis, H. Cowt., 427. 
collumianus, Stimps., 419. 
crinitus, Dana, mentioned, 422. 
cylindricus, Kingsley, 
tioned, 422. 
dasycheles, H. Cout., 426. 
edwardstz, Couch, 395. 
frontalis, H. If, Edw., sp. dub., 
425, 
gracilipes, Stimps., mentioned, 
426. 
gracilis, Heller, 419. 
var. Alluaudi, H. Cout., 


men- 


419, 
hailstonei, H. Cout., 419; men- 
tioned, 418. 


hippothoé, de Man, 427. 
insignis, Heller, 426. 
lanceloti, H. Cout., 426. 
leptochirus, H. Cowt., 427. 
longecarinatus, Hilgend., 426. 
lutini, H. Cowt., 423. 
macrochirus, Richters, 419. 


444 


Alpheus malhaensis, H, Cout., 419. 
megacheles, Hazlstone, 418, 

419. 

microstylus, H. Cout., 423. 

obeso-manus, Dana, mentioned, 
422. 

ovaliceps, H, Cout., 423. 

pachychirus, Stemps., 425; men- 
tioned, 425. 

pacificus, Dana, 427. 

paracrinitus, M/%ers, mentioned, 
427. 

var. bengalensis, 1. Cout., 
mentioned, 427. 

paraculeipes, H. Couwt., 423; 


mentioned, 424. 
paradentipes, H. Cout., 419. 
paragracilis, H. Cout., 419, 421. 
paralcyone, H. Cout., 424. 
paralpheopsides, H. Cout., 427. 
parvirostris, Dana, 427. 
percyi, H. Cout., 426. 
phrygianus, H. Cout., 423. 
pinnophylax, Otto, 390. 
providencei, H. Cout., 424. 
rostratipes, Pocock, 419, 420, 

421. 
scriptus, Risso, 362. 
seurati, H. Cout., 419. 
sp. (2), 419, 421. 
splendidus, H. Couwt., 426. 
stanleyi, H. Cout., 423. 
staphylinus, 7. Cout., 418. 
strenuus, Dana, 427. 
angulatus, H. Cout., 427. 

styliceps, H. Cout., mentioned, 
423. 
superciliaris, H. Cout., 425. 
ventrosus, H. M. Hdw., 419. 
Ambarvalia, Distant*, 306. 
pyrops, Distant*, 306. 
Amphibrachium _ euplectelle 
(Schulze), mentioned, 216. 
infestans, Dendy, nom. tent., 
217. 
Amphigona pusilla, Hardl., 197. 
Amphinema, Haeck., sp. (2), 170, 
172, 181. 
Awmphioxus, Yarrell, 23. 
Amphipalemon, Borrad., 406-408; 
mentioned, 342, 398, 405. 
cooperi, Borrad., 407. 
gardineri, Borrad., 407, 408. 
willeyi (Borrad.), 407. 
Amphogona apsteini (Vanhéffen), 
Wl, 1725 19% 


INDEX 


Anchistia, Dana, 329, 359. 
aesopia, Bate, 354. 
amboinensis, de Man, 366. 
americana, Kingsley, 371. 
amethystea, Heller, 370. 
aurantiaca, Dana, 376. 
brachiata, Stimps., 370. 
brocku, de Man, 363. 
dane, Stimps., 370. 
edwardst, Pauls., 371. 
elegans, Pauls., 371. 
ensifrons, Dana, 370. 
ensifrons, de Man, 370, 371. 
gracilis, Dana, 363. 
grandis, Stimps., 370. 
inequimana, Heller, 369. 
korn, Lo Bianco, 354. 
longimana, Dana, 370. 
notata, Heller, 363. 
petitthouarsiz, de Man, 369, 

370. 
var. spinigera, de Man, 
369. 
scripta, Heller, 362. 
spinigera, Ortm., 380. 
tenella, S. J. Sm., 363. 
tenuipes, Holmes, 372. 
Anchistioides, Pawls., 405, 406. 
Anchistioidide, 405—408. 


| Anchistus, Borrad., 330-338, 342- 


350, 387-389. 
armatus (H. IM. Edw.), 389. 
biunguiculatus, Borrad., 388. 
inermis (A/iers), 388. 
miersi (de Man), 324, 388. 
mirabilis (Pesta), 388, 389. 
spinuliferus (MMers), 388. 
Ancyclocaris, Schenkel, 327-338, 
346, 350, 355. 
aberrans (Wobil2), 355, 356. 


brevicarpalis, Schenkel, 355, 
356. 
hermitensis (Rathbun), 355, 


356. 
latirostris (Lenz), 355, 356. 
Anemonia sulcata, WM. Hdw. & 
Haime, 15. 
Annelids, mentioned, 238. 
Anthomedusa, Browne (2), 185. 
Anthomeduse, 170, 173-185. 
Anthophyllum, Ehrb., 58. 
cuspidatum, Dana, 59. 
fasciculare, Ehrb., 59, 61. 
hystrix, Dana, 59. 
musicale (Ehrb.), 59, 62. 
spherula, Ehrb., 64, 65. 


Antipathes, Pallas, mentioned, 12. 
Aphaniptera, 161. 
Aphrastrea (WM. Hdw. & Haime), 
122. 3 
deformis (Zam.), 122. 
Aquelicium, Distant*, 289, 290. 
brunnescens, Distant*, 290. 
elegantulum, Distant*, 289. 
typicum, Distant*, 289. 
Arete, Kinb., mentioned, 420. 
equalis, H. Cout., 413. - 
indicus, H. Cowt., 413. 
Argonauta, Zinn., mentioned, 436. 
Armilustrium, Distant*, 295. 
gardineri, Distant*, 296. 
scotti, Distant*, 296. 
Aselgeia, Walk., mentioned, 276. 
Aselgeoides, Distant*, 275. 
insularis, Distant*, 276. 
Astacus tyrrhenus, Petagna, 328, 
390. 
Asteropus, Sollas, 251-252; men- 
tioned, 235. 
haeckeli, Dendy, 251, 252. 
simplex (Carter), 226, 251. 
Astrea = Astrea. 
Astreide possessing distinct Coral- 
lites, by G. Matthai, 1-140. 
Astrangia asiatica, Mich., 97. 
solitaria, Zeswewr, mentioned, 2. 
Astrea, Lam., 38, 49, 72, 77, 78, 
115. 
abdita, Lam., 91, 93. 
afinis, Gard., 89, 114. 
ananas, Lam., 79, 84. 
var. stellis amplioribus, 
Lam., 87. 
annularis, Lam., 76. 
var. 2, Lam., 103. 
annuligera, M. Edw. & Haime, 
103. 
argus, Lam., 75. 
coarctata (Duchass. & Mich.), 


104. 

dane (M. Edw. & Haime), 
87. 

deformis, Ehrb., 79, 83, 116, 
122. 


deformis, Lam., 79, 122. 

denticulata (Ell. & Sol.), 113, 
114. 

denticulata, Lam., 79, 83. 

dipsacea, Lam., 100, 101, 102, 
110, 112. 

doreyensis, M. Edw. & Haime, 
91. 


Astrea forskaliana, M. Edw. & 

Haime, 54. 

Sragilis, Gard., 87. 

fusco-viridis, Quoy & Gaimard, 
91. 

halicora, Ehrb., 106, 107. 

heliopora, Lam., 72, 74. 

hemprichu, Ehrb., 110, 111. 

hombront, Verr., 107. 

interstincta, Oken, 43. 

laperousiana, M.Kdw. & Haime, 
103. 

lobata, Gard., 102. 

magnifica, Dana, 91. 

melicerum, Khrb., 95, 97. 

microphthalma, Lam., 41, 43. 

ocellina, Dana, 41. : 

okent, Gard., 84, 86, 87. 

ordinata, Verr., 87. 

pallida (Dana), Gard., 113, 
114, 

pandanus, Dana, 91. 

pectinata, Gard., 121. 

pentagona, Ehrb., 91, 93. 

pleiades, Lam., 102. 

puteolina, Gard., 80. 

retiformis, Lam., 118, 119. 

rotulosa, Lam., 84, 86, 87. 

rotumana, Gard., 114. 

solidior, M. Edw. & Haime, 
105. 

speciosa, Dana, 89, 91. 

spongia, Khrb., 118. 

tesserifera, Ehrb., 109, 110. 

versipora, Lam., 103. 

Astroides calycularis, I, Edw. & 
Haime, 3. 

Astropeplus pulcher, Sollas, 252, 253. 

Athanas djiboutensis, H. Cout., 413. 

Athysanus, Burm., 316. 
frontalis, Distant*, 316. 
insularis, Distant, mentioned, 

314. 

Atolla, Haeck., 203-204. 
bairdii, Fewkes, 203. 
chuni, Vanhéffen, 203. 
verrillii, Fewkes, 203, 204. 
wyvillei, Haeck., 171, 172, 203, 

204. 

Aulactinia, Verr., 28. 

Aulocalyx, Schulze, 211. 
irregularis, Schulze, 211-214. 
serialis, Dendy*, 211-214. 

Aurora, Sollas, 242-251. 
aurora, Hentschel, 

247, 250, 251. 


243, 244, 


INDEX 


Aurora cribriporosa, Dendy*, 226, 
235, 243-249, 259. 


distincta (Thiele), 245, 249, 
259. 

globostellata (Carter), 243, 245, 
247. 

membranacea (Hentschel), 243, 
251, 


providentiz, Dendy*, 226, 243, 
245-247, 249. 

reticulata, Dendy*, 244. 

rowi, Dendy*, 226, 243, 245, 
249-251. 

sterrastrea, Row, 243, 245, 251, 
259. 

Automate salomoni, H. Cout., 413. 


Balanophyllia regia, Jowrd., 12, 14. 
Balclutha, Kirkp., 318. 
chersonesia, Distant*, 317. 
varicolor, Distant*, 318. 
Baryastrea = seq. 
Baryastrea, M. Edw. & Haime, 
66. 
chrenbergana, M. Edw. & 
Haime, 69, 70. 
solida, M. Edw. & Haime, 69, 70. 
transversa, M. Edw. & Haime, 
60. 
Benthoteuthide, 431, 432. 
Betzeus utricola, Richters, 425. 
Borradaile, L. A., Pontoniine, 323— 
396 ; Carides, 397-412. 


_Bougainvillia, Less., mentioned, 170. 


fulva, Agass. & Mayer, 170- 
172, 178, 179. 
Brixia, Stal, 284, 285; mentioned, 
276. 
mahensis, Distant*, 284. 
stellata, Distant*, 284. 
Browne, E. T., Medusz, 169-210. 
Bunodes, Grosse, 22, 23. 
thallia, Gosse, 8. 
Bythoscopus, Germ., 
tioned, 307. 
bimaculicollis, Stal, 309. 
indicus, Distant, 308. 


men- 


308 ; 


Callianassa thyrrhenus, Risso, 390. 

Calpanthula, van Beneden, ftnote, 
16. 

Campanulina, Hincks, mentioned, 
170. 

Cancer custos, Forsk., 391. 

Caneirona, Distant, 276, 277. 

indica, Distant, ftnote, 276. 


SECOND SERIES—ZOOLOGY, VOL. XVII. 


445 


Caneirona maculipennis, Distant*, 
277. 

Carides from the Western Indian 
Ocean, by L. A. Borradaile, 
397-412. 

Carmarina hastata, Haeck., 199. 

Carmaris rosea, Agass. & Mayer, 
199. 

Carmentalia, Distant*, 296. 

biformis, Distant*, 296. 

Caryophyllia (pars), Lam., 58. 

astreata, Lam., 64, 65. 

Jasciculata, Lam., 59. 

musicalis, Lam., 62, 64. 

smithii, Stokes & Brod., ftnote, 
59. 

Cassiopea andromeda var. maldi- 
vensis, Browne, 172, 207. 

Catharius advena, Reitter, 145 

Sascipennis, Reitter, 146. 

Cephalopoda, by G. C. Robson, 429— 
442; distrib., 429. 

Cercopide, 306-307. 

Cerianthula, van Beneden, ftnote, 16. 

Charybdeidee, 202. 

Charybdea, Pér. & Les., sp., 171, 
172, 202. 

Chelusa seychellensis, Distant, men- 
tioned, 314. 

Chiroteuthide, 431, 434. 

Chiroteuthis exophthalmica, Chun, 
431, 434. ; 

Chondrilla, Schmidt, 267-269 ; men- 
tioned, 230. 

australiensis, Carter, 226, 267- 
269. 

grandistellata, V’heele, 245, 268, 
269. 

mixta, Schulze, 226, 267-269. 

sacciformis, Carter, 226, 245, 
268, 269. 

Chondrosiide, 226, 267-269. 

Chunoteuthis, G. C. &obs.*, 431, 
432. 

minima, G. C’. Robs.*, 431, 432. 

Chypastrea oblita, Duchass. & Mich., 
39, 69, 70. 

Cicada pulverulenta, Distant, 275. 

Cicadidee, 274-275. 

Cixius, Zatr., mentioned, 276. 
Cladocera, M. Edw. & Haime, men- 
tioned, 33. 

arbuscula 
tioned, 2. 

Cladopsammia rolandi, Lac.-Duth., 

ftnote, 31. 


(Lesweur), men- 


o7 


446 


Claviscopulia intermedia, Schulze, 
208. 
Clusivius, Distant*, 277. 
spectabilis, Distant*, 277. 
Celida indica, Walk., 315. 
Cenopsammia, Uv, Edw. & Haime, 
ftnote 3, 7, 9, 11, 12, 13, 14, 16, 
ftnotes 17 & 18, 29, 31. 
Coleoptera: Cucujide, Cryptopha- 
gide, by A. Grouvelle, 141-156. 
Coléura seychellensis (?), 162. 
Colonial. Astreeide possessing dis- 
tinct Corallites, by G. Matthai, 
1-140. 
Colpophyllia gyrosa (Hil. & Sol.), 
mentioned, 2. 
Colydium frumentarium, Fabr., 
144. 
Comanthus annulatus(?), mentioned, 
364. 
sp., mentioned, 377. 
Commolenda, Distant, mentioned, 
276. 
Conchodytes, Peters, 330-350, 392— 
394; mentioned, 325, 328. 
biunguiculata (Pauls.),392,394. 
domestica (Gibbes), 392-394. 
margarita (Verr.), 392, 394. 
meleagrine, Peters, 324, 334, 
ftnote 337, 392, 393. 
tridacne, Peters, 324, 334, 
-ftnote 337, 392, 393. 
Consualia, Distant*, 304. 
robusta, Distant*, 304. 
Coppatias albescens, Row, 2538, 254. 
distinctus, Thiele, 243. 
amconditus, Tops., 253. 
johnstoni, Tops., 253. 
Coralliocaris, 327-352, 
381-386. 
agassizi, H. Cout., 386. 
atlantica, Rathbun, 382, 385. 
aualitica, Vobili, 385. 
brevirostris, Borrad., 382, 384. 
camerani, Vobili, 382, 383. 
graminea (Dana), 324, 383. 
hecate, Vobihi, 382, 385. 
inequalis, Ortm., 383. 
japonica, Orim., 324, 337, 341, 
383, 384. 
lamellirostris, Stimps., 384. 
lucina, Vobilt, 383, 384. 
macrophthalma (H. IZ. Hdvw.), 
324, 337, 382, 383. 
nudirostris (Heller), 324, 382, 
384, 


Stumps., 


INDEX 


Coralliocaris quadridentata, Rath- 
bun, 382. 
rathbuni, Distant*, 385. 
rhodope, Mobili, 332, 385. 
superba (Dana), 337, 383, 384. 


var. japonica, Ortm., 


384. 
tridentata, Miers, 386. 
tridentata, Rathb., 385. 
truncata, Rathb., 382, 385. 
Coronate, 202—204. 
Corymorpha, Sars, 173. 
nana, Alder, 173. 
nutans, Sars, 173. 
Corynactis viridus, Adlm., 14. 
Coutiére, H., Alpheidze, 413-428. 
Coutierea, Nobili, 329, 330, 332, 
345, 347, 349, 350, 386. 
agassizi (H. Cout.), 386. 
Cranchiide, 431, 434. 
Crangon, Fabr., 410. 
Crangonide, 411. 
Cryptamorpha desjardinsi (Guér.), 
142, 150. 
fasciata, Wollaston, 146. 
muse, Wollaston, 150. 
Cryptophagide, 154-156. 
Cryptophagus adventa, Waltl., 145. 
guerini, Alib., 145. 
striatus, Rouget, 145. 
Ctenocephalus felis (Bouché), 161. 
Cucujide, 143-144. 
Cunina lativentris (Gegenbd.), 202. 
sp., 171, 172, 201. 
Curiatius, Distant*, 285. 
insignis, Distant*, 285. 
Cylicia, M. Hdw. & Haime, 31, 
32. 
Cyphastrea = seq. 
Cyphastrea, Klunz., 37-48, 70, 77; 
mentioned, 117. 
aspera, Quelch, 43, 44. 
bottce = seq. 
bottat, M. Edw. & Haime, 39, 
40, 69, 70. 
brueggemanni, Quelch, 41, 43. 
capitata, Stud., 39, 41. 
chaleidicum, Klunz., 29, 41-- 
48, 68. 
dana, M. Edw. & Haime, 41. 
Jorskaelana, Gard., 39. 
Sorskaliana, Vaughan, 39. 
gardineri, Matthai*, 48. 
hemprichana, Gard., 47, 48. 
incrustans, Klunz., 39. 


maldivensis, Gard., 45, 48. 


Cyphastrea microphthalma (Lam.), 
41, 43-48. 

muelleri, M. Edw. & Haime, 
43, 44, 45. 

muellert, Ortm., 45, 48. 

(2) ocellina, M. Edw. & Haime, 
41. 

ocellana, Stud., 41, 46. 

pleiades (EU. & Sol.), 39. 

savignyt, M. Edw. & Haime, 


41-47. 

serailia (Forsk.), 7, 31, 39-47, 
70, 93. 

suvadive, Gard., 41, 42, 45, 
46, 47. 


Cyteesis, Hschsch., 177-178. 
herdmani, Browne, 177, 178. 
macrogaster, Haeck., 177. 
nigritina, Haeck., 177. 
pusilla, Gegenb., 177. 
tetrastyla, Hschsch., 170, 171, 

Tit, TUR. 
vulgaris, Agass. & Mayer, 177. 
vulgaris, Bigelow, 177. 


Dagama, Distant, mentioned, 317. 
Daradascoides, Distant*, 286. 
mahensis, Distant*, 286. 
Daradax, Walk., mentioned, 286. 
Deferundata, Distant*, 295. 
aldabrana, Distant*, 295. 

Dendroclava[dohrnii], Weissm., 181. 

Dendrophagus sutwralis, White, 150. 

Dendrophyllia gracilis, , Hdw. & 
Haime, 14. 

Dendy, A., Hexactinellid Sponges 
(Triaxonida), 211-224; Homo- 
sclerophoraand Astrotetraxonida, 
225-271. 

Dennisia, Norman, 359. 

sagittefera, Norman, 362. 
Derbide, mentioned, 291. 
Dercitopsis, Dendy, 228-230. 

ceylonica, Dendy, 228, 229, 

230. 

clathrata (Kirkp.), 229. 

mammillaris (Lendenf.), 229. 

minor, Dendy*, 226, 229-230; 

mentioned, 267. 

oxeata (Tops.), 229. 

Dermestes swrinamensis, Linn., 144. 

seadentatus, Fabr., 144. 

Desmocaris, Sollaud, 326-328. 

Diastra sterrastraea, ow, 235, 243. 

Dichoceenia stokesi, WM. Hdw. & 
Haime, mentioned, 2. 


Dipleurosoma, Aael Boeck, men- 
tioned, 190. 

Diploastrea, Matthai*, 37, 72. 

heliopora (Zam.), 24, 72; men- 
tioned, 35, 75. 

Dipsasirea, Blainv., 77, 115. 
deformis, Blainv., 122. 
denticulata, Blainv., 79. 
Javosa, Blainv., 112. 
solida, Blainy., 117. 
versipora, Blainv., 103. 

Diptera Pupipara, 161-167. 

Distant, W. L., Rhynchota, Part II, 

Suborder Homoptera, 273-321. 

Donatia, Vardo, 260-267. 
cliftoni (Bowerb.), 265. 
ingalli (Bowerb.), 226, 261, 

263-265. 
japonica (Sollas), 226, 261-265. 
lyncurtum, Auct., 226, 262, 264. 
lyneurium, Wardo, 226, 260, 
261. 
parvistella, Baer, 262. 
robusta, Dendy, 265. 
seychellensis( Wright), 226,261, 
264, 265. 
stella-grandis, 
266. 
tylota, Hentschel, 266. 

Donatiide, 226, 259-267. 

Dorypleres incrustans, Tops., 253. 
biangulata, Lindgren, 253, 254. 

Dragmastra, Sollas, 237-239. 
lactea (Carter), var. mauritiana, 

Dendy*, 226, 238, 239. 


Dendy*, 226, 


Fichinastrea, Blainv., 48. 
gemmacea, Blainv., 54. 
rosularia, Blainv., 50. 

Kchinophyllia, Klwnz., 49. 

Echinopora, Lam., 27, 37, 38, 48— 

58; mentioned, 5, 118. 

aspera (M. Edw. & Haime), 49. 

carduus, Klunz., 54, 58. 

concamerata, Klunz., 54,56, 57. 

ehrenbergi, M. Edw. & Haime, 
54, 56, 58. 

flexuosa, Verr., 50, 51. 

Sruticulosa, Klunz., 54, 56, 58. 

gemmacea (Lam.), 50, 53, 54— 
58; mentioned, 5. 

helli, Rousseau, 52, 54, 57. 

hemprichi, M. Edw. & Haime, 
54, 56. 

hirsutissima, M. Edw. & Haime, 
50-55, 57; mentioned, 5. 


INDEX 


Kchinopora horrida, Dana, 50, 51. 
lamellosa (Hsper), 22, 50-58. 
magna, Gard., 49. 
reflexa, Dana, 50. 
ringens, Dana, 52, 54. 
rosularia, Dana, 50-54. 
rousseaur, M. Edw. & Haime, 54. 
solidior, M. Edw. & Haime, 52— 

56. 
tertia, Gard., 52, 54. 
undulata, Dana, 50. 
KEchinothrix turcarum (?), 
tioned, 386. 

Ecionemia, Bowerb., 241, 242. 
acervus, Bowerb., 241. 
carteri, Dendy, 226, 242. 
laviniensis, Dendy, 226, 242. 

Edwardsia, Quatrefages, 22, 23. 

Elidiptera madagascariensis, Sign., 

= 29, 

Emporius, Ganglbauer, 145. 

Enoploteuthid, 430, 432. 

Hogypona walkeri, Kirkp., 314. 

Equirria, Distant*, 290, 291. 
phalaena, Distant*, 290. 

Erylide, 226, 256-259. 

Erylus, Gray, 256-259. 
carteri, Sollas, 259. 
cylindrigerus, [dley, 259. 
lendenfeldi, Sollas, 226, 257, 

258; mentioned, 230. 
proximus, Dendy*, 226, 258, 
259. 
Kuphyllia glabrescens, Bourne, 9, 11, 
14; mentioned, 2. 

Euphysa, orbes, 173. 
aurata, Yorbes, 173. 
tetrabrachia, Bigelow, 173. 

Euphysora, Maas, 173-177. 
bigelowi, Maas, 170-174. 
tetrabrachia, Bigelow, 173. 
valdivie, Vanhéfien, 173, 175. 

Euplectella aspergillum, Owen, 217. 

Eurete erectum, 7. L. Schulze, 218. 

Luryplatus, Motsch., 150. 

Euryrhynchus, Miers, 329. 

Explanaria, Lam., 38, 48. 
annularis, Ehrb., 76. 
argus, Ehrb., 75, 76. 
aspera, M. Edw. & Haime, 

49. 
galaxia, Khrb., 43, 44. 
gemmacea, Lam., 52, 54, 56. 
var. stellis comosis, Lam., 


men- 


52, 
hemprichi, Ehrb., 54, 56. 


AA7 


Pavastrea magnifica, Blainv., 91. 
Favia (Oken), ftnote 10, 36, 37, 
38, 77, 78, 90, 94, 114-117, 122; 
mentioned, 3, 5. 
abdita (HU. & Sol.), 34, 36, 91, 
93, 108, 110, 116. 
acropora (Linn.), 79, 102. 
adduensis, Gard., 97, 112. 
afinis, Gard., 79, 83, 89. 
amicorum, M. Edw. & Haime, 
79, 83, 84. 
amplior, M. Edw. & Haime, 86, 
87, 90. 
ananas, Hil. & Sol., 27, 98. 
aspera, M. Edw. & Haime, 80, 
83. 
bertholleti (Valenc.), 82, 83, 94, 
95, 112, 113, 116. 
bowerbanki, I. Hdw. & Haime, 


113. 

cavernosa, Klunz., 83, 84, 85, 
89, 90. 

clouei, WZ. Hdw. & Haime, 80, 
83, 89, 90. 

complanata, Hhrb., 84, 109, 110, 
111. 

deformata, M. Edw. & Haime, 
80, 83. 

denticulata, Ehrb., 79, 80, 83, 
84, 86, 117. 


dipsacea (Zam.), 102. 
doreyensis, WZ. Hdw. & Haime, 
13, 36, 53, 81-88, 101, 105, 
116; mentioned, 74. 
ehrenbergi, Klunz., 80-84, 106, 
107. 
—— yar. laticollis, Klunz., 83. 
var. sulcata, Klunz., 83. 
favosa (HII. & Sol.), 79, 112. 
favus (Morsk.), 16, 26, 79, 83, 
84, 92, 93, 96, 113, 116, 117; 
mentioned, 5. 


fragum (Hsper), 113; men- 
tioned, 2. 

geoftroyi, M. Edw. & Haime, 
80, 83. 


halicora (Zhrb.), 106, 107, 116. 

hawatiensis, Vaughan, 96. 

hemprichii (Zhrd.), 110-112. 

hirsuta (I. Edw. & Haime), 
79, 100, 109. 

hombroni (Rousseaw), 107. 

hululensis, Gard., 53, 87, 100. 

jacquinott, M. Kdw. & Haime, 
79, 80, 83. 

laccadivica, Gard., 84, 85. 


57—2 


448 


Favia laxa (Klunz.), 99, 105. 
lobata, Klunz., 102, 103. 
microphthalma, Ehrb., 39, 41. 
okeni, M. Edw. & Haime, 89, 

90; mentioned, 5. 
parvimurata, Gard., 113. 
pentagona (sper), 92, 95, 97, 

116, 119. 
radiata, Oken, 89. 
rotulosa, Bhrb., 86, 87, 88, 115. 
rousseaut, M. Edw. & Haime, 

79, 83. 
savignyt, M. Edw. & Haime, 

80, 83, 102; mentioned, 5. 
solidior (I. Edw. & Haime), 

105. 
tubulifera, Klunz., 89, 90, 91. 
urvilleana, WZ. Hdw. & Haime, 

86. 
uva, Ehrb., 89, 90. 
vasta (Klwnz.), 79, 108, 110. 
versipora, Khrb., 34, 79, 80, 82, 

105. 
wakayana (Gard.), 34, 106. 

Feronia spinifera, Veach, 162. 

Fescennia, Stal, 288. 
aurea, Distant*, 288. 
bimaculata, Distant*, 288. 

Flabellum, Zesson, 17, 29, 31, 32, 

ftnote, 34; mentioned, 7. 

Flatoides, Guér., 300. 
protea, Distant*, 300. 

Fordicidia, Distant*, 290, 291. 
robusta, Distant*, 291. 

Fulgoride, 275. . 

Fungia (Lam.), 14, 29. 


Galaxea (Oken), 20, 35, 37, 38, 58— 
66. 

aspera, Quelch, 60, 62. 

astreata, IZ. Edw. & Haime, 65. 

bougainvillei (Blainv.), 59. 

cespitosa, Dana, 60, 62. 

clavus, M. Edw. & Haime, 62, 
64. 

cuspidata, Oken, 59. 

ellist, M. Edw. & Haime, 60,61, 
62. 

esperi, Sche., 2. 

explanata, Quelch, 65. 

fascicularis (Limn.), 9, 20, 22, 
24, 59-65, 85. 

fragilis, Quelch, 62. 

heterocyathus, Ortm., 62, 64. 

hexagonalis, I. Hdw. & Haime, 
59, 65. 


INDEX 


Galaxea irregularis, M. Edw. & 
Haime, 60—62. 

lamarcki, M. Edw. & Haime, 
64, 65. 

laperouseanea (Ml. Edw. & 
Haime), 59, 65-66. 

musicalis (Linn.), 32, 61-67. 

organum, Oken, 64. 

pauciradiata (Blainv.), 59. 

tenella, Briigg., sp. dub., 62, 64. 

Galboa typica, Dist., mentioned, 
ftnote, 308. 

Geodia, Lam., 254-256. 

auroristella, Dendy*, 226, 254— 
256. 

canaliculata, Schmidt, 243. 

carteri, Sollas, 243. 

globostellifera (Carter), men- 
tioned, 256. 

stellosa, Dendy, 244. 

variospiculosa, Dendy, 244. 

Geodiide, 226, 254-256. 

Geryones mexicana, Agass. & Mayer, 
199. 

Geryonia hexaphylla, Maas, 199. 

proboscidalis (Forsk.), 171,172, 
199. 
Glyphocrangon, MW. Hdw., 410, 411. 
ceca (Wood-Mason), 411. 

Glyphocrangonide, 411. 

Gnathodus, Fieb., 318. 

Gnathophyllide, 408-410. 

Gnathophyllum, Latr., 398, 408, 
409, 410. 

americanus, Latr., 409. 
fasciolatum, Séimps., 409, 410. 
tyrrhenus, Desm., 390. 

zebra, Richters, 409. 

Goniastreea = seq. 

Goniastrea (1. Hdw. & Haime), 
ftnote 10, 12, 37, 38, 77, 78, 82, 
87, 115-122. 

bournoni, M. Edw. & Haime, 
118, 120. 

cerium (Dana), 120. 

coronalis, Quelch, 120, 121. 

eximia, Gard., 118. 

favistella (Dana), 93, 116. 

Javus, Klunz., 96, 116-119. 

grayi, M. Edw. & Haime, 120, 
121. 

halicora (Hhrb.), 117. 

f. obtusa, Gard., 106. 

var. acuta, Klunz., 106— 

109, 116. 


laxa, Quelch, 95, 97, 116, 117. 


Goniastrea multilobata, Quelch, 116. 
pectinata, Klunz., 13, 34, 116, 
118, 120-121. 
planulata, . Hdw. & Haime, 
116, 121, 122. 
quoyt, M. Edw. & Haime, 120, 
121. 
retiformis (Zam.), 14, 20, 36, 
51, 118-120. 
rudis, M. Edw. & Haime, 95, 
97, 116. 
serrata, Ortm., 84, 87, 116. 
seychellensis, Klunz., 80, 83, 
116, 117. 
solida (IZ, Edw. & Haime), 13, 
117-121. 
Grouvelle, A., Coleoptera: Cucujide, 
Cryptophagide, 141-156. 
Gypona prasina, Walk., 314. 
Gyractis, Bovert, 25, 35. 


Halicreas papillosum, Vanhaffen, 
171, 172, 195-196. 
Hapalips, Reitter, 142. 
championi, Grouwv.*, 142, 154, 
155. 
scotti, Growv.*, 142,144,155, 156. 
Haplaxius, Fowler, mentioned, 279. 
Harpiliopsis, Borrad.*, 329-338, 
341-343, 347, 349, 350, 379-80. 
beaupresi (Audouin), 324, 338, 
379. 
depressus (Stimps.), 324, 379, 
380. 
Harpilius, Dana, 327-332, 337, 338, 
342, 343, 347, 349, 350, 380, 381. 
beawpresir, Heller, 380. 
consobrinus, de Man, 380, 381. 
depressus, Stimps., 380. 
gerlachei, Vobilz, 380, 381. 
tmermis, Miers, 388. 
lutescens, Dana, 337, 341, 380, 
381. 
miersi, de Man, 388. 
spinuliferus, Miers, 388. 
Heliastrea, M. Edw. & Haime, 49, 
72, 77-79. 
acropora, M. Edw. & Haime, 
103, 119. 
annularis, M. Edw. & Haime, 
76, 77, 115. 
annuligera, M. Edw. & Haime, 
103, 104. 
cavernosa (Esper), 75, 76, 79. 
conferata, M. Hdw. & Haime, 
75, 76, 79. 


Heliastrea forskelana, M. Edw. & 
Haime, 54, 58; mentioned, 5. 

gigas, M. Edw. & Haime, 75,79. 

heliopora, M. Edw. & Haime, 
72, 74; mentioned, 4, 105. 

lamarckana, M. Edw. & Haime, 
75, 76, 79. 

laperouseana, M. Edw.& Haime, 
103, 104. 

quadrangularis, M. Kdw., 79. 

radiata, M. Kdw., 79. 

solidior, M. Edw. & Haime, 


105, 106. 
stellulata, M. Edw. & Haime, 
76, 77, 79. 
Heliotites interstincta, WW. Hdw. & 
Havme, 45. 


Hensenanthula, van Beneden, 16. 
Heterocarpus affinis, Borrad., 399. 
unicarinatus, Borrad., 399. 
Heterocyathus quicostatus, J, 
Edw. & Haime 7, ftnote 13, 14. 
Heterorete, Dendy*, 214. 
pulchra, Dendy*, 211, 214. 
Heterotiara, Maas, 183. 
anonyma, Maas, 183. 
minor, Vanhéffen, 170,171, 183. 
Hexactinellid Sponges (Triaxonida), 
by A. Dendy, 211—224. 
Hippoboscide, 161-162. 
Hippolysmata, Stimps., 403; men- 
tioned, 397. 
kikenthali (de Man), 403. 
vittata (Stumps.), 403. 
Hippolyte gibberosus, M. Edw. & 
Haime, 401. 
kiikenthalt (de Man), 403. 
marmoratus, M. Edw. & Haime, 
401. 

Hippolytide, 401-404. ~ 
Homosclerophora and Astrotetrax- 
onida, by A. Dendy, 225-271. 
Hydnophora, Fischer de Waldh., 

ftnote, 19. 
Hydromeduse, 170, 171, 173. 
Hymenocera, Latr., 398, 405, 408, 
409, 410. 
ceratophthalma, Balss, 408, 409, 
410. 
elegans, Heller, 410. 
nipponensis, De Haan, 391. 


Idiocerus, Lewis, 307. 
scotti, Distant*, 307. 
Igerna, Kirkp., 308. 
bimaculicollis, Jacobi, 309. 


INDEX 


Iguvium, Distant*, 287-288. 

albomaculatum, Distant*, 287— 
288. 
Ino, Cast., 150. 
Inopeplini, 150-151. 
Inopeplus, Smith, 142, 150. 
mimetes, Growv.*, 142, 151. 
pictus, Cast., 142. 
Trene, Bedoé, spp., 170, 171, 172, 187. 
Isaurus asymmetricus, Hadd. & 
Shackl., 16. 

Isophyllia dipsacea, Dana, men- 
tioned, 2. 

Isops membranacea, Hentschel, 242, 
243. 

Ivinga typica, Distant, 293. 


Jaspis, Gray, 252-254. 
biangulata, Thiele, 253, 254. 
johnstonii (Schmidt), 226, 253— 

254. j 

Jassus, Fabr., 315, 316. 
deplanatus, Spangb., 315. 
determinatus, Distant*, 316. 
indicus, Distant, 315. 


Ketumala, Distant, 299-300. 

rubromarginata, Distant*, 299— 
300. 

Kolla, Distant, 311-312. - 
funeralis, Distant*, 312. 
seychellensis, Distant*, 312. 

Kosalya, Distant, mentioned, 286. 

Kronos, Distant*, 307-308. 
typicus, Distant*, 308. 


Lemophleus, Cast., 141, 143. 
mirus, Growv., 141, 142, 143. 
propior, Grouv.*, 141, 142, 143, 

144, 147. 
proximus, Growv., 144. 

Laodice, Less., sp., 172, 186. 

Latridius museorum, Ziegler, 145. 

Latrunculia, Barboza du Bocage, sp., 

mentioned, 238. 
Leander, Desm., mentioned, 330, 
339, 341, 404. 
debilis, Stimps., 397, 404, 405. 
gardineri, Borrad., 404. 
serratus, Borrad., 339. 
“ Leaping squid” (Stenoteuthis bar- 
tramt), 434. 
Leptastrea = seq. 
Leptastrea (I, Hdw. & Haime), 37, 
38, 66-72, 75; mentioned, 5. 
bottar, Klunz., 69, 70, 71. 


449 


Leptastrea ehrenbergana, M. dw. 
& Haime, 68, 69, 71. 
immersa, Klunz., 68, 69, 70. 
inequalis, Klunz., 69, 70. 
roissyana, WM. Hdw. & Haime, 
22, 36, 61, 66-72. 
solida, WW. Hdw. & Haime, 69, 
7, Til 
transversa, Klunz., 67, 68. 
Leptochela robusta, Stemps., 398. 
Leptomedusa, Browne, No. 1, 190; 
No. 2, 191; No. 3, 191. 
Leptomeduse, 170, 186-191. 
Leucandra australiensis, Dendy, 
mentioned, 216. 
phillipensis, Dendy, mentioned, 
216. 
Leuckartiara, Hartl., 181-182. 
gardineri, Browne*, 170, 172, 
181. 
Ligur uvee (Borrad.), 397, 401. 
Limentinus, Distant*, 316. 
aldabranus, Distant*, 316. 
Liocranchia gardineri, G'. C. Robs.*, 
431, 434. 
reinhardti, Pfef:, mentioned, 
435. 
Lipeurus subsignatus, Gebel, 161. 
Liriope, Zess., 170, 197-198. 
sp.? 171, 198-199. 
tetraphylla (Cham. & Lys.), 171, 
172, 198. 
Lodoicea sechellarum, Zabzll., men- 
tioned, 155. 
Loligo, Zam., mentioned, 429. 
pealeii, Zes., mentioned, 433. 
Lollius, Stal, 293, 294. 
atromaculatus, Distant*, 294. 
furcifer, Stal, 294. 
gratiosus, 
294. 
virescens, Distant*, 294. 
Lynchia maura (Bigot), 162. 
Lysmata affinis (Borrad.), 402-403. 


Melic., mentioned, 


chiltoni, Kemp, mentioned, 
402 

seticauda (isso), mentioned, 
402. 


Lysmatella, Borrad., 397, 403-404. 
prima, Borrad., 404. 


Macropsis indica, Leth., 308. 
Madrepora, Linn., 33, 35. 
abdita, Ell. & Sol., 91, 93. 
acropora, Linn., 102. 
ananas, Ell. & Sol., 84, 98, 99. 
57—3 


450 


Madrepora annularis, Hil. & Sol., 
76. 
cavernosa (Forsk.), 75, 79, 82, 
84, 90; mentioned, 89. 
chaleidium, Forsk., 41, 42. 
cusprdata, Esper, 59. 
denticulata, Ell. & Sol., 79, 
83. 
divergens, Forsk., 62. 
Jascicularis, Linn., 59, 62. 
Javosa, Esper, 91, 112, 113. 
favus (Morsk.), 79, 82, 83, 95, 
117, 118. 
merustans, Forsk., 39, 41. 
intersepta, Esper, 75. 
interstincta, Kisper, 43, 45. 
lamellosa, Esper, 50. 
musicalis, Linn., 62. 
organum, Pallas, 64. 
pentagona, Hsper, 95. 
pleiades, HU/. & Sol., 39, 77. 
radiata, Esper, 89. 
rotulosa, Ell. & Sol., 78, 84, 
87, 115. 
serailia, Forsk., 39, 40, 43, 45. 
solida, Forsk., 117, 118. 
stellulata, E7/. d Sol., 76 
wva, Esper, 89. 
Madreporaria, Classific. discussed, 
2-5, 34. 
Magog sacciformis, Sollas, 268, 269. 
Maiestas, Distant*, 312, 313. 
illustris, Distant*, 313. 
Mallophaga, Aphaniptera, and Dip- 
tera Pupipara, by H. Scott, 161— 
167. ; 
Manicina, W/. Hdw. & Haime, 28, 
29. 
areolata (Zinn.), 7; mentioned, 
2. 
Margaritophora fimbriata (?), men- 
tioned, 394. 
Marygrande, Pesta, 387. 
mirabilis, Pesta, 349, 389. 
Matsumurana, Distant*, 317. 
facialis, Distant*, 317-318. 
Matthai, G., Colonial Astraeide pos- 
sessing distinct Corallites, 1-140. 
Matutinus, Distant*, 278. 
opulentus, Distant*, 278. 
Meandrina labyrinthica (HU. & 
Sol.), mentioned, 2. 
Meduse, by E. T. Browne, 169- 
210. 
Meenoplus atrovenosus, Leth., 291. 
Meleagrina, Zam., mentioned, 393. 


INDEX 


Merhippolyte orientalis, de Man, 403. 
Mesonema, Lschsch., 188. 
celum pensile, Modeer, 188. 
macrodactylum, Maas, 189. 
pensile (Modeer), 170, 171, 
188-189. 
Metalpheus, (?) H. Cout.*, 419. 
Metapenzeus commensalis (?), 343. 
Metridium, Oken, mentioned, 23. 
dianthus, Oken, ftnote, 17. 
marginatum, WM. Hdw., 23, 
ftnote, 25. 
Mimotelliz, Mats., mentioned, 317. 
Modeeria multitentacula, Fewkes, 
180. 
Molge alpestris (Zam.), mentioned, 
437. 
aspera (Gray), mentioned, 437. 
cristata (Zam.), mentioned, 
437. 
montana (?), mentioned, 437. 
Monanus concinnulus (Walk.), 141, 
142, 146. 
denticulatus, Grouv., 141, 142, 


145. 
ornatus, Grouv.*, 141, 142, 
144, 146. 


Mussa corymbosa, Dana, 2, 9. 
Mycedium okeni, WU. Hdw. & Haime, 
13, 17, 26, 49. 
Myopsida, d’Orb., mentioned, 429. 
Myriastra, Sollas, 235-237. 
cavernosa, Dendy*, 226, 236— 
237. 
parva (Row), 226, 235, 
simplicifurca, Sollas, 236. 


Narcomedusz, 171, 199-202. 
Nausithée punctata, K6lliker, 171, 
172, 202. 
Nehela, Buch. White, 
mentioned, 307. 
aterrima, Distant*, 311. 
bimaculicollis, Distant*, 309. 
conspicua, Distant*, 310. 
elegantula, Distant*, 309, 310. 
Distant*, 310 


308-311; 


flavolineata, 
311. 
lineoligera, Distant*, 310. 
scutellata, Distant*, 310. 
spectabilis, Distant*, 309. 
Neoprivesa, Distant*, 297. 
fuscovaria, Distant*, 297. 
Nika processa, Bate, 410. 
Nikoides maldivensis, Borrad., 398, 
411. 


? 


Nilaparvata, Distant, 303, 304. 
mahensis, Distant*, 304. 


| Nisia, Melich., 291-293. 


atrovenosa, Melich., 291-292. * 
fuscofasciata, Distant*, 292, 
293. 
maculosa, Distant*, 292, 293. 
sulphurata, Distant*, 292. 
thoracica, Distant*, 293. 
Normania tenuilaminaris, Sollas, 
230. 
Northea, Hook. f., sp., mentioned, 
279, 315. 
Nycteribia, Zatr., 163-167. 
bellardii, Rondant, mentioned, 
163. 
fryeri, H. Scott*, 163-167. 
stichotricha, Speiser, 163. 


| Nycteribiide, 162-167. 


Obelia, Péron d: Lesuewr, men- 
tioned, 170. 
Oceania nutricula, MeCrady, 180. 
Ocypode ceratophthalmus (Pallas), 
332. 
Cidipus, Dana, 381; mentioned, 
328. 
dentirostris, Pauls., 383. 
graminea (Dana), 383. 
macrophthalmus, Dana, 383. 
superbus, Dana, 383. 
Olfersia maura, Bigot, 162. 
Oliarus, Stal, mentioned, 276. 
Olindias singularis, Browne, 171, 
172, 192. 
tenuis, Browne, 192. 
Ommatostrephidee, 431, 433. 
Oncopsis nigritus, Melich., 309. 
Onychoteuthide, 430, 431. 
Onychoteuthis, Lechtenstern, 
431. a 
Opiconsiva, Distant*, 301-303. 
balteata, Distant*, 302. 
colorata, Distant*, 301-302. 
derelicta, Distant*, 303. 
fuscoraria, Distant*, 301. 
gloriosa, Distant*, 302-303. 
insularis, Distant*, 303. 
modesta, Distant*, 303. 
Orbicella, Klunz., 49, 66, 72, 77, 
78, 79; mentioned, 3. 
acropora, Linn., 102, 103. 
annularis, Dana, 76, 77; men- 
tioned, 2. 
annuligera, Gard., 103, 104. 
borradailei, Gard., 84, 85. 


430, 


Orbicella botiar, Gard., 69. 
coronata, Gard., 106. 
curta, Gard., 106. 
ehrenbergana, Gard., 67, 69. 
Sorskalana, Kilunz., 54, 58. 
Junafutensis, Gard., 105. 
helipora, Gard., 105. 
ammersa, Gard., 70. 
klunzingeri, Gard., 69. 
lasca, Klunz., 84-86, 99-100. 
lobata, Marenz., 102, 103. 
mammillosa, Klunz., 5, 54, 58. 
minikowensis, Gard., 2, 72, 74. 
orton, Gard., 102. 
rotumana, Gard., 106. 
solidior, Gard., 105. 
stellulata, Dana, 76. 
versypora, Gard., 104. 
wakayana, Gard., 104, 105. 
Ormensis madagascariensis, Stal, 
299. 
Oryzephilus surinamensis (Linn.), 
141, 142, 144. 
Osaka hyalina, Distant, 299. 
relata, Distant, 299. 
Oulastrea, M. Edw. & Haime, 77. 
Ovactis brasiliensis, Had., 16. 


Pachastrella, Schinidt, 230. 
crassiuscula, Wendenf., 230, 
232. 
tenuilaminaris (Sodlas), 
230-232. 
Pachastrellide, 226, 230-234. 
Pachynus, Stal, 308. 
bimaculicollis, Stal, 309. 
Pachyopsis chlorophana, Melich., 
308. 
Paganalia, Distant*, 314. 
virescens, Distant*, 314. 
Palemon, Fabr., 328, 342. 
beaupresii, Aud., 379. 
debilis, Dana, 404. 
marmoratus, Olivier, 401. 
petitthouarsti, Aud., 369. 
Palemonella, Dana, 327-338, 342— 
350, 356-359. 
affinis, Zehnter, 357, 358. 
amboinensis, Zehnter, 357, 358. 
batei, Borrad.*, 333, 357, 358. 
biunguiculata, Vobil2, 357. 
elegans, Borrad., 323, 357, 
359. 
gracilis, Pauls., 359. 
laccadivensis, Alcock & Anders., 
345, 357, 358. : 


226 


? 


INDEX 


Palemoneila longirostris, Borrad., 
323,-357, 359. 
orientalis, Dana, 356-3857, 358. 
orientalis, Rathbun, 358. 
rathbunensis, Borrad.*, 357, 
358. 
tenuipes, Dana, 345, 357, 358. 
tridentata, Borrad., 323, 357, 
359. 
yucatanica, Ives, 357, 358. 
Palemonetes, Heller, 329. 
Palemonide, 404—405 ; sub-families, 
326-328, 404. 
Palemonopsis willeyi, Borrad., 405. 
Pamendanga, Distant, mentioned, 
287. 
Pandea, Less., juv., 170, 171, 182. 
conica (Quoy & Gawm.), men- 
tioned, 183. 
Pandalide, 398-400. 
Pandalus gracilis, Borrad., 398. 
Pandanus Hornei, Balf. 7, men- 
tioned, 299. 


Pantachogon apstein, Vanhoffen, 
197. 
rubrum, Vanhoften, 171, 172, 


195. 

Parakosalya, Distant*, 286-287. 
insularis, Distant*, 287. 

Paralimnus, Matsum., 320. 
silhouettensis, Distant*, 320. 

Parasirea = seq. 

Parastrea, M. Edw. & Haime, 77. 
afinis, M. Kdw. & Haime, 79. 
amicorum, M. Edw. & Haime, 

79. 
amplior, M. Edw. & Haime, 84. 
ananas, M. Edw. & Haime, 84. 
clouei, Valenc., 89. 
deformata, M. Edw. & Haime, 


79. 

denticulata, M. Edw. & Haime, 
79. 

doreyensis, M. Edw. & Haime, 
84. 


hombroni, Rousseau, 107. 
lobata, M. Edw. & Haime, 102. 
radiata, M. Kdw, & Haime, 89, 
90. 
rotulosa, M. Edw. & Haime, 84. 
rousseaui, M. Edw. & Haime, 


79. ° 

savignyi, M. Edw. & Haime, 
79. 

urvilliana, M. Edw. & Haime, 
84, 


451 


Paratypton siebenrocki, Balss, ft- 
note, 323. 
Pasipheide, 398-400. 
Peachia, Gosse, 22. 
hastata (Grosse), 8. 
Pelagia, Péron & Lesuewr, 170, 204— 
206. 
flaveola, 
206. 
flaveola, Mayer, 205. 
panopyra (Péron & Les.), 171, 
204, 206. 
papillata, Haeck., 205. 
sp.? A., 171, 205. 
sp.? B., 206. 
tahitiana, Agass. & Mayer, 205, 
206. 
Pelias, Roux, 359. 
amethysteus, Roux, 370. 


Eschsch., 171, 204 


? 


scriptus, Roux, 362. 
Periclimenzus, Borrad., 329, 330, 
338, 342-350, 377-379. 
fimbriatus, Borrad., 324, 378, 
379. 
robustus, Borrad., 324, 378. 
Periclimenes, Costa, 327-333, 336- 
352, 359-377. 
aftinis, Borrad., 324, 367, 372; 
mentioned, 373. 
amboinensis (de Man), 365, 


366. 

americanus (Kingsley), 368, 
371. 

amethysteus (Azsso), 328, 360, 
370. 


amymone, de Man, 368, 371. 

aurantiacus (Dana), 349, 360, 
376. 

borradailet, Nobili, 364. 

borradailei, Rathbun, 324, 331, 
369, 372, 376. 

brachiatus, Stimps., 369, 370. 

brevinaris, Nobili, 331, 364. 

brocketti, Borrad., 324, 368, 
374, 


brocki (de Many 394, 361, 
363. 

ceratophthalmus, Borrad., 324, 
332, 365. 

commensalis, Borrad., 361, 
364. 

compressus, Borrad., 324, 333, 
368, 373. 


cornutus, Borrad., 324, 365. 
danz (Stimps.), 367, 370. 
demain, Kemp, ftnote, 323. 


452 


Periclimenes denticulatus, Mobzli, 
341, 368, 372. 
dubius, Borrad., 324, 367, 373. 
edwardsi (Pauls.), 366, 371. 
elegans (Pauls.), 367, 371; 
mentioned, 373. 
ensifrons (Dana), 331, 367, 
370. 
ensifrons, de Man, 371. 
frater, Borrad., 324, 333, 361, 
364. 
gorgonidarum (Badlss), sp. dub., 
376. 
gracilis (Dana), 362, 363. 
grandis (Sézmps.), 367, 370. 
hermitensis, Rathbun, 376. 
hertwigi, Balss, 376. 
holmesi, Wobili, 368, 372. 
incertus, Borrad., 324, 362, 
364. 
insignis, Costa, 362. 
kolumaduluensis, Borrad., 324, 
331, 369, 376. 
korni (Lo Bianco), 354. 
lifuensis, Borrad., 366, 371. 
longimanus (Dana), 368, 370. 
nilandensis, Borrad., 324, 367, 
372. 
notatus (Heller), 362, 363. 
nove-zealandie, Borrad., ft- 
note, 323. 
parasiticus, Borrad., 362, 363. 
parvus, Borrad., 362, 363. 
petitthouarsi (Awd.), 341, 352, 
366, 369. 
var. denticulata, Nobili, 
372. 
var. spinifera, de Man, 
369. 
potina, Nobili, 362, 363. 
pottsi, Borrad., 369, 374. 
pusillus, Rathbun, 362, 363. 
rotumanus, Gorrad., 368, 371. 
seriptus (Asso), 328, 361, 362. 


seychellensis, Borrad., 324, 
369, 375. 

soror, Wobili, 361, 362; men- 
tioned, 364. 


sp., de Man, 376. 

spiniferus (de Man), 324, 331, 
341, 352, 366, 380. 

suvadivensis, Borrad., 324, 369, 
375. 

tenellus, S. J. Sm., 345, 361, 
363. ; 

tenurtpes, Borrad., 372. 


INDEX 


Periclimenes tenuipes (Leach), sp. 
dub., 376. 
vitiensis, Borrad., 324, 367, 371. 
Perigonimus, Sas, mentioned, 170. 
Peripatus, Lands., 23. 
Periphragella elise, Marshall, 213. 
Peyerimhoff, P. de; Description 
de la Larve et de la Nymphe 
de Prostominia convexiuscula, 
Grouv., 156-159. 
Phialidium, Zewck., mentioned, 191; 
spp., 170, 171, 172; sp. A., 186; 


sp. B., 187. 
Philonexis, d’Orb., mentioned, 346. 
Pheenicopterus minor, (Geoffroy, 
mentioned, 161. 
antiquorum, Zemm., men- 
tioned, 161. 


Phromnia rubra, Sign., mentioned, 
299. 
Phyllangia americana, UM. Hdw. & 
Haime, mentioned, 2. 
Phyllognathia, Borrad., 332, 409, 
410. 
ceratophthalma (Balss), 409. 
Phymasirea = seq. 
Phymastrea, M. Edw. & Haime, 77, 
78, 79. 
aspera, Quelch, 94, 95, 
valenciennensti, M. Edw. & 
Haime, 94, 95. 
Pilochrota (2) lactea, Sollas, 238. 


Placospongia, Gray, mentioned, 
243. 
Plakinastrella clathrata, Kirkp., 
228. 


copiosa, Schulze, 228. 
mammillaris, Lendenf., 228. 
oxeata, Tops., 228. 

Plakinide, 226, 228-230. 

Plakortis, Schu/ze, mentioned, 229. 

Plesiastrea = seq. 

Plesiastrea, M. Edw. & Haime, 77. 
haeckeli, Briigg., 95, 98. 
indurata, Verr., 98, 99, 105. 
peront, M. Edw. & Haime, 98, 

99. 


quatrefagesana, M. Hdw. & 
Haime, 103, 104. 

urvillet, M. Edw. & Haime, 
76, 77. 

versipora, M. Kdw. & Haime, 
103, 104. 


Pochazia, Am. Serv., mentioned, 294. 
Polypodide, 431, 435, 
Polypus, Schneid., 436, 437. 


Polypus arborescens, Hoyle, 431, 
438. 
fontanianus, d’Orb., 431, 437; 
distrib., 429 ; mentioned, 438, 
440. 
gardineri, Hoyle, 431, 437, 438; 
mentioned, 440. 
granulatus (Zam.), 431, 440, 
441. 
horridus (d’Orb.), 431, 439. 
horsti (Joub.), mentioned, 440, 
44], 
L., Hoyle, 431, 440. 
P., G. C. Robs.* (2), 431, 440. 
rugosa, Bosc, mentioned, 440. 
venustus, Mang, 431, 439. 
vulgaris (Lam.), mentioned,439, 
Pontonella, Heller, 394. 
glabra, Heller, 395. 
Pontonia, Latr., 328-340, 343, 348— 
350, 352, 389-392. 
armata, H. M. Edw., 389. 
ascidicola, Borrad., 390, 391. 
biunguiculata, Pauls., 394. 
brevirostris, Miers, 390, 391. 
californiensis, Rathbun, 389, 
391. 
custos, Guér., 390. 
dentata, Richters, 380. 
diazone, Joliet, 391. 
domestica, Gibbes, 393. 
flavomaculata, Heller, 390, 391. 
grayi, Rathbun, 390, 391. 
macrophthalma, H. M. Edw., 
383. 
maculata, Stimps., 393. 
maldivensis, Borrad., 387. 
meleagrine, Bate, 393. 
mexicana, Guér., 390, 391. 
minuta, Baker, 389, 392. 
nipponensis, De Haan, 390, 
391. 
parasitica, Roux, 390. 
phallusice, Marion, 391. 
pinne, Orim., 389, 391. 
tridene, Dana, 393. 
tyrrhena, Latr., 390. 
tyrrhena (Petagna), 390, ftnote 
337. 
unidens, Kingsley, 391. 
Pontonides, Borrad.*, 329-333, 342, 
348-350, 387. 
maldivensis, Borrad.*, 324, 387. 
Pontonie enflée, H. M. Edw., 393. 
Pontoniine, by L. A. Borradaile, 
323-396. 


Pontoniopsis, Borrad., 329-332, 338, 
341, 346, 350, 377. 
comanthi, Borrad., 377. 
Porites, WZ. Edw. & Haime, 33, 35. 
Prionastrea = seq. 
Prionastrea, M. Edw. & Haime, 3, 
77, 78, 115. 
abdita, M. Edw. & Haime, 31, 
32, 91, 93, 97. 
aspera, Quelch, 94, 95. 
australensis, M. Edw. & Haime, 


94, 95. 
crasstor, M. Edw. & Haime, 
91, 93, 106. 


echinata, Gard., 91. 

Javosa, M. Hdw. & Haime, 112, 
113. 

flexuosa (Dana), 93. 

Jusco-viridis, Gard., 91, 92. 

gibbosa, Klunz., 91, 93. 

gibbosissima, M. Edw. & Haime, 
95, 97. 

halicora, M. Edw. & Haime, 
80, 83, 94, 95. 

hemprichi, M. Kdw. & Haime, 
110, 112. 

hirsuta, Gard., 101. 

magnifica, M. Edw. & Haime, 
91, 93. 

magnistellata, M.Kdw.& Haime, 
80, 83, 112, 113. 

melicerum, M. Edw. & Haime, 


OH, G7, OE. 

michelint, M. Edw. & Haime, 
109. 

obtusata, M. Edw. & Haime, 
Gil- OB 

pentagona, Klunz., 93, 95, 96, 
97. 

profundicella, M. Edw. & 
Haime, 117. 


purpurea, Gard., 91. 

quoyi, M. Edw. & Haime, 91,93. 

robusta, Stud., 91, 93. 

rousseaut, M. Edw. & Haime, 
79, 94. 

seychellensis, M. Edw. & Haime, 
79, 80, 83, 121, 122. 

spinosa, Gard., 80, 111, 112. 

sulfurea, M. Edw. & Haime, 
il, Obi. 

suvadive, Gard., 80, 81. 

tenella, Dana, 114. 

tesserifera, M, Edw. & Haime, 

- 109. 
vasta, Klunz., 108, 109. 


INDEX 


Privesa, Stal, 297-298. 
fryeri, Distant*, 297. 
melanaria, Distant*, 298. 
punctifrons, Sign., 298. 
Privesana, Distant*, 298-299. 
infusca, Distant*, 299. 
Proboscidactyla gemmifera(Fezwkes), 
184. 
ornata, McCrady, 184. 
sp., 170, 172, 185. 
tropica, Browne, 170, 172, 184. 
Processa processa (Bate), 410, 411. 
Processide, 410-411. 
Prostominia, Revtter, 152-154. 
convexiuscula, Growv.*, 
152-154, 156-159. 
lewisi, Rettter, 142. 
scotti, Grouv.*, 142, 152-153. 
simoni, Growv., 142. 
Prostominini, 152-154. 
Psammastra conulosa, Kieschnick, 
239. 
Psammeecini, 147-150. 
Psammeecus, Latr., 142, 147-150. 
desjardinsi, Guér., 150. 
letulus, Growv.*, 142, 148. 
nitescens, Growv.*, 142, 149. 
reitteri, Grouv., mentioned, 149. 
simoni, Growv., 142, 147. 
sp., 142, 147. 
Pseudino, Fairm., 150. 
Pseudolfersia, Coquillet, 162. 
spinifera (Leach), 162. 
Pseudophanus signatus, 
150. 
Pteropus aldabrensis, 7rwe, 162. 
edwardsi, Geoff:, 162. 
Ptyelus mahei, Distant, 306. 
Pulex felis, Bouché, 161. 
Pundaluoya, Kirkp., 300-301. 
pulchella, Distant, 301. 
simplicia, Distant, 300. 


142, 


Leconte, 


Ramella tubulosa, Schulze, 219, 220. 
Regulus crinitus, Dana, 399. 
Rhabdastrella distincta, Thiele, 235. 
Rhabdodragma, Dendy*, 239. 
conulosa, Dendy*, 226, 239. 
Rhaphidhistia spectabilis (2), men- 
tioned, 245. 
Rhizostoma andromeda var. maldi- 
vensis, Browne, 171. 
Rhizostome, 207. 
Rhopalonema, 
195. 
ceruleum, Haeck., 193, 194. 


Gegenb., 193-194, 


453 

Rhopalonema funerarium, Van- 
héffen, 193, 194. 

velatum, Gegenb., 170, 172,193, 
194, 

Rhynchota, Part II, Suborder Ho- 
moptera, by W. L. Distant, 273— 
321. 

Ricania, Germ., mentioned, 294. 

Ricanoptera, Melich., mentioned, 
294. 

Robson, G. C., Cephalopoda, 429- 
4492, 

“Rotteken’s muscle,” 17. 


Sagartia, Gosse, 23, 30. 
spongicola, Glosse, 23. 

Sagartide, 30. 

Sanidastrella coronata, Tops., 240. 

Sarcinula, Lam., 58. 
astreata, Blainv., 64. 
ellissti, M. Kdw. & Haime, 60. 
erecta, M. Edw. & Haime, 62. 
JSascicularis, M. Edw. & Haime, 

59. 
hystrix, M. Edw. & Haime, 60. 
irregularis, M. Edw. & Haime, 
60. 
musicalis, Blainy., 62. 
organum, Lam., 64, 65. 
Saron marmoratus (Olivier), 397, 
401. 
neglectus, de Man, 401. 
Sarostegia, Zops., 217-222. 
oculata, Tops., 211, 215, 217, 
219-222. 
Saxicave, mentioned, 238. 
Sceurgus unicirrhus, 7iberi, 431, 
435. 

Scaphoideus, Uhiler, 318. 
seychellensis, Distant*, 320. 
tessellatus, Distant*, 319. 
vagans, Distant*, 319. 

Scaphophyllia lobata, Stud., 116. 
cylindrica, M. Edw. & Haime, 

116. 
Sclerothammopsis compressa, Wilson, 
219. 
Scott, H., Mallophaga, Aphaniptera, 
and Diptera Pupipara, 161-167. 

Scyphomeduse, 171, 202-207. 

Semeostomezx, 204. 

Senecio sechellensis, J. G. Baker, 

mentioned, 285, 293. 
Sepia, Linn., mentioned, 429, 
Sepioteuthis (D’Orb.) spp., men- 
tioned, 434. 


454 


Siderastrea radians, Vaughan, men- 
tioned, 9. 

Sidonops oxyastra, Lendenf., men- 
tioned, 256. 

Silvanini, 144-147. 

Silvanophleus, Sharp, 143. 

Silvanus advena, Hrichs., 145. 

bicornis, Erichs., 144. 


hebetatus, Growv., 141, 142, 
145. 

mercator, Fauy., 144. 

scuticollis, Walk., 141, 142, 
145. 


signatus, Frauenf., 146. 
surinamensis, Latr., 141, 144. 
triangulus, Reitter, 149. 
Sminthea, Gegend., 195. 
eurygaster, Gegenb., 171, 172, 
194-195. 
Solenastreea = seq. 
Solenastrea, M. Edw.& Haime, 33, 38. 
bournoni, M. Edw. & Haime, 
41, 42. 
bowerbankui, M. Edw. & Haime, 
41, 42. 
chalcidicum, M. Edw. & Haime, 
41, 42, 44. 
ellisit, Duchass. & Michel., 39. 
JSorskaliana, M. Edw. & Haime, 


43, 44, 

gibbosa, M. Edw. & Haime, 41, 
42. 

hemprichiana, M. Hdw. & 


Haime, 41, 42, 48. 
hyades (Dana), 2. 
micans, Duchass. & Michel., 39. 
sarcinula, M. Edw. & Haime, 
39, 40. 
Solmaris lenticula, Haeck., 200. 
Say, Lei, Wy, Ise. 
Solmundella, Haeck., 200-201. 
bitentaculata, Vanhéffen, 200- 
201. 
mediterranea (d/iiller), 
172, 200-201. 
Sponges, Hexactinellid, 211-224; 
Homosclerophora and Astrote- 
traxonida, 225-271. 
Stauria astreiformis, J Hdw. & 
Haime, 113. 
Steenstrupia, Mayer, 173. 
bigelowi, Mayer, 174. 
normani, Browne*, 171, 173, 
174-175. 
rubra (forbes), 173. 
tetrabrachia (Bigelow), 173. 


il, 


INDEX 


Stegopontonia, Wobili, 329, 331, 
332, 347, 349, 350, 386. 
commensalis, Wobili, 386. 
Stelletta awrora, Hentschel, 242. 
ewastrum, Carter, 257, 259. 
globostellata, Carter, 242, 247. 
herdmani, Dendy, 235. 
lactea, Carter, 237, 238. 
pathologica, Schmidt, 232. 
reticulata, Carter, 242. 
stemenst, Keller, 250. 
tuberosa, Hentschel, 236. 
Stellettide, 226, 234-254. 
Stellettinopsis simplex, Carter, 251, 
252. 
Stenoteuthis bartrami (Lesueur), 
431, 433; distrib., 429. 
Stephanocenia, M. Edw. & Haime, 
33, 75, 78. 
dendroidea, M. Edw. & Haime, 
75. 
intersepta (Esper), 74-75. 
maldivensis, Gard., 96. 
michelini, M. Edw. & Haime, 
75. 
Stephanocora, Ehrb., 48. 
hemprichi, Ehrb., 54, 56. 
Stevensonia, J. Duwnec., sp. men- 
tioned, 156. 

Stichopathes, Brook, finote, 31. 

Streblide, absence noted, 163. 

Stryphnus, Sod/as, mentioned, 235. 

Stylinide, ftnote, 35. 

Synalpheus biunguiculatus, Stumps., 
416; mentioned, 417. 

var. pachymeris, H. Cout., 
417. 

charon, Heller, 416. 

fossor, Pauls., 416. 

gravieri, H. Cout., 414. 

hastilicrassus, H. Cout., 417. 

levimanus, 1.Cout.,mentioned, 
418. 

—— haddoni, H. Cout., men- 
tioned, 417. 

longicarpus, Herrick, 418. 

lophodactylus, H. Cowt., 417. 

merospiniger, H. Cout., 415. 

metaneomeris, HW. Cout.*, 414. 

var. streptodactylus, H. 
Cout.*, 414. 

neomeris, de Man, mentioned, 
414. 

neomerts, H. Cout., 413, 414. 

nilandensis, H. Cout., 415. 

oxyceros, H. Cout., 416. 


Synalpheus otiosus, H. Cowt., 415. 
pachymeris, H. Cout.*, 417. 
var. Caryadosi, H. Cout., 
417. 
paraneomeris, H. Cout., 415. 
var. praslini, H. Cout.*, 
415. 


var. Seychellensis, H. 
Cout.*, 415. 
sladeni, H. Cout., 417. 
sp., Bate, 414. 
streptodactylus, de Man, 414. 
trionyx, H. Cout., 416. 
triunguiculatus, Pauls., 416. 
tumido-manus, Pauwls., 417. 
Syneoryna, JZhrenb., 
170. 


mentioned, 


Taonidium, Pfeffer, sp., 431, 435. 


suhmii (Hoyle), mentioned, 
435. 

Taphozous mauritianus, Geoffroy, 
163, 167. 


saccolaimus, Zemm., 167. 
Teleoteuthis, Verr., sp., 430, 431. 
Telephanus fasciatus, Redtenb., 

150. 

pictus, Cast., 151. 

Tethea ingalli, Bowerb., 264. 

robusta, Bowerb., 267. 
Tethya, Auct., 260-267. 

asbestella, Lam., 260. 

cranium, Lam., 260. 

ingalli, Sollas, 264, 265. 

japonica, Lindgren, 262. 

lacunata, Lam., 260. 

lyncurium, Lam., 260, 261, 
262, 263. : 
var. 6, Dendy, 264. 

seychellensis, Sollas, 265. 
Tethyade = seq. 

Tethyide = Donatiide, 259-267. 
Tetractinellida, 227. 

Tetraxonida, 226, 227-269. 
Tettigonia jactans, Walk., 315. 
Thalassianthus, Lewck., 35. 
Thalassocaris affinis, Borrad., 400. 

crinitus (Dana), 399, 400. 

lucidus (Dana), 400. 

maldivensis, Borrad., 400. 
Thor (Heller), 397, 401, 402. 

jloridanus, Kingsley, 402. 

maldivensis, Borrad., 401-402. 

paschalis (Heller), 397, 402. 
Tiara, Wagner, mentioned, 181. 

conica, Less., 183. 


Tiaropsis rosea, Agass.& Mayer, 170, 
172, 186. 
Tozeuma armatum, Pawls., 402. 
Trachomeduse, 171, 192-199. 
Trachynemaeurygaster, Haeck., 194. 
mammeeforme, Haeck., 194. 
Trachypora, Verr., 49. 
Trapezia, Latr., mentioned, 330. 
Tridacna gigas, Brug., mentioned, 
389. 
spp., mentioned, 388, 393. 
Tridacnocaris, Nobili, 387. 
Tridacophyllia (pars), Blainv., 48. 
Tubastrea, Blainv., 77. 
Tubiclava, Al/m., 181. 
Turbinaria, Oken, 41. 
Turris neglecta, Less., 179. 
Turritopsis, WcCrady, 179-181. 
nutricula, McCrady, 170, 171, 
172, 179, 180. 
var. pacifica, Maas, 179. 
pacifica (Maas), 179. 
polycirrha, Keferstein, 179,180- 
181. 

Tylana, Sta, mentioned, 293. 
Typton, Costa, 328-333, 341, 343, 
346, 349, 350, 394-395. 
bouvieri, obi, 394, 

ftnote, 349. 
spongicola, Costa, 394-395. 
spongiosus, Bate, 395. 


395, 


Ugyops, Guér., 305-306. 
facialis, Distant*, 305. 


INDEX 


Ugyops senescens, Distant, 305. 

seychellensis, Distant*, 305. 
Ujna, Distant, 313. 

flavidipes, Distant*, 313. 
Ulastrea, M. Edw. & Haime, 77. 
Uleiotini, 143-144. 
Ulundia, Distant, 299. 

madagascariensis, 

299. 
Urocaridella, Borrad., 327-342, 346, 
348, 350, 352. 

Borrad., 


Distant*, 


gracilis, 

353. 
Urocaris, Stimps., 325, 329-333, 
337, 338, 341-354. 

esopius (Bate), 354. 

indica, Kemp, ftnote, 323. 

infraspinis, Rathbun, 353, 354. 

korni (Zo Bianco), 354. 

longicaudata, Stimps., 323, 345, 
353, 354. 

longipes, Stemps., 354. 

psamathe, de Man, 323, 353, 
354. 

sp., 323. 


323, 352- 


Verschaffeltia, H. Wendl., sp. men- 
tioned, 156. 
splendida, mentioned, 144, 147. 
Vioa johnstonii, Schmidt, 252, 253. 
schmidti, Carter, 253. 
Voleanalia, Distant*, 279-284. 
atrostriata, Distant*, 280. 
atrovaria, Distant*, 280. 


ERRATUM. 


p. 376, line 16, for (Dorna) read (Dana). 


455 


Volcanalia capitata, Distant*, 283- 
284. 

cardui, Distant*, 281. 
designata, Distant*, 281. 
fumosa, Distant*, 282. 
modesta, Distant*, 283. 
picturata, Distant*, 281-282. 
typica, Distunt*, 279-280. 
uniformis, Distant*, 283. 


Walkeri, Kirkp., 314. 

Willsia, Huxley, sp., 184. 

Wormia ferruginea, Bazll., men- 
tioned, 153, 156, 293. 


Xenia hicksoni, Ashw., ftnote, 31. 

Xenosceleni, 154-156. 

Xenospongia johnstonii, Lendenf., 
253. 


Yanga andriana, Distant, 275. 
seychellensis, Distant, 275. 
Yodomia, Lebwohl, 232-234. 
ijimai, Lebwohl, 232, 234. 
perfecta, Dendy*, 226, 232- 
234. 


Zanclea, Gegenb., 185. 

juv., 170, 172, 176. 

orientalis, Browne*, 176. 

spp., 176, 177. 
Zoanthus, Cuv., 22. 

coppingeri, Hadd. & Shack., 16. 
Zooxanthelle, 32. 


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completed in 30 Vols., and a few entire sets are still for sale. Only certain single volumes, or parts to 
complete sets, may be obtained at the original prices. The price of the Index to Vols. 1-25 is 8s. to 
the public, and 6s. to Fellows; to Vols. 26-30, 4s. to the public, and 3s. to Fellows. 

The Second Series of the 'ransactiOns is divided into Zoological and Botanical sections. ‘lhe prices 
of the Zoological parts of those which have been published are as undermentioned :— 


SECOND SERIES.—ZooLoey. Srconp SpRiEs.—ZooLtocy (continued), 
When Price to the Price to When Price to the Price to 
Volume Published Public Fellows Volume Published Public Fellows 
AB Bs. is oe 1Sey Als iG) ous yds Big. “de 
Ieeearts! eoVallee aSib= 09) so Om Ome Gh) M86 XI. Part I. cow RO sae) 44 Osun @. & @ 
Payah WL non IG ces © 6S Won O B W 
Tie Parts le XVI 1879-88 2. 7 1 Oe. a U8, 5; Pass UL oc TOO. © 6 O.. 0 2 
Pay I, 25 MOO so © 1° Os. O D © 
Ill. Parts I. VI. 1884-88 ...5 18 0...4 8 6 Day WW ie 11909) 0) 2k ee OR 
Peyte Wil, oso) GMO) Eso OY TL PO) aon @ SB 
; rts) J.-L. = oak ae iad 
Io emia ESI OT deletes es) Be Part Wille ge 190 «0 5 0 ey Oleenme 
W. Parts oN es04...6 10 @ic.24 1h 9 Part VIII. ... 1911...0 4 0...0 3 0 
Pari UX, oe 7 1901 2.20) DaiGi a Oz eNO 
VI. Parts 1-VIII. 1994-99...617 6.5 3 8 Bath Me Uva ae Oe eee 
[Pein ee LIS IBWsoo OY VAN Oy cag (> 3} 
VII. Parts L-XI. 1896-1900...6 7 9...4 15 9 Bane MIs (--- MOUS e105 SEO to ery 
(In progress.) 
WAUUL TREES I SKIQUL OOOO... 6 ME Sono BTS OM ae I ee YO seh 8 Ma OW 
ani : Part le ae  DOOW a: 1. 42 10S OF 18" 30 
TX, Part “La 1903 2.0 9 00°68 9 Pre Wl I O18 Ox. OB © 
Part II ... 1903° ..0 8 0..0 6 0 Part’ IV. ds: 1909) Sos tOe 0) Semon 
Part II. ... 1908 ...1 4 0...018 0 Part V. Index 1909 ...0 5 0..0 3 9 
Part IV. ... 1904 0) 6) VO) 5 0) 4516 
Part V. & 1902 a6.0 8 Oce@ & @ | MMU Wea It =. OO9) enw 4s OF 5 OSG 
Part VI. ... 1904 oo WW) 1 Occe W) 4b een Ub, coos CHOY co Bo Bs OW soa th NB 8 
Part VII. ... 1904 oo WO Osea OM Zh &: ever JU ooo) ICI) Sos ) Kh OC oe @ A © 
Pays WIN oon OOS F501) Oa20 7 GB Part IV. Index 1910 ...0 5 0...0 3 9 
Revny IDS, ccc LGXOS “0 6 0...0 £ 6 4 
Part XI. nee 1907 Hoo (Oye 11) 0 ae 0 9 0 Part ne oo UWI o65 Il 0 0. eee 0) s; (0) 
Part XIV. Index 1907 20; 43) OO) 42) 23 NV Parte. le’ a NOI Ae MOO +5 alumi 
tb We =. 1902)... ily 40 eee OMISINO 
yoy Ba Iho. TOOL OA Oe Ore 8 fase Ul ae . 
Part Dll. .3. a912) 51 6 y2Ohs O19 16 
Rantewlile) sen 1904 ssanO) 1Si OO) 6) 0) awk JAW 1913 0-12 0 0 9 @ 
Peay WG 56 UNS 5 OO Oo. © BY) a Sa 8 saad 
Pant IVa. eee 1905) O10 10) ee O Gs aXeVile tart 1. | 9 ce llO)IS eer oak) ere melts aan 
Part V. ... 1906...0 7 6...0 5°3 Part: IL.) ce: vlOUS =. I) WO Pee O mts io 
Part Vib “pe 1906-3. 10) 310. OF 253 Peay UN doy NOB) oo O 1 O06 O Y 
Payay WAU, coc GO ono M 8B Oa O 4 fs Pap ING ao, GE ce WO O 5 1 BB 
Part VIII. ... 1907 ...0 4:50...0 3 0 Part V. Index1914...0 3 0...0 2 3 
Part DX. 2) 909) a elee al OMe Ol anno 
Pate Oe ea oO) Oa O BO | QVlb Renu IK see NOM sos BO OD coo 1 IO © 
(In progress.) | Pera lly mecca aS Gon IL 68) “Osco @ aly 


2nd Ser. ZOOLOGY. | [VOL. XVII. PART 3. 


THE 


TRANSACTIONS - 


OF 


THE LINNEAN SOCIETY OF LONDON. 


THE PERCY SLADEN TRUST EXPEDITION 


TO 


THE INDIAN OCEAN IN _ 1905, 


UNDER THE LEADERSHIP OF 
Mr J. STANLEY GARDINER, M.A. 


VOLUME VI. 


Reports Nos. 7—9 of this volume; Nos. 112—114 of the whole series 


LONDON: 


PRINTED FOR THE LINNEAN SOCIETY 
BY J. B. PEACE, M.A., AT THE UNIVERSITY PRESS, CAMBRIDGE. 
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE, PICCADILLY, W. 1. 
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW, E.C. 4. 


August 1917. 


LINNEAN SOCIETY OF LONDON. 


MEMORANDA CONCERNING TRANSACTIONS. 


The First Series of the ‘Transactions, containmg both Botanical and Zoological contributions, has been 
completed in 30 Vols., and a few entire sets are still for sale. Only certain simgle volumes, or parts to 
complete sets, may be obtained at the original prices. The price of the Index to Vols. 1-25 is 8s. to 
the public, and 6s. to Fellows; to Vols. 26-30, 4s. to the public, and 3s. to Fellows. 


The Second Series of the ‘I'ransactions is divided into Zoological and Botanical sections. he prices 


of the Zoological parts of those which have been published are as undermentioned :— 


SECOND SERIES.—ZooLoey. Seconp Serius.—ZooLoey (continued). 


When Price to the Price to When Price to the Price to 
Volume Published Public Fellows Volume Published Public Fellows 
2) 8 dh é& 85 th 33 85 a9 G5 Gh 
i Pacis LG ISTO) STO Wa 0 BP 20 Paw I NOS =. OD 4 O@ineO@ B @ 
’ Part II. Wo. 8 O-. 0 & O 
il, Paris LOMWIWUL IS7OS89 coo 7 IT Ooo D 18 & Part III. 1509 ... © 8 Ou. O 4 GB 
A Part IV. IOS so OU Os O 9 © 
Ill. Parts I-VI. 1884-88 ...5 18 0...4 8 6 Pacterye 1965". Onlomto canna 
Part VI. MNO ao OL Oo O 8 3 
: gy Wm, sae EAL 
ING Barts le Ue So 8 ec on Oe Nae Part VIL. Oe» Oo. 0 8 o 
V. Parts 1-XI. 1888-94...610 6..417 9 Part Ve a Oe a 
Part IX. While) BY Gis 0 2 © 
VI. Parts L—VIII. 1894-99...617 6...5 8 3 Ean eure oe Deh 
Part XI. MONE co 2 Dun O 3 O 
Vile Parts 1X, 1896-1900 2607 Set ta 99 Part XII. 1913 ..0 8 0...0 6 0 
(In progress.) 
Wilh, Peric ILOOUIL MOOG... 6 We Yo. B 2 S| UU Penh IL NOG ae Os. 1 1 O 
Part IL. CO eek ee ees O 13° O 
IBS, Pant Tk IE 20 2 OoaO 6° O Part III. 1908 .. O16 0. 012)" 
Part II. IE Oo) 8 OLnO E Part 1V. WO ae Lal Ol. 1 96 
Part ILL. WEY ae A Ose O18. O Part) Va Index 1909) 2 0) 5 ieO;eerOunoinO 
Part IV. 1902) 06 20: OR , 
Part V. ih sO Cue oO 2B ape he em meee O28 68 
Part VII. inv 6.0 6 O40 A 6G Bani TIT TOO SOO eo ae 
Part 1X. IOS ee OG Oca) 4G | sony, Be IL IG) oo LUO a 1 8 6 
Part X. 1906 ...0 12 0..0 9 0 Part II. MOM cco LO Ose O15 © 
Part XI. NOD? oo O@ IF Oo. 9 @ Part IIL. Cie teed Oka tO 
Part XII. 1907 ..0 3 0...0 2 3 Part) IV. Index 1912.00) uo Oe. 10) edu 
Part XIII. NG PO. Bo Ow. O 26 
Part XIV.Index1907 ...0 3 0...0 2 3 | XV. Part I TOd2) seo L ad Oe glee 
Part II. eel OAL O18 Oo 
X. Part I. 1004 ok © 2B Ooo O® QB 8 Part III. 11D co 1 O Ou O 1. 6 
Parcel: 190A oc © 8 O20 B O Part IV. IONE 20 OIA Oo OM 2 O 
Part II. 19057-7780) 9) OF Oe xevile Part Le 1a Oe @ 19 -O 
Part IV. USO) ere NOLO Okc Oe ie Part II. iehiepeee a all OPecs Ol © 
Part) Ve IQS coo O % Goo D H & Psyes IU0L 191 6 O19 O..0 9 @O 
Part VL. MeO OED aaa 8 Part IV. (Cee Ws Qe. t 2 8 
Part VII. 190% ee 0 ee Oe eee Part V. Index 1914.0) 3 OlNO; ama 
Part VILL. IVs OL Oca O B O 
Part IX. TO) ot Oo O 1 O [AAU em NE esac a Ui oep KD (0 
Part X. TN Ass OAD ce O 8 © Part II. IS eet ES OW) ace OUT. og 
(In progress.) Part III. UHM ban I BD W250 O 18 9 


° ond Ser. ZOOLOGY. ] 


r+ 
- Me \ 
\5 Xx \ 


wxbAD 


[VOL. XVII. PART 4. 


THE 


TRANSACTIONS 


OF 


THE LINNEAN SOCIETY OF LONDON. 


THE PERCY SLADEN TRUST EXPEDITION 


TO 


THE INDIAN OCEAN IN 1905, 


UNDER THE LEADERSHIP OF 


Mr J. STANLEY GARDINER, M.A. 


VOLUME VI. 


Reports Nos. 10 and 11 of this volume; Nos. 115 and 116 of the whole series. 
Also Titlepage, Contents, and Index to Vol. XVII. 


KRY 


LONDON : 
PRINTED BY J. B. PEACE, M.A., AT THE CAMBRIDGE UNIVERSITY PRESS. 
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE, PICCADILLY, W. 1 
AND BY LONGMANS, GREEN, AND CO., PATERNOSTER-ROW, E.C. 4 


March 1921. 


@ 


LIBRARYisy 


+) by 
PED ui oun) iy nae 
J Su oe 


Lek 


YO a ee ie 
mn (ia 
eh. A a 


PANG ER SAG ic 


‘i 


WW ay ee 0 


LINNEAN SOCIETY OF LONDON. 


MEMORANDA CONCERNING TRANSACTIONS. 


The First Series of. the ‘Transactions, containing both Botanical and Zoological contributions, has been 
completed in 30 Vols., and a few entire sets are still for sale. Only certain single volumes, or parts to 
complete sets, may be obtaimed at the original prices. The price of the Index to Vols. 1-25 is 8s. to 
the public, and 6s. to Fellows; to Vols. 26-30, 4s. to the public, and 3s. to Fellows. 


The Second Series of the Transactions is divided into Zoological and Botanical sections. The prices 
of the Zoological parts of those which have been published are as undermentioned :— 


SECOND SERIES.—ZooLoey. SECOND SERrES.—ZooLoey (continued). 
Whe: Price to th Price t : . F 
Volume Published Tee : Wellows Volume Published a eo Fellows 
eo G3 Gy 9) be Gh 8a th 8 & @, 
I, lewis ISIN WSS son 8 NWO - Oo B WY EB XG ear tale o05 JIBXOS) non O 4b Oo @ & O 
Bn Barty eye 01909) 22. 10008) 20 OGIO 
If. Parts I.—XVIII. 1879-88 ...7 17 0...5 18 5 | Porte lly sos NE oe) LA Ole v8 
| amg WW cos IO 3. OI Oa.O OD ©O 
i “Parts [=Vi. 1884=88'...5 18 0 ..4 8 6 Part V. Beet OOON ce (Osa) Ome rtOr nae 
IVa Barts) LN wt8eG288\4..37) 8 022) 110 EAE om TE os OU aoe O88 
leans WINS ceo IOI G8 O B Oa. O 8 9 
Ve) Parts) LOX 1888-94"... 6 10° 6... 4717. 9 Part VII. ... 1911 ...0 4 0..0 3 0 
| Bante Xe oe Sy Oil. ORM Oi Gy Ome w LO 
VI. Parts I.—VIII. 1894-99...6 17 6...5 3 3” Part X. ... 1911 ...0 2 6..0 2 0 
Path DIPS eo alee e() ch 0) cas O & WO 
VII. Parts L-XI. 1896-1900...6 7 9...415 9 Part XI... 1913 ...0 8 0...0 6 0 
(In progress.) 
Wild, Pais TCE TOMOMO LLG A ce Be Cle em) LY ea Eee I @) see TL 
Pam 0 dan IO one HE EO nce O 1S 
IDG Remy IE Boo” IBS coe) YW @ 6  ¥) leat IU see! IDO ca ONG O se W 19 
jPearh JU sae IGOR) eb OP SW cc. © 1G Part DVin 11909) co Ome ee ae 
Partie se G03 cee 14 SOm ON mS 20 Part V. Index 1909 ...0 5 0... 0 38 
Part af Oeics OE ONS aie Ape ee As Mics © 18 
Part a Oe On O) 10 TAO ano Part 11. Saeolo eo) 3600 ear 
Part ee TOU ete 1D Osi Ue @ Part) Ul | eo ee 90) 6 ONG NO menOM 
Part VU. ... 1904 ..0 6 0...0 4 6 Reha ING Ina MEN Ge O H O on O 8 
Part VIII. ... 1904 oO MY Os 7 6 
Part IX. ... 1905 oO 8 On. & 8 XO, lemg IL ee OO), csc eles: Oey lee 
Part X. ... 1906 ~ O01 On.0 9 0 Peyne I aco TIO see lO 2 OM cen ©: 1K 
Pais YO o2, 1907 oO 1 O.0 9 © Part lees OUT oxo (8) RO ee ve 
Parte eeeeLo0T) ee Ouss! On Omen «8 Part IV. Index 1912..0 5 0..0 38 
Paras XG0UL cos ION cco @ OB Mos 4 6 AWE Te eraE Th cage CIO) Gan Ih OO) oka Th ail 
Part XIV. Index 1907 oO) 8s -'Occo, 0) B98} Perpe Ue sue IGG gop zt ©. 3 © 1S 
ein JOU sae I ee Th OB © coe © NY 
Xe Part) 1: GOA ee VO woe ON ss On ih S Day IN Lae) OHS 4. O12 Oo. @ 9 
FOU ES ee UES Boh) a On aR XOVle) arte, Uk) 2 eee) sO Sap nsl ARn Omeeen Ones 
Part Mis ee 1905, 4.201890" 0l- 20416 09 
Bart ae 5s SSS ee sO pee Oat 
Piya IY, cas. IICKO too (UO Ooo O 7 G : , 
Peis JU0R sus 9 IONS 6, O 1A O so. O © 
Peyhe Wau eee USKNG cos OP MB coe) GS 
F [Deyn lier ou) GR coe TIO) SO) coe vil) 
Part) Vile eee 0G Ones Oe Omme2 ine 3) Peet Vane icnel ye ee 
Parc oville se TOOT OMI OR ONDE 5 ee at : ae ce 
Pamu WAU soe TOO? coo © 2 Oven © 8 @ | MVM Wate Myo MGI 1 A O O coo i 10 
yD ei TIE) cog who ak Ors GO 15 © Part H. ... 1916 ...1 3 0...017 
Burt X. | ... 2 ROLLE OMe mcr OMnBEO Part I ... 1917..1 5 0... 0 18 
(In progress.) | Painh J0VG 3 WOR oe OI Ooo O 


Sco COMO Con Cy eon (Os Sl Ca OC COMO cy O om Co Sco Cre omc) 


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