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MEMOIRS 


of the 
INDIAN MUSEUM 


Vol. IV. 


An Account of the 
CRUSTACEA STOMATOPODA 
OF THE INDO-PACIFIC REGION 


based on 


THE COLLECTION IN THE INDIAN MUSEUM. 


By 
SPANIEN SIE NPS BEATS, | 
Assistant Superintendent, Indian Museum, 123 2 q 8 Ze 


with which are issued 
ILLUSTRATIONS of the ZOOLOGY of the R.IM.S.S. ‘INVESTIGATOR’ 
under the command of 


Commander ©. S. HICKMAN, R.I.M. 


Crustacea Stomatopoda, Plates I—X. 
Issued by order of Captain WALTER LUMSDEN, R.N., 


Director, Royal Indian Marine. 


PUBLISHED BY ORDER OF THE TRUSTEES OF THE INDIAN MUSEUM. 


Calcutta : 
PRINTED AT THE BAPTIST MISSION PRESS. 


SEPTEMBER, 1013. 


Price, with plates, J'ifteeu Rupees. 


MEMOIRS 


of tbe 


INDIAN MUSEUM 


Vol. IV, ert. 


An Account of tbe 


CRUSTACEA STOMATOPODA 


OF THE INDO-PACIFIC REGION 


based on 


THE COLLECTION IN THE INDIAN MUSEUM. 
By 


STANLEY KEMP, B.A., 


Assistant Superintendent, Indian Museum, 


with which are issued 


ILLUSTRATIONS of the ZOOLOGY of the R.IM.S.S. ‘INVESTIGATOR ? 


under the command of 


Commander C. S. HICKMAN, R.I.M. 


Crustacea Stomatopoda, Plates I—X. 
Issued by order of Captain WALTER LUMSDEN, R.N., 


Director, Royal Indian Marine. 


PUBLISHED BY ORDER OF THE TRUSTEES OF THE INDIAN MUSEUM. 


Calcutta : 
PRINTED AT THE BAPTIST MISSION PRESS. 


SEPTEMBER, 1013. 


Price, with plates, Fifteen Rupees. 


m 


US VER 


Miscellaneous Zoological Publications. 


Rs. As. | Rs. As. 
Account of the Deep-sea Brachyura collected by | Echinoderma of the Indian Museum: Littoral 
the R.I.M.S. ‘‘ Investigator.’’ By A. Alcock, | Holothurioidea collected by the R.LM.S. 
M.B., C.M.Z.S. . 67708 | ‘‘ Investigator.’’ By R. Koehler and C. Vaney 2 o 
Account of the Deep sea Madreporaria collected Echinoderma of the Indian Museum: Deep-sea 
by the R.I.M.S. ‘‘ Investigator.’’ By A. Alcock, | Ophiuroidea collected by the R.I.M.S. ‘‘ Investi- 
M.B., C.M.Z.S. ue fie 4 0 gator.’’ By R. Koehler ae 10 © 
Account of the Triaxon (Hexactinellid) sponges Echinoderma of the Indian Museum : Shallow- 
collected by the R.I.M.S. ‘‘ Investigator.”’ By water Ophiuroidea collected by the R.I.M.S. 
F. E, Schulze, Ph.D., M.D. .. 16 0 ‘“ Investigator.’’ By R. Koehler 35 VOR ao) 
Netount or the Aleyonarians kollected by ‘He Echinoderma of the Indian Museum, Part V: 
R.I.M.S. ‘‘Investigator.’’ PartI. By ]J. Arthur An account of the Deep-sea Asteroidea col- 
Thomson, M.A., and W. D. Henderson, M.A., lected by the R.I.M.S. ‘‘ Investigator.’’ By R. 
B.Sc. : Are to Koehler .. 12 0 
11 a b th Echinoderma of the Indian Museum, Part VA An 
or = ‘ ee 5 ans € ai ee ik Be account of the Shallow-water Asteroidea. ByR. 
: - Koehler ©... 20 © 
Thomson, M.A., and J.J. Simpson, M.A., B.Sc. 20 0 
Aids to the identification of Rats connected with Echinoderma of the Indian Museum, Part VII: 
Plague in India. By W. C. Hossack, M.D dE The Crimoids of the Indian Ocean. ByA. ETS Clark 2000 
ad ise onthe Asiatic Beetles in the Indian Figures and Descriptions of nine Species of Squil- 
Museum. PartI. Family Carabidæ, Subfamily lidæ from the Collection of the Indian Museum. 
Nas $ 9 è B Wood-M EAS et dited b 
Cicindelinæ. By N. Annandale, D.Sc., and W. ‘es Re N SLB a EMZ. a CH CORRE % A 
GC a fl Indian Grnciicent Part Te a ae Guide to the Zoological Collections exhibited in 
atalogue o — = 


tion the Brachyura Primigenia. By A. Alcock, | oe es > Sn en Museum. el vs 

MD, Fe Guide to the Zoological Collections exhibited in 
Catalogue of Indian Decapod Crustacea. "Part Fe — the Bish ‘Gallery eee Ne aa Nec ns By 

Brachyura. Fasciculus II.—Indian Freshwater A: Alcock MBJCM.ZS. 2 ies 

Crabs— Fotamonide. “By A 7 lese, SET: E., | Guide to the Zoological Collections exhibited: in the 

M.B., LL.D., ERS. SO Invertebrate Gallery of the Indian Museum. 
Catalogue of the Indian Decapod Crustacea. Part By A. Alcock, M.B., CMZS. (Out of print.) 

ee on cee a pee Cea By Guide to the Zoological Collections exhibited in the 

oy te ee = B EN, Reptile and Amphibia Gallery of the Indian Mu- 


Catalogue = the dan Decapod Crustacea. Part seum. By A. Alcock, MB, CM.Z.S. (Out of 


III.—Maerura. Fasciculus I.— The Prawns of 


print.) 
the Peneus Group. By A. Alcock, M.B., LL.D., Hand List of Mollusca in the Indian Museum 
F.R.S., CLE. de ESSEN Parts I and ce and Fasciculus E. By G. Nevill, 
Catalogue of Indian Deep-sea Crustacea : Decapoda | CM.ZS., etc. Index Parts I and II. By W. 
Macrura and Anomala in the Indian Museum. | Theobold 2 


By A. Alcock, M,B., LL.D., C.M.Z.S.. BR MO © 


Illustrated Catalogue of Asiatic Horns and 
Catalogue of Indian Deep-sea Fishes in ie Indian 


Antlers in the Indian Museum. By T. Bentham 2 o 


Museum. By A. Alcock, M.B., C.M.Z.S. Fax List of Batrachia in the Indian Museum. By W. L. 
Catalogue of Mammalia in the ae Museum, : Sclater, M.A., F.Z.S. 5 2 EEE NO! 

Part I. By J. Anderson, M.D., LL.D., F.R.S. _ List of Birds in the Indian Museum. Part I.— 

Part II. By W. L. Sclater, M.A., F.Z.S. 2230,20 Corvide, Paradiseidæ, Ptilonorhyuchide and 
Catalogue of Mantodea in the Indian Museum, Crateropodidæ. By F. Finn, B.A., FZS. LE TO 

Parts I and II. By J. Wood-Mason, F.Z.S.,ete. 2 0 List of Snakes in the Indian Museum. By W. L. 
Catalogue of Moths of India, Parts I to VII. By Sclater, M.A., F.Z.S. 1220 

E. © Cotes and C. Swinhoe, F.L.S., F.Z.S., etc. 5 2 Monograph of the Asiatic Chiroptera and Catalogue 
Echinoderma of the Indian Museum: Account of Se | of the Species of Bats in the Indian Museum. 

the Deep-sea Holothurioidea collected by the | By G. E. Dobson, M.A., M.B., F.R.S. 3x0 

R.I.M.S. ‘‘ Investigator.’’ Er R. Koehler and Monograph of the Oriental Cicadidæ, Parts I to : 

C. Vaney .. ne pitt), XO) VII. By W.L. Distant, FES: 45 2 3 14 


The above can be obtained from the Superintendent of the Indian Museum, Calcutta, and from Messrs. Ptiediandet 
& Sohn, 11, Carlstrasse, Berlin. 


Other Publications edited and sold by the Superintendent of the Indian Museum (also 
obtainable from Messrs. Friediander & Sohn) issued by the. Director 
of the Royal Indian Marine. 


Illustrations of the Zoology of the R.I.M.S. ‘‘ Investigator,’’ 1892. Fishes, Plates I to VII. Crustacea, Plates 
I to V, 1894. Fishes, Plates VII to XIII. Crustacea, Plates VI to VIII. Echinoderma, Plates I to III, 1895. 
Echinoderma, Plates IV and V. Fishes, Plates XIV to XVI. Crustacea, Plates IX to XV, 1896. Crustacea, 
Plates XVI to XXVII, 1897. Fishes, Plate XVII. Crustacea, Plates, XXVIII to XXXII. Mollusca, Plates I to 
VI, 1898. Fishes, Plates XVIII to XXIV. Crustacea, Plates XXXIII to XXXV. Mollusca, Plates VII and VIII, 
1899. Fishes, Plates XXV and XXVI. Crustacea, Plates XXXVI to XLV, 1900. Fishes, Plates XXVII to 
XXXV. Crustacea, Plates XLVI to XLVIII. Index, Part I, 1901. Crustacea, Plates XLIX to LX. Mollusca, 
Plates IX to XIII, 1902. Crustacea, Plates LVI to LXVII. Crustacea, Plates LXVIII to LXXVI. Fishes, Plates 
XXXVI to XXXVIII, 1905. Crustacea (Malacostraca), Plates LXXVII to LXXIX. Crustacea (Entomostraca), 
Plates I and II. Mollusca, Plates XIV to XVIII, 1907. Fishes Plates XXXIX to XLIII. Crustacea (Entomostra- 


ca), Plates III to V. Mollusca, Plates XIX and XX, 1908.—Re. 1 per plate. Mollusca, Plates XXI to XXIII, 1909.— 
As. 6 per plate. 


AN ACCOUNT 
OF 


THE STOMATOPODA OF THE INDO-PACIFIC REGION. 


By STANLEY Kemp, B.A., 
Assistant Superintendent, Indian Museum. 


Plates I-—X. 
CONTENTS. 

Page 

Introduction 5 LE ce RT ae He cas NOT 
Systematic account se + ze er de on He ES 
Squilla 2: a de a Sc Er 53516 
Pseudosquilla #3 Ps de N: ae o% sc CA 
Lysiosquilla ae int Ws sis à 3 2109 
Coronida BR ae >. Hr Se he 2.729 
Odontodactylus BR 58 oe oe 50 50 Bp tS} 
Gonodactylus aye a de a 30 oe .. 145 
Addendum ce 5d 23 ae a on = co 205 
List of species known from the Atlantic, Mediterranean and Pacific Coast of America .. 2200 
Bibliography 5 = ce = 58 AR 08 .. 206 
Index De SE ne 10 50 6 ae so Bus 

INTRODUCTION. 


The history of the collection of Stomatopoda in the Indian Museum dates from 
some twenty years ago when the late Mr. J. Wood-Mason, at the expenditure of 
much time and energy, devoted himself to the acquisition of a representative series 
of Indo-pacific species To the small nucleus collection then existing he was able to 
add numbers of known and unknown forms, not only from the coasts of India, but 
from New Zealand, Australia, Hongkong, Singapore and Mauritius. 

Mr. Wood-Mason died in 1893, just after completing the arrangement and 
classification of a large part of the collection. It was hoped that among his papers 
material for a full account of the species would be found, but this unfortunately did 
not prove to be the case. A number of figures accompanied by a few rough labora- 
tory notes alone were avdilable and although his successor, Col. Alcock, was able to 
edit from these an account of nine species, no connected report on the whole collec- 
tion has hitherto appeared. Since 1895, when Wood-Mason’s descriptions were 


2 Memoirs of the Indian Museum. Doc IV, 


published, very considerable additions have been made, largely owing to the pains- 
taking efforts of succeeding Surgeon-Naturalists on the R.I.M.S.S. ‘Investigator.’ 

As may be imagined the task of writing an account of this collection, already 
to a large extent arranged and classified by Wood-Mason, was one of no especial 
difficulty : the manuscript names with which many of the species were supplied have 
been a fruitful source of guidance. It was my original intention to restrict the 
present paper to an account of the Indian species, but it soon became apparent that 
the collection was sufficiently representative to justify work on more ambitious lines, 
and I have conséquently attempted to give a descriptive catalogue of all adult 
Stomatopoda from Indo-pacific waters, indicating as far as a study of the literature 
enables me, the chief points of those species which I have not been able to examine 
myself. Owing perhaps to their peculiar habits many species of Stomatopoda are 
so rare that it is only by abstraction from the work of previous writers that a concise 
account of the forms occurring in a given area can be prepared. 

Under the somewhat loose term ‘‘ Indo-pacific ’’ I include all localities from Suez 
and S. Africa to Australia, New Zealand, Oceania and Japan. The Pacific Coast of 
America is omitted ; for the species of Stomatopoda known from this region show 
little affinity with those of the area defined above and have, moreover, already 
been fully dealt with by Bigelow. 

It is not necessary to treat the history of our knowledge of the recent Stomato- 
poda at any considerable length. H. Milne-Edwards in 1837 gave an account of the 
few species known in his day, and this was succeeded in 1849 by De Haan’s admirable 
chapter on the Japanese species in Siebold’s Fauna Japonica. After the latter work 
our knowledge did not increase very rapidly ; in 1880 Miers was able to comprise 
within the limits of two short papers descriptions of all the forms then known and up 
to the present day this revision has remained the principal work of reference to the 
Indo-pacific forms. Since 1880 a great number of papers dealing with Stomatopoda 
have appeared, scattered through numerous scientific periodicals, and among these 
important contributions to our knowledge of the Indo-pacific species are to be found 
ia the writings of Brooks, de Man, Henderson, Nobili, Hansen, Miiller, Borradaile, 
Tattersall and Fukuda. 

Of publications not dealing directly with Indo-pacific species Bigelow’s valuable 
systematic account of the American forms (1894) and Giesbrecht’s very complete 
monograph of those found in the Mediterranean (1910) will be found indispensable 
to all working at this group of Crustacea. 

At the present day the following one hundred and thirty-nine species and varieties 
of adult Stomatopoda are known.' ‘The names of those which have not been found in 
the Indo-pacific region are printed in italics; an asterisk (*) indicates that examples 
have been available for examination and a dagger (+) implies that the type specimens 
have been seen. 


' Not including Squilla minor, Jurich, and Leplosquilla schmeltzii, A. Milne-Edwards, which are 
based on very immature specimens. 


1913.| S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 


aculeata, Bigelow. 
alba, Bigelow. 
+ *annandalei, Kemp. 
*armata, Milne-Edwards. 
*biformis, Bigelow. 
+ *boops, Kemp. 
chlorida, Brooks. 
*costata, De Haan. 
+ *decorata, Wood-Mason. 
*desmaresti, Risso. 
*dubia, Milne-Edwards. 
*empusa, Say. 
fasciata, De Haan. 
+ *foveolata, Wood-Mason. 
+ *gibba, Nobili. 
+ *gilesi, Wood-Mason, MS. 
+ *gonypetes, Wood-Mason, MS. 
gracihpes, Miers. 
+ *hieroglyphica, Kemp. 
+ *holoschista, Wood-Mason, MS. 
intermedia, Bigelow. 
+ *interrupta, Wood-Mason, MS. 
+ *investigatoris, Lloyd. 
*laevis, Hess. 
*lata, Brooks. 
*latreiliei (Eydoux and Souleyet). 
*leptosquilla, Brooks. 
*mantis, Latreille. 


Oo 


SQUILLA: 


mantoidea, Bigelow. 
*massavensis Kossmann. 
*microphthalma, H. Milne-Edwards. 
miles, Hess. 
*multicarinata, White. 
neglecta, Gibbes. 
*wepa, Watreille: 
*oratonia, De Haan. 
T° do Var penpensan Kemip: 
pallida, Giesbrecht. 
*hanamensis, Bigelow. 
parva, Bigelow. 
*polita, Bigelow. 
*prasinolineata, Dana (Ives)'. 
prasinolineata, Miers'. 
quadridens, Bigelow. 
*quinquedentata, Brooks. 
*raphidea, Fabricius. 
rotundicauda, Miers. 
*yugosa, Bigelow. 
*scorpio, Latreille. 
+* do. var. immaculata, Wood-Ma- 
son, MS. 
+ *stridulans, Wood-Mason. 
+ *supplex, Wood-Mason. 
+ *tenuispinis, Wood-Mason. 
+ *wood-masoni, Kemp. 


PSEUDOSQUILLA. 


cevisit (Roux). 
*ciliata (Fabricius). 
dofleini, Balss. 
empusa (De Haan). 
ferussacı (Roux). 
lessoni (Guérin). 


megalophthalma, Bigelow. 
*oculata (Brulle). 
*ornata, Miers. 

oxythyncha, Borradaile. 
*pilaensis, de Man. 

*stylifera (H. Milne-Edwards). 


1 See p. 201. 


4 Memoırs of the Indian Museum. 


[Vor. IV, 


LYSIOSQUILLA. 


*acanthocarpus, Miers. 
armata, Smith. 
biminiensis, Bigelow. 

do. var. pacificus, Borradaile. 
capensis, Hansen. 
crassispinosa, Fukuda. 
decemspinosa, Rathbun. 
desaussurei, Stimpson. 
digueti, Coutière. 
eusebia, Risso. 
excavatrix, Brooks. 
glabriuscula, Lamarck. 
+ *insignis, Kemp. 
latifrons, De Haan. 


Te GO. 


*maculata (Fabricius). 

var. sulcirostris, Kemp. 
mataguesensis, Bigelow. 
miersi, De Vis. 

+ *multifasciata, Wood-Mason. 
occulta, Giesbrecht. 
plumata, Bigelow. 
polydactyla, v. Martens. 
scabricauda (Lamarck). 
scolopendra (Latreille). 
septemspinosa, Miers. 

+ *spinosa (Wood-Mason). 

+ *tigrina, Nobili. 
vicina, Nobili. 


CORONIDA. 


bradyt (A. Milne-Edwards). 


+ *multituberculata (Borradaile). 


*trachura (v. Martens). 


ODONTODACTYLUS. 


*brevirostris (Miers). 
carinifer (Pocock). 
cultrifer (White). 
elegans (Miers). 
hanseni (Pocock). 


havanensis, Bigelow. 

*japonicus (De Haan). 

latirostris, Borradaile. 

*scyllarus (Linnaeus). 
+ *southwelli, Kemp. 


GONODACTYLUS. 


*acanthurus, Tattersall. 

acutirostris, de Man. 
brevisquamatus, Paulson. 

brooksi (de Man). 
*chiragra (Fabricius). 

f * do. var. platysoma, Wood-Mason. 
*demani, Henderson. 
do. var. espinosus, Borradaile. 

dor var. spinosus, Bigelow. 
drepanophorus, de Man. 

ectypus (Müller). 
*excavatus, Miers. 

festae, Nobili. 


fimbriatus, Lenz. 
folini, A. Milne-Edwards. 
+ *furcicaudatus, Miers. 
*glaber (Lenz). 
*olabrous, Brooks. 
*graphurus, Miers. 
+ *glyptocercus, Wood-Mason. 
guerini, White. 
*herdmani, Tattersall. 
hystrix (Nobili). 
+ *nefandus, Kemp. 
*oerstedi, Hansen. 
*pulchellus, Miers. 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 5 


*spinosissimus, Pfeffer. tanensis (Fukuda). 
spinoso-carinatus, Fukuda. trispinosus, Dana. 
stoliurus (Müller). tuberosus, Pocock. 


The species known from the Atlantic and Mediterranean and from the Pacific 
Coast of America are separately listed at the end of this paper with a few of the more 
important references and synonyms (p. 200). 

The exact relations of the Stomatopoda with other groups of Crustacea present 
a difficult problem. The existing forms show a high degree of specialization coupled 
with several features that at first sight seem primitive. Of the latter the most striking 
is the freely movable ophthalmic and antennular somites, but this is now regarded 
as a secondary character and must also be taken as evidence of specialization. 
With the views which Calman has expressed on the affinities of the group I am in 
entire agreement and his conclusion (1909, p. 331) that it seems most probable that 
“the Stomatopoda are a lateral offshoot from the main stem of the Malacostraca, of 
which, in the absence of connecting links, it is as yet impossible to determine the 
exact relations,” will, I believe, meet with general acceptance. 

The oldest known fossil Stomatopoda have been found in the Jurassic beds of the 
Solenhofen and have been referred to the genus Sculda. Little information of the 
really satisfactory nature is available concerning these forms; but there is, I believe, 
enough to warrant the creation of a separate family for their reception (see p. I5). 
The species are for the most part ornamented with an elaborate sculpture in no respect 
less remarkable than that of some of the existing forms of Squilla and Gonodactylus, 
though of an entirely different type, and it is clear that they also must have attained 
a high degree of specialization. It is to be hoped that specimens will soon be found 
which will provide satisfactory material for a study of the appendages. In parti- 
cular, information regarding the form of the second thoracic appendage, which attains 
such a monstrous development in the Squillidae, would be most valuable. This limb 
has been found in an excellent state of preservation in fossil examples of Squilla, but 
it has never been satisfactorily demonstrated in Sculda. Seeing that in the former 
genus it appears to be well adapted for preservation in stratified rocks it is difficult, 
on the view that it possesses a similar development, to explain its absence in the 
latter. 

Species of Squilla have been obtained in the Cretaceous and in the London Clay 
and other tertiary deposits, and in all structural details which have been studied they 
appear to bear a close resemblance to existing representatives of the genus. 

Brooks held that the forms which he included under the genus Protosquilla (here 
regarded merely as a section of Gonodactylus) were the most primitive that persist ; 
but this view has been severely criticized by Hansen and is not now generally 
accepted. My own observations lead me to conclude that it is among the species of 
Squilla that the most primitive forms are to be sought ; but the precise relationships 
of the several genera are by no means easily traced. 

The genera fall without difficulty into two groups, the one comprising Squilla, 


6 Memoirs of the Indian Museum. No IV, 


Pseudosquilla, Lysiosquilla and Coronida, while the other includes Odontodactylus and 
Gonodactylus. The condition of the ‘ischio-meral’ articulation of the second 
maxillipede or raptorial claw is the character on which this primary division is based. 
This character, indeed, appears to me to possess an importance which has not hitherto 
been appreciated and it shows that the affinities of Coronida, usually regarded as a 
close ally of Gonodactylus, have been to a great extent misunderstood. 

In the attempt to frame diagnoses of the four genera comprised in the first section 
of the family considerable difficulties are encountered, for although each genus 
has indeed a very distinct facies, several isolated forms exist which differ widely from 
the more typical members in structures that are either aberrant or annectant. A 
careful examination of species belonging to these four genera has not yielded me 
material for making any improvement in the classification hitherto adopted. The 
system employed is certainly unsatisfactory, but the groups are for the most part so 
readily distinguished that, even in the absence of good morphological characters, I 
cannot believe that their separation is unwarranted. 

The existing species of Stomatopoda are for the most part restricted to the 
warmer waters of the globe, but a few forms, such as Squilla mantis and S. desmaresti, 
have been found as far north as the British coasts, while others, such as Pseudosquilla 
stylifera, Squilla aculeata, S. gracilipes and S. armata, penetrate as far south as Chili 
and Patagonia, the last being also known from S. Africa and from New Zealand. 

Stomatopoda are exclusively marine, but one form, Squilla scorpio var. immacu- 
lata, extends for a considerable distance up the Ganges delta and must at times be 
brought in contact with water of quite low salinity. 

The large majority of the species are littoral or inhabit moderate depths; only 
the following eleven have been found below the 100 fathom line :— 


Squilla armata. Squilla massavensis. 
Squilla biformis. Squilla stridulans. 

Squilla gilesi.' Squilla tenuispinis. 
Squilla investigators. Lystosquilla insignis. 
Squilla leptosqulla. Lysiosquilla maiaguesensis. 


Odontodactylus havanensis. 


The greatest depth at which any Stomatopod has been found is 370—41g fms. 
( S. leptosquilla— Investigator ’ ). 

Stomatopods are essentially burrowing animals and many of the characteristic 
features of the order are adaptations for this purpose ; but they are not fossorial in 
the sense in which this term is generally used among the Decapoda. The curious 
form and geniculation of the second maxillipedes must be admirably suited to an 
animal which lies in wait for its prey at the mouth of a burrow, while the small 
carapace and loose segmentation of the body-somites enables it to reverse its position 
in a narrow tunnel without i due aley: The peculiarly m modified branchial m is 


Gone in in a trawl fished me ae of 80 and 110 on 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 7 


also in all probability an adaptation to this special mode of life, and it may be noticed 
that among the Decapoda the attachment of the branchial plumes to the swimmerets 
is known in one aberrant genus, Callianidea, the species of which also appear to make 
burrows. 

It is to be regretted that so little is yet known of the habits of the Indo-pacific 
Stomatopoda. Brooks’ brilliant researches on a few Atlantic species and Giesbrecht’s 
more recent investigations of the Mediterranean forms have supplied information of 
great value, but no complete account of the life-history of any Indo-pacific species 
has yet appeared. While it is probable that most of the Squillidae have their own 
specific idiosyncrasies, it seems almost certain that there are certain broad distinc- 
tions in habits between at least some of the different genera of the family. Sgwilla 
appears to prefer a bottom of mud, sand, or sand and shells, and though it evidently 
constructs burrows it must often wander from them, for it is constantly captured in 
the light nets used by fishermen on different parts of the Indian coast. ZLysiosquilla 
on the other hand is in my experience very rarely found, though in reality it may 
perhaps be quite abundant. The species of this genus seem to live in deep burrows 
which they leave only on rare occasions, and to this fact may be attributed the 
scarcity of specimens in collections and the large number of forms which are known 
only from single individuals. The species of the genera Pseudosquilla, Odontodactylus 
and Gonodactylus seem for the most part to inhabit rough ground ; they are often 
found in abundance on oyster-beds and on coral reefs. Brooks has observed that 
the Atlantic Gonodactylus oerstedi burrows into solid coral or limestone rock and it is 
probable that the same is the case with the Indo-pacific species of the genus.' The 
members of the genera Pseudosquilla and Odontodactylus possibly have similar habits, 
but of this nothing is known. 

It is only in very rare instances that other living organisms are found attached 
to Stomatopoda, but five different forms, recently identified and recorded by Dr. N. 
Annandale and Mr. H. B. Preston, have been discovered on material in the Indian 
Museum. 

Squilla holoschista.—Several small undeterminable Balanı were found attached 

to the dorsal surface of the carapace (Annandale, Rec. Ind. Mus., vii, 1912, 
p. 124). 
Squilla sp.°—A few immature barnacles of the genus Dichelaspis (probably 
D. warwickir) were found attached to the pleopods (Annandale, loc. cit.,p. 124). 
Squilla investigatoris.—On the telsons of two of the type specimens of this species 
colonies of Platypolyzoon investigatoris, a new genus and species of Polyzoon, 
were obtained (Annandale, loc. cit., pp. 124—126, text-fig.). 

Squilla fasciata.—On the carapace, raptorial claws and telson of a specimen ‘ent 

us by Prof. K. Kishinouye, colonies of the Polyzoon Triticella korenii, G. O. 
Sars, were found (Annandale, loc. cit., p. 124). 


1 See addendum, p. 197. 
? The specimen was laid aside, prior to identification, and could not subsequently be discovered. 


8 Memoirs of the Indian Museum. [Vor. IV, 


Gonodactylus chiragra.—Two individuals of this species bore on the under surface 
of the thorax and between the pleopods examples of Epistethe gonodactyh, a 
hitherto unknown genus and species of Gastropod Mollusc (Preston, Rec. Ind. 
Mus., vii, I9I2, pp. 126, 127, text-figs.). 

Coutière (1905) gives an interesting account of Lysiosgunlla digueti, a species which 
lives commensally along with a Polynoid in the burrows of Balanoglossus. 

The natives of Bengal for the most part classify marine animals according to the 
estimation in which they are held as articles of liet. For this purpose they consider 
Stomatopods worthless ; they term them chingri-poka (prawn-insects) or darıd-poka 
(séa-insects) in contradistinction to chingri-mu.hli (prawn-fish), 7.e. Penaeidae and 
Palaemonidae. It is, however, possible that they are used for food in some parts of 
the Indian Empire: in the Museum collection is a large example of Squilla raphidea 
which was purchased in the market at Akyab in Arakan. 

Of the one hundred and thirty-nine species and varieties of Stomatopoda which are 
known ninety-seven have been found within the limits of the area dealt with in this 
paper, while fifty-four have been taken on the coasts of the Indian Empire and of 
Ceylon. 


The tables’ on pages 10 and 11 will give a general idea of the known distribution of 
the species in the Indo-pacific area. 


Only a very few forms found in this region have been discovered also in the 
Atlantic or on the West Coast of America :— 


Squilla armata oe .. Chili and Patagonia. 
Pseudosquilla ciliata .. Florida, Bermuda, Porto Rico. 
Pseudosquilla oculata .. Canaries, Madeira. 
Pseudosquilla stylifera .. California. 

Lysiosguilla maculata Wiest lidies: 


In addition Lysiosquila biminiensis, represented in the Indo-pacific by the form 
pacificus, is found at the Bimini Islands, Bahamas, and the Atlantic Gonodactylus 
oerstedi is so closely allied to the Indo-pacific G. chiragra that some will perhaps 
prefer to regard it merely as a subspecies. 

The figures on page 12 are intended to elucidate some of the terms employed in 
this memoir which might otherwise prove confusing. In such matters as the nomen- 
clature of the various carinae of the carapace, abdomen and telson no exact unifor- 
mity is to be found in the literature and I have consequently adopted those which 
appear to be most suitable. 

The form of the eye is often of great importance in systematic work. The 
relation of the corneal and peduncular axes, whether at right angles or oblique, 
affords a valuable criterion and scarcely calls for any explanation (see Bigelow, 1594, 
text-fig. 14, p. 522). In certain species, more particularly in the case of the .closely 


! It will be noticed that the divisions into which the Indo-pacific area has been grouped: for the 
purposes of this table are by no means equal. But the majority of the specimens comprised in the 
collection under examination are from Indian localities and it has been thought best to ewer the distribu- 
tion on these coasts in somewhat fuller detail. ; 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 9 


allied forms belonging to the oratorıa section of the genus Squilla, the precise size of 
the cornea is of great importance ; but inasmuch as its breadth becomes proportion- 
ately less as the animal increases in size, the character must be used with caution. 
The number of times in which the breadth of the cornea is contained in the median 
length of the carapace affords a ready means of comparing the size of the eye in 
allied species, and the figure so obtained I have designated the “‘ corneal index.’ ’ 

The mandibular palp, by the number of segments which compose it and by its 
presence or absence, affords useful taxonomic indications ; but there is reason to 
believe that the importance of the“character is by no means equal in the different 
genera.' In Sqwilla, for instance. its value is purely specific, in Gonodactylus it is of 
considerable assistance in breaking the genus into groups, while in the case of Pseudo- 
squilla stylifera it seems likely that geographical races can be separated by its means. 
The mandibular palp may easily be examined without dissection by deflecting one of 
the great raptorial claws. The palp will then be seen directed forwards and lying 
close up against the side-wall of the carapace (text-fig. 5, p. 12). 

The precise relations which the six segments that compose the thoracic limbs of 
the Stomatopoda bear with the seven (or eight) recognized in other Malacostraca are 
as yet unknown. I have used the familiar terms ischium, merus, carpus, propodus 
and dactylus purely for the sake of convenience: it must not be thought that by 
such application any homology with the Decapod limb is intended. 

In descriptions of species of Squilla it has been found convenient to note the 
presence or absence of the terminal spines on the abdominal carinae in tabular form. 
Squiila nepa, for instance, has the following spine-formula : — 


Carinae. Abdominal somites. 
Submedian te a ate a A5 0: 
Intermediate a er NA) SA NS AIG: 
Lateral 08 > + Sonn (Li) ZA 
Marginal a pe ir Ye Bie By ETES 


By this it is to be understood that the submedian carinae of the last three 
somites terminate in spines. The intermediate carinae of the last four somites and 
the lateral carinae of the last five also have a spinous ending, while, in addition, 
the intermediate carinae of the second somite and the lateral carinae of the first 
somite (figures enclosed in brackets) end in spines in some, but not in all, of the 
specimens examined. All the marginal carinae are produced as spines: the marginal 
carina of the last somite does not exist in any species of Stomatopod and the spine 
which is found at the postero-lateral angle of that somite is a continuation of the 
lateral carina: 

31 Brooks considered that the form of the processes on the endopodite of the first 
abdominal appendage of the male would be found to yield much assistance in the 
taxonomic treatment of the group. In my experience, however, this is not the case 
and I have consequently omitted reference to this appendage in the systematic 
account: of the species. (Cont. on p. 14.) 


! Further reference will be found under the separate generic headings. 


IO Memoirs of the Indian Museum. [Morrie 


I ee | oa 8 
Sole ole i) = VA 
ee 2 
SPECIES. 2 88 8 | S| lee E 
5 m | ot) | v 2 a Sue 3 satires 
5 S 28 a | a | . |A De 08 a Cl 4 A = (ss Ÿ 
DOME) MS Se ESS CNE Se een 
NUS EME aaa 
Dans | EP GO| Ole Re) MEN | as, | es |e © | © 
M4 |A = faye et) ©) (Eur EST a SO) ima | =e || © 14 
| | | | 
SQUILLA. PN eae | | 
S. latreillea oo | 6x | x X 3 X | 
S. decorata 5% | A a ME Aes ILL 
S. gibba xe Sea tee RE ARR tele Ne MOREL ES GER | 
S. microphthalma a eat OMS reg ses De Rn MO lar le 
S. chlorida st BE PSN QE ei Sa eet eto a ee (a | | 
S. rotundicauda .. SN | ay PO LE ite, Ap eae tl eg RE ee) | 
S. fasciata as Sa ear i ROG a emmy HU a || 
S. miles a“ rea ht ice | AS AR ANR AR LS ER RAT SAR RE TERRES ER 
S. lata sche Ree Er Se meee ee NE APT EE x | | 
S. gilesi és UE | Pete aul eee eat lee | | 
S. armata Bi ee | ER tele a a a OG, ce ae Mal N 
S. scorpio se CU] x | X | x | 
do., var. immaculata | x ene dl) 
S. mantoidea rel ea] x | | 
S. leptosquilia sa Rage ENE Ne ees (EDR Ce RS eg SU eae Ne | 
S. tenuispinis | | | De ECO oS | | 
S. laevis sa eau aes A ell a "i Nel eal |e ee rx | | 
S. quinquedentata SATIN BN LE ee ae | | | 
ee ae Boy le, 0 I | x leon ee x : x x | Sy | 
S. foveolata SR) CARRE en ne X Bn | 
S. nepa x RPI ER A SE Ex X Sa | ? 
S. holoschista  .. en N | | Xen ox | | 
S. oratoria ie ne Be | | + XD CN ER oa Eee EX 
do., var. perpensa Fe er a TS | DE NO Res 28 
S.interrupta  .. eo Te AE NES Ex ne de UE Gay so. | 
S.wood-masom .. SEE en: x WE aes Gh ems leo | 
S. massavensis .. Beatie x | | | | 
S.stridulans .. al re hoc a Een 6-0 Se | 
S. investigatoris a | Bana haa lax | | 
S. supplex As es N; EC | | | 
S, costata à N el ee RP ra | Ie hearst had | X | 
S. multicarinata .. I IA Nie ISA iia ln RX x xd | 
S. vaphidea ES D X al MIE EE A XS DE X x | 
S. annandale .. a Ei sl X | 
| | | | | 
PSEUDOSQUILLA. ESP | | Im | 
Ar | EC | 22 KNIE ET ie a oo 
. ornata $3 A reenact eens latent || as oar Noes RE de 28 
P. oxyrhyncha eos A eer ein a Mires. PE ES SR Ge. x 
P. oculata N a ne m AU ee LS AA EN Me X 
P. megalophthalma Sean | | | | 
P. empusa ll ule MA TER Ar | ee a x 
P. dofleini a ls Men eno RE er 
P. pilaensis an ie OX ae Ne saint ES NAS RAU IS 0X | | 
P. stylifera a vos slice) de IMD eon ces lea plan PE N | I EX 
| | 
LYSIOSQUILLA. | | | | 
L. maculata oe FAR PCR ye | RER axe x | Se KURT X IE < | 
do., var. sulcwostris .. | ..\ | | | | X | | 
L. miersi ie at | | | | x | 
L. capensis aa el sean | | | | | 
L. crassispinosa a ee Re uns He ARRET Re rater ee,” 1% CRE EC 
| | 


1913.| 


S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 


| Japan. 


| Australia. 


2: 


| Oceania. 


II 
J 
i=} 
sv) 
"Ss 
(DJ 
N 
= 
® 
A. 
x 


+ | © | = [a] zZ, 
gro. a la te 
ale os | bo 
SPECIES. USE & a dE le 
a a | One) D 1 ei gel 
cD) © a WX a a a leo) ~¢ ia 5 | a ‘An 
D | OVP a oe NT ENS ORNE al eS 
rg re los A fe) aS [ss] ga OH <q a 
DGSE 3 | S| Pliediedles| S| à 
Slee | À |et\om ja iq | & | D 
L. spinosa SE RT RE 
Ib acanthocar pus | RE aX CR NE EX 
L. multifasciata .. Bealls? X Me x 
L. biminiensis subsp. pacificus | ae 
ke tigrina ae Ppl | x 
L. vicina Be | 
L. insignis | : | x 
L. latifrons Ihr 
CORONIDA. | 
C. trachura eee | ee abate ae de ee Has RES 
C. multituberculata DRE on Re ete En tee RE US OR AE 
ODONDODACTYIEUS | 
O. scyllarus 50 Belle au CR, RENE" een RES QUES 
O. cultrifer nae Sa | real] 28188 tb ER en ee Re NE ER by << 
O. carinifer oC SOS er ial ss TE MR ee N LES Er | | x 
O. elegans % So X | | 
O. japonicus a. Meal ete ress eK eves a oe | teed re je 
O. hansent HB SR a a aces Ete a RR re Re EX 
O. latirostris De 8 Im EE | | | | 
O. southwella GE creel weston LEE utes | pete N a RE x | 
O. brevirostris  .. Sic (ser SN | CAN ME PERS GR SE CH | 
GONODACTYLUS. IS es | 
G. chiragra a BE I ESG XGA | a ee OX Xa EX 
do., var. platysoma AR a ee er Ne) 
G. acutirostris .. ne ee 1% 
G. demani oe FAT TAN N RE LS 
do., var. spinosus AN DRAN u CN ee bts | | 
Ds, SVE\ite CSDILOSUS © SBA ass eos || acs! NEC Fee lon lise | 
G. glabrous ere BEE OSes EX EIN EEK KEN EEE lid, IC A| EX Enr 
G. graphurus .. I se Sa Ra ar RE Pe ARR El Ba 
G. herdmani ie Bea a ee. CREER CERTES rn | | 
G. drepanophorus. DRE tg lar seal PRES Re | fie ax 
G. brevisquamatus ala. | 
G. fimbriatus haecceal| so. Gate « 
G. spinoso-carinatus Se 
_G. furcicaudatus 2 ae | ae 
G. pulchellus IEEE < Kae OX: | 
G. nefandus + iio) | EE | | RE RE EX 
G, trispinosus .. ace ni AU Te PEL a ; 
G. tuberosus en RE Rn Re el ae Nox 
G. glaber : Se BA SPORE en ES RS Coal inert lex 
G. tanensis oc I So ee | laren Ne ci lt ILES: 
G. stoliurus a GA | cee ES es tl ea LACS ea E00 < 
G. ectypus : DU DER Ne X set eth >, Gay | | 
G. glyptocercus Ramesh ae ee | | KK nal: 
G. excavatus as Mea ers | ee. eb silos te 1x 
G. brooksi 2 Sl RS a RON a a GE US 
G. hystrix 88 a er Ne ers el Re nee 
G. spinosissimus Oy ioe, Geel ems CANONS er I < | | 
G. guerini as ne tear) (teeta lier Sa ett a UE | 
G. acanthurus .. I) AE nee 


xxx 


I2 


Memoirs of the Indian Museum. 


IVO IV, 


KOCH. S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 
913 § 
EXPLANATION OF TEXT FIGURES. 
Fig. 1. Rostrum, carapace, exposed thoracic somites and first abdominal somite 
(Squilla). 
V-VIII. Fifth to eighth thoracic somites. 
I. First abdominal somite. 8. Cervical groove. 
I. Rostrum. 9. Posterior reflected part of marginal carina. 
2. Antero-lateral angle of carapace. 10. Lateral processes of 5th thoracic somite. 
3. Anterior bifurcated portion of median carina. ıI. Median ) I hoe 1 
dc os» 
5. Lateral fcarna of carapace. 13. Intermediate 
a latente De aie \earinae of abdominal somites. 
7. Gastric groove. 15. Marginal 
Fig. 2. Raptorial claw or second thoracic appendage (Sguilla). 
I. Ischium. 3. Carpus. 
2. Merus. 4. Propodus. 
5. Dactylus. 
Fig. 3. Last two abdominal somites and telson (Squilla). 
VII, VIII. Seventh and eighth abdominal somites. 
1. Median 7. Submedian | 
2. Submedian | 8. Intermediate > teeth of telson. 
3. Intermediate \ carinae of abdominal somites. g. Lateral | 
4. Lateral 10. Submedian | 
5. Marginal II. Intermediate >» denticles of telson. 
6. Median carina of telson. 12. Lateral | 


St ony ESS) NES 


I. First thoracic appendage. 
. Mandibular palp. 


. First intermediate 
. Second intermediate 


13. Prae-lateral lobe or denticle of telson. 


Fig. 4. Last abdominal somite and telson (Odontodactylus). 


Submedian \ 8. Intermediate 


f carinae of last abdomi- 9 First lateral 


: i f telson. 
| aha > Carinae of telson 


10. Marginal 


. Lateral / II. Second lateral 

. Median 12. Submedian ) 

. Submedian carinae of telson. 13. Intermediate teeth of telson. 
. Second submedian 14. Lateral ) 


Fig. 5. Lateral view of carapace, raptorial claw deflected (Squilla). 


3. Third thoracic appendage. 
4. Base of raptorial claw. 


I4 Memoirs of the Indian Museum. Nora 


Keys have been provided to facilitate the determination of the species. Those 
of Pseudosquilla, Coromda, Odontodactylus and Gonodactylus embrace all species 
hitherto known, but those of Squilla and Lysiosquilla have reference only to the 
Indo-pacific forms. 

In the course of the preparation of this report I have received much assistance 
from museums and institutions in other countries. Through the good offices of Dr. 
W. T. Calman I have obtained from the Trustees of the British Museum the loan of 
a most valuable series of Squilla belonging to the S. nepa and S. oratoria groups. 
In many other matters also I am under a great debt of obligation to Dr. Calman ; he 
has always been ready to place his very special knowledge of Crustacea at my dis- 
posal and he has spared neither time nor trouble in obtaining for me particular 
items of information from the collections in his charge. To Mr. J. C. Moulton I am 
indebted for the loan of a small but very valuable collection from the Sarawak 
Museum. Mr. T. Southwell has kindly presented us with a fine series of Stomatopoda 
from the Ceylon pearl banks, and I wish also to thank Dr. Pearson for sending me, on 
loan, the entire collection in this group from the Colombo Museum. 

It is probable that before this paper is published Mr. Patience’s account of the 
Stomatopoda collected in Portuguese E. Africa and the Mergui Archipelago by 
Messts. Simpson and Rudmose Brown, will have appeared. Mr. Patience has kindly 
sent me examples of nearly all the species of the order which he had in his 
hands, and in so doing he was good enough to give me permission to make any 
reference to them that I desired. In addition I have received specimens of several 
scarce and interesting species from Prof. Kishinouye, Dr. Chilton, Mr. David Stead _ 
and Mr. Alan Owston and, through the kindness of Mr. Doncaster, I have been 
enabled to examine the type specimens of Coronida (Squilla) multituberculata. 

In India itself I have been fortunate in obtaining the co-operation of the Bombay 
Natural History Society, which, in past years, has been instrumental in acquiring for 
us many species of crustacea from the west coast of India, and, in addition to the 
members of the staff of the Indian Museum, Mr. J. Hornell at Tuticorin and Mr. T. 
H. Hill at Cuddalore and Porto Novo have both made special collections on my behalf. 

The plates illustrating this memoir are issued as ‘‘ Illustrations of the Zoology of 
R.I.M.S.S. ‘Investigator.’ ‘” The figures were all drawn under my supervision by 
Babu S. C. Mondul, Artist to the Marine Survey. The care which he has taken in 
this work and the fidelity of his delineation has, I feel, added materially to the value 
of the report. 

Unless otherwise stated the type specimens are preserved in the Indian Museum. 


LOS) S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 15 


SYSTEMATIC PART. 
STOMATOPODA, 


The Stomatopoda belong to the sub-class Malacostraca and constitute the only 
order of the last of the four divisions into which the second series of the sub-class is 
divided: This division is known by the name Hoplocarida, and is defined by Calman 
(1909) in the following terms :— 

“ Carapace leaving at least four of the thoracic somites distinct ; two movable 
segments are separated from the anterior part of the head, bearing respectively the 
pedunculate eyes and the antennules ; protopodite of antenna of two segments ; 
mandible without lacinia mobilis ; posterior thoracic limbs with protopodite of three 
segments (the relation of the segments of the anterior thoracic limbs to those of the 
limbs in the other divisions is doubtful) ; an appendix interna on pleopods; hepatic 
caeca much ramified ; heart much elongated, extending through thoracic and abdomi- 
nal regions ; spermatozoa spherical; development with metamorphosis, a free-swim- 
ming nauplius stage is not certainly known.’’ 

Since the time when it was discovered that the forms classed as Erichthidae 
were nothing more than larval stages, the order Stomatopoda has been held to com- 
prise only a single family, the Squillidae. It seems, however, that the species 
belonging to the fossil genus Sculda, Munster, are distinguished from the recent 
forms by features sufficiently important to entitle them to separate family rank and 
it may be expected that, with the acquisition of more perfect specimens, other 
characteristic differences will be disclosed. In the present state of our knowledge the 
families may be separated from one another as follows :— 


I. Exopodite of uropod composed of a single segment an .. SCULDIDAE 
(fossil). 
II. Exopodite of uropod composed of two quite distinct segments .. SQUILLIDAE 


(fossil and recent). 


In the Sculdidae the spines on the margin of the telson appear to have all been 
movable and in this respect differ from those of recent Stomatopoda, while from the 
figures which Kunth gives (1870) it seems that only three thoracic segments are 
left uncovered by the carapace. The form of the raptorial claw is unfortunately 
unknown,' but that the limb is far less conspicuous than in the Squillidae may be 
surmised from the fact that, though no trace of it can be discovered in Sculda, it has 
been found in the much scarcer fossil Squilla.” 


' Kunth (1870) is of the opinion that the raptorial claw is traceable in one of his specimens of 
Sculda pennata in which the under surface is shown, but in his figure (pl. xvii, fig. 2) this is far from 
obvious; it seems more probable that the part referred to is merely a portion of the reflexed lateral 
margin of the carapace. 

* Squilla lewisi, Woodward, Q. J. Geol. Soc., xxxv, 1879, pl. xxvi, fig. 4. 


16 


Memoirs of the Indian Museum. OPEN 


Family SQUILLIDAE. 


To this family all recent Stomatopoda are referred, along with a few fossil species 
from the London clay and from the Cretaceous rocks of Lebanon, and Necroscilla 
wilsont, Woodward, a form of doubtful affinities described from a fragment obtained 


in the middle coal-measures. 
The existing forms may be classed under six genera which are distinguished 


as follows :— 


I. Articulation between merus and ischium of raptorial claw terminal 
(normal) ; merus grooved inferiorly for reception of propodus throughout — 
its length ; propodus finely pectinate or with a series of fixed spines along 
outer edge of dorsal surface; dactylus rarely inflated at base, 
Carapace with well-marked carinae; cervical groove defined across 
dorsum of carapace; first five abdominal somites with longitudinal 
carinae (raptorial dactylus not inflated at base) ae 3 .. Squilla, p. 16. 
Carapace without carinae ;! cervical groove not extending across 


I 


dorsum of carapace; ! first five abdominal somites without longitu- 


dinal carinae.! 


T. 


Abdomen usually compressed ; raptorial dactylus not inflated at 

base with two, rarely three, teeth on inner margin,” or unarmed; 

telson with sharp median carina and (in adults) with other 

carinae on either side BR ah ms .. Pseudosquilla, p. 94. 
Abdomen depressed; raptorial dactylus not inflated at base 

with atleast four teeth on inner margin”; telson without median 

carina, often smooth or with a transverse circlet of spines .. Lysiosquilla, p. 100. 
Abdomen depressed ; raptorial dactylus with three teeth on 

inner margin”; telson closely studded with fine spinules or large 

tubercles, with or without a pair of submedian carinae.. .. Coronida, p. 129. 


II. Ischio-meral articulation of raptorial claw situated at a point anterior 
to proximal end of merus, which consequently extends backwards con- 


siderably beyond the joint; ventral surface of merus grooved and 

hollowed for reception of propodus for not more than three-quarters its 

length ; dactylus inflated at base. 

A. Dactylus of raptorial claw with teeth on its inner margin .. .. Odontodactylus, p. 133. 


B. Dactylus of raptorial claw without teeth on its inner margin .. Gonodactylus, p. 145. 


1793: 
1825. 
1837. 
1841. 


Genus Squilla, J. C. Fabricius. 


Squilla, J. C. Fabricius, Ent. Syst., ii, p. 5II (partim). 

Squilla, Latreille, Encycl. Méthod., X, p. 467 (partim). 

Squilla, Squilles fine-tailles, 2nd sect., H. Milne-Edwards, Hist. Nat. Crust., II, pp. 518, 520. 
Clorida, Eydoux and Souleyet, Voy. de la ‘ Bonite,’ Zool., i, Crust., p. 264. 


| Except in the Mediterranean Pseudosquilla ferussaci. I have seen no examples of this very rare 
and interesting Mediterranean species. It appears to combine the telson of a Pseudosquilla with many 
of the characteristic features of Sqwilla, but is perhaps rather more nearly allied to the former genus than 


to the latter. 


For an account and figures of the species, see Giesbrecht, 1910, p. 34, pl. iv, figs. 37-48. 


? Excluding the terminal tooth, 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 17 


1849. Squilla, sect. Raphideae, De Haan, in Siebold’s Fauna Japonica, Crust., p. 221. 
1852. Squilla, Dana, U.S. Explor. Exped., Crust., p. 618. 

1880. Squilla, Miers, Ann. Mag. Nat. Hist. (5), V, p. 16. 

1880. Chloridella, Miers, ibid., p. 13. 

1880. Leptosquilla, Miers, ibid., p. 12. 

1886. Squilla, Brooks, Voy. H. M. S. ‘Challenger,’ XVI, Stomatop., p. 23. 

1890. Pterygosquilla, Hilgendorf, Sitz.-ber. Ges. naturf. Freunde, Jahrg. 1890, pp. 172, 187. 
1894. Squilla, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 509. 

1894. Pterygosquilla, Bigelow, ibrd., p. 492. 

1894. Leptosguilla, Bigelow, tbid., p. 492. 

1895. Squilla, Hansen, Isop., Cumac. u. Stomatop., Plankton-Exped., p. 69. 

1903. Chloridella,’ Rathbun, Proc. U. S. Nat. Mus., XXVI, p. 54. 

1910, Squilla, Stebbing, Ann. S. African Mus., VI, p. 405. 


Carapace narrower in front than behind, with conspicuous gastric and cervical 
grooves, the latter sharp and continuous across mid-dorsal area ; longitudinal carinae 
usually distinct, never wholly absent ; antero-lateral angles usually spinous. Cornea of 
eyes bilobed. Mandibular palp three-segmented or wholly absent. Merus of raptorial 
claw articulating terminally with ischium ; ventral surface of merus longitudinally 
hollowed throughout its length ; upper margin of propodus finely pectinate, or (more 
rarely) with a series of long stiff spines ; dactylus not inflated at base,armed with teeth 
on inner margin. Shorter ramus of last three thoracic appendages unjointed, or with 
an imperfect suture separating short basal and long linear distal parts. Free thoracic 
and abdominal somites depressed, with distinct longitudinal carinae. Telson of good 
length, with a median carina, and smooth, pitted, tuberculate or carinate on either side. 
Distal margin with three pairs of large teeth, submedians occasionally with movable 
tips ; one or more submedian denticles, several (always more than four) intermediate 
and one lateral. Ventral process of uropods terminating in two sharp spines, inner 
longer than outer and nearly always bearing a small lobe on its external margin. 

Well-marked secondary sexual distinctions occur in some species. The modifica- 
tions, which are found only in the adult male, affect the structure of the raptorial 
claw or the development of the carinae of the abdominal somite and telson. 


Miers recognized a distinct genus under the name Chloridella, comprising those 
species which possess very small eyes ; but Brooks has shown that these forms are 
linked to the more typical representatives of the genus by such species as S. /ata and 
S. fasciata. Hansen regards Hilgendorf’s Pterygosquilla as a synonym of Squilla and 
identifies P. laticauda, the only known species of the former genus, with the earlier 
Squilla gracilipes of Miers. Leptosquilla appears to be founded only on a late post- 
larval stage, and, although it cannot be identified with any known adult form, there 
is little doubt that it may safely be referred to this genus. 

Squilla, as has already been mentioned, is the oldest established genus of Stomato- 
poda at present living and probably contains the most primitive species of the order ; 


! Miss Rathbun considers that the name Squilla is preoccupied, but this view is not accepted by 
Stebbing (1910). 


18 Memoirs of the Indian Museum. | [Vor. IV, 


it comprises a large number of species inhabiting all the warm and temperate waters of 
the globe. Comparison of the species from the Atlantic, Mediterranean and W. Coast 
of America with those from the Indo-pacific seems to show that evolution has on the 
whole taken a distinct and separate course in the latter area. In the Indo-pacific a 
single species, Squilla mantoidea, is the sole representative of the section to which the 
vast majority of Atlantic, Mediterranean and W. American forms belong—a section 
which includes all the common species in those localities.! On the other hand the 
Indo-pacific section that surpasses all others in point of individual abundance and 
comprises some fourteen” species, is represented in the other regions only by the West 
Indian Squilla alba. The ‘ Chloridella’ group, moreover, is found only in the Indo- 
pacific. 

One species only, Squilla armata, an isolated form wholly restricted to southern 
latitudes, is common to both areas. 


A peculiar structural feature of the genus and one that, hitherto, has largely 
escaped attention,is the complete suppression of the mandibular palp in certain species. 
Hilgendorf first noticed its absence in the form which he called Pterygosquilla laticauda 
and, more recently, Jurich has made a similar discovery in the case of Squilla lepto- 
sguilla. Among the species which I have been able to examine eight show no trace of 
this appendage :— 


Squilla gibba, Nobili. Squilla scorpio, Latreiïlle. 
Squilla armata, Milne-Edwards. Squilla laevis, Hess. 

Squilla leptosquilla, Brooks. Squilla hieroglyphica, Kemp. 
Squilla tenwispinis, Wood-Mason. Squilla costata, De Haan. 


Squilla leptosquilla and S. tenuispinis are closely allied forms and the same is the 
case with S. /aevis and S. hieroglyphica. 

In Decapoda the structure of the mandibular palp, its presence or its absence, is 
recognized as a valuable clue to the affinities of the various species and in many cases 
it affords almost the sole distinction between genera.. It is clear that its importance 
is far less in the Squillidae. Sguilla gibba by many evident external characters takes 
its place in the ‘ Chloridella’ section of the genus, though it is perhaps a somewhat 
aberrant member thereof ; S. scorpio finds distant allies in S. data and S. gilesi; S. 
laevis and S. hreroglvphrca seem to form a small subsection of the S. nepa and quinque- 
dentata group, while S. costata by the number of carinae on its abdomen appears to be 
related to S. multicarinata. Squilla armata seems to be an isolated species and the 
same is the case with S. leptosquilla and tenuispims, which are restricted to deep water. 

It seems then that the palpless species found in the Indo-pacific fall into six sep- 
arate classes, which show no trace of common ancestry distinct from that of the other 
species of the genus, and the inference that the palp has been suppressed on no less 
than six separate occasions in the ontogeny of the Indo-pacific species can scarcely 
be avoided. 


' Such as S mantis, empusa, dubia, prasinolineata and panamensis. 
* S. laevis, quinquedentata, nepa, oratoria, stridulans. etc. 


1913. | S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 19 


It will be noticed that the palp in this genus is either well formed and composed 
of three segments, or wholly absent ; no instances of a one or two-segmented palp 
have been observed. | 


The inter-relationships of the Indo-pacific species cannot be traced with any great 
degree of satisfaction. S. vaphidea and S. annandalei, in the possession of large fixed 
spines on the penultimate segment of the raptorial claw, stand apart from all the rest, 
and the latter fall into two groups characterized by the structure of the fifth thoracie 
somite. The first of these groups contains the small-eyed species originally placed 
in. a separate genus, but linked to the more normal types by such forms as 5. chlorida, 
S. fasciata, S. lata and S. gilesi. It also contains the highly-specialized deep-water 
section, comprising S. leptosguilla and tenuispinis and a few isolated species, S. scorpio, 
S. armata and S. mantoidea. The second group, which in point of individual abund- 
ance greatly outweighs the first, contains a number of closely-allied species that fall 
into three sections, typified respectively by S. laevis, S. quinquedentata and S. nepa. 
In addition it includes four highly-specialized forms, S. investigatoris, S. costata, S. 
multicarinata and S. supplex. The last is perhaps a remote connexion of S. daevis ; 
the other three appear to be early offshoots of the original nepa and quinquedentata 
stock ; but in each of them specialization has progressed on different lines. | 


The species of Squilla for the most part inhabit shallow water and seem to prefer 
sandy or muddy ground. They construct burrows ; but as they frequently leave them 
in pursuit of prey, they are often taken in abundance in fishermen’s nets. Giesbrecht 
(1910) gives a valuable account of the bionomics of Squilla mantis in the Mediterranean. 


Brooks (1886, p. 168) states that ‘‘ Squwilla stridulates by rubbing the serrated 
spine of the swimmeret across the serrated ridge on the ventral surface of the telson. 
The noise which is thus made under water can be clearly heard above the surface.’’ 
This observation was made on Squilla empusa. Giesbrecht (1910, p. 201) found that 
Squilla mantis produced sound in a similar way, but only when it was grasped with a 
pair of forceps. From the structure and relative position of the parts the method of 
sound-production is not very clear ; but most Indo-pacific species possess a lateral 
carina on the under surface of the telson as in S. empusa and S. mantis, and it is prob- 
able, therefore, that stridulation is effected in the same way: my efforts to induce 
living Squilla to produce sound have, however, proved quite unsuccessful. Wood- 
Mason unfortunately gives no reason for his choice of stridulans as.a specific name and 
his specimens do not seem to possess any special structural distinctions which would 
furnish an explanation. 


KEY TO THE INDO-PACIFIC SPECIES OF Squilla. 
The following key to the Indo-pacific species of Squilla is based on adult 
characters ; great caution should be used when employing it for specimens under 
40 mm. in length. 


I, Upper margin of propodus of raptorial claw with a series of fine and 
even pectinations in addition to a few movable spines at proximal end. 


D 
© 


Memoirs of the Indian Museum. 


A. Lateral margin of fifth thoracic somite, seen in dorsal view, composed 
of a single process, spinous or subacute. 
I. Cornea of eyes very small, its breadth less than greatest breadth 
of stalk. 
A. First five abdominal somites with faint but distinct sub- 
median carinae. 

I. Mandibular palp present; telson with numerous granular 
ridges or scattered tubercles on either side of median 
erest. 
A. Lateral process of fifth thoracic somite directed 

outwards, those of two succeeding somites an- 

teriorly rounded; inferior surface of telson smooth 
on either side of post-anal carina 
B. Lateral process of fifth thoracic somite bent ob- 
liquely forwards, those of two succeeding somites 
acutely pointed anteriorly; inferior surface of telson 
usually with tubercles or granular ridges on either 
side of post-anal carina .. a5 : 

II. Mandibular palp absent; telson without dorsal tuber- 

cles: in adult male with a pair of huge smooth curved 

ridges terminating on intermediate marginal teeth 

B. First five abdominal somites without trace of submedian 
carinae. 

I. Antero-lateral angles of carapace sharply spinous. 

A. Breadth of cornea about one-third of total length of 

eye; lateral process of fifth thoracic somite short 
B. Breadth of cornea about one-half of total length of 
eye; lateral process of fifth thoracic somite long... 

| II. Antero-lateral angles of carapace rounded 
II. Cornea of eyes small or large, its breadth always greater than 
greatest breadth of stalk. 
A. Numerous sharp longitudinal carinae on either side of median 
crest of telson. 

I. Antero-lateral angles of carapace spinous; raptorial 
dactylus with six teeth'; no submedian carinae on 
first five abdominal somites .. Fe 

II. Antero-lateral angles of carapace not spinous; raptorial 

dactylus with four teeth!; submedian carinae on all 
abdominal somites a De 
B. No longitudinal carinae on either side of median crest of telson. 

I. Raptorial dactylus with more than four teeth!; telson 
with short (normal) marginal teeth. 

A. Median carina of carapace, when present, not bifur- 
cated anteriorly. 

I. Median carina of carapace absent; mandibular 
palp present; raptorial dactylus with six 
teeth!; lateral processes of sixth and seventh 
thoracic somites posteriorly rounded. 

a. Submedian carinae absent from at least first 


[Vor. IV, 


. latreillei, p. 24. 


. decorata, p. 27. 


. gibba, p. 28. 


microphthalma, p. 31. 


chlorida, p. 33. 


. rotundicauda, p. 33. 


. fasciata, p. 34. 


. miles, p. 36. 


! Including the terminal tooth. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 


three abdominal somites; no marked 

secondary sexual distinctions ; 

b. Submedian carinae present on all abdom- 

inal somites; telson of adult male differ- 

ing greatly from that of female 

2. Median carina of carapace absent; mandibular 

palp absent; raptorial dactylus with seven to 

nine (rarely six) teeth! ; lateral processes of 

sixth and seventh thoracic somites spinous 

posteriorly 5 . ; 

3. Median carina of carapace present; mandibular 

palp absent; raptorial dactylus with five 

teeth!; lateral processes of sixth and seventh 
thoracic somites posteriorly rounded. 

a. Lateral process of fifth thoracic somite with 

a large black dorsal spot : 

b. Lateral process of fifth thoracic somite 

without a black spot oe as 

B. Median carina of carapace sharply defined and an- 
teriorly bifurcated [raptorial dactylus with six 
teeth! ; lateral processes of sixth and seventh 
thoracic somites sharply acute posteriorly | 

II. Raptorial dactylus with only four teeth!; submedian 
and intermediate marginal teeth of telson very long 

[mandibular palp absent]. 

A. Median and lateral carinae sharp and distinct in an- 
terior half of carapace; lateral process of fifth 
thoracic somite directed straightly outwards 

B. Median and lateral carinae entirely absent from 
anterior half of carapace; lateral process of fifth 
thoracic somite trending obliquely forwards 

B. Lateral margin of fifth thoracic somite, seen in dorsal view, composed 
of two distinct processes, usually of a more or less antrorse ante- 
rior spine and a rounded posterior lobe, rarely of two sharp spines. 
I. First five abdominal somites each with eight longitudinal carinae. 

A. Lateral margin of sixth thoracic somite not bilobed; mandi- 
bular palp absent. 
I. Anterior bifurcation of median carinae of carapace sharp 
and distinct ; raptorial dactylus with six teeth ! 
II. Anterior bifurcation of median carinae of carapace absent ; 
raptorial dactylus with five teeth ! ; 
B. Lateral margin of sixth thoracic somite bilobed, anterior lobe 
rarely (2.e. in S. boops) small and inconspicuous; mandi- 
bular palp present. 
I. Raptorial dactylus with only five or six teeth. ! 

4. Raptorial dactylus with only five teeth.! 

I. Rostrum with median carina; breadth of cornea 
more than one-third of median length of cara- 


la p. 37. 


. gilesi, p. 29. 


. armata, p. 4I. 


. Scorpio, p. 42. 


do. var.immaculata, p. 45. 


. mantoidea, p. 45. 


.. leptosquilla, p. 46. 


. tenwispinis, p. 48. 


. laevis, p. 49. 


. Mmeroglyphica, p. 51. 


| Including the terminal tooth, 


D: 
D 


Memoirs of the Indian Museum. 


pace; lateral margin of seventh thoracic somite 

not bilobed 2 08 : 

2. Rostrum without median carina; breadth of 
cornea much less than one-third of median 
length of carapace; lateral margin of seventh 
thoracic somite distinctly bilobed. 

a. Cornea set almost transversely on eye- 
stalk; outer inferior margin of raptorial 
merus terminating in a sharp tooth; an- 
terior lobe of seventh thoracic somite short 

b. Cornea set obliquely on eyestalk ; outer in- 
ferior maigin of raptorial merus not ter- 
minating in a tooth; anterior lobe of 
seventh thoracic somite well-developed, 
more than half as long as posterior lobe .. 

B. Raptorial dactylus with six teeth.! 

I. Cornea not wider than eyestalk; rostrum with 
short median carina; surface of carapace and 
abdomen with a coarse mesh-like reticulation 

2. Cornea very clearly wider than eyestalk; ros- 
trum without median carina; surface of cara- 
pace and abdomen smooth or rugose, never 
reticulate. 

a. Cornea set transversely on eyestalk. 

i. Posterior half of median carina of cara- 
pace, anterior to cervical groove, 
simple; submedian carinae of fourth 
abdominal somite ending in spines .. 

ii. Posterior half of median carina of cara- 
pace, anterior to cervical groove 
finely bicarinate throughout almost 
its entire length; submedian carinae 
of fourth abdominal somite not ending 
in spines 50 oe 

b. Cornea set very obliquely on eyestalk. 
i. Lateral carinae of first five abdominal 

somites simple—not bicarinate. 
a. No rows of tubercles on either side 
of median crest of telson. 

(i) Anterior breadth of carapace 
less than half its median 
length, including rostrum; 
anterior bifurcation of medi- 
an carina of carapace rarely 
altogether obsolete ; anterior 
margin of ophthalmic somite 
convex, truncate or emargin- 
ate; dorsal surface of cara- 


[Vor. IV, 


. boops, p. 55. 


quinquedentata, p. 52. 


gonypetes, p. 54. 


foveolata, p. 58. 


nepa, p. 60. 


. holoschista, p. 64. 


1 Including the terminal tooth. 


1913.] 


S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 


pace and abdomen rare- 

ly smooth, never highly 

polished. 

(a) Median carina of cara- 
pace sharp and distinct 
throughout its course; 
dorsal carina of rap- 
torial carpus with 3-5 
tubercles .. 3 

(8) Median carina of carapace 
interrupted at base of 
anterior bifurcation !, 
dorsal carina of rap- 
torial carpus with less 
than three tubercles. 

* Dorsal carina of rap- 
torial carpus entire, 
but terminating ab- 
ruptly before reach- 
ing anterior margin ; 
margin of longer 
spine of bifurcate 
process of uropods 
in front of external 
lobe very clearly 
concave : 

**Dorsal carina of rap- 
torial carpus with 
‘two prominent tu- 
bercles; margin of 
longer spine of 
bifurcate process of 
uropods in front of 
external lobe con- 
WSS 06 ch 


(ii) Anterior breadth of carapace 


less. than half its median 
length, including rostrum ; 
anterior bifurcation of medi- 
an carina of carapace alto- 
gether obsolete; anterior 
margin of ophthalmic somite 
with a minute median point ; 
dorsal surface of carapace 
and abdomen smooth and 
highly-polished 


23 


. oratoria, p. 66. 


. do. var. perpensa, p. 70. 


interrupta, p. 72. 


. wood-masont, p. 74. 
B. Distinct rows of tubercles on either 


side of median crest of telson .. massavensis, p. 76. 


' In rare instances the whole anterior bifurcation is obsolete as in S. wood-masont. 


24 Memoirs of the Indian Museum. 


ii. Lateral carinae of first five abdominal 

somites bicarinate 

II. Raptorial dactylus with ten to eighteen (usually an 

to seventeen) teeth ! re ; 

Il. First five abdominal somites each with more than eisht longi- 
tudinal carinae. 

A. Lateral process of sixth thoracic somite not bilobed; first 
five abdominal somites witb a median carina, making nine 
in a transverse series [mandibular palp present; raptorial 
dactylus with five teeth || ; 

B. Lateral process of sixth thoracic somite bilobed; numerous 
sharp carinae on abdominal somites. 

I. Carapace with normal number of carinae and with 
numerous short ridges and tubercles; abdomen tuber- 
culate laterally with numerous carinae in middle line; 
mandibular palp absent; raptorial dactylus with six or 
seven teeth ! : Sig : 

II. Entire surface of carapace and en multicarinate ; 

mandibular palp present; raptorial dactylus with five 

Lee baw aemare . 

II, Upper margin of propodus of pir claw ith an series “al pectina- 

tions replaced by stiff spines; spines well-spaced, large and small alter- 

nating more or less regularly (raptorial dactylus with nine, rarely eight, 
teeth). ! 

A. Antennular peduncle shorter than rostrum and carapace combined; 
lateral margin of sixth thoracic somite not distinctly bilobed; sub- 
median carinae of fifth abdominal somite not ending in spines; ulti- 
mate segment of outer uropod suffused with black only on its inner 
longitudinal half 00 56 ays 5 

B. Antennular peduncle fully as long as rostrum and carapace combined; 
lateral margin of sixth thoracic somite distinctly bilobed ; submedian 
carinae of fifth abdominal somite ending in spines; ultimate segment 
of outer uropod jet-black with a white midrib 


[VORA 


. stridulans, p. 78. 


. investigatoris, p. 80. 


. supplex, p. 82. 


. costata, p 84. 


. multicarinata, p. 86. 


. vaphidea, p. 88. 


. annandalei, p. 92. 


ı. Squilla latreillei (Eydoux and Souleyet). 


Plate I, figs. I-4. 


1841. Ciorida latreillei, Eydoux and Souleyet, Voy. de la ‘ Bonite, Zool., I, Crust., p. 265, pl. v, 


figs. 2-5. 
1880. Chloridella latreillei, Miers, Ann. Mag. Nat. Hist. (5), V, p. 15. 
1894. Squilla latreillei, Bigelow, Proc. U. S. Nat. Mus. XIV, p. 500. 


1895. Chloridella latreillei, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 6, pl. iv, figs. 6-13. 


The dorsal surfaces of the carapace and abdomen are smooth and without trace 
of punctuation. The carapace is more or less triangular in shape ; anteriorly it is a 
little more than half as broad as long, while its greatest width measured in front of the 
postero-lateral angles is about equal to its median length. Except for an indistinct 


! Including the terminal tooth. 


1013.| S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 25 


ridge running close to the inner edge of the gastric groove, the only carina on the cara- 
pace is a short portion of the marginal. It is invisible anteriorly, but defines the 
posterior edge on either side and is curved internally, extending forwards for a short 
distance on the surface of the carapace. The sharp teeth at the antero-lateral angles, 
owing to the strong convexity of the anterior margin, fail to reach the level of the 
rostral base. 

The rostrum is smooth above and at the base fully twice as broad as its median 
length. The lateral margins are usually a trifle upturned and converge rapidly to a 
broad, rounded apex. 

The eyes (fig. 3) are very broadly pyriform, little more than one and a half times 
as long as broad, and do not nearly reach to half the length of the basal segment of the 
antennular peduncle. The very small bilobed cornea is only about half the width of 
the greatly swollen median portion of the stalk. The inner margins of the stalk are 
straight, flat, and in close juxtaposition for three-quarters of their basal length ; the 
external margins are feebly keeled. 

The antennular peduncle is as long as, or a trifle longer than, the carapace. The 
two basal segments of the antennal endopodite are very long and reach as far as the 
apex of the scale. 

The mandibular palp is composed of three segments. 

The outer inferior margin of the merus of the raptorial claw is bluntly rounded 
distally. The dorsal carina of the carpus is sharp and entire, but terminates abruptly 
before reaching the distal edge. Externally the carpus is grooved and obscurely 
ridged. The propodus has three proximal movable spines, the second being the 
shortest, and a series of fine pectinations along the margin opposed to the dactylus. 
The dactylus bears four or five teeth including the terminal one, the fifth or proximal 
tooth, when present, is usually small' and lies close up against the next of the series. 
The outer margin of the dactylus is strongly convex and bears a rounded prominence 
at its base. 

The last three thoracic somites bear intermediate carinae ; submedian carinae may 
also be found, but as a rule are very obscure or wholly absent on all the somites except 
the last. The fifth somite is produced laterally to a short and sharp point directed 
almost transversely outwards (fig. 4) ; the pair of spines found on the inferior surface 
in certain allied species is obsolescent or entirely missing. The lateral margins of the 
sixth and seventh somites are slightly upturned and truncately rounded ; those of the 
eighth project anteriorly in the form of a small acute or subacute lobe. 

The abdominal somites are greatly depressed. The first five bear four pairs of 
longitudinal carinae of which the submedian and intermediate, though always clearly 
visible, are faintly marked. All the somites, except the first and last, bear a small 
transversely-grooved tubercle in the middle line. The six carinae of the sixth somite 
are strongly swollen in the adult male and, in both sexes, the submedians are slightly 


' The dactylus of a male specimen figured by Wood-Mason (1895, pl. iv, fig. 9) is quite abnormal 
in the extreme reduction of {wo proximal teeth. 


26 Memoirs of the Indian Museum. LWoiG. JOY, 


convergent posteriorly and the intermediates more or less curved inwards. The follow- 
ing abdominal carinae end in spines ' :— 


Carinae. Abdominal somites. 
Submedian Ei aie 3 6. 
Intermediate .. ae ar 5,6. 
Lateral IR 00 06 5,6. 
Marginal Le ss de (2) (3) 4, 5. 


The telson (figs. I, 2) is considerably broader than long and is strongiy convex 
dorsally. The median carina is distinct, very obscurely notched at the base, and in 
most cases terminates in a short spine; on either side of it there are a number of 
ridges which are usually broken up into tubercles. Close to the median carina on each 
side a row of from three to six tubercles is usually found, the two rows converging 
posteriorly and meeting in a small tubercle immediately behind the distal end of 
the carina. Beyond this there are a number of oblique ridges—in some cases very 
obscure, in others sharp and broken up into a series of tubercles—and, in addition, 
several isolated tubercles or transverse tuberculate ridges may usually be seen close to 
the proximal margin. The six marginal teeth are acute but short ; the submedian pair 
is provided with minute movable tips, but these in most well-grown specimens are 
broken off. The outer edges of at least the intermediate pair are serrate and a small 
blunt praelateral lobe is usually distinct. There are one or two pairs of submedian 
denticles, four to seven intermediate, and one lateral, and these like the primary teeth 
frequently possess serrate margins. The ventral surface of the telson is quite smooth 
except for a short, but well-marked, post-anal crest. 

In adult males (fig. 2) the median carina is rather more swollen than in the female 
(fig. 1) and there is an enormously inflated triangular swelling at the base of each mar- 
ginal tooth. The tubercles, moreover, are generally more rounded and less conspicuous 
than is the case in the other sex. 

Wood-Mason (1895, pl. iv, figs. 8, 12) has given two figures of the telson of this 
species and two additional figures will be found on Pl. I, figs. 1 and 2. The latter, 
which were drawn from specimens obtained in the Persian Gulf, differ considerably 
from the more normal types figured by Wood-Mason, that of the female, in particular, 
showing quite an extreme tubercular development. 

The peduncular segment of the outer uropod (figs. I, 2) is provided with a single 
dorsal spine near its articulation with the exopodite. The inner spine of the ventral 
bifurcate process is more than twice the length of the outer and bears a well-developed 
lobe on its external aspect. Internally the process is armed with five to eight sharp 
spines. The spines on the outer margin of the basal segment of the exopod are flat- 
tened and curved and the ultimate is fully two and a half times as long as the next of 
the series. 


! This tabular method has been adopted for showing the position of the spines on the abdominal 
somites. Numbers in brackets indicate that spines occur on these somites in some specimens, but not 
in all. (See p. 9.) 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 2 


The colour of spirit specimens is not very striking. The posterior margins of the 
carapace and of the last three thoracic and first five abdominal somites are narrowly 
bordered with black pigment and frequently a small black spot is visible on the dorsal 
surface of the eyestalk, touching the cornea. 


The eight specimens in the Indian Museum are registered as follows :— 


au Sandheads, Ganges Delta. A. J. Milner. Io’, 61 mm. 
en Sandheads, Ganges Delta. J. Barnet. 19, 49 mm. 
> Sandheads, Ganges Delta. W. M. Daly. Id, 7I mm. 
22" Orissa Coast, Bengal, 13 fms. ‘ Investigator.’ Id, 46 mm. 
> Vizagapatam Coast, Madras. ‘Investigator.’ 12,64 mm. 
se Madras. (purchased). 192,62 mm. 


Bes Persian Gulf, 25 fms. 
28°59 N., 50°5’E. ‘Investigator.’ IEG 5 1 Oy By 7/6} tawbutle 


In addition to the above records Squilla latreillei is known only from Singapore 


(Eydoux and Souleyet). 


2, Squilla decorata (Wood-Mason). 
Plate I, figs. 13—16. 


1875. Clorida decorata, Wood-Mason, Proc. As. Soc. Bengal, p. 231, reprinted in Ann. Mag. 
Nat. Hist. (4), XVII, p. 263 (1876). 

1880. Chloridella decorata, Miers, Ann. Mag. Nat. Hist. (5), V, p. 15. 

1895. Chloridella decorata, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 9, pl. iv, 


figs. 14-17. 

This species may be readily distinguished from the preceding by the following 
characters :— 

I. The lateral process of the fifth thoracic somite consists of a long and sharp spine 

which is strongly bent forwards (fig. I4). 
2. The inferior margin of the same somite bears on either side a small acute tooth. 
3. The lateral margin of the seventh somite is rounded behind, but in front is 
produced as a sharp spine (fig. 14). 

4. Ihe inferior surface of the telson bears a number of short carinae, which are 
more or less broken up into tubercles, on either side of the well-marked 
post-anal carina (fig. 16). 

The rostrum is short, much wider than long, and its upturned lateral margins 
converge to a broadly-rounded apex. ‘The eyes (fig. 13) are similar to those of S. lat- 
rellei, but the stalk is rather less swollen ; the inner margins are flattened and in 
juxtaposition for about three-quarters of their basallength. There is a three-segmented 
mandibular palp. ‘The raptorial claw also resembles that of the preceding species ; in 
all the specimens examined the dactylus bears five teeth and the proximal tooth is well 
separated from the next of the series. 

The lateral process on the anterior part of the eighth thoracic somite is sharply 


28 Memoirs of the Indian Museum. vor 100, 


acute and directed forwards, more so than in any example of S. Zatreillei (cf. figs. 4 and 
I4). Submedian carinae are present on all the abdominal somites and are also distinct 
on the last three segments of the thorax. The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian a Fe ah 6. 
Intermediate ge Do oe (SIMS AC: 
Lateral 3% 50 Sie (3) 4,5, 6 
Marginal do 50 00 (2) 85 aly 5 


The submedian carinae of the first five somites are strongly divergent posteriorly 
(fig. 15). On the sixth somite between the submedian and intermediate carinae are 
numerous well-marked vermiculate elevations. 

The upper surface of the telson (fig. 15) is closely covered with tubercles, which 
in most cases form definite rows. The telson of the type specimen has already been 
figured by Wood-Mason (1895, pl. iv, fig. 16), that of another example is illustrated 
on Pl. I, figs. 15, 16. The marginal teeth are well developed and without serrations 
on their edges. There are two submedian denticles, seven intermediate and one lateral. 
The inner margin of the bifurcate process from the base of the uropod bears from seven 
to thirteen sharp spines (thirteen in the type). The lobe on the outer margin of the 
longer spine is large and distinct. 


The specimens show no characteristic colouring, but, in one example, the black 
spot on the eyestalk already noticed in the preceding species is very distinct. Accord- 
ing to Wood-Mason the colour of one of the specimens in life was ‘ delicate salmon- 
pink, darkest in the mid-dorsal line.’ 

There are four specimens of this species in the Indian Museum :— 


3086 


=; Port Blair, Andamans. J. Wood-Mason. 19, badly mutilated, ca. 75 mm. TYPE. 
m Ye River entrance, Burma. ‘Investigator.’ 19, 49 mm. 
> Off Irrawaddy Delta. 
15°29’ N., 94°55’ E. ‘Investigator.’ 19,70 mm. 
ms Off Akyab, Arakan Coast, 
34 fms., 19°56’ N., 92°32’ E. ‘Investigator.’ RO 29 Vi 


In the last and smallest of the above specimens the spinous process at the antero- 
lateral angles of the seventh thoracic somite is less well developed than in the others, 
though still clearly distinct from S. Jatreiller in this respect. The telson shows only 
faint traces of rows of tubercles on its dorsal aspect, and on its ventral surface none 
whatever can be distinguished. In other details this young example agrees well with 
the larger individuals. 

Only the above specimens are known. 


3. Squilla gibba, Nobili. 
Plate I, figs. 5—12. 
1903. Squalla (Chloridella) gibba, Nobili, Boll. Mus. Torino, XVIII, no. 447, p. 30; p. 31, fig. 3. 
This species is allied to the two preceding, but may be distinguished from them 
by the following characteristics :— 


oz. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 29 


1. The median carina of the carapace is well marked and there are in addition 
traces of intermediate and lateral carinae. There is a small subsidiary spinule 
below the stout spine which defines the antero-lateral angles (fig. 7). 

2. The rostrum bears a distinct longitudinal carina in its anterior half. 

3. The eyestalks reach beyond the middle of the basal segment of the antennular 
peduncle. 

4. The two basal segments of the endopodite of the antenna are short and do not 

reach to half the length of the antennal scale. 

. The mandibular palp is entirely absent. 

6. The lateral margin of the fifth thoracic somite is produced as a broad falcate 
process, which is very strongly bent forwards (much more so than in S. 
decorata) and closely resembles that of S. scorpio ; the lateral margins of the 
two succeeding somites are more narrowly rounded than in either of the 
preceding species (fig. 5). 

7. The dorsal surfaces of the free thoracic and first five abdominal somites are 
finely rugose. 

8. The dorsal surface of the telson, in the female (fig. 12), is finely rugose on either 
side of the median carina, but is without any trace of tubercular ridges. In 
the adult male (fig. 11) the surface is smooth, but a huge curved and rounded 
ridge, higher than the median carina, extends from the vicinity of the antero- 
external angle to the apex of the submedian teeth. In both sexes the post- 
anal crest is represented only by a single prominent tubercle. 

g. The inner margin of the bifurcate process from the base of the uropod is finely 
serrate, not spinous. 


UL 


The rostrum in this species is rather longer than in S. latreillei and decorata ; it is 
only a little broader than long and its lateral margins are upturned. The cornea of 
the eyes is also rather larger than in the two preceding species and the inner margins 
of the stalk, though considerably flattened, do not lie close against one another.‘ The 
antennular peduncle is shorter than the carapace, excluding the rostrum. 

The dorsal crest of the carpus of the raptorial claw (figs. 8, 9) terminates in a 
subacute point and is sometimes obscurely notched near the middle. The propodus 
of the adult male (fig. 8) is very broad distally, though normally developed in the 
female (fig. 9). The dactylus is, in the male, externally convex and slightly swollen 
at the base ; in the female it is less swollen and the outer margin is very distinctly 
sinuous. There are five dactylar teeth (including the apical one) and these are much 
longer and more slender in the female than in the male. As in S. decorata all the teeth 
are well separated from one another. 

In addition to the falcate lateral process, the fifth thoracic somite bears a blunt 
forwardly-directed tooth on either side of its inferior surface (fig. 10). The lateral 


! It is the inner margin of the stalk that is dilated in this species and not the outer, as in S. 
latreillei, S. decorata and S. microphthalma. The figures are all of right eyes; compare fig. 6 with figs. 3, 
13 and 17. 


30 Memoirs of the Indian Museum. More 


margins of the two succeeding somites are more narrowly rounded than in S. Zatrerller 
and the anterior spinous process found on the seventh somite of S. decorata is absent 
(fig. 5). The eighth somite is produced laterally as a rounded anterior lobe. 

Faint submedian and intermediate carinae are present on the last three thoracic 
and on all the abdominal somites. The submedians are parallel and not posteriorly 
divergent as in the preceding species. On the 3rd, 4th and 5th somites there are ex- 
tremely faint traces of median tubercles. The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian SF hee a ae 6. 
Intermediate ER a oe 5. ©: 
Watenalguee ba Sa ms 510: 
Marginal.. ar BE So BSAA BA Os 


The lateral carinae are semi-obsolete except on the Ist somite. The submedian 
and intermediate carinae of the 6th somite are greatly swollen in the adult male. 

The form of the telson is astonishingly different in the two specimens examined. 
In the smaller femaie example (fig. 12), the median carina is obscurely notched near 
the base and terminates in a short spine. ‘The dorsal surface on either side is very 
finely rugose, but is otherwise smooth except for the lateral edge which is upturned 
proximally and for obscure elevations at the base of the submedian and intermediate 
teeth. The usuai three pairs of marginal teeth are well developed and those of the 
submedian pair appear to have had movable tips. There are two or three submedian 
denticles, six intermediate and one lateral. ‘The praelateral denticle is missing. In the 
adult male, the type (fig. 11), the base of the submedian spines is greatly inflated, 
while from the intermediates two huge, swollen and internally curved ridges, higher 
than the median carina, extend backwards nearly as far as the antero-lateral corners. 
The submedian and intermediate spines are very blunt and the lateral spine and denticle 
are represented by two small nicks in the margin. There are three or four submedian 
and six or seven intermediate denticles. The median carina is obscurely notched at the 
base, the distal spine is non-existent and on either side, in the hollows between the 
three prominences, there are obscure indications of a longitudinal row of pits. The 
lateral margin is strongly upturned proximally. 

The bifurcate process from the base of the uropod resembles that of the preceding 


species, except that the spines on the inner margin are replaced by a row of fine 
serrations. 


The colouring of spirit specimens does not seem distinctive. 


There is one specimen in the Indian Museum :— 


= Buntal or Pulo Burong, Borneo. Sarawak Museum. 12, 48 mm. 


This individual was presented by Mr. J. C. Moulton, to whom I am also indebted 
for the opportunity of examining and figuring Nobili’s type specimen, the only other 


example known. ‘The latter, a male 76 mm. in length, was found at Pulo Burong, 
Borneo. 


I913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 31 


4. Squilla microphthalma, H. Milne-Edwards. 
Plate I, figs. 17—20. 


1837. Squilla microphthalma, H. Milne-Edwards, Hist. Nat. Crust., II, p. 523. 
1841. Clorida microphthalma, Eydoux and Souleyet, Voy. de la “Bonite,’ Zool. I, Crust., 
pp. 264, 266. 
1849. Squilla microphihalma, de Haan. in Siebold’s Fauna Japonica, Crust., p. 221. 
? 1880. Chloridella microphthalma (depressa), Miers. Ann. Mag. Nat. Hist. (5), V, p. 14, pl. ii, 
figs. I—4. 
1882. Chloridella microphthalma, Haswell, Cat. Australian Crust., p. 207. 
1894. Squilla microfhthalma, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 509. 
1895. Chloridella microphthalma, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 8, pl. iv, 
figs. I—5. 
1898. Chloridella microphlhalma, de Man, Zool. Jahrb. Syst., X, p. 691, pl. xxxviii, fig. 76, a. 
1904. Squilla microphthalma, Jurich, Stomatop. deutsch. Tiefsee-Exped., p. 368, pl. xxvi, fig. I. 


This species and the two following may easily be distinguished from the three 
preceding by the complete absence of submedian carinae on the first five abdominal 
somites and by the proportions of the eyes. The latter are decidedly longer than in 
the species of the /atreillei group and the cornea, through narrower than the inflated 
basal portion of the stalk, is much broader (fig. 17). 

The following appear to be the principal characters of S. microphthalma :—The 
dorsal surface of the carapace and abdomen is quite smooth. The carapace, which 
does not possess median or lateral carinae, bears an extremely close resemblance to 
that of S. Zatreillei, but is rather narrower in front. The antero-lateral angles are 
produced as sharp spines. The rostrum is smooth above, as long as, or a trifle longer 
than wide and its lateral margins which are not upturned meet in an acute but rounded 
apex: 

The eyes (fig. 17) are narrowly pear-shaped and reach to at least two-thirds the 
length of the basal segment of the antennular peduncle. The cornea is two-thirds the 
maximum width of the stalk and its greatest breadth is about one-third the length 
of the whole organ. The internal margins of the stalk are not distinctly flattened and 
in juxtaposition as in S. Jatreille:. The antennular peduncle is about as long as the 
carapace, excluding the rostrum. The two basal segments of the antennal endopodite 
are long, reaching almost to the apex of the scale. The mandibular palp is present and 
is composed of three segments. 

The raptorial claw is almost precisely similar to that of S. /atreillei, but the ex- 
ternal margin of the dactylus is very feebly sinuous. The latter segment bears four 
teeth including the apical one, and in some specimens a very small additional tooth is 
present, proximal in position and lying close up against the next of the series. 

The last three thoracic somites bear faint intermediate, but no submedian, carinae. 
The fifth does not possess a pair of inferior spines but is produced laterally to a very 
short acute point, which is directed almost straight outwards (fig.18). The lateral 
margins of the two following somites are broadly rounded; that of the eighth is 
produced anteriorly as a rounded lobe, 


D 


Memoirs of the Indian Museum. Morte 


Oo 


The abdominal somites bear faint intermediate, lateral and marginal carinae, but 
the submedian are entirely absent except on the last. The following carinae end in 
spines :— 


Carinae. Abdominal somites. 
Submedian Le ma = 6. 
Intermediate = as 5, 6. 

Lateral .. Sa os ae 5, 6. 
Marginal.. Se a Be (3) (4) 5. 


The dorsal ornamentation of the telson is variable and, both in general features 
and in its sexual differences, presents a close resemblance to that of S. /atreillei. The 
figures on Pl. I, figs. 19, 20, will convey a better idea of the characters than a long 
description. The submedian teeth have minute movable tips and there are two 
submedian denticles, six or seven intermediate and one lateral. The under surface of 
the telson is smooth except for an obsolete post-anal carina. 

The process from the base of the uropod bears a series of six or seven sharp spines 
on its inner margin. Of the two primary spines the inner is nearly twice the length of 
the outer and bears a rounded external lobe near its middle point. 


The smallest specimen in the Indian Museum, a female 33 mm. in length, differs 
from the foregoing description in having the rostrum distinctly broader than long, in 
having three submedian and five intermediate denticles on the telson, in having the 
outer margins of the intermediate teeth serrate, and in having the oblique rows of 
tubercles on either side of the telson more conspicuous. In other characters, however, 
it agrees so well with typical examples that there can be little doubt that it belongs 
to the same species. 

The specimen from Australia which Miers doubtfully referred to S. microphthalma 
in some respects resembles this small individual ; but the rostrum, according to Miers’ 
figure, is still broader and the inner margin of the ventral process of the uropods is 
stated to be armed only with a series of small spinules, as in C. decorata. On the 
acquisition of further specimens from the Australian coast it will perhaps appear that 
this form is distinct, and in this case the name depressa, suggested by Miers, is to be 
employed. 


The colour of spirit specimens is not very striking. The lateral and posterior 
margins of the carapace and the posterior margins of the last three thoracic and five 
abdominal somites are rather broadly defined by black pigment. 


There are four specimens of Squilla microphthalma in the Indian Museum :— 
3313 


7 Karachi. Karachi Museum. Io, 52mm. 
= Karachı. al me OA TN 
= Bombay. Bombay Nat. Hist. Soc. 19,50 mm. 
= Madras Coast. (no history). 19,33 mm. 


In addition to the above records S. microphthalma is known from ‘les cötes 
.de l’Inde” (Milne-Edwards) and from Zanzibar, 33 fathoms (Jurich). The speci- 


1013.] S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 33 


men described by Miers! and referred to above was found at Port Essington, N. Aus- 
tralia. 


5. Squilla chlorida, Brooks. 


1886. Squilla chlorida, Brooks, Voy. H.M.S. ‘Challenger,’ XVI, Stomatop., p. 40, pl. ii, 
figs. I—5 


1894. Squilla chlorida, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 510. 


This species is only known from the type specimen. According to Brooks’ des- 

cription and figures it differs from S. microphthalma only in the following particulars :— 

I. The carapace is rather broader anteriorly and possesses faint lines indicating 
lateral carinae. 

2. The cornea of the eyes is considerably more expanded than in C. microph- 
thalma ; its greatest breadth is about one half the total length of the organ, 
but is, however, decidedly less than the maximum width of the stalk. 

3. The lateral process of the fifth thoracic somite is a long, straight, outstanding 
spine. 

4. The postero-lateral angles of the first four abdominal somites are obtuse. 


The dactylus of the raptorial claw possesses five well-separated teeth. The telson 
is smooth below and bears numerous scattered rounded tubercles on either side of the 
dorsal carina. ‘There are three or four submedian denticles, six or seven intermediate 
and one lateral. 

One specimen, a male 40 mm. in length, was found by the ‘ Challenger ’ Expedi- 
tion at Amboina, China Sea, in 15 fathoms. 


The specimen which Lanchester has recorded under the name of Chloridella chlorida, 
Brooks, from Kelantan, Malay Peninsula (Proc. Zool. Soc., 1001, p. 554) does not 
appear to belong to this species. I,anchester notes that submedian carinae occur on 
all the abdominal somites. This would seem to indicate that the specimen should be 
classed with the /atreille: group, but it is not clear to which, if any, of the three species 
it should be referred. 


6. Squilla rotundicauda (Miers). 


1880. Chloridella rotundicauda, Miers, Ann. Mag. Nat. Hist. (5), V, p. 15, pl. ii, figs. 5, 6. 
1894. Squilla rotundicauda, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 509. 


Miers’ description of the unique specimen runs thus :— 

““ This species is nearly allied to C. microphthalma, but differs as follows :—The 
rostrum is somewhat more elongated; there is no spine at the antero-lateral angles 
of the carapace. ‘The carinae of the first to fifth postabdominal segments are very 
faintly indicated ; those of the sixth segment are strongly defined. The median carina 


be erroneous. 


. 34 Memoirs of the Indian Museum. Worse 


of the terminal segment is thickened and obtuse posteriorly ; the lateral marginal 
spines are obsolete, and the intervening denticles are small and not acute. The eye- 
peduncles have their inner margins straight and are convex only on their outer surface.’ 

S. rotundicauda may be distinguished from all the species of the microphthalma 
and /atreillei group by the absence of the spines at the antero-lateral angles of the cara- 
pace. The posterior thickening of the median carina of the telson (in the female) 
also appears to be a noteworthy feature. 

It should be noticed that the rostrum of Miers’ specimen of S. microphthalma 
seems to have been unusually short. 

The type specimen, a female 70 mm. in length, was found at Formosa. 


7. Squilla fasciata, de Haan. 
Plate I, figs. 21—23. 
1844 ? Squilla fasciata, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. 4. 
1840. Squilla fasciata, De Haan, 1bid., text, p. 224. 
1880. Squilla fasciata, Miers, Ann. Mag. Nat. Hist. (5), V, p. 20. 
1886. Squilla fasctata, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 37, pl. ii, fig. 8; 
pl. iii, figs. 4,5. 
1893. Squilla fasciata, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 473. 
1894. Squilla fasciata, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 510. 
1903. Chloridella fasciata, Rathbun, Proc. U. S. Nat. Mus., XXVI, p. 54. 
1907. Chloridella fasciata, de Man, Trans. Linn. Soc. Zool. (2) IX, p. 440. 


The dorsal surface of the carapace and abdomen is smooth and highly polished. 
The carapace, measured behind the antero-lateral angles, is half, or rather more than 
half, its length, excluding the rostrum! (fig. 21). The lateral margins are not angulated 
in front of the postero-lateral corners. Between the gastric grooves and the margins, 
in the anterior half of the carapace, there is on each side a shallow groove ; the median 
and intermediate carinae are entirely absent” and the lateral and marginal are only 
visible in the posterior quarter of the carapace. The antero-lateral angles are pro- 
duced as strong spines that reach almost or quite as far as the level of the rostral base. 
The rostrum is smooth dorsally, scarcely broader than long, and its lateral margins, 
which are not upturned, converge to a rather sharply rounded apex. 

The cornea of the eyes is not large but is very clearly wider than any part of the 
stalk ; the greatest breadth (across the cornea) is equal to about three-quarters of the 
total length. The corneal and peduncular axes are very slightly oblique. The anten- 
nular peduncle is about as long as the carapace, excluding the rostrum. The two 
basal segments of the antennal endopodite are short and do not reach much beyond 
the middle of the scale. The mandibular palp is present and three-segmented. 

The outer inferior margin of the merus of the raptorial claw terminates in a 
rounded angle ; the carpus has a groove and keel on its external aspect and its dorsal 


! Brooks’ figure is, I believe, incorrect in showing the carapace so very much narrowed anteriorly. 
* In fig. 21 the representation of a groove on either side of the gastric grooves erroneously bears 
some resemblance to an intermediate carina. 


1013.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 35 


carina is entire and ends in a sharp spine before reaching the distal margin. The 
propodus has the usual three movable spines and row of pectinations along the edge 
opposed to the dactylus. The dactylus bears six teeth, including the apical one, and 
the outer margin, which is obscurely angled posteriorly, is not, or is only very feebly, 
sinuous. 

There are no submedian carinae on the last four thoracic somites (fig. 21). The 
intermediates are present and on the fifth somite each is oblique and is continuous 
with the posterior margin of the single sharp spine which forms the lateral process. 
This spine points forwards and a little downwards and beneath it on the inferior 
surface of the somite there is on each side another sharp spine, which is directed 
straight downwards. ‘The lateral margins of the sixth and seventh somites are more 
or less truncate and rather sharply rounded posteriorly. 

The first five abdominal somites are remarkable for the entire absence of the sub- 
median carinae. These, however, are conspicuous on the sixth somite and the inter- 
mediates, laterals and marginals are well developed on all. The following carinae end 
in spines :— 


Carinae. Abdominal somites. 
Submedian a à À 6. 
Intermediate 2 ss ER (3) 4, 5, 6. 
TKateraleer ve a ae 394, 560: 
Marginal 7022 3,4159. 


The telson (fig. 23) ! is as broad as, or a trifle broader than long. The median carina 
is well marked ; it is notched near the base and ends in a sharp spine which over- 
hangs two small tubercles. On either side there is another longitudinal carina about 
three quarters the length of the median and external to these there are several other 
shorter carinae, six of which reach to the apices of the sharp primary marginal teeth. 
There are three or four submedian denticles, six to eight intermediate and one lateral. 
The praelateral denticle is present. On the ventral side of the telson the post-anal 
carina is distinct and on either side of it there are, in larger specimens, two other 
carinae which are more or less interrupted. 

The basal segment of the uropod terminates in a sharp dorsal spine which projects 
over the articulation of the exopodite. The bifurcate process bears from nine to twelve 
spines on its inner margin. The inner primary tooth is almost twice the length of the 
outer and bears a prominent external lobe at about its middle point. The first 
segment of the exopodite is considerably longer than the second. 

The spirit specimens are closely covered with brownish and black chromatophores, 
which, by their absence, define an interrupted and irregular pale transverse band in 
the posterior third of the carapace. The hinder edges of the abdominal somites may 
be suffused with black between the lateral and intermediate carinae. The exopodite of 
the uropod is quite black except for the anterior half of the proximal segment and the 
apices of the endopodite are similarly coloured. 


! In the Andaman specimen, from which this figure was drawn, the telson is damaged and abnormal 
on the left side. 


36 Memoirs of the Indian Museum. Orne 


According to a colour-sketch kindly made for me from the smallest specimen by 
Capt. R. B. Seymour Sewell, the median part of the carapace is, in life, dark brown 
throughout its length. This patch of colour encloses a large, pale, oval spot in the 
anterior half and further back is very much narrowed. The carinae of the abdomen 
are brown, and this colour also defines the position of the non-existent median and 
submedian carinae. The eyes are golden yellow. The marginal spines of the telson 
and the distal portions of both inner and outer uropods are deeply suffused with 
reddish purple. 


There are three specimens of S. /asciata in the Indian Museum :— 


pe Port Blair, Andamans. (Purchased). 19,68 mm. 
St Of Tavoy Point, 50 fms., 13°29’ N., 97°30’ E. ‘ Investigator.’ 19,40 mm. 
Se Port Maria, Elphinstone I., Mergui Archipelago ‘ Investigator.’ 19,50 mm. 


Through the kindness of Prof. K. Kishinouye I have been able to compare these 
specimens with an example from Japan : the specimens are in close agreement. Prof. 
Kishinouye informs me that the species is very rare in Japanese waters. His example, 
a male, 79 mm. in length, is encrusted with a Polyzoon, Triticella korenii, Sars (see 
p. 7) and was found in the Bay of Tokio. 

Previous records of Sguilla fasciata are not numerous. It is known from Japan, 
Inland sea and other localities (De Haan, Brooks, de Man, Rathbun) and in Chinese 
waters from Chusan, 10—14 fathoms, and Holothuria Bank, 34—36 fathoms (Po- 
cock). 


8. Squilla miles, Hess. 
1865. Squilla miles, Hess, Arch. f. Naturgesch., XXXTI, i, p. 169, pl. vii, fig. 21. 
1880. Squilla miles, Miers, Ann. Mag. Nat. Hist. (5), V, p. 17. 
1882. Squilla miles, Haswell, Cat. Australian Crust., p. 207. 
1887. Squilla miles, de Mau, Zool. Jahrb. Syst., II, p. 714. 
1894. Squilla miles, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 509. 


I have seen no examples of this fine species. It appears to be allied to the pre- 
ceding, but is readily distinguished by the following characters :— 


I. The antero-lateral angles of the carapace are not spinous. 

2. The rostrum is somewhat longer than broad. 

3. ‘The cornea of the eye is not so distinctly wider than the stalk. It is set 
obliquely ; but the obliquity is the reverse of that usually found, the inner 
margin of the stalk being shorter than the outer. 

4. The dactylus of the raptorial claw bears only four teeth, including the terminal 
one. 

5. 'The lateral margin of the fifth thoracic somite is bluntly rectangular, not 
spinous and antrorse as in the preceding species. 


1 For this information I am indebted to Dr. W. T. Calman, who, at my request, made an 
examination of the specimen in the British Museum. 


1013.| S. KEMP : Crustacea Stomatopoda of the Indo-Pacific Region. 37 


6. Submedian carinae are present on the last three thoracic and on all the abdo- 
minal somites', and on the sides of each between the lateral and intermediate 
carinae there is a flattened triangulate prominence. 

7. The telson bears a great resemblance to that of S. fasciata, but there are two 
or three long continuous carinae on either side of the median crest in place of 
the one found in that species. According to Hess’ figure the first and third 
of these carinae are connected proximally, while the second and third fuse 
posteriorly and are continued as a singie keel to the apex of the submedian 
tooth. On the surface of the telson outside these carinae there is an oblique 
row of short ridges or tubercles. The submedian teeth have movable tips and 
are placed close together, the margin occupied by the intermediate denticles 
being thus much longer than in S. fasciata. ‘There are three or four minute 
submedian denticles, nine or ten intermediate and one lateral. The praelateral 
denticle appears to be absent. 


The type of Squilla miles, which measures 172 mm., was found at Sydney (Hess) 
and the only other known example, a male about 90 mm. in length, is recorded from 
Victoria (Miers). 

9. Squilla lata, Brooks. 
Plate Be 11229. 
1886. Squilla lata, Brooks, Voy. H.M.S. ‘ Challenger’, XVI, Stomatop., p. 34, pl. iii, figs. 1—3. 
1804. Squilla lata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 510. 

This species is allied to S. fasciata, but is easily distinguished by the absence of 
carinae on either side of the median crest of the telson. 

The whole dorsal surface is smooth and polished. On the carapace the median 
carina is wholly absent, though the small mid-dorsal pit is visible ; fine intermediate 
carinae are found on either side of the gastric groove, but the laterals and marginals 
only exist in the extreme posterior part of the carapace behind the cervical groove. 
The lateral margin is very obtusely angled in front of the rounded postero-lateral 
corners and the antero-lateral angles are produced to a sharp spine that does not reach 
the level of the rostral base. The anterior margins on either side of the rostrum aze 
distinctly sinuous. 

The rostrum is about one and a half times as long as broad, its lateral margins 
are sinuous—concave anteriorly—and converge to a narrow rounded apex. In its 
anterior half it may bear a very feeble longitudinal ridge, but it is never definitely 
carinate. 

The eyes are elongated. The cornea is distinctly wider than the stalk ; its breadth 
is about two-thirds the total length of the whole organ ; the corneal and peduncular 
axes are a trifle oblique. The antennular peduncle is shorter than the median length 
of the carapace. The mandibular palp is present and is three-segmented. 

The dorsal carina of the raptorial carpus terminates abruptly before reaching 


! It seems that Hess, when he mentioned three longitudinal carinae on the abdomen, did not reckon 
the marginal. 


38 Memoirs of the Indian Museum. Doig. 20%, 


the anterior margin. The dactylus bears six teeth including the terminal one : the 
outer margin is convex and bears an acute lobe at the base. 

The lateral process of the fifth thoracic somite is, in the Indian specimens, rather 
longer than is indicated in Brooks’ figure ; it is almost straight and trends obliquely 
outwards and forwards. The pair of inferior spines are sharp. The lateral margins 
of the two succeeding somites are broadly rounded, those of the eighth somite are 
lobed anteriorly. ‘The intermediate carinae of the free thoracic somites are distinct, 
but the submedians are either wholly absent or are represented only by obsolete traces 
on the last two somites. 

There are no submedian carinae on the first three abdominal segments, obsolescent 
indications of them may, however, be found on the fourth and fifth somites ; on the 
sixth they are invariably present and well developed. The intermediate, lateral and 
marginal carinae are, as is usual, quite distinct. The following abdominal carinae end 
in spines :— 


Carinae Abdominal somites. 
Submedian À I ER 6. 
Intermediate 2th se a (3) 2; Bin On 
Lateral + soe + 2 A, By Ob 
Marginal at ee 5 EA) Bo 4, So 


These figures refer to the Indian specimens ; Brooks remarks that in the type 
spines are found only on the last somite. 

The dorsal surface of the telson is finely rugose and, on either side of the median 
crest, is obscurely marked with faint parallel furrows. The marginal teeth are well 
developed and there are one to three! submedian denticles, six to eight intermediate 
and one lateral. In the Indian examples the outermost median denticle is borne 
on the inner face of the intermediate tooth and projects distally beyond its fellows. 
There is no praelateral denticle. The post-anal carina is short but distinct. 

The bifureate process of the peduncular segment of the uropods bears six to eleven 
sharp teeth on its inner margin and on the outer face of the longer spine there is a large 
and very conspicuous rounded lobe (fig. 24). 

The two largest specimens examined are males ; but, except for a feeble swelling 
at the base of each intermediate marginal denticle of the telson, no distinctive sexual 
modifications are to be found. 

The Indian examples of S. lata differ from Brooks’ account in the possession of a 
dorsal spine at the distal end of the peduncle of the uropods and in a few other details 
noticed above, the most important being the greater abundance of spines on the 
abdominal somites. Should the distinctions prove constant it may be necessary to 
separate specimens from the Bay of Bengal under a distinct subspecific name. 


Spirit specimens exhibit no characteristic colouring. 
The three examples in the collection are registered as follows :— 


15079 
10 


Gulf of Martaban, Burma; 53 fms., 
14°38’12” N., 96°24’30” E. ‘Investigator.’ IG, 29 ,64—76 mm. 


! Two or three in the Indian examples, a single broad lobe in the types. 


Tore] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 39 


The only other known specimens of S. data are those on which Brooks founded 
his original description. They were found in the Arafura Sea, south of New Guinea, 
in 49 fathoms, and the largest measured 82 mm. in length. 


10, Squilla gilesi, Wood-Mason, MS. 
Plate II, figs. 25—27. 


1908. Squilla gilest, Lloyd, Rec. Ind. Mus., II, p. 33 (sine desc.). 
1911. Squilla giles’, Kemp, Rec. Ind. Mus., VI, p. 95. 


Squilla gilesi is an extremely close ally of S. /ata and it is possible that it will 
eventually prove to be nothing more than a race or variety of that species. It is 
distinguished only by the following characters :— 


1. The intermediate carinae of the carapace are obsolete and are represented only 
by a pair of feeble and smoothly-rounded elevations. The lateral margins 
of the rostrum are in most cases less decidedly sinuous (fig. 26). 

2. Submedian carinae are quite distinct on the last two thoracic (fig. 26) and on 
all the abdominal somites. 

3. The inner margin of the bifurcate process of the uropods bears only a series of 
serrations, though one or two of the ultimate members of the series may be 
spinous (fig. 25). The lobe on the outer face of the longer tooth is small. 

4. Secondary sexual characters are strongly marked. In adult males the distal 
end of the raptorial propodus is proportionately much broader than in the 
female, the dactylus is slightly dilated at the base, the teeth are shorter and 
the outer margin more strongly convex. In this sex also the median carina 
of the telson is greatly inflated, especially at its anterior and posterior ends 
and the greater part of the distal margin is strongly swollen (fig. 27). 


The last character though only available in males, seems none the less to have 
considerable importance. Brooks has noted that the sexes in S. /ata are alike and in 
the examples in the Indian Museum scarcely any differences are perceptible even in 
large examples over 70 mm. in length. In S. giles’, on the contrary, the modifica- 
tions in the telson and raptorial claw are quite striking even in specimens only 55 mm. 
in length. 

In other respects S. gi/esi bears an exceedingly close resemblance to the Indian 
examples of S. /ata. ‘The spines on the abdominal somites are sometimes a little more 
numerous :— 


Carinae. Abdominal somites. 
Submedian .. ie arg 2 6. 
Intermediate Le ks ee (2) 35 2 55 Oe 
Lateral a dh 5% sa (08) (A). Bs Ay B5 Os 
Marginal Er ses Se er (Ze 


There are one to three submedian denticles, five to eight intermediate, and one 
lateral. 


40 Memoirs of the Indian Museum. Morenve 


. À very small specimen (°-), a male 19 mm. in length, taken in company with 
large typical examples, is in an early post-larval stage. The spines at the antero- 
lateral angles of the carapace are not developed, submedian carinae are absent except 
on the last somite, and on the margin of the telson there are numerous minute sub- 
median denticles and ten or eleven intermediate. A similar but slightly larger example 
(1), measuring 23°5 mm., is referred with some doubt to this species. The denticles 
on the margin of the telson are, as might be expected, fewer in number than in the 
small example, but in the bifurcate process of the uropods the lobe on the outer face 
of the long tooth is very much larger than in that specimen or in adults. In both 
these individuals the inner edge of the process is finely serrate. 


Spirit specimens of S. gilesi show no characteristic colouring. 


The following examples are in the Indian Museum :— 


7514-18 
—m Off Madras Coast: 80—110 fıns., 


TASTSETSENN 8018300 ‘ Investigator.’ 1,04 mm; 49, 72—82 mm. 
ANDRES: 
Madras Coast. ‘ Investigator.’ Ig,235 mm. 
= Off Orissa Coast; 19° 49° N., 
86° 43’ E. ‘ Investigator.’ a ©), ie iaahoor, 
m N.E. Bay of Bengal; 65 fms., 
200184 IN, CO” SOY 12, “ Investigator.’ 43, 19—59 mm. 


7519 


10 


Persian Gulf; 35 fms., 
HO AN HO Ilo, SYS GA” SO" 18, ‘ Investigator.’ 19,56 mm. 


[Squilla minor, Jurich.] 
1904. Squilla minor, Jurich, Stomatop. Deutsch. Tiefsee-Exped., p. 364, pl. xxv, figs. 4, 44. 


Allied, apparently, to the two preceding species is Jurich’s Sqwlla minor, des- 
cribed from two extremely small specimens (16°5 and Ig’5 mm. in length) from 50 
metres near Zanzibar. According to the description and figures these examples differ 
from Brooks’ adult S. /ata in the following characteristics :— 


I. There are no intermediate carinae on the carapace and no submedian carinae 
on any segment of the post-abdomen except the last. 

2. The rostrum is much wider, a little broader than long. 

3. The eyes are extremely large, their length being about one-third the length of 
the carapace and rostrum. 

4. There is a prominent outstanding lobe at the proximal end of the external 

margin of the raptorial dactylus. 
. The spine on the ventral surface of the fifth thoracic somite is unusually well- 
developed ; it extends outwards and is visible in dorsal view. 

6. There are three or four submedian denticles on the telson and on either side of 
the median crest there is an oblique row of six short elevations which appear 
to correspond with the interspaces between the impressed lines in S, /ata. 


O1 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. AI 


7. The lobe on the outer margin of the longer spine of the bifurcate process of 
the uropods is much larger. 
Some at least of these features are such as one would expect to find in young post- 
larval Squilla, but it is, I believe, impossible to be certain of the relations of Jurich’s 
species until adult specimens from the E. African Coast have been described. 


11, Squilla armata, H. Milne-Edwards. 
Plate II, figs. 28, 29. 


1837. Squilla armata, H. Milne-Edwards, Hist. Nat. Crust., II, p. 52T. 

1849. Squilla armata, Nicolet, in Gay’s Hist. fisica de Chile, Zool., III, p. 223. 
1849. Squilla armata, De Haan, in Siebold’s Fauna Japonica, Crust., p. 221. 5 
1879 Squilla armata, Kirk, Trans. N. Zealand Inst., XI, p. 401. 

1880. Squilla armata, Miers, Ann. Mag. Nat. Hist. (5), V, p. 26. 

1886. Squilla armata, Filhol, Miss. de l’ile Campbell, III, 2e, p. 435. 

1891. Squilla armata, Chilton, Trans. N. Zealand Inst., XXIII, p. 60. 

1801. Squilla armata, A. Milne-Edwards, Mission à Cap Horn, VI, p. 53, pl. vii. 
1894. Squilla armata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 515, figs. 9, Io. 
1900. Squilla armata, Whitelegge, Mem. Australian Mus., IV, p. 190. 

1902. Squilla armata, Stebbing, S. African Crust., II, p. 45. 

IgII. Squilla armata, Chilton, Trans. N. Zealand Inst., XLIII, p. 135, figs. I, 2. 
1911. Squilla armata, Chilton, Rec. Canterbury Mus., I, p. 306. 


The dorsal surface is smooth. The breadth of the carapace is greater than half 
its median length ; the median and intermediate carinae are entirely missing. The 
lateral carinae are distinct, but the marginals are only visible in the posterior quarter. 
The antero-lateral angles are produced to a short broad spine which reaches about to 
the level of the rostral base. The rostrum is about as long as wide ; it has no dorsal 
carina and its lateral margins converge to a broadly rounded apex (fig. 28). 

The dorsal processes of the ophthalmic somite differ noticeably from those of any 
other Indo-pacific species of the genus. They consist of a pair of sharp forwardly- 
directed spines widely separated from one another (fig. 28). The anterior edge of the 
ophthalmic somite terminates in two sharp points separated by a semicircular emargina- 
tion. The cornea of the eyes is greatly expanded ; its breadth is almost or quite equal 
to the length of the whole organ. The corneal and peduncular axes are oblique. 

The mandibular palp is entirely absent. 

The dorsal carina of the carpus of the raptorial claw terminates in a sharp tooth 
just behind the anterior margin (fig. 29). The dactylus bears seven, eight or nine 
(rarely six) teeth including the terminal one and its outer margin is convex with a 
small acute lobe at the base. | 

On the thoracic somites submedian and intermediate carinae are distinct. The 
lateral process of the fifth somite consists of a large tooth directed straightly outwards . 
the two spines on the inferior margin are distinct. The lateral margins of the two fol- 
lowing somites are rounded in front, but posteriorly each is produced to an acute point. 

On the abdominal somites the usual four pairs of carinae are well defined. The 
following terminate in spines :— 


42 Memoirs of the Indian Museum. Nor LOY , 


Carinae. Abdominal somites. 
Submedian er I es 6. 
Intermediate an i. UD (2) hes AS nor 
Lateral .. Br ne de Io By Sp Cp Sy Oo 
Marginal .. Re of Na T2 NS; ANS: 


In addition, on the posterior margin of the fifth abdominal somite between the 
submedian and intermediate carinae, there is a small group of one to four spines. 
These spines do not occur in any other species of Squilla. 

The dorsal surface of the telson is finely rugose, but is otherwise almost smooth 
on either side of the median crest. Faint traces of symmetrical rows of pits are occa- 
sionally visible. The six marginal teeth are well developed and the submedian pair 
possesses movable tips. On the inner face of the submedian spines there is a broad 
lobe which is sometimes separated from the spine by an incision and so becomes a 
submedian denticle. ‘The intermediate denticles vary in number from nine to twelve 
and there is one lateral. The praelateral denticle is absent. 

The inner margin of the ventral process of the peduncular segment of the uropods 
is finely serrate. The lobe on the outer edge of the longer spine is large in the two 
specimens examined ; but, according to Bigelow, is proportionately small in very large 
examples. 

Both the individuals in the Indian Museum are female. According to Bigelow 
there are no marked secondary sexual differences. 


Squilla armata seems to be an isolated form and does not appear to possess’ any 
close affinity with any other species of the genus. 


The two specimens examined are registered as follows :— 
7228"? New Zealand. Chas. Chilton. 29 , 64 and 80 mm. 


10 

The species is restricted to southern latitudes. It has been recorded from the 
Gulf of St. George on the Atlantic coast of Patagonia, from Cape Horn and from the 
Pacific coast of Patagonia and Chili (Milne-Edwards, Nicolet, Bigelow) and in these 
localities Bigelow notes that it is found between 51 and 122 fathoms. It is also known 
from New Zealand (Kirk, Chilton), from New South Wales (Whitelegge) and from 
S. Africa, off Cape Point Lighthouse, in 45 fathoms (Stebbing). 


12. Squilla scorpio, Latreille. 
Plate eo 


1825. Squilla scorpio, Latreille, Encycl. Method, X, p. 472. 

1837. Squilla scorpio, H. Milne-Edwards, Hist. Nat. Crust., II, p. 522. 

1847. Squilla scorpio, White, List Crust. Brit. Mus., p. 84. 

1849. Squilla scorpio, De Haan, in Siebold’s Fauna Japonica, Crust., p. 221. 

1880. Squilla scorpio, Miers, Ann. Mag. Nat. Hist. (5), V, p. 18, pl. ii, fig. 7. 
1892. Squilla scorpio, de Man, in Weber’s Zool. Ergebn. Nied. Ost-Ind., II, p. 518. 
1893. Squilla scorpio, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 453. 

1894. Squilla scorpio, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 510. 

1901. Squilla scorpio, Lanchester, Proc. Zool. Soc., II, Pp. 554. 


| S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 43 


The dorsal surface of the carapace is smooth and as a rule highly polished ; the 
abdomen is very obscurely rugose. The breadth of the carapace, measured behind 
the antero-lateral angles, is barely half its length, excluding the rostrum. ‘The median 
carina is distinct in front of the cervical groove (fig. 30), but is obsolete anteriorly. 
The small median dorsal pit is clearly visible and is placed nearer to the cervical groove 
than to the anterior margin. The intermediate carinae are well-marked ; the lateral 
also are well-developed anteriorly and posteriorly, but are usually obsolete in the 
middle. The antero-lateral angles terminate in stout spines which, owing to the strong 
convexity of the anterior margin, fail to reach the level of the rostral base. Beneath 
each antero-lateral spine there is a small blunt or subacute lobe. The postero-lateral 
angles of the carapace are rounded. | 

The rostrum is a trifle longer than broad ; its straight lateral margins are not up- 
turned and converge to a rounded apex. Dorsally the rostrum is smooth except for 
a sharp and distinct median longitudinal carina in its distal half. 

The eyes are small and elongated ; the breath of the cornea is about equal to the 
length of the stalk (cornea excluded), and the corneal and peduncular axes are decid- 
edly oblique. The antennular peduncle is short, scarcely two-thirds the length of the 
carapace. The two basal segments of the antennal endopodite are also short, the distal 
one not reaching to half the length of the scale. The mandibular palp is wholly 
absent. 

The outer inferior margin of the merus of the raptorial claw is bluntly rounded 
anteriorly. The carpus bears the usual groove on its external aspect and its dorsal 
carina is entire and terminates acutely before reaching the distal margin. The distal 
end of the propodus is considerably broader in the male than in the female. The 
dactylus bears five teeth including the apicalone. The outer margin is furnished with 
a small proximal lobe ; it is convex in the male but distinctly sinuous in the female. 

There are faint submedian and strong intermediate carinae on the last three 
thoracic somites. The fifth somite bears a pair of broad tooth-like lobes inferiorly ; the 
lateral lobe consists of a single strong falcate process terminating in a sharp forwardly- 
directed spine. An oblique carina, which commences on the dorsal aspect of the 
somite in the position usually occupied by the intermediate carina, extends down- 
wards and is continuous with the posterior margin of the process. ‘The antero-lateral 
margins of the fifth somite are elevated and form a distinct ridge. The lateral margins 
of the sixth and seventh somites are rounded; the eighth projects as a blunt 
anterior lobe. 

The first five abdominal somites possess four pairs of longitudinal carinae, but the 
submedians are not very strongly marked. The two median pairs of carinae on the 
sixth somite are distinctly swollen in the male. The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian ar oe a 6. 
Intermediate ar ER Ab BOs 
latéral" pi Pe bo 5,6. 


Marginal .. ats 50 (a) (4) P53 ,0- 


A4 Memoirs of the Indian Museum. Vor; IV, 


The telson has a well-marked median longitudinal carina which is usually ob- 
scurely notched at the base ; it terminates in a small spine overhanging a blunt tu- 
bercle. On either side of this carina the telson is smooth but, when examined with a lens, 
is seen to be very finely rugose, and occasionally faint traces of symmetrically arranged 
rows of pits are visible. On the margins, the submedian, intermediate and lateral 
teeth are well developed, as is also the small praelateral denticle ; each forms the 
termination of a short blunt carina. There are one or two submedian denticles, three 
to six intermediate and one lateral. 

In the telson of the adult male the median carina, especially in its posterior half, 
and the carinae which abut on the marginal spines, are considerably more swollen than 
in the female, but the interspinous marginal areas are never dilated as they are in 
S. gilest and some other species. 

The bifurcate process from the base of the uropod is finely serrate internally. The 
inner spine is much longer than the outer and beats a prominent external lobe a little 
beyond its middle point. 


Spirit specimens invariably possess a large and very distinct black patch on each 
lateral process of the fifth thoracic somite (fig. 30). There is also a black transverse 
patch on the second abdominal somite and a suffusion of the same colour at the distal 
end of the basal segment of the outer uropod. 

Freshly preserved examples are pale in colour with a thin sprinkling of black 
chromatophores. The median and submedian carinae of the carapace, abdomen and 
telson are bright red. The eyes are black with a yellowish stalk. Frequently there 
is a row of four or five black spots on either side of the median crest of the telson ; 
and the swollen bases of the marginal teeth are dusky. The colouring noticed in 
spirit specimens is, of course, very distinct in fresh examples. 


Most of the specimens of Squilla scorpio in the Indian Museum are referred to the 
var. immaculata. Only the following typical examples have been examined :— 


Se Madras. (Purchased). 18,19, 60 and 76 mm. 
7032-5 .' 
aa uvicoriny, Ss diary cr kun J. Hornell. 53,82, 41-57 mm. 

10 À 

= Bombay. Bombay Nat. Hist. Soc. 23, 86 and 93 mm. 

= Karachi? Karachi Museum. 19,97 mm. 


I have also examined one other specimen kindly lent me by Mr. J. C. Moulton— 


Tambak, Borneo. Sarawak Museum. 19,00 mm.! 


This species has been previously recorded from N. Australia (Miers) ; Celebes 
(de Man); the Malay Peninsula (Lanchester) ; Pondicherry (Latreille) and Madras 
(Henderson). 


1 In this specimen the right raptorial dactylus is malformed, doubtless owing to an injury. The 
teeth are twisted and distorted and are six in number. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 45 


var. immaculata, Wood-Mason, MS. 
Plate eee 


This form differs from typical S. scorpio in the entire absence of the black spot 
on the lateral processes of the fifth thoracic somite (fig. 31), and with this colour dis- 
tinction the following minor structural differences are correlated :— 


I. The rostrum is not longer than wide and its apex is rather broader than in 
_ typical specimens. 

2. The median carina of the carapace is sharp and distinct for a longer distance 
in front of the mid-dorsal pit, but terminates more or less abruptly before 
reaching the anterior margin. 

3. The lateral carinae of the carapace are usually distinct throughout their course. 

4. The lateral margins of the sixth and seventh thoracic somites are slightly more 
upturned than in typical specimens and the process on the anterior part of 
the eighth somite is a trifle more acute. 

The secondary sexual differences are as well marked as in the typical form. 

The specimens in the Indian Museum are registered as follows :— 


a“ E. of Terribles, Arakan Coast; 13 fms. ‘ Investigator.’ Id, 48 mm. 
= ‘ Bay of Bengal.’ V. Ball. Id,5o mm. 
ae ‘Calcutta.’ J. Wood-Mason and V. Ball. 83,72, 4I—95 mm. 
= Pratapnagar, Calcutta. B. L. Chaudhuri. id, 87 mm. 
TES Ghaga Nata, Mutlah R. near Calcutta 

: (brackish water). S. B. Nath. 148,82 , 69—86 mm. 
ze Saugor I., M. of Hushli R. J. Munro. 1d', 87 mm. 

in Hugli Estuary. Mus. Collr. 23,19, 51—64 mm. 
3%! Port Canning, Ganges Delta. R. D’Cruz. 19, 87 mm. 
zung Karachi. (no history.) 1,06 mm. 
en (Locality unknown). 83,82, 40—64 mm. 


The variety has never been taken in company with typical examples. The speci- 
men from Pratapnagar was found in brackish water and this was probably also the 
case with those labelled ‘ Calcutta.’ 


13. Squilla mantoidea, Bigelow. 


1893. Squilla mantoidea, Bigelow, John Hopkins Univ. Circ., no. 106, p. 101. 
1894. Squilla mantoidea, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 521, figs. 13, 14. 


Bigelow gives the following diagnosis of this species :—‘‘ Eyes triangular, but 
with the corneal axis at right angles to the peduncular one ; dactylus of raptorial claw 
with six teeth, outer margin not sinuate ; rostrum subquadrate, carinate ; carapace 
with five carinae, the median one bifurcated, and with strong anterior lateral spines ; 
lateral spine of the fifth thoracic segment short, straight, acute and flattened obliquely, 
lateral processes of the next two segments strongly produced and acute ; submedian 
carinae on thoracic and abdominal segments without spines, except the sixth abdo- 
minal ; telson with a crest and a long ventral keel, twelve or more lines of pits on each 


46 Memoirs of the Indian Museum. [Vor.. IV, 


side, six marginal spines ; denticles 5-6, II-I2, and r.’’ For further details Bigelow’s 
long and detailed description should be consulted. 

Squilla mantordea is probably, as Bigelow has suggested, a near ally of:the Medi- 
terranean S. mantis ; the mandibular palp, however, has not been examined. From 
all Indo-pacific species with a single lateral process on the fifth thoracic somite it is 
easily distinguished by (1) the sharp anteriorly-bifurcated carina of the carapace and 
(2) by the large eyes with the cornea set transversely on the stalk. 

S. mantordea is known only from a single specimen, a female 120 mm. in length, 
from Borneo (Bigelow). 


14. Squilla leptosquilla, Brooks. 
1886. Squilla leptosquilla, Brooks, Voy. H.M.S. ‘Challenger,’ XVI, Stomatop., p. 30, pl. 1, 
figs. I, 2. 
1894. Squtlla leptosquilla, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 510. 
1899. Squilla leptosquilla, Alcock and Anderson, Ann. Mag. Nat. Hist. (7), III, p. 292. 
1904. Squilla leptosquilla, Jurich, Stomatop. deutsch. Tiefsee-Exped., VII, p. 370, pl. xxv, 
figs. I, 2 (including var. dentata, p. 372). 


The dorsal surfaces of the carapace and abdomen are finely rugose, but when 
dried appear polished. The carapace measured behind the antero-lateral angles is 
considerably less than half its median length. The median carina is clear and distinct 
in front of the cervical groove, but is absent anteriorly for about half the distance in 
front of the small dorsal pit. The intermediate and lateral carinae are sharp and very 
conspicuous and the lateral margin is not angled posteriorly. Antero-laterally the 
carapace is produced as a short spine that reaches nearly as far as the level of the 
rostral base. 

The rostrum is a trifle longer than wide and reaches to the base of the ophthalmic 
somite. The lateral margins, which are not upturned, are rounded and converge to a 
subacute apex. In the middle of the dorsal surface there is a short but distinct median 
longitudinal carina. 

The eyes are rather small. The breadth of the cornea is about equal to the greatest 
length of the whole organ and the corneal and peduncular axes are very oblique. The 
antennular peduncle is as long as the rostrum and carapace combined. The two basal 
segments of the antennal endopodite are short and do not reach to half the length of 
the scale. 

There is no palp on the mandible. 

The outer inferior margin of the merus of the raptorial claw is broadly rounded. 
The carpus is grooved and ridged externally and the carina on its dorsal aspect is 
entire and terminates acutely close behind the distal margin. The outer end of the 
propodus is a little broader in the male than in the female and the pectinate margin is 
rather noticeably sinuous. The dactylus bears four slender teeth including the apical 
one and its outer margin is evenly convex in both sexes and projects as a small but 
prominent lobe near the articulation of the propodus. 

The last four thoracic somites possess well-marked submedian and intermediate 


1913.) S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 47 


carinae ; the latter are very sharp on the anterior portions of the sixth and seventh 
somites and, in the fifth, are frequently produced as a small acute outstanding pro- 
cess. The fifth somite bears a pair of teeth on its inferior margin and, on each side, 
extends laterally as a single long and straight lobe, which ends acutely, but does not 
trend forwards as in many of the allied species. The lateral margins of the sixth and 
seventh somites consist of a single lobe, which is obliquely truncate and sinuous ante- 
riorly and terminates posteriorly in a sharply acute point. 

The abdominal somites are broad and greatly depressed. The first five possess 
eight well-marked longitudinal carinae and on each of them, except the first, a small 
but conspicuous transversely-grooved median tubercle is present. The last somite has 
six carinae and the spines that terminate the submedian and lateral pairs are very 
long, especially the latter. The complete spine formula is :— 


Carinae. Abdominal somites. 
Submedian i ae ae 6. 
Intermediate bya sa Hele ah (1) WZ Rae Aeon O: 
Lateral .. a se Rn 234 50% 
Marginal 00 sa do A Sp Che, Fe 


The telson is very characteristic and shows little resemblance to that of any of 
the previously described species. Its peculiar appearance is due to the enormous 
development of the marginal and dorsal teeth, to the great elevation of the median 
carina and to the very deep incision between the submedian teeth. The form of the 
telson is moreover extremely broad and, if the marginal teeth are omitted, is only 
about half as long as wide. The dorsal carina is very high and very sharp, the basal 
notch is obsolete and the distal spine is strong and sharp, extending to fully half the 
length of the submedian teeth. The latter are quite two-thirds as long as the rest of 
the telson and are widely divergent posteriorly. On their inner margins they bear 
from ten to eighteen minute spinules, which are closely set proximally. ‘The inter- 
mediate teeth are even longer than the submedian and are somewhat inturned, espe- 
cially in the male : there are nine to twelve intermediate denticles, one or two of 
which are frequently situated on the outer margin of the submedian spines. In the 
female there is a small lateral denticle and the lateral tooth is long, reaching nearly to 
half the length of the intermediate. In the male the lateral tooth is short and greatly 
swollen at the base, completely obscuring the lateral denticle. There is also, in this 
sex, a well-marked swelling at the base of the intermediate teeth and this extends 
inwards along the interspinous margin. The mid-dorsal carina is not dilated in the 
male. The post-anal carina is short but distinct. 

The bifurcate prolongation from the base of the outer uropod is composed of two 
very long and slender spines. The inner, which does not greatly exceed the outer, is 
very nearly half the length of the entire segment and bears in the distal third of its 
external margin only a minute rudiment of the usual lateral lobe. The outer uropod 
is not more than two and a half times as long as wide in the adult male; in a female 
of the same length it is much narrower, about three times as long as wide. 

Two female specimens have the denticles on the telson arranged as in Jurich’s 


48 Memoirs of the Indian Museum. | Vor. IV, 


var. dentata ; another is on one side typical and on the other of the dentata form. The 
differences do not, in my opinion, deserve the distinction of a varietal name. 

In spirit specimens light chestnut brown colouration defines all the carinae of the 
carapace and the submedian and intermediate carinae of the abdomen. ‘There are also 
in some specimens rather conspicuous brown markings on each of the abdominal somites 
between the intermediate and lateral carinae. On either side of the median carina of 
the telson there is a large and conspicuous, oval, chestnut brown spot: the two are 
confluent posteriorly beneath the base of the distal spine. 

There are five specimens in the Indian Museum, registered as follows :— 


2314-5 


10 NE WAR NISSAN OS OAS“ 1B, x65 ae: | 


u Andamans. ( 14°13’ N., 93°40’ E. 370—419 fms. j 


‘ Investigator.” 23,29, 68—131 mm. 


ES. of Port Blair, Andamans. 188—220 fms., “Investigator.” 19, 89 mm. 
11°31/407 N., 92°46/40” E. 
_ S. leptosquilla has also been recorded from the Celebes sea near the Philippine Is., 
115 fathoms (Brooks), and from the vicinity of the Nicobars, 296 metres (Jurich). 


15 Squilla tenuispinis, Wood-Mason. 
Plate III, figs. 32—34. 


1891. Squilla tenuispinis, Wood-Mason, Ann. Mag. Nat. Hist. (6), VII, p. 271. 
1894. Squilla tenuispinis, Alcock, Ann. Mag. Nat. Hist. (6), XIII, p. 400. 


This species is very closely allied to S. /eptosquilla, but may be distinguished from 
it by the following characters :— 


‘1. The dorsal surfaces of the carapace and abdomen are smooth, polished and 

without trace of rugosity. 

2. The median carina of the carapace is obsolete and is entirely absent in front 

of the small dorsal pit (fig. 32). | 

3. The intermediate carinae of the carapace are completely absent and the laterals 

are only visible in the extreme posterior part of the carapace (fig. 32'). 

4. The rostrum only reaches to three-quarters the length of the antennular somite. 
It is broader than long, bluntly rounded anteriorly, and shows only the faintest 
indications of a median carina. 

The eyestalks are rather more swollen. 

The lateral process of the fifth thoracic somite trends obliquely forwards. 

7. The telson is rather broader and the mid-dorsal carina is much less elevated 

(figs. 33, 34). 


Several male specimens of comparatively large size were examined, but the margins 
of the telson are not LE as strongly swollen as in large S. leptosquilla of the same 


© 1 


! The Be is erroneous in ae intermediate carinae on the carapace. Anterior to the cervical 
groove the regions external to the gastric grooves are evenly rounded; only at the extreme posterior end 
of the indicated carina is there the faintest trace of a keel. ‘Ihe median carina, also, does not extend 
as far forwards as is shown. 


1913.] S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 49 


sex and the lateral tooth though shorter and stouter than in the female is much longer. 
There are ten to fourteen fine submedian denticles, nine to twelve intermediate and 
one lateral. The spines on the abdominal somites have the same distribution as in 
S. leptosquilla and, as in that species, there is no mandibular palp and the dactylus 
of the raptorial claw bears four teeth including the apical one. 

The preserved specimens show no trace of colour ; the large brown spots on the 
telson, which are invariably found in S. leptosquilla, seem to be absent. Wood-Mason 
remarks that the colour in life is deep pink. 


There are eleven specimens in the Indian Museum registered thus :— 


DE Off Ganjam Coast, Madras; 98—102 fms., 


18°40’ N., 84°46’ E. “Investigator.” I¢, 62mm. TYPE. 

a2 Off Cheduba, Arakan Coast, Burma. “Investigator.” 19,35 mm. TYPE. 
nz Off Masulipatam, Madras; 95 fms., 

15°56'20” N., 81°26’10” E. ‘ Investigator.” 28,45, 46 mm. 
ee Off Masulipatam, Madras; 240 fms., 

15°56’50” N., 81°30°30” E. ‘ Investigator.’ Id, 375 mm. 
T4 Off Nellore Coast, Madras; 112 fms., 

14555” N., 80°20’50” E. ‘ Investigator.” 30,29, 4I—58 mm. 
fe Off Andaman Is.; 188—220 fms., 

IT 37 40° N., 92° 46°40” E. ‘ Investigator.’ I 9,50 mm. 


Only the above specimens are known. 


16, Squilla laevis, Hess. 
Plate III, figs. 35—37. 
1865. Squilla laevis, Hess, Arch. f. Naturgesch., XXXI, i, p. 170, pl. vii, fig. 22. 
1887. Squilla laevis, de Man, Zool. Jahrb., Syst., II, p. 715. 
1894. Squilla laevis, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 511. 
1898. Squilla laevis, Stead, Zoologist (4), II, p. 211. 

The carapace is smooth or very feebly rugose and its breadth behind the antero- 
lateral spines is more than half its median length, including the rostrum. ‘The lateral 
margin is concave and is produced to a blunt angular point immediately in front of 
the rounded postero-lateral angles. The median carina is distinct' and is anteriorly 
bifurcated for about one-fifth its length anterior to the well-marked cervical groove 
(fig. 35). The small mid-dorsal pit is conspicuous, as are also the intermediate and 
lateral carinae. The rostrum is subquadrate, considerably wider than long ; its lateral 
margins converge to the rounded antero-lateral angles and the distal edge is truncate 
or broadly convex. Dorsally it bears a sharp carina in its distal half and a pair of 
other carinae running close to the lateral margins. 

The cornea of the eye is wider than the stalk and is set obliquely on it; its 
breadth is clearly less than the whole length of the organ. The ophthalmic somite 
projects between the base of the eyestalks in the form of a very prominent subquad- 


! Except in a large male specimen in which it is worn away and obsolescent. 


50 | Memoirs of the Indian Museum. IVOrANE 


rate lobe which is slightly emarginate at the apex (fig. 35). The antennular peduncle 
is shorter than the median length of the carapace, excluding the rostrum. The mandi- 
bular palp is entirely missing. 

The outer inferior margin of the merus of the raptorial claw (fig. 36) is bluntly 
rectangular at its distal extremity. The carpus is deeply grooved externally and 
the dorsal carina, which is high and entire, terminates abruptly before reaching the 
anterior edge. The propodus possesses three movable spines at the base of its pecti- 
nate margin and the opposite or inferior edge is, in both sexes, remarkable for its sharp 
carina which is produced as a stout tooth at the distal end; this tooth is not found in 
any other species of Squilla. The dactylus itself is convex or feebly sinuous externally 
and on its inner margin bears six curved teeth including the apical one (fig. 36). 

The dorsal surfaces of the exposed thoracic somites and abdomen are finely rugose. 
The lateral margin of the fifth thoracic somite consists of a sharp anterior forwardly- 
curved spine and a posterior lobe which is rather narrowly rounded at the apex. In 
the possession of a bilobed margin on this somite, S. /aevis, along with fifteen other 
species hereinafter described, differs conspicuously from any of the forms previously 
mentioned. The pair of inferior spines found in most of the preceding species is 
entirely absent. The lateral processes of the two succeeding somites are composed of 
a single lobe, obliquely truncate in front and terminating in an acute but non-spinous 
postero-lateral angle. The eighth somite is provided with a sharp antero-lateral tooth 
(figs. 35, 37). The spines on the abdomen are disposed as follows :— 


Carinae. Abdominal somites. 
Submedian Er ae Be 5 © 
Intermediate Br Er à Bia Os 
Wateraleer a = oe (3) 4, 5, ©. 
Marginal Se de 50 16 Ay By Che. Sha 


The median carina of the telson is only very feebly notched at the base and the 
symmetrical rows of pits are distinct on eitherside. There are two or three submedian 
denticles, six to eight intermediate and one lateral. There is a carina at the base of 
each marginal tooth and the carina which runs along the lateral edge of the telson, in 
its anterior third, ends in a blunt praelateral denticle. There is no post-anal carina. 

The inner spine of the bifurcate process of the uropod is twice the length of the 
outer and bears a small lobe on its outer edge just behind its middle point. The basal 
segment of the exopod is only a trifle longer than the ultimate segment and bears 
seven movable spines on its external margin. 

Secondary sexual distinctions are slight. In the adult male the raptorial pro- 
podus is slightly more dilated than in the female ; and the dactylus in the former sex 
is of a rather more clumsy build and has shorter and stouter teeth. 

The preserved specimens show no characteristic colouration. 

There are five examples of the species in the Indian Museum :— 

TS New South Wales. D. G. Stead. 3¢, JI—IIg mm. 


7511—2 


nn Port Jackson, New South Wales. Australian Museum. 29, 99 and 105 mm. 


Squilla laevis is at present only known from the coasts of New South Wales ; 
it has been recorded from Port Jackson (Stead) and Sydney (Hess). 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. SI 


17. Squilla hieroglyphica, Kemp. 
Plate III, figs. 33—41. 
IgII. Squilla hieroglyphica, Kemp, Rec. Ind. Mus., VI, p. 96. 

This species is closely allied to the preceding, but differs from it in the following 

characters :— 

I. The median carina of the carapace is distinet in front of the small mid-dorsal 
pit, but the anterior bifurcated part is entirely absent. The lateral margins 
are not angled in front of the rounded postero-lateral corners and the anterior 
width is only about half the median length, including the rostrum (fig. 38). 

2. The rostrum is as long as wide and its strongly convergent lateral margins meet in 
a narrow rounded apex. Inits carination it resembles that of S. laevıs (fig. 38). 

3. The cornea of the eyes is set transversely on the stalk. The anterior margin 
of the ophthalmic somite is broadly convex and does not project as a pro- 
minent lobe between the base of the eyestalks (fig. 38). 

4. The propodus of the raptorial claw does not possess a spine at the distal end 
of its inferior margin. The dactylus is armed with only five teeth including 
the terminal one (fig. 39). 

5. The anterior lateral process of the fifth thoracic somite is broader and shorter, 
and the posterior process of this somite and the lateral margins of the two 
succeeding somites are more broadly rounded. ‘The lateral portions project 
almost horizontally outwards and do not reach downwards as much as in the 
preceding species ; this feature, taken in conjunction with the slightly greater 
proportional length of the free thoracic somites, is responsible for the widely 
different aspect of these parts in lateral view (cf. fig. 37 x 13 and fig. 40 x 5). 

6. The following abdominal carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian de de oF 5, 6. 
Intermediate id Ba As 45,0: 
Tateral .. oe 5% Sf Bs dl) Ee CE 
Marginal ar Le ir 25 Bo do Fo 


7. On the telson there are five submedian denticles, ten to twelve intermediate 
and one lateral. There is no praelateral denticle. On the ventral surface there 
is a sharp and long post-anal carina (fig. 41). 

8. There are eight or nine movable spines on the outer margin of the basal joint 
of the exopodite of the uropod. 


The mandibular palp, as in S. /aevis, is entirely missing. 


The unique example of S. hieroglyphica is of a dark yellow colour and appears to 
have been preserved in spirit for many years ; jet-black markings are, however, dis- 
tinct and on the telson seem to be quite characteristic. The surface of the carapace 
and abdomen is very sparsely covered with large black chromatophores which are 
aggregated and sharply define the outer margins and median carina of the rostrum, 
the anterior portion of the gastric groove and the posterior margin of the segments of 


52 Memoırs of the Indian Museum. Vom 


the post-abdomen except the first and last. There are three prominentspots arranged 
in the form of a triangle on each eyestalk. The peculiar arrangement of the chromato- 
phores on either side of the telson is shown in fig. 41 ; it differs entirely from that 
seen in any other species. 

The solitary specimen (°,) is a female 53 mm. in length. Unfortunately no 
information concerning the locality at which it was taken is available, though there 
can be little doubt that it was found within the area with which this paper is concerned. 


18, Squilla quinquedentata', Brooks. 
1886. Squilla quinquedentata, Brooks, Voy. H.M.S. ‘Challenger,’ XVI, Stomatop., p. 26, pl.i, 
ers joe wl, ile, Où 
1894. Squilla quinquedentata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 511. 

The surface of the carapace is smooth, highly polished and very finely punctate- 
The abdomen also presents a polished appearance, but is rather more coatsely pitted. 

The breadth of the carapace, measured behind the antero-lateral angles, is about 
half its length excluding the rostrum. The median carina is well-marked in front 
of the cervical groove. Anterior to the small dorsal pit it is distinct fora distance equal 
to the length of the rostrum, but of the anterior bifurcated portion only the faintest 
traces exist. Behind the cervical groove the median carina is sharp ; it is bifurcated 
for the greater part of its length and ends posteriorly in a prominent rounded tubercle, 
The intermediate and lateral carinae are well developed. ‘The antero-lateral angles are 
produced to short spines that barely reach to the level of the rostral articulation ; the 
posterior angles are sharply rounded. 

The rostrum is subquadrate, longer than wide, with its upturned lateral margins 
only very slightly convergent. Dorsally it is finely punctate and bears a smooth, flat, 
median tubercle. 

The eyes are not large; the breadth of the cornea is about equal to the total length 
and is not more than one-fifth the length of the carapace. The corneal and peduncular 
axes are very slightly oblique and the stalk is somewhat dilated externally. The 
anterior margin of the ophthalmic somite is evenly convex. The antennular peduncle 
is shorter than the carapace excluding the rostrum. The mandibular palp consists 
of three segments. 

The merus of the raptorial claw is armed with a stout spine at the end of its outer 
inferior margin. ‘The carpus is grooved and ridged externally and the dorsal carina 
is entire and terminates abruptly before reaching the anterior edge. The propodus 
is not broadened distally in the adult male. The dactylus is armed with five teeth 
including the apical one ; the external margin is rather strongly sinuous and is very 
obtusely angled near its basal articulation. 

Submedian and intermediate carinae are distinct on the last three thoracic somites. 
The lateral margins of the fifth somite are bilobed ; the anterior lobe consists of a 
sharp forwardly-directed spine, the posterior is shorter and subacute. ‘There are no 
inferior spines on this somite. ‘The sixth and seventh somites are also bilobed later- 


! See addendum, p. 195. 


1913. | S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 53 


ally ; in each the posterior lobe is acute and very much broader than the anterior. In 
the sixth somite the anterior lobe is narrow, acute, and a trifle shorter than the posterior ; 
in the seventh it is very short and bluntly rounded. The lateral process on the anterior 
part of the eighth somite is sharply acute. 

There are, as usual, four pairs of longitudinal carinae on the first five abdominal 
somites and three on the last, and on all the somites except the first and the sixth there 
is a small transversely-grooved median tubercle. The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian ae ts oe So: 
Intermediate ie ar. ge AUS SO 
Lateral RS EN: ait 425510. 
Marginal ma as py By Bo Aly 5 


The median carina of the telson is well marked ; it is obscurely notched at the 
base and terminates in a sharp spine overhanging a small tubercle. The dorsal sur- 
face on either side is finely rugose and is impressed with a number of parallel and 
oblique rows of pits. These are sometimes confluent and form distinct grooves. The 
six marginal teeth are well developed, the intermediate pair being sometimes slightly 
inturned. The praelateral denticle is distinct and at its base and at the base of each of 
the teeth there is a short rounded carina. There are three submedian denticles, seven 
or eight intermediate and one lateral. The post-anal carina is well developed. 

The bifurcate process from the base of the uropods is finely serrate along its inner 
margin. The inner spine is about twice the length of the outer and bears a prominent 
external lobe near its middle point. 

The two male specimens do not show any marked secondary sexual modifications. 


This species bears a strong superficial resemblance to S. wood-masoni, but may 
at once be distinguished from it and from all other species of the oratoria group by 
the form of the eyes and by the number of teeth on the raptorial dactylus. 


As in the specimen described and figured by Brooks there is a well-defined dark 
patch of colour at either end of the median ridge of the telson. In one of the Indian 
specimens the posterior margins of the post-abdominal segments are defined by black 
pigment and, in addition, in the posterior part of each somite except the first and last, 
there is a dark patch which extends laterally as far as the longitudinal grooves between 
the submedian and intermediate carinae. There is also on each of the first five abdomi- 
nal somites a dark sinuous line immediately above the lateral carinae. 


There are two specimens of S. quinquedentata in the Indian Museum— 


T Bombay. Bombay Nat. Hist. Soc. IS, 103 mm. 
7520 
10 


Balasore Bay, Orissa coast; 15 fms. ‘Golden Crown.’ Io, cd. 115 mm. (damaged). 


Hitherto this species was known only from the type specimen, a male 136 mm. in 
length, found by the ‘ Challenger’ Expedition in the Arafura Sea, south of New 
Guinea, in 28 fathoms. 


54 Memoirs of the Indian Museum. Io I, 


19. Squilla gonypetes, Wood-Mason, MS. 
Plate IV, figs. 42—44. 
1893. Squilla affinis, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 474 (partim), 
1908. Squilla gonypetes, Lloyd, Rec. Ind. Mus., II, p. 33 (sine desc.). 
1911. Squilla gonypetes, Kemp, Rec. Ind. Mus., VI, p. 96. 


This species is very closely allied to S. quinquedentata but may be distinguished 
by the following characters :— 


1. The rostrum is distinctly longer and its upturned lateral margins are rather 
more strongly convergent. 

. The eyes are larger, the breadth of the cornea is about one quarter the length 
of the carapace and the corneal and peduncular axes are strongly oblique 
(fig. 42). 

3. The antennular peduncle is a little longer than the carapace excluding the 

rostrum. 

4. The outer inferior margin of the merus of the raptorial claw is not produced 
as a spine; the external margin of the dactylus shows only the very feeblest 
traces of sinuation (fig. 43). | 

5. The lateral lobes of the sixth thoracic somite are about equal in length ; the 
anterior one is apically truncate and is not very much narrower than the 
posterior which terminates acutely. Both lobes of the seventh somite are 
acute, the anterior being more than half the length of the posterior (fig. 42). 

6. There are more spines on the abdominal somites. The following carinae 
end in spines :— 


ND 


Carinae. Abdominal somites. 
Submedian Re ee os 5, On 
Intermediate ae Be Ar 2 ANS 10: 
Lateral .. EN a Sd Do 35 Lip 5, Op 
Marginal “te Bs es T2: 9: 4 05 


In respect of the punctuation and carination of the carapace and abdomen and 
in the number of teeth on the raptorial dactylus, this species bears the closest resem- 
blance to S. guinquedentata and, like it, possesses a three-segmented mandibular palp. 

The telson (fig. 44) also is closely similar, but the marginal teeth are longer and 
the outermost submedian denticle is often distinctly larger than any of the rest. The 
lobe on the outer aspect of the longer spine of the uropodal process is rather unusually 


prominent. ’There are three or four submedian denticles, six or seven intermediate 
and one lateral. 


Though so nearly allied to S. guinguedentata, there is, I believe, no doubt that S. 
gonypetes is a distinct species. The greater obliquity of the corneal portion of the eyes 
and the different form of the lobes on the margins of the thoracic somites are charac- 


ters which preclude the suggestion that the specimens are merely young examples of 
the form described by Brooks.! 


| Among the species belonging to the S. oraloria group it is noticeable that in very young specimens 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 55 


The colouring of the telson is much the same as in the preceding species, but that 
of the abdomen is different. The proximal half of the second abdominal somite bears 
a prominent transverse black patch extending laterally as far as the sinuous groove 
between the submedian and intermediate carinae and there is also, on the posterior 
half of the fifth abdominal somite external to each submedian carina, a large well 
defined quadrate black spot (fig. 44). 

Four specimens in the Indian Museum agree in possessing the above characters :— 


aus Off Andaman Is.; 60 fms. ‘Investigator.” 1¢,50mm.; 19,36mm. TYPES. 
3859 Off Cheduba, Arakan Coast; 7 fms. ‘Investigator.’ 13,49 mm. 


aie 
#27 Persian Gulf; 47 fms., 26°24’ N., 56°2’ BE. ‘ Investigator.” 12,26 mm. 


10 

I also refer to this species a single specimen which differs from the rest in having 
the anterior bifurcated part of the rostrum fine but distinct, and absent at the point 
of junction with the single posterior part of the carina (as in S. interrupta) ; the ros- 
trum also is subtriangular, the margins distinctly more convergent than in typical gony- 
petes and the intermediate carinae of the third and the laterals of the fourth abdominal 
somites do not terminate in spines. In other respects the specimen bears the closest 
resemblance to those listed above. 


7536 
10 


Off Vizagapatam Coast, Madras; 20 fms. ‘Investigator.’ 19, 47 mm. 


Two specimens from the British Museum collection (recorded by Pocock under 
the name of Squilla affinis) have also been examined, and one, which is identified with 
some doubt, kindly lent by Mr. A. Patience : — 


Holothuria Bank, China Seas, 53 fms. H.M.S. ‘ Penguin.’ 22,25 and 37 mm. 
Mergui Archipelago. Simpson and Rudmose Brown. I, 33 mm. 


Two of these three specimens have lost their raptorial claws and in them the 


median carina of the carapace resembles that of the example from the Madras Coast. 
The other individual (China Seas, 25 mm.) is perfect and is quite typical. 


20. Squilla boops, Kemp. 
Plate IV, figs. 45—47. 

1911. (May) Squilla boops, Kemp (sphalm. boopis), Rec. Ind. Mus., VI, p. 97. 

1011. (August) Squilla quadraticauda, Fukuda, Annot. Zool. Japon., VII, p. 287, pl. xi, figs. 3—5. 

tg11.! Squilla quadraticauda, Fukuda, Döbuts. Z. Tokyo, XXIII, p. 174, fig. 1. 
This very distinct species may be readily distinguished from both the preceding 

by the following. characters :— 

I. The rostrum bears a sharp median longitudinal carina. 


2. The eyes (fig. 45) are enormously swollen and expanded ; the breadth of the 
cornea is considerably more than one-third the length of the carapace and 


the cornea of the eye is a trifle more transversely placed than in adults, while there is very little differ- 
ence in the form of the thoracic lobes. 

1 I have not seen this publication and have no knowledge of the precise date at which it was issued. 
The paper is, I believe, in the Japanese language. 


56 Memoirs of the Indian Museum. (Wor, ION, 


there is a prominent lobe on the external aspect of the eyestalk. The corneal 
and peduncular axes are very oblique. 

3. The antero-lateral angles of the carapace are produced as acute points which 
project outwards and downwards, scarcely reaching at all forwards beyond 
the adjacent anterior margin. 

4. The lateral processes of the fifth thoracic somite consist of a sharp and very 
slender anterior spine, which is oblique but is not strongly curved forwards 
as in most allied species, and a very short acute posterior process. The ante- 
rior process of the sixth somite has the form of a very small acute lobe which 
at its base is not more than one-fifth the width of the broad but acute posterior 
process. The seventh somite is not bilobed laterally but is angled acutely 
behind and rectangularly in front (fig. 45). 


For the rest the principal characters of the species are as follows :— 

The whole surface of the carapace and abdomen is smooth, highly polished and 
without trace of punctuation. The carapace is broad anteriorly; its breadth mea- 
sured behind the antero-lateral angles greatly exceeds half its length, including the ros- 
trum. The median carina is entirely absent in front of the small dorsal pit but other- 
wise resembles that of S. quinquedentata. The lateral and intermediate carinae are well 
marked. The rostrum is broader than long and its upturned lateral margins converge 
to a broad, evenly rounded apex. The anterior margin of the ophthalmic somite is 
sharply pointed between the base of the eyestalks. The antennular peduncle is about 
as long as the carapace. 

The outer inferior margin of the merus of the raptorial claw (fig. 46) is distally 
rounded. The dorsal carina of the carpus is high and terminates in a strong lobe-like 
tooth behind which a minute tubercle is visible. The dactylus, as in the two preced- 
ing species, possesses five teeth including the apical one ; the outer margin is very feebly 
sinuous and is produced almost rectangularly near its articulation with the propodus 
(fig. 46). 

The submedian carinae are rather faint on the abdomen and on the free thoracic 
somites are semi-obsolete. The intermediate carinae are distinct on the thoracic and 
abdominal somites. The following abdominal carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian Le, ibe A 5,6. 
Intermediate BS ae Br 2 Bo Alp 55 Ce 
Lateral .. 3 ae sven iy 2, SA 5 0; 
Marginal Ars HR =f ily 25 Bo Aly Fe 


The telson is almost precisely similar to that of S. guinquedentata, but the teeth 
are rather longer and the intermediate pair, in the type specimen, are not inturned. 
There are three or four submedian denticles, eight intermediate and one lateral (fig. 47). 

All the median parts of the carapace, abdomen and telson are covered with very 
small grey chromatophores. ‘These are not sufficiently crowded to detract from the 
general yellowish appearance of the spirit specimen, but are closely aggregated and 
form faint transverse lines at the posterior margins of the abdominal and last three 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 57 


thoracic somites. The posterior half of the fifth abdominal somite between the marginal 
and lateral carinae and the outer posterior angles of the sixth somite are black. 

The only differences that I have been able to detect between the specimen which 
Fukuda has described under the name of Squilla quadraticauda and the type of S. 
boops are that in the former example the rostrum is more triangular in outline, the 
intermediate carinae of the second and the lateral carinae of the first abdominal somites 
do not end in spines, and the submedian pair of marginal spines of the telson are more 
divergent and the intermediate pair more convergent. Also, according to Fukuda’s 
figure, the anterior margin ofthe ophthalmic somite is not pointed in the Japanese indivi- 
dual, the antennular peduncle is longer and the teeth of the raptorial dactylus shorter. 

Fukuda’s specimen is much smaller than that from Indian waters and some of the 
differences noticed may be merely growth-stages. The Japanese example may possibly 
represent a distinct variety, but there can be little doubt that both individuals must 
be referred to a single species. 


— Gulf of Martaban, Burma; 67 fms. 14°26’ N., 96°23’ E. ‘Investigator. 19,8q9mm. TYPE. 


The example described by Fukuda under the name of S. quadraticauda is a female, 
40 mm. in length. It was obtained at Matsuwa, Sagami Province, Japan. 


DAE SOPH CIES OR THES. NEPA GROUP. 


This group may be briefly defined as containing those species of Squilla which 
have a series of fine pectinations along the upper edge of the propodus of the raptorial 
claw, six teeth on the raptorial dactylus (including the terminal one), bilobed lateral 
margins to the fifth and sixth thoracic somites and eight longitudinal carinae on the 
first five segments of the abdomen. 

Until comparatively recently all the species which agreed in the above characters 
were included under a single specific name, Squilla nepa. Bigelow, however, in 1894, 
redefined Berthold’s S. affinis (=S. oratoria, De Haan) and pointed out the features 
by which it might be distinguished from S. nepa. Shortly afterwards two other species, 
S. foveolata and S. stridulans, were described by Wood-Mason and during the last few 
years Nobili and Balss have insisted on the claims of Kossmann’s S. massavensis to 
specific recognition. 

But even now, the characters of some of the forms are very imperfectly known, 
and in examining the material at my disposal I have been led to describe three new 
species and one new variety, all of which appear to be far from uncommon in Indo- 
pacific waters. Fortunately, I am able to introduce these new forms with a very con- 
siderable degree of confidence. This is due in the first place to the large and valuable 
series of specimens in the Indian Museum and secondly to the numerous examples from 
other sources which I have been able to examine. In particular, mention must be 
made of the fine series kindly lent by the Trustees of the British Museum. This collec- 
tion, which contains numbers of specimens recorded by Miers, Brooks, Henderson and 
Pocock, has enabled me to trace the synonymy of some of the species in a way that 


58 Memoirs of the Indian Museum. [Vora ave 


would otherwise have been impossible and has proved of inestimable value in checking 
and correcting the preliminary results arrived at from a study of the material in the 
Indian Museum. Moreover, coming as they do from many widely distant localities, 
I have been able to obtain a very good estimate of the distribution of the various forms. 
In all nearly 800-specimens have been examined. 

To many, the characters of the species which I have described below will appear 
extremely trivial, and it must be admitted that much more work is necessary before 
their exact relations can be decided in a really satisfactory manner. The point, how- 
ever, which I specially desire to emphasize, which has been more and more impressed 
upon me as additional collections came under examination, is that in Indian waters 
there exist seven perfectly distinct forms that may always be separated from one 
another with the greatest facility. Four of these, which I have called S. oratoria 
var. perpensa, S. interrupta, S. wood-masom and S. massavensis, are very close allies 
of S. oratoria, s.s.,a species which does not seem to occur in its typical form westwards 
of the Philippines. Some will perhaps be inclined to regard them as varieties or sub- 
species, but this is a point of little moment so long as their structure and distribution 
are understood. 

It is only with great difficulty that young specimens (sometimes as much as 
40 mm. in length) can be determined. In several cases I have been obliged to defer 
any attempt at precise identification, but I do not doubt that if sufficiently extensive 
series were available it would be possible to separate the different forms in a satis- 
factory manner at all stages. 

I have found myself unable to determine the precise position of the form which 
Nobili recognized under the name Sguilla affinis var. intermedia.' The original speci- 
mens were obtained at Nias and Singapore and, on comparing them with ‘ S. affinis’? 
from the latter locality, Nobili notes that the external margin of the raptorial dactylus 
is more strongly sinuous and the eyes a little smaller. The dorsal carina of the carpus 
of the raptorial claw is without tubercles. It is possible, and even probable, that the 
form will prove identical with one of those described in this paper,’ but without 
knowledge of the characters afforded by the median carina of the carapace, by the 
lateral lobes of the free thoracic somites, and by the basal process of the uropods, no 
satisfactory conclusions can be reached. I have tried to obtain a loan of the original 
specimens, but have been unable to discover where they are preserved. 


21. Squilla foveolata, Wood-Mason. 
Plate IV, fig. 48. 
1895. Squilla foveolata, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 2, pl. ii, fig. 1. 
The dorsal surface of the carapace and abdomen is very coarsely and deeply 
impressed. This is specially marked on the antero-lateral parts of the carapace and 


! Nobili, Boll. Mus. Torino, XVIII, 1903, No. 455, p- 30. 
? These specimens are perhaps Squilla oratoria var. perpensa and S. interrupta. 
° As a specific name intermedia is preoccupied by Bigelow’s S. intermedia from the American Coast. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 59 


over all the median portions of the carapace and abdomen. On these areas the 
punctuation is so coarse and so close that the surface stands up as a sharp mesh-like 
reticulation which obscures the median and submedian carinae (fig. 48). 

The breadth of the carapace measured behind the antero-lateral angles is about 
half its length, excluding the rostrum. The median carina, which owing to the reti- 
culation of the surface can only be traced with difficulty, extends throughout the 
length of the carapace and is bifurcated anteriorly for about half its length in front 
of the small but distinct dorsal pit; it is also deeply bifurcate posterior to the sharply- 
defined cervical groove. The lateral and intermediate carinae are more distinct. The 
antero-lateral angles are produced as sharp spines that fail to reach the level of the 
rostral base and the lateral margins are obtusely angled in front of the postero- 
lateral corners. 

The rostrum is fully as broad as long and its lateral margins are upturned and 
converge to an evenly rounded extremity. The dorsal surface is rugose and bears a 
sharp median carina in its anterior two-thirds. 

The eyes are very small. The corneais about as wide as the basal portion of the 
stalk and the internal margin of the latter is dilated and ridged. The corneal and 
peduncular axes are very slightly oblique.' The ophthalmic somite is produced ante- 
riorly as a short rectangular process between the bases of the eyestalks. The anten- 
nular peduncle is unusually long, greatly exceeding the combined length of the cara- 
pace and rostrum. ‘The dorsal processes of the antennular somite are pitted and 
reticulate dorsally. The mandibular palp is present and is composed of three segments. 

The outer inferior edge of the merus of the raptorial claw is pitted externally 
and is distally rounded. There are two parallel and connected ridges forming an H- 
figure on the outer aspect of the carpus. The dorsal carina is entire and terminates 
abruptly before reaching the anterior margin. The propodus bears the usual three 
movable spines and the row of pectinations on the margin opposed to the dactylus. 
The latter is provided with six slender teeth including the apical one and the 
external margin, which shows only the faintest traces of sinuation, is bluntly angled 
proximally. 

The last three thoracic somites (fig. 48) are furnished with submedian and inter- 
mediate carinae, the former being very indistinct. The lateral margin of the fifth 
somite is deeply bifurcated and consists of two subequal spines, the anterior of which 
is sharper and situated on a lower plane than the posterior and points almost straight 
forwards. The sixth thoracic somite is deeply bilobed laterally; the lobes are greatly 
expanded, coarsely pitted and are of about equal length and breadth: the anterior is 
distally truncate while the posterior is rounded at the apex. In the seventh somite 


! The eye in S. foveolata is very peculiar. In other species of Squilla the long axis of the cornea is 
usually in the same plane as the longest transverse axis of the stalk, but in the present case the cornea is 
twisted and its inner end is, in relation to the stalk, subdorsal. The cornea, as mentioned above, is set 
rather obliquely on the stalk; but this obliquity is the reverse of that found in most other species, 
for the inner margin of the stalk is a trifle shorter than the outer. 


60 Memoirs of the Indian Museum. Mora 


the anterior process is acute and is much shorter and narrower than the rounded 
posterior one (fig. 48). 

Four pairs of longitudinal carinae are present on the first five abdominal somites 
and three pairs on the sixth. ‘The submedians, on all except the last somite, are 
almost lost in the coarse reticulation of the surface. There is an obscure median 
tubercle on the 2nd, 3rd, 4th and 5th somites. 

The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian 2: Se is Sn Aa 35 Os 
Intermediate 2: ER Ro 
Lateral de a bo (As By Zp Bo Oe 
Marginal a a nie Tinie oredr oe 


The telson is considerably longer than broad. The median carina is not notched 
at the base and terminates in a short spine which overhangs a small tubercle. On 
either side the surface is finely rugose; it bears a longitudinal line of small tubercle 
and beyond this several oblique rows of symmetrically-disposed pits leading to the 
interspaces of the intermediate denticles. |’These pits are very deep and often con- 
fluent so that the spaces between them are well defined and appear as blunt ridges. 
Six sharp marginal teeth are present, each of which forms the termination of a 
sharp dorsal carina. There is no praelateral denticle. There are three (rarely two) 
submedian denticles, six to eight intermediate and one lateral. The post-anal crest 
is often broken up into a number of irregular tubercles. 

The bifurcate process from the base of the uropod is finely serrate internally. 
The inner spine is nearly twice the length of the outer and bears externally, at about 
its middle point, a prominent rounded lobe. ‘The paddle or outer segment of the 
exopodite is distinctly shorter than the basal segment. 

The preserved Indian specimens of S. foveolata show no trace of their original 
colour; the Chinese examples are of a uniform blue-green tone. 


This beautifully sculptured species appears to be very rare. Nine specimens 


from the following localities are in the Indian Museum :— 
3332 


> Hongkong. G. Dennys. 19,09,77—108 mm. 
TYPES. 
m™ Vé River entrance, Tennasserim, Burma. ‘ Investigator.’ 19,76 mm. 
ee Off Amherst I,  Tennasserim, Burma. ‘ Investigator.’ 1?, 67 mm. 


6 
Only the above specimens are known. 


22. Squilla nepa, Latreille' (Bigelow) = 
Plate IV, fig. 49. 
1825. Squilla nepa, Latreille, Encycl. Méthod., X, p. 471. 
1845. Squilla nepa, Berthold, Abhandl. Gess. Wiss. Göttingen, III, p. 29, pl. iii, figs. 3-5. 


1849. Squilla nepa, De Haan, in Siebold’s Fauna Japonica, Crust., p- 22T. 
1861. Squilla edwardsii, Giebel, Zeitschr. Ges. Naturwiss., XVIII, p. 320. 


1 See addendum, p. 195. 


TOT S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 61 


1880. Squilla nepa, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 25, 458, pl. ii, fig. 13 (partim). 
1893. Squilla nepa, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 452 (partim). 

1893. Squilla nepa, Bigelow, John Hopkins Univ. Circ., No. 106, p. 102. 

1894. Squilla nepa, Bigelow, Proc. U.S. Nat. Mus, XVII, p. 535, fig. 21. 

1908. Squilla hemischista, Wood-Mason, MS., Lloyd, Rec. Ind. Mus., II, p. 32 (sine desc.). 


Squilla nepa was only clearly distinguished from S. oratoria in 1893. ‘The precise 
identity of the specimens recorded in the following papers is uncertain :— 
- 1796. Cancer (mantis) digitalis, Herbst, Krabben u. Krebse, II, p. 92, pl. xxxiii, fig. 1. 

1837. Squilla nepa, H. Milne-Edwards, Hist. Nat. Crust., II, p. 522. 

1839. Squilla nepa, Randall, Journ. Acad. Sci. Philadelphia (1), VIII, p. 147. 

1847. Squilla nepa, White, Hist. Crust. Brit. Mus., p. 83. 

1859. Squilla nepa, Nicolet, in Gay’s Hist. fisica de Chile, Zool. III, p. 224. 

1865. Squilla nepa, Heller, Reise ‘Novara’ Exped., Crust., p. 124. 

1876. Squilla nepa, Miers, Cat. Crust. N. Zealand, p. 89. 

1882. Squilla nepa, Haswell, Cat. Australian Crust., p. 208. 

1886. Squilla nepa, Filhol, Miss. de l’île Campbell, III, 2e, p. 435. 

1887. Squilla nepa, Walker, Journ. Linn. Soc., XX, p. 113. 

1892. Squilla nepa, de Man, in Weber’s Zool. Ergebn. Nied. Ost.-Ind., II, p. 518. 

1892. Squilla nepa, Thallwitz, Abhandl. Kon. Mus. Dresden, No. 3, p. 55. 

1001. Squilla nepa, Lanchester,! Proc. Zool. Soc., p. 553. 

1906. Squilla nepa, Lanchester,’ Fascic. Malayenses, Zool., III, p. 133. 


In the following papers reference is probably made to the true S. nepa, but there 


is a possibility of confusion with S. holoschista :— 
1898. Squilla nepa, de Man, Zool. Jahrb. Syst., X, p. 693. 
1899. Squilla nepa, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 275. 
1900. Squilla nepa, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 519. 
1901. Squilla nepa, Lenz, Zool. Jahrb. Syst., XIV, p. 477. 
1903. Squilla nepa, Nobili, Boll. Mus. Torino, XVIII, No. 452, p. 23, and No. 455, p. 38. 
1906. Squilla nepa, Tattersall, Ceylon Pearl Oyster Rep., V, p. 166. 
1908. Squilla nepa, Stebbing, Ann. S. African Mus., VI, p. 44. 
IgIo. Squilla nepa, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 571. 


The dorsal surface of the carapace and abdomen is finely rugose ; especially so on 
the median area of the latter. The carapace measured behind the antero-lateral angles 
is rather less than half its length, including the rostrum. ‘The median carina of the 
carapace (fig. 49) is sharp and distinct throughout its course. It is bifurcated in front 
for half, or rather more than half, its length anterior to the cervical groove, and the small 
median dorsal pit is enclosed in the base’of the fork. Behind the cervical groove it is 
bifurcated for the greater part of its length and projects as a prominent tubercle in 
the middle of the posterior margin. The intermediate and lateral carinae are distinct. 
The spines at the antero-lateral angles are very strong and project far beyond the level 
of the rostral articulation ; the postero-lateral angles are rounded. 

The rostrum is a little broader than long and its upturned lateral margins are rather 
strongly convergent. Dorsally it sometimes bears an obscure median tubercle and 
this, in one very large specimen, forms an obsolete median carina. 


! Lanchester does not seem to have been aware of Bigelow’s important paper of 1894. 


62 Memoirs of the Indian Museum. [Vou. IV, 


The eyes are not greatly expanded. The cornea is much broader than the stalk 
and its greatest breadth is distinctly less than the length of the whole organ. The 
corneal and peduncular axes are at right angles to one another and the anterior margin 
of the ophthalmic somite is produced and at the apex truncate or slightly emarginate. 
In very young specimens, about 30 mm. in length, the corneal index ' is approximately 
45 ; it increases as the animal grows and is about 7'0 in examples between 80 and 
90 mm. in length, while in the largest individual examined it is as much as 80. The 
antennular peduncle is shorter than the carapace excluding the rostrum. The mandi- 
bular palp is composed of three segments. 

The outer margin of the merus of the raptorial claw terminates anteriorly in a 
sharp tooth. The carpus is grooved and carinate externally and its dorsal keel is 
furnished with two tubercles, of which the distal is often bilobed ; more rarely there 
are three distinct tubercles. The merus is somewhat expanded at its distal end in the 
adult male and bears three movable spines and a row of fine pectinations on the margin 
opposed to the dactylus. The dactylus is provided with six curved teeth including 
the apical one ; the external margin is obtusely angled at the base and is strongly 

inuous in both sexes. The proximal part of the dactylus is slightly swollen. 

The last three thoracic somites are furnished with sharp submedian and intermediate 
carinae. The fifth somite is bilobed laterally and does not possess any spines on the 
inferior margin. The anterior process consists of a very sharp antrorse spine; the 
posterior process, which is not so sharp, is directed straight outwards and is less than 
half the length of the anterior. The two marginal lobes of the sixth somite are almost 
equal in length, but the anterior is narrower than the posterior and is obliquely trun- 
cate at the apex. Both lobes of the seventh somite are acute, but the anterior is very 
much shorter and narrower than the posterior. 

There are four pairs of sharp longitudinal carinae on the first five abdominal 
somites and three pairs on the sixth. On the 2nd, 3rd, 4th and 5th between the 


submedian carinae there is a small elongated and transversely-notched median tubercle. 
The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian ? a5 ge 2050: 
Intermediate a os aa (A) BoA Big Oe 
Lateral } a (Q) 2, Bp Als 35 Oe 
Marginal bh is oe ey Ay D Aly So 


The telson bears a close resemblance to that of the species belonging to the quin- 
quedentata group. ‘The median carina is sharp and is very obscurely notched at the 
base ; it terminates in a short spine which overhangs a single blunt tubercle. The 
oblique rows of pits leading to the interspaces of the intermediate marginal denticles 
are distinct and there is also a single longitudinal row on either side of the median 
carina. The submedian, intermediate and lateral teeth are well marked and behind 


1 The corneal index :—the number of times the breadth of the cornea is contained in the median 
length of the carapace, excluding the rostrum. (See p. 9.) 


* Very rarely, in large specimens, the submedian carinae of the third abdominal somite end in spines. 


1013.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 63 


the last there is a small praelateral denticle. This denticle and each of the teeth forms 
the termination of a short but well-marked longitudinal carina. There are three 
or four (rarely two) submedian denticles, seven to nine (rarely six) intermediate and one 
lateral. The post-anal carina is well marked. 

The bifurcate process of the uropod is finely serrate internally. ‘The inner spine 
is fully twice the length of the outer and bears a small rounded external lobe near the 
distal end of its proximal half. The ultimate segment of the exopodite is almost 
equal in length to the basal segment. 


To this species I refer several extremely young individuals, between 20 and 30 
mm. in length, in which the spines at the antero-lateral angles of the carapace are not 
developed. In these examples the corneal index varies from 3°6 to 4'5, the corneal 
and peduncular axes are very slightly oblique and the median carina of the carapace is 
obsolete anteriorly. 


The general colour of living specimens of this species is usually smoky-grey. All the 
catinae of the carapace, abdomen and telson are defined by very dark grey pigment ; 
on the second abdominal somite there is frequently a dark dorsal patch and a 
similar patch is occasionally found on the fifth somite. The eyes are bright trans- 
lucent green, darker at the inner dorsal end of the cornea. The last three segments of 
the raptorial claw are pure white, those of the appendages of the last three thoracic 
somites and portions of the uropods are greenish yellow. The ventral surface is 
uniformly pale. 

The dark grey colouration mentioned above appears to persist fairly well in 
alcohol, but is sometimes entirely absent. Several specimens from Tuticorin, received 
in formalin shortly after capture, showed a wholly different type of colouring. There 
was no trace of dark pigmentation and the whole dorsal surface along with the basal 
parts of the appendages was suffused with rosy red, tending to dull purple on the 
telson and to a deep bluish purple on all the segments of the uropods except the outer 
segment of the exopod. The median and submedian carinae were red and the an- 
tennular peduncle purple. 


There are two hundred and fifteen specimens in the Indian Museum from the 


following localities :— 


> Hongkong. G. Dennys. Id ,12, 99 and 129 mm. 
a Singapore ? Raffles Museum. Id, 119 mm. 
ran F. Stoliczka. 18,32, 84—118 mm. 
3060 

sa Sandheads, Ganges Delta. J. Barnett. AS, JO, SES un 


N. Annandale, F. H. Gravely, } 


10 0 . Ba 
a Fini, ses Corse S. W. K. and J. Caunter. 189,169, 55—101 mm. 


4 
= > Off Orissa Coast, 7—8 fms. ‘ Investigator.’ 15S, 24% , 59—105 mm. 
3462—79 

9 


64 Memoirs of the Indian Museum. 


Off Ganjam Coast, Madras, 
7—9 fms. 


Madras. 73—91 fms. 


5781-97 ? 
9 a Off Vizagapatam Coast, 
A Madras. 


‘ Investigator.’ 


‘ Investigator.’ 


| ‘ Investigator.’ 
< Madras Museum. 
| (purchased.) 


[Vor. IV, 


Oe AIO 22—93 mm. 


73,132, 64—100 mm. 


348 , 309 ,26—133 mm. 


3461-6 
"#6 Cuddalore and Porto Novo, T.H. Hill. 58,69, 59—158 mm. 
S. India. 
SM Cochin, S. W. India. F. Day. 48,49 ,56—67 mm. 


( Bombay Nat. Hist. Soc. 


| H. P. le Mesurier. 23,22,57—166 mm. 


Bombay. 


=o. Karachı. Karachi Museum. 14,57 mm. 


7307—10 
10 


a Tuticorin, S. India. 


I have also examined the following specimens from other sources :— 


50,1292, 58—149 mm. 
GS LO 5 Sue, an 


(no locality). 
J. Hornell. 


SARAWAK MUSEUM. 
Buntal and Burong Is., Borneo. 
CoLOMBO MUSEUM. 
Ceylon. 
TRIVANDRUM MUSEUM. 
Thirkaunapuzha, Travancore. R.S.N. Pillay. GE 
In the present confused state of our knowledge it is scarcely necessary to enumerate 
the many localities from which S. nepa has been reported. It is clear from the series 
in the Indian Museum that the species is distributed over a very wide area ranging 
from Hongkong to Karachi. Most of the records subsequent to 1894 may very reason- 
ably be regarded as correct and, if this should prove to be the case, the known distri- 
bution extends from Honolulu (Lenz) to Madagascar (Lenz) and Durban (Stebbing). 


23. Squilla holoschista', Wood-Mason, MS. 
Plate IV, figs. 50—53. 
1893. Squilla nepa, Henderson, Trans. Linn. Soc., Zool. (2), V, p. 452 (partim). 
1908. Squilla holoschista, Lloyd, Rec. Ind. Mus., II, p. 32 (sine desc.). 
1911. Squilla holoschista, Kemp, Rec. Ind. Mus., VI, p. 97. 
This species is very closely allied to the preceding, but may readily be distinguished 
from it by the following well-marked characteristics :— 
I. The median carina of the carapace in front of the cervical groove 1s bifurcated 
anteriorly for less than one third of its length and is finely bicarinate through- 
out the greater part.of its entire extent (fig. 51). 
2. The lateral margins of the rostrum are rather more strongly convergent an- 
teriorly and the apex is more rounded (fig. 50). 


Id,32, 58—11I7 mm. 
23,29, 84—145 mm. 


19,70, 84 mm. 


! See addendum, p. 195. 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 65 


3. The corneal portion of the eyes is less expanded ; in specimens from 80 to go 
mm. in length the corneal index is about 9’0 whereas it is about 7:0 in 
examples of S. nepa of a similar size. 

4. In the raptorial claw the spine which terminates the outer inferior margin of 
the merus is smaller and in the adult male the distal end of the propodus is 
very broad and the external margin of the dactylus is only very feebly 
sinuous (fig. 52). 

5. The submedian carinae of the fourth abdominal somite never end in spines : — 


Carinae. Abdominal somites. 
Submedian ae Ks es 5, 6. 
Intermediate He it of (3) 4, 5, 6. 
Lateral .. ae ac 6 M) (@) By 25 Sy ©. 
Marginal Se =: Gb DRANG: 


Not infrequently the marginal teeth of the telson are a trifle shorter than in the 
preceding species and the dorsal rows of pits are less strongly marked. There are two 
to four submedian denticles, eight to ten intermediate, one lateral, and one praelateral. 


Living specimens of S. holoschista closely resemble S. nepa in colour, but the dark 
transverse patches on the second and fifth abdominal somites are absent and the general 
tone of all the other markings is greenish rather than smoky-grey. This greenish 
colouring is especially distinct on the median and marginal carinae of the telson. The 
yellowish pigment has the same distribution as in S. nepa. 

I have examined one hundred and three specimens of this species :— 

a“ Sandheads, Ganges Delta. A. J. Milner: Id, 76 mm. 


7301—6 
10 
8042 
16 


( N. Annandale, ) 


a ur | F. H. Gravely and 8. W.K. | 


53,52, 76—00 mm. 


a Off Vizagapatam Coast, Madras. | : , 
‘ Investigator. Id, 32, 73—93 mm. 
= 74—9 {ms. 
9 2 
3069 
sa | Madras. Madras Museum and purchased, 53, 129 , 49—00 mm. 
am Madras. Madras Museum. TS LZ C5906 Tats 
TYPES. 
u Cuddalore and Porto Novo, I, Jet, til 194,219, 63—-95 mm. 
S. India. 
a Ceylon. Colombo Museum. 14,29,70—85 mm. 
70) (no locality). 29,49, 67—83 mm. 
British Museum. 
Madras. J. R. Henderson. Id, 64 mm. 
(1893, sub S. nepa.) 
Rameswaram I., G. of Manaar. J. R. Henderson. Te, 77 mm: 
Trivandrum Museum, 
Cape Comorin. : 19, 76 mm. 


Distribution. —East Coast of India from the Ganges Delta to Ceylon. 


66 Memoirs of the Indian Museum. Nora 


24. Squilla oratoria, de Haan. 
Plate V, figs. 54-56. 


1844 ?! Squilla oratoria, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. 2. 
1845. Squilla afinis, Berthold, Abhandl. Gess. Wiss. Göttingen, III, p. 26, pl. iii, figs. I—2. 
1849. Squilla oratoria, De Haan, in Siebold’s Fauna Japonica, Crust., p. 223. 
1880. Squilla nepa, Miers, Ann. Mag. Nat. Hist. (5), V, p, 25 (partim). 
1886. Squilla nepa, Brooks, Voy. H.MS. ‘ Challenger,’ XVI, Stomatop., p. 25. 
1803. Squilla affinis, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 494 (partim). 
1893. Squilla affinis, Bigelow, John Hopkins Univ. Cric., No. 106, p. 102. 
1894. Squilla afims, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 538, fig. 22. 
1907. Chloridella afinis, de Man, Trans. Linn. Soc. Zool. (2), IX, p. 430. 
1908. Squilla oratoria, Stebbing, Ann. S. African Mus., VI, pp. 44, 45. 
Squilla mauritiana, Wood-Mason, MS. 
1908. Squilla ovatoria, Lloyd, Rec. Ind. Mus, II, p. 33. 


The identity of the specimens recorded in the following papers is uncertain ; some 

very probably belong to one or other of the new forms described below :— % 

1865. Squilla oratoria, Heller, Reise ‘ Novara’ Exped., Crust., p. 124. 

1898. Squilla affinis, de Man, Zool. Jahrb., Syst. X, p. 693. 

1899. Squilla affinis, Nobili, Ann. Mus. Civ. Genova, (2), XX, p. 275. 

1900. Squilla afınis, Nobili, Ann. Mus. Civ. Genova, (2), XX, p. 510. 

1901. Squilla affinis, Nobili, Boll. Mus. Torino, XVI, No. 397, p. 14. 

1902. Squilla affinis, de Man, Abhandl. Senck. Ges. Frankfurt, XXV, p. 911. 

1G03. Chloridella affınis, Rathbun, Proc. U. S. Nat. Mus., XXVI, p. 55. 

1903. Squilla affinis, Nobili, Boll. Mus. Torino, XVIII, No. 455, p. 38. 


Squilla oratoria along with its variety perpensa and the allied species interrupta, 
wood-masoni, stridulans and massavensis form a very compact group ; all may be sepa- 
rated at a glance from S. foveolata, nepa and holoschista by the large eyes, the cornea of 
which is set very obliquely on the stalk. 

The special characters by which S. oratoria is distinguished from the other closely 
allied forms are :— 

1. The dorsal surface of the carapace and abdomen is strongly punctate and 

never presents a polished appearance. 

2. The median carina of the carapace is sharp and distinct throughout its course 

and is bifurcated in front for only about one quarter its length anterior to 
the cervical groove (fig. 54). 

3. The breadth of the carapace measured behind the antero-lateral angles is less 

than half its length, including the rostrum. 

4. The rostrum is subquadrate and its lateral margins, which are not infrequently 

upturned, are slightly convergent anteriorly. 


! I am not certain of the exact date when the atlas to the Fauna Japonica was published (in the 
copy consulted the plates are bound up with the text and are undated), but I have accepted Stebbing’s 
statement (1908) that, as De Haan himself contended in 1849, it antedates Berthold’s paper. 


1913.| S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 67 


5. The corneal index varies from 3'8 in a specimen only 31°5 mm. long to 60 in 
the largest examples. In specimens between 80 and 90 mm. in length it 
varies from 4°I to 4:3. 

6. The anterior margin of the ophthalmic somite between the bases of the eye- 
stalks is squarely truncate or emarginate, never pointed. 

7. The outer inferior margin of the merus of the raptorial claw is produced to a 
sharp and prominent spine. 

8. The dorsal carina of the raptorial carpus bears from three to five sharp tuber- 
cles in large specimens' ; in smaller examples (50-80 mm.) these are frequently 
represented merely by more or less obscure lobes, while in very young indivi- 
duals (30-60 mm.) the dorsal margin is often entire. The outer margin of the 
dactylus is distinctly sinuous. 

9. The anterior lobe of the lateral margin of the sixth thoracic somite is parallel- 
sided and is much narrower but scarcely shorter than the posterior lobe 
(fig. 54). 

10. The following abdominal carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian ? ae a as 540: 
Intermediate Bc Le SE (2) (3) 4, 5, 6. 
Lateral .. ce os a (2) 35 4, 5, 0. 
Marginal ae ni LE DEAN HE 


11. There are four, less commonly three or five, submedian denticles, seven to 
nine, less commonly six or ten, intermediate and one lateral. 
12. In the bifurcate process from the basal segment of the uropods, the margin 


anterior to the small lobe on the outer edge of the longer spine is always 
strongly concave. 


The colour of spirit specimens is not characteristic. 


Squilla oratoria is, in my experience, much the most variable of the species in the 


group to which it belongs. In practice, however, the sharp and continuous median 
carina of the carapace and its very short anterior bifurcation will, in nearly every 
instance, suffice to distinguish it. The principal features in which the specimens differ 


from the account given above are noticed under the separate records. 
In the Indian Museum are nineteen specimens :— 


oe Yokohama. Berlin Museum. 19, 186 mm. 
EL Nagasaki. F. C. Dale. 19,160 mm. 
3330 9829— 33 
DE DR 4 ( Hongkong Museum. 
ssn | Hongkong. le. Donne 38,89 ,55—146 mm. 


' The tubercles are not sufficiently sharp or distinct in fig. 55. 


3 In four specimens the submedian carinae of the fourth abdominal somite end in spines (see notes 


under the several records). 


68 Memoirs of the Indian Museum. Nyon, I, 


8153 
6 
3394 —6 


i 


Hongkong. Hungerford. 30,12, 64—113 mm. 


4269 
4 


Mauritius. ! (Purchased.) Id ,19, 116, 126 mm. 
(S. mauritiana, Wood-Mason, MS.) 


These specimens differ from typical examples in having the punctuation of the dorsal surface 
very poorly developed. The rostrum also is longer (about as long as broad) and its lateral 
margins are very strongly upturned; the anterior lateral process of the sixth thoracic somite 
is very clearly shorter than the posterior and the submedian carinae of the 4th abdominal 
somite and the lateral carinae of the Ist and 2nd end in spines. 


Thanks to the courtesy of the Trustees of the British Museum, I have also been 
able to examine the following very interesting series of thirty-one specimens :— 


Hawaiian Is. Hawaiian Govt. from Internat. I2, ca. 120 mm. 
Fish. Exhib. of 1883. 


Except that the carpus of the raptorial claw bears only two tubercles and obscure traces of a 
third on its dorsal margin, this specimen is almost the exact counterpart of the two peculiar 
examples recorded from Mauritius. 


Honolulu reefs, Hawaiian Is. ‘ Challenger.’ | 19, 146 mm. 
(Brooks, 1866, sub S. nepa.) 


Except that the intermediate and lateral carinae of all the abdominal somites end in spines this 
example is precisely similar to the preceding. One raptorial claw is missing and in the other, 
which has been broken off and subsequently redevelopel, the dorsal carina of the carpus (as 
in young specimens) terminates abruptly at its distal end but is otherwise entire. 


Yokohama. H. Batson Joyner. 43, 147—177 mm. 
(Miers, 1880, sub S. nepa.) 
Yokohama. ‘Challenger.’ Id ,140 mm. 
(not mentioned by Brooks.) 
Pokoska, Japan, 5—25 {ms. ‘ Challenger.’ 19,145 mm. 
(Brooks, 1886, sub S. nepa.) 

Kobe, Japan. ‘ Challenger.’ 19,77 mm. 
(Brooks, 1886, sub S. nepa.) 

Inland Sea of Japan, 15 fms. ‘Challenger.’ 19,40 mm. 
(Brooks, 1886, sub S. nepa.) 

Inland Sea of Japan. R. Gordon Smith. 33, 71—138 mm. 

Inland Sea of Japan. (de Man, 1907, sub C. affınıs.) IG, 45 mm. 


De Man mentions that the submedian spines of the telson probably possess movable tips, but 
this, I believe, is due merely to their partial fracture. No such character is shown in other 
examples of a similar length. 


! The localities of specimens obtained by purchase are frequently inaccurate and, though I have no 
special reason to distrust this label, I am none the less inclined to regard the record with suspicion. 
Before we can accept the enormous increase in the known geographical range which is implied, fresh 
specimens either from Mauritius or from localities lying between it and the China Seas must be examined, 
for S. oraloria in the strict sense appears to be entirely absent from the coasts of British India. In the 
peculiar varietal characters which are noticed the specimens agree closely with two examples from 
Hawaii. 


Ton. S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 69 


Cheefoo, China. Swinhoe. Id, 76 mm. 

(Miers, 1880, sub S. nepa.) 
Shanghai. Swinhoe. 

(Miers, 1880, sub S. nepa.) 23,12, 89—103 mm. 
Chusan, China. P. W. Bassett Smith. 33,42, 315—88 mm. 


(Pocock, 1893, sub S. affinis.) 


The two largest specimens are typical, but in three of the small ones the tubercles on the dorsal 
aspect of the carpus are undeveloped. 


Foochow, China. F. W. Styan. 39, 78—87 mm. 
In the laigest specimen the median carina is not continuous, but shows a small gap on either 
side near the base of the anterior bifurcation. In one of the smaller specimens also the right 
limb of the bifurcate part is finely interrupted near its anterior end. 


Amoy, China. Stevens. IG, I, IOI, 108 mm. 
(Miers, 1880, sub S. nepa.) 

The dorsal punctuation of the carapace and abdomen is very poorly developed. In the larger 
specimen the anterior half of the bifurcate portion of the median carina of the carapace is 
entirely missing and a wide gap separates the base of the forked portion from the simple 
posterior part; in the smaller specimen the carina is, as usual, continuous. In both there 
are only very feeble indications of the tubercles on the dorsal aspect of the carpus (considering 
the size of the specimens these should be well developed), the anterior lateral process of the 
6th thoracic somite is noticeably shorter than the posterior and the anterior process of the 
7th somite, which in other specimens is acutely produced, is represented merely by a blunt 
obtuse and rounded angle (fig. 56). The spinulation of the abdomen is normal. 


Philippine Is. BE Jo Van, IG, 81 mm. 
(Miers, 1880, sub S. nepa.) 


In this specimen the median carina of the carapace is interrupted at the base of the bifurcate 
part as in var. perpensa and S. interrupla. For the rest it very closely resembles the 
Hawaiian examples and those reported from Mauritius, though the rostrum is a trifle shorter. 
The submedian carinae of the last three, the intermediate carinae of the last five, and the 
lateral carinae of all six abdominal somites are produced posteriorly in the form of spines. 


The following specimens from other sources have also been examined :— 


Prof. K. Kishinouye. 


Bay of Tokio, Japau. , “ Most common and widely | 33,22, 38—168 mm. 
distributed.’’ K. K. 


Mr. Alan Owston. 
Okinose, Sagami, Japan. 14,19, 150 and 165 mm. 


It is possible that the specimens labelled Mauritius, along with those in the British 
Museum collection from the Hawaiian and Philippine Is., deserve to be separated from 
the rest under a distinct varietal or subspecific name, but the material available is not 
sufficient to justify such a course. It should, however, be remembered that these 
examples differ from those found on the coasts of China and Japan in the longer rostrum 
with strongly upturned lateral margins and in the more complete spinous formula 


70 Memoirs of the Indian Museum. [Vor. IV, 


on the abdomen. ‘The shortness of the anterior lateral process of the sixth thoracic 
somite is paralleled in the two remarkable specimens from Amoy. 

Squilla oratoria is evidently an abundant species in Chinese and Japanese waters, 
but is perhaps scarce in other localities. Apart from the record from Mauritius, which, 
for the present at least, may be regarded as doubtful, it is only known from the 
Hawaiian and Philippine Is. The probability of confusion with one or other of the 
succeeding forms is so great that little reliance can be placed on the localities given in 
the literature. 

var. perpensa', Kemp. 
Plate V, figs. 57-59. 
1880. Squilla nepa, Miers, Ann. Mag. Nat. Hist. (5), V, p. 25 (partim). 
1884. Squilla nepa, Miers, Voy. H.M.S. ‘ Alert’, p. 298. 
1888. Squilla nepa, de Man, Journ. Linn. Soc., XXII, p. 295. 
1893. Squilla affinis, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 474. 
Squilla interrupta, Wood-Mason, MS. (partim). 
1908. Squilla interrupta, Lloyd, Rec. Ind. Mus. ii, p. 32 (sine desc.). 
1911. Squılla oratoria, var. perpensa, Kemp, Rec. Ind. Mus., VI, p. 98. 

Under this name I separate the form of Squwilla oratoria which occurs on the coasts 
of British India. In these localities the variety exists to the complete exclusion of the 
typical race, but both forms occur together on the coasts of S. China. 

The var. perpensa may be easily distinguished from the typical form by two 
characters :— 

I. The median carina of the carapace is always interrupted,and wholly absent for 

a short space, at the base of the anterior bifurcation (fig. 57). The two arms 
of the bifurcate portion are frequently fine and obscure and, in a few speci- 
mens, quite obsolete. 

2. The carpus of the raptorial claw has a sharp elevated carina on its dorsal aspect 

which terminates abruptly before reaching the anterior margin ; apart from 
this, there is no trace whatever of any dorsal tubercle (fig. 58). 

The variety also differs from most examples of the typical form in the more quad- 
rate shape of the rostrum, the apex of which is squarely truncate or even a trifle con- 
cave. The lateral lobes of the sixth somite sometimes bears the closest resemblance 
to those of oratoria, s.s., more usually the anterior is broader, more pointed and less 
parallel-sided. The anterior lobe of the seventh somite is usually clearly longer than 
in the allied form (fig. 57). 

The following abdominal carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian aks ae Bib 5,6. 
Intermediate is aie ee 4, 5, 6. 
Lateral or af 2 SAS (Oy 
Marginal ae as = 142,9 A555: 


On the telson there are three or four submedian denticles, six to nine intermediate 
and one lateral. 


! See addendum, p. 196. 


1913.] S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 71 


It will be noticed that under the records of typical S. oratoria are included four 
examples from the British Museum collection in which the median carina of the carapace 
is partially or completely discontinuous. There is reason to believe that one of these 
examples (that from Amoy) is wholly abnormal ; the others are in this respect, and in 
this respect only, intermediate between the typical form and the variety. 

In the entire absence of tubercles on the dorsal carina of the carpus, the var. 
perpensa resembles some very young individuals of 5. oratoria, s.s., but in the former 
the character persists throughout life and in all examples except the four mentioned 
above it is found in association with a discontinuous median carina. 

The Indian specimens present scarcely any perceptible difference from those 
found in S. China, except that in examples from the latter locality the breadth of the 
cornea is perhaps a trifle less and the dorsal punctuation frequently rather more pro- 
nounced. 


The colour of spirit specimens is not characteristic. In recently caught examples 
from Tuticorin the posterior margins of the last three thoracic and allthe abdominal 
somites are defined by dull greenish pigment. The submedian post-abdominal carinae 
are of a similar colour, as are also those of the carapace and the median and marginal 
carinae of the telson. The spines of the telson and of the ventral process of the uropod 
are tipped with bright rosy red and in many specimens suffused patches of the same 
colour are visible on the abdomen between the submedian, intermediate and marginal 
carinae and on the free thoracic somites external to the intermediates. The outer sur- 
face of the raptorial merus is also rosy red and traces of the same tint are frequently 
visible on the carapace. The inner half of the outer uropod is dusky and the eyes are 
brown. 


I have examined one hundred and fifty-two specimens of S. oratoria var. perpensa, 
and in this series the constancy of the characters noted above is most remarkable. 
In the Indian Museum are one hundred and forty-six examples :— 


—— Hongkong. Hongkong Museum. 83, 169, 55—107 mm. 
4851—72 
iss Hongkong. G. Dennys, 184, 229, 67—98 mm. 
10 
DE Hongkong. R. Hungerford. 58,52,66—IoI mm. 
© Mergui Archipelago. Mus. Collr. 19,70 mm. 
(de Man, 1888, sub S. neha.) 
a Of Irrawaddy Delta, 20 fms. ‘ Investigator.’ 30,22, 35—76 mm. 
175,2 200NE, 94550 E: 
5769 
N Sandheads, Hugli Delta. A. J. Milner. 22,57 and 60 mm. 
mon 
a Madras Coast. (no history.) 19,12,65 and 79 mm. 


7036—41 
10 


ws Tuticorin, S. India. J. Hornell. 22, 409 , 39—75 mm. TYPES. 


7521—4 


72 Memoırs of the Indian Museum. [NOE EVE 


The following two small specimens may probably be referred to this species; 
they differ, however, from the other examples in having the lateral margins of the 
rostrum rather strongly convergent :— 


as Port Blair, Andaman Is. J. Wood-Mason. Ig, 26 mm. 
4520 
10 


Persian Gulf, 53 fms., ‘Tnvestigator.’ I? ,3I mm. 
26° 20’ N., 53° 54’ E. 


The following specimens from the British Museum collection have also been 
examined :— 
Holothuria Bank, China Seas, P. Basset Smith. Id, 43 mm. 


(Pocock, 1893, sub S. affinis.) 
The anterior bifurcated part of the median carina of the carapace is obsolete. 


Kowlson Bay, Hongkong. P. Basset Smith. Id, 44mm. 
(Pocock, 1893, sub S. affinis.) 
Port Darwin, N. Australia. ‘ Alert.’ Id, 61 mm. 


(Miers, 1884, sub S. nepa.) 


Singapore. S. F. Flower. Id, 92 mm. 


The var. perpensa is distributed over an area extending from Hongkong and N, 
Australia to the Persian Gulf, entirely replacing the typical form in the western part 
of its range. I have seen no specimens from Japan or the northern coasts of China. 


25. Squilla interrupta, Wood-Mason, MS. 
Plate V, figs. 60—62. > 


? 1852. Squilla oratoria, Dana, U.S. Explor. Exped., Crust., I, p. 621. 
1893. Squilla affinis, Henderson, Trans. Linn. Soc., Zool. (2), V. p. 453 (Partim). 
Squilla interrupta, Wood-Mason, MS. (partim). 
1908. Squilla interrupta, Lloyd, Rec. Ind. Mus., II, p. 32 (partim; sine desc.). 
1911. Squilla interrupta, Kemp, Rec. Ind. Mus., VI, p. 98. 

This species differs from S. ovatoria,s.s. in the following constant characteristics :— 

1. The median carina of the carapace is interrupted at the base of the anterior 
bifurcation as in 5. oratorıa var. perpensa (fig. 60). % 

2. The rostrum is subquadrate ; its anterior margin is truncate (sometimes even a 
trifle concave) and its lateral margins are only very slightly convergent ante- 
riorly (fig. 60). 

3. The eyes are distinctly smaller than in S. oratoria or its variety, specimens of 
similar size being compared. In the smallest examples in the collection 
the corneal index is about 5:0, in the largest about 6:1, while in individuals 
between 80 and 90 mm. in length, the figure usually falls between 5'2 and 5°5. 

4. The carpus of the raptorial claw is constantly provided with two, and only two, 
dorsal tubercles (fig. 61). 

5. In the bifurcate process from the base of the uropod the margin of the longer 
spine anterior to the external lobe is never concave; it is occasionally feebly 
sinuous, but in nearly every instance is definitely convex (fig. 62). 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 73 


Squilla interrupta differs from the var. perpensa in regard to characters 3, 4 and 5 
(supra). The anterior margin of the ophthalmic somite is usually definitely emarginate, 
never pointed. ‘The lateral processes of the sixth and seventh thoracic somites gener- 
ally bear a close resemblance to those of S. oratoria var. perpensa, and the same number 
of spines on the abdominal somites and of marginal denticles on the telson are found. 
The outer margin of the dactylus of the raptorial claw is more strongly sinuous than 
in either of the preceding forms, but hardly so much so as in S. nepa. 

The colour of spirit specimens does not appear to be characteristic. 

This is one of the commonest species of Squilla in Indian waters. Except that 
in three examples there are exceedingly feeble indications of a third tubercle on the 
dorsal aspect of the raptorial carpus (on one side only), the one hundred and twenty- 


two specimens examined are absolutely constant in exhibiting the characters noted 
above. 


The Indian Museum contains the following series :— 


PES Hongkong. Hongkong Museum. Id, £2,100 and 103 mm. 
#9 Hongkong. G. Dennys. 43,42, 74—102 mm. 
we Singapore ? Raffles Museuin. 34, 84—103 mm. 
> Camorta, Nicobars. F. Stoliczka. 19,81 mm. 
= Akyab, Arakan Coast. W. Dodgson. 12,58 mm. 
7982—91 . § Commissioners of a's 
Be Sandheads, Hughli Delta. À dut Plone | 58,52 ,44—109 mm. TYPES. 
705 
4 
3065—6 
ms Sandheads, Hughli Delta. Commissioners of ( 38,1092, 63—106 mm. 
: H.M. Pilot Brigs. ) 
1284—6 
9 
es Ghapa Natta, Mutlah R., near S. B. Nath. 1, 84 mm. 
Calcutta. 
=°=2 Mouth of R. Hughli. J. Munro. 73,82, 48—105 mm. 
“° Off Mutlah Light, Hughli R. G. M. Giles. 19,9 mm. 
~*~ Puri, Orissa Coast. F. H. Gravely. 19,19,70 and 72 mm. 
Far Off Ganjam Coast, ‘ Investigator.’ 43, 56—85 mm. 
Madras, 7 fms., 
>= Off Vizagapatam Coast, ‘Investigator.’ Id, 61 mm. 
Madras, 74—9 fms. 
7557 
| an. Bombay Nat. Hist. Soc. 49,172, 47—119 mm. 
com 
TE Panvel Creek, Bombay J. Caunter. 43,52, 68—90 mm. 
won Karachi ? Karachi Museum. 1,19, 85 and 93 mm. 
| . 
5. - Karachi Museum | 
5 \ Arabian Sea. À Rs i 
sil | and A. O. Hume: | een 


a Persian Gulf. F. H. Townsend. 19,067 mm. 


74 M emoirs of the Indian Museum. [Vol. IV, 


7282-3 


10 Persian Gulf, 13 fms., ‘Investigator.’ 261 9 2 EH 02 m. 
29° 20’ N., 48° 47’ E. 
— Persian Gulf, 25 fms., ‘ Investigator.’ AS 22, WSO nam. 
28° 59 N., 50° 5 E. 
a Koweit, Persian Gulf. ‘ Investigator.’ EE 5 Say Malo 
7  (locality unknown.) S. MeLachlan. LOST: 


The following specimens from other sources have also been examined :— 


BRITISH MUSEUM. 


S. Fo: mosa. Formosa Museum. 13,00 mm. 
Singapore. Bedford and Lanchester. 29,08 and rot mm. 
Sunderbunds, Ganges Delta. F. Day. 23,42, 42-66 mm 


SARAWAK MUSEUM. 

Buntal and Burong Is., Borneo. 32, 37—73 mm. 
BomBAy NATURAL HISTORY SOCIETY. 

Bombay. 23, 80 and 120 mm. 


The range of S. snterrupta consequently extends from Formosa and Hongkong 
westwards to the Persian Gulf. 


26. Squilla wood-masoni, Kemp. 
Plate V, figs. 63—65. 


1880. Squilla nepa, Miers, Ann. Mag. Nat. Hist. (5), V, p. 25 (partum). 

1893. Squilla afinis, Henderson, Trans. Linn. Soc. Zool. (2), V, p 453 (partem). 
Squilla polita, Wood-Mason, MS. (nec Bigelow). 

1908. Squilla polita, Lloyd, Rec. Ind. Mus., II, p. 33 (sine desc.). 

1911. Squilla wood-masont, Kemp, Rec. Ind. Mus., VI, p. 99. 


Squilla wood-masom may be distinguished from the three preceding forms by the 
use of the following characters :— 


1. The dorsal surface of the carapace and abdomen is smooth, highly polished and 
without trace of punctuation. 

The carapace is shorter and broader ; its breadth behind the antero-lateral 
angles is almost invariably more than one-half of its length in the median line, 
including the rostrum (fig. 63). | 

The anterior bifurcate portion of the median carina of the carapace is entirely 
obsolete, though its position is often marked by dark lines of pigment. 

4. The eyes are large; the corneal index varies from 3:0 in young examples to 4:2 
in large specimens. In examples between 80 and 90 mm. in length it falls 
between 3°4 and 3:9 The anterior margin of the ophthalmic somite is strongly 
convex and is produced to a small but quite distinct median point. 

5. The carpus of the raptorial claw somewhat resembles that of S. oratoria, s.s. It 
is armed dorsally with two or three tubercles, and in addition the extreme 
distal edge is usually produced to a blunt tooth-like process (fig. 64). 

6. The dactylus of the same limb is strongly sinuate externally. The whole seg- 
ment is of a more clumsy build than in the preceding forms ; the teeth on the 


D 


Oo 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 75 


inner margin are shorter and stouter and the incisions between them are less 
deeply cut. 

7. The margin of the bifurcate process of the basal segment of the uropod, anterior 
to the lobe on the outer face of the longer spine, is sharply concave as in 
S. oratoria ; thelobe itself is, however, much smaller and less conspicuous than 
in that apecies or than in S. interrupta (fig. 65). 


In this species, also, the cervical groove of the carapace is exceptionally deeply- 
cut and the spines at the antero-lateral angles are (in adults) unusually small. The 
rostrum is subquadrate with its lateral margins slightly convergent anteriorly. The 
lateral processes of the thoracic somite are decidedly shorter than in the two preceding 
species, especially the anterior one on the fifth somite. The following abdominal 
carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian Fe a be 5, 6. 
Intermediate sh oe PE Bo Abe By Oc 
Lateral .. ae ee a Dy Be fhe (6 
Marginal Bs a ae T2 NO NA AS: 


The post-anal carina on the ventral surface of the telson is, as a rule, longer than 
in the preceding species and on the margin there are two or three submedian denticles, 
eight to eleven intermediate and one lateral. 

In a very young example from Muscat! (345 mm.) the lateral margins of the 
rostrum are more strongly convergent anteriorly than in adults and the tubercles on 
the raptorial carpus are less distinct. In this example, moreover, the carapace is not 
so wide as in adults, the breadth being a trifle less than one-half the length including 
the rostrum. The great breadth of the carapace is, in Indian waters, a very marked 
character of this species and is quite constant in all examples except the two immature 
individuals mentioned above. Two large specimens from Zanzibar are, however, wholly 
abnormal in this respect, for, in both, the breadth of the carapace, as in the preceding 
species, is very clearly less than half the length including the rostrum. In other respects 
they agree closely with the remaining specimens, and there is little doubt that they 
may safely be referred to S. wood-masoni. Should further collections show that the 
character is constant in specimens found on the E. African Coast the creation of a 
distinct subspecies will, I believe, be justified. 

Of this species fifty-four specimens have been examined. The following are in the 
Indian Museum :— | 


MS Hongkong. Hongkong Museum. 39, 60—131 mm. 
BB Port Jackson, N. S. Wales. Australian Museum. 19,144 mm. 
a Pondicherry. (Purchased.) 19, 107 mm. : 
3% Madras. (Purchased.) 53,32, 89—I09 mm. 
: TYPES: 


! This specimen was identified prior to the examination of a series of 5: massavensis. It is not 
improbable that this example along with that from Aden should more properly be referred to that 
species. 


76 Memoirs of the Indian Museum. [Vol. IV, 


3078 | 
5 
3087 N Madras Museum. } 


: Madras, 83, 65—99 mm. 


ee = | (purchased.) | 
5 Off Madras Coast, 23 fins. ‘ Investigator.’ 19, IoI mm. 
14°4'57” N., 80°20’50” Is. 
= Orissa Coast, 7—8 fins. ‘Investigator.’ 13, 98 mm. 
7296—9 { N. Annandale, | 
“| Puri, Orissa Coast. F. H. Gravely and | 93,132, 50-90 mm. 
10 j J. Caunter. 
© Sandheads, Ganges Delta W. M. Daly. 19,69 mm. 
Ces Gite, F. H. Townsend. 1,52 mm. 


10 
The following specimens from the British Museum collection have also been ex- 
amined :— 


Singapore. S. F. Flower. Id,77 mm. 

Rameswaram I., G. of Manaar. J. R. Henderson. 14, 89 min. 
(Henderson, 1893, sub S. affinis.) 

Madras. F. Day. IS , 106 mm. 
(Henderson, 1893, sub S. affinis.) 

Muscat.! J. B. Miles. Id, 345 mm. 

Aden, 5—8 fins.! Capt. Shotland. Id, 41 mm. 

Zanzibar. Col. Playfair. 13,12,75 and 105 mm. 


(Miers, 1880 sub S. nepa.) 


According to the above records this species extends over an area ranging from 
Hongkong and the Australian Coast to the Persian Gulf, Aden and Zanzibar. 


27. Squilla massavensis, Kossmann. 
1880. Squilla massavensis, Kossmann, Zool. Ergebn. Roth. Meer., III, p. gg. 
1906. Squilla massavensis, Nobili, Ann. Sci. Nat., Zool. (9), IV, p. 340. 
1010. Squilla massavensis, Balss, Denk d. math -naturwiss. Klasse d. Kais. Akad. Wiss., Wien, 
LXXXV, p. II, text-figs. a, c, d. 

I refer to this species nine specimens obtained by Mr. F. H. Townsend in the 
Gulf of Oman and the Persian Gulf. The largest is only 108 mm. in total length and 
is consequently much smaller than Kossmann’s type (140 mm.). 

Squilia massavensis is evidently an extremely close ally of S. wood-masom and 
like that species may be distinguished from S. oratoria and S. interrupta by the dac- 
tylus of the raptorial claw (see Balss, 1910, text-fig. 3), which is strongly sinuous 
externally and provided with short teeth, while other less conspicuous characters are 
the size of the cornea—the corneal index is about the same as in S. wood-masoni— 
the deep-cut cervical groove and the extreme reduction in size of the lobe on the 
outer aspect of the longer of the two spines forming the basal process of the uropod. 

From S. wood-masoni the species may be distinguished by the following char- 
actes 


1. The dorsal surfaces of the carapace and abdomen do not present a polished 
appearance. 


! The specimen was identified prior to the examination of a series of S. massavensis. Itis not im- 
probable that it should more properly be referred to that species. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. ZN, 


2. The anterior breadth of the carapace is less than half the median length, 

including the rostrum. 

3. The rostrum is much more strongly narrowed anteriorly and is proportionately 

a trifle longer. 

4. The anterior lateral process of the sixth thoracic somite is usually a little 

longer and narrower. 

5. The telson shows important distinctions. It has the same series of impressed 

pits as are found in other species of the group; but, in addition, is provided 
with rows of tubercles on either side of the middle line converging to a 
point beneath the apex of the spine which terminates the median carina. 
This feature is possessed by no single one of its allies. The tubercles are 
only developed in well-grown specimens and cannot be detected in the two 
smallest examples which measure 45 and 49 mm. The single row which can 
be seen in a specimen 65 mm. in length is a prominent feature of all the 
other larger examples. In very large individuals, such as those described 
by Kossmann and Nobili and figured by Balss (1910, text-fig. 4), three rows 
of tubercles are found, and in these the difference between S. massavensis 
and its allies is very pronounced. In the largest specimen from the Persian 
Gulf the row nearest the median carina is well developed, while the other 
two are quite rudimentary. 

There are two or three sharp tubercles on the dorsal aspect of the carpus and the 
number of spines on the abdominal somites is the same as in S. wood-masoni. In 
the specimens examined there are two to four (usually three) submedian denticles, 
seven to nine intermediate and one lateral. Nobili mentions that in the specimen 
which he examined there are six intermediate and two (!) laterals. In the largest 
male the carinae of the marginal telson teeth and the bases of the intermediate 
denticles are noticeably inflated, more so than in any other species of the same group. 

It must be confessed that had the two smallest individuals (under 50 mm. in 
length) been found alone, and not taken in company with larger examples, the greatest 
difficulty would have been experienced in their identification. From S. wood-masoni 
of similar size, they are distinguished only by the rostrum, which is more strongly 
narrowed anteriorly, and by the different proportions of the carapace. To young 
S. oratoria var. perpensa the resemblance is even closer ; the form of the rostrum and 
the total absence of the anterior bifurcation of the median carina of the carapace! 
afford almost the only means of distinction, for in immature S. massavensis the char- 
acteristics of the raptorial propodus and carpus are not developed and the lobe on 
the larger spine of the basal process of the uropod is conspicuous. 

According to the authors who have previously described this species, the anterior 
bifurcation of the median carina of the carapace is present, but this feature is not 
shown in any of the specimens I have examined. Nobili (1906) considers that the 
two small tubercles found on the last five abdominal somites between the interme- 


| The presence of the anterior part of this bifurcation is not absolutely constant in S. oratoria var. 


perpensa. 


78 Memoirs of the Indian Museum. [Vows iva 


diate and lateral carinae constitute a valuable character of the species. They are, 
indeed, rather exceptional in their development in large individuals, but more or less 
evident traces of them can usually be seen in large examples of the allied forms. 
Balss (1910) in comparing this species with S. nepa notes a distinction in the carinae of 
the carapace, those of the intermediate pair being confluent anteriorly with the 
laterals in S. nepa, whereas they end abruptly in S. massavensis. The condition of 
these carinae in S. oratoria and other large-eyed species of the group is, however, 
identical with that found in S. massavensis, while the confluence of the two carinae 
in S. nepa is not a constant character. 

The colouring of spirit material is not striking; but there are large dark grey 
patches on the second and fifth abdominal somites—a feature not found in S. wood- 
masont or in S. oratoria vat. perbensa. 

For the opportunity of examining a series of this scarce species I am indebted 
to the Secretary of the Bombay Natural History Society, who has kindly forwarded 
to me eight specimens, 457 108 mm. in length obtained by Mr. F. H. Townsend in 
the Gulf of Oman, lat. 25° 21’ N., long. 58° 07’ E., at a depth of 175 Du The 
only other example I have seen is registered as follows :— 

a Persian Gulf. F. H. Townsend. [6 , 52 mm. 

Prior to an examination of the other specimens I had identified this individual as S. wood- 
masoni, while remarking on the difference in the proportions of the carapace and rostrum. On 
re-examination I find slight but distinct traces of tubercles on the telson. ‘The two small 


specimens in the British Museum collection, recorded on p.76 as S. wood-masoni, should 
also, in all probability, be referred to this species. 


The precise locality of the type specimen, which was obtained in the Red Sea, is 
not given. Nobili has recorded the species from Massouah and Djibouti, while the 
specimen determined by Balss was found at Suez. 


28. Squilla stridulans, Wood-Mason. 
Plate V, fig. 66. 
1894. Squilla stridulans, Wood-Mason, Ann. Mag. Nat. Hist. (6), XIII, p. 400. 
1895. Squilla stridulans, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 5, pl. ii, fig. 3; 
pl. iii, fig. 1. 
Squilla Stridulans may be readily distinguished from the six preceding forms by 
the following characters :— 
1. The lateral process of the fifth thoracic somite can scarcely be termed bilobed : 
it consists of two sharp and subequal spines (fig. 66). 
2. The lateral carinae of the first five abdominal somites are sharply bicarinate 
throughout their length. 


For the rest the principal characters of the species are as follows -— 

The carapace is narrow anteriorly and its breadth behind the antero Jateral angles 
is less than half the median length excluding the rostrum. The median carina is 
sharply defined throughout its course and is bifurcated in front for more than one-third 
its length anterior to the cervical groove, the junction of the two carinae being quite 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 79 


close to the small mid-dorsal pit. ‘The lateral margin is obtusely angled in front of 
the rounded postero-lateral corners as in S. foveolata. Both carapace and abdomen 
are deeply and rather coarsely punctured. 

The lateral margins of the rostrum are upturned and converge to a truncate apex. 
The corneal portion of the eyes is greatly expanded and is set obliquely on the stalk. 
The corneal index varies from 3°7 to 4-2 and in one specimen, which is quite excep- 
tional among Indian examples, is as much as 4:9. The anterior margin of the 
ophthalmic somite is rounded. 

The outer inferior margin of the merus of the raptorial claw terminates in a rounded 
angle, and is not toothed. The dorsal carina of the carpus is entire or with a very 
obscure tubercle and ends abruptly before reaching the anterior margin. ‘The outer 
margin of the dactylus is evenly convex throughout its length and the teeth on its inner 
margin are long and slender. 

The anterior of the two spines on the margin of the fifth thoracic somite is 
situated at a lower level than the posterior. The lateral lobes of the sixth somite are 
about equal in length and breadth ; the posterior lobe is sharply pointed, the anterior 
more or less obliquely truncate. Both processes of the seventh somite are acute, the 
anterior being both shorter and narrower than the posterior. 

As already mentioned the lateral carinae of the first five abdominal somites are 
sharply bicarinate. This unusual feature is very distinct in adults and the narrow 
area between the two carinulae is coarsely pitted; in very young individuals the 
bicarination is inconspicuous. The following abdominal carinae end in spines :— 


Carinae. Abdominal somites. 

Submedian a oe a 5,6. 

Intermediate LE BE Be Bo Aly Sy (Ob 

Lateral ie ae set palin 2 RS AGG 0: rn 
Marginal SE en de 1203 4a Ss 


The marginal teeth of the telson are unusually long and the denticles are excep- 
tionally numerous. Of the latter there are five or six submedians, twelve to sixteen 
intermediate,’ and one lateral. 

In the bifurcate process from the base of the uropods, the lobe on the outer margin 
of the longer spine is small or obsolete in adults, but is usually quite distinct in-very 
small specimens. 

In very young examples, also, the median projection on the posterior margin of the 
carapace is exceptionally well developed and the rostrum, as noticed in small indivi- 
duals of some of the preceding species, is more strongly narrowed anteriorly. 


A single very large specimen from Japan, the only individual known from any 
locality outside the Bay of Bengal, differs from the Indian examples in the ms 
particulars :— 

1. The carapace is distinctly broader ; its breadth at de antero-lateral angles 

is es half the median leach, The median carina of the carapace iS 


! In one specimen, ‘Given appears to be abnormal in this Fee, there are only ten intermediate 
denticles. 


80 Memoirs of the Indian Museum. Wor. nV; 


bifurcated anteriorly for only two-sevenths of its length in front of the cervi- 
cal groove ; the distance between the small mid-dorsal pit and the base of 
the fork is more than half the length of the bifurcated part and is far greater 
than in typical examples. 
2. The rostrum is proportionately broader and its apex is more rounded anteriorly. 
3. The corneal and peduncular axes of the eye seem to be a trifle less oblique and 
the corneal index, as might be expected in so large a specimen, is greater : 
5/3- 
In other respects, however, the specimen agrees very closely with typical examples. 
As regards colour, spirit specimens are as a rule darker than those of the allied 
species but the general plan is closely similar. The longitudinal grooves of the 
carapace, the lateral margins of the rostrum, and the posterior margins of the last 
three thoracic and first four abdominal somites are rather broadly marked with black 
pigment. ‘The median carina of the carapace is also defined by similar pigment, but 
much less distinctly. The abdominal somites are rather thickly strewn with black 
chromatophores, which in most cases form decided patches along the inner aspect of 
each lateral carina. These patches are specially well-marked on the fifth somite and 
there is also in most cases, more particularly in young specimens, a pair of dark spots 
in the middle of the second somite. ‘The pits and carinae on the dorsal surface of the 
telson are defined by black pigment and the uropods are faintly suffused with the same 
colour. 
I have been unable to find any structural character which will account for the 
name which Wood-Mason chose for this species (see p. IQ). 


There are thirty-five specimens of S. stridulans in the Indian Museum :— 


Ee Nellore Coast, Madras; 80o—ı1o fms., ‘Investigator.’ 43,52, 69—100 mm. 
TA TONI N (HOS 963) BO? 18, 
== Masulipatam Coast, Madras; 95 fms.,  ‘Investigator.” 62, 41—92 mm. 
15° 56° 25” N., 81° 26’ 10” E. 
Ze Masulipatam Coast, Madras; 240 fms., “Investigator.’ 29, 48 and 74 mm. 
i 15° 56’ 50” N., 81° 30’ 30” E. 
IT? Orissa Coast; 68 fms. ‘ Investigator.’ 19,19,52 and 97 mm. 
TYPES. 
MS Orissa Coast; 68 fms. ‘ Investigator.’ 30,122, 38—86 mm. 
7685 Misaki, Japan. A. Owston. Tg, 144 mm. 


10 


Only the above specimens are known. 


29, Squilla investigatoris, Lloyd. 
Plate VI, figs. 67, 68. 


1907. Squilla investigatoris, Lloyd, Rec. Ind. Mus., I, p. Io. 
1908. Squallu investigatoris, Lloyd, Rec. Ind. Mus., II, p. 29, pls. ii and iii. 
This species is readily distinguished from its allies by the unusually large number 
of teeth on the dactylus of the raptorial claw. Tn other respects it bears a close resem- 
blance to the species of the preceding group. 


IgI3.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacifie Region. 81 


The dorsal surface is smooth, highly polished and without trace of punctuation. 
The carapace measured behind the antero-lateral angles is about half its length includ- 
ing the rostrum. The median carina is distinct posteriorly, but is almost or entirely 
obsolete in front of the small mid-dorsal pit. Frequently two feeble ridges are visible 
near the frontal margin—the vestiges of the anterior bifurcation. Intermediate and 
lateral carinae are distinct. The antero-lateral angles are produced as short spines 
which fail to reach the level of the rostral base. The rostrum is subquadrate, its basal 
breadth being equal to its median length. The lateral margins are scarcely upturned 
and converge very slightly to a broad rounded apex. 

The cornea of the eyes is greatly expanded ; its breadth is about one-third the 
median length of the carapace. The corneal and peduncular axes are very slightly 
oblique. The anterior margin of the ophthalmic somite is sometimes straight or 
slightly convex, but in most cases is produced to three small but distinct points. The 
mandibular palp is three-segmented. 

The outer inferior margin of the merus of the raptorial claw is bluntly angled at 
its distal end. The dorsal carina of the carpus is furnished with from two to four sharp 
teeth. The dactylus bears thirteen' to eighteen teeth including the terminal one, this 
number far exceeding that found in any other species of the genus. The outer margin 
is slightly sinuous and is obtusely angled behind. 

The last three thoracic somites bear the usual submedian and intermediate carinae. 
The lateral margin of the fifth thoracic somite is bilobed ; the anterior lobe consists 
of a sharp spine directed obliquely forwards ; the posterior lobe is straight, sharply- 
acute, and only about half its length. Both lobes of the sixth somite are pointed; 
the anterior is a little shorter than the posterior and is only about half its width. The 
posterior lobe on the seventh somite is subacute ; the anterior is very small and in 
some cases almost obsolete. 

There are, as usual, eight longitudinal carinae on the first five abdominal somites 
and six on the sixth. The following carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian ae Sie Æ (5) 6. 
Intermediate iu Me a Ro tly By ©. 
Lateral =e He T2) 40508 
Marginal st a Br 12,9, 4,5. 


The telson closely resembles that found in the six preceding species. The median 
carina is notched near the base and the intermediate marginal teeth are sometimes 
slightly inturned. There are three or four (rarely five’) submedian denticles, eight 
to eleven intermediate and one lateral. The small praelateral lobe is conspicuous. 

The bifurcate process of the uropods is serrate on its internal margin. The usual 
lobe on the outer edge of the longer spine is distinct. 


The dorsal surfaces of the carapace, antennules, antennae, raptorial merus, abdo- 


' One specimen has only ten teeth on the left dactylus. 
? In several cases the submedian denticles are abnormally developed; in one specimen eight were 
observed. 


82 Memoirs of the Indian Museum. [WIG ION, 


men and telson are in most cases thickly strewn with small black chromatophores, 
which give the animal a decidedly dusky appearance. The chromatophores are aggre- 
gated on the longitudinal carinae and on the posterior margins of the abdominal and 
last three thoracic somites. ‘The telson is often very densely pigmented all over except 
in the vicinity of the marginal denticles. On the uropods the endopod and basal seg- 
ment of the exopod are suffused with black at their distal ends and the proximal part 
of the ultimate exopodal segment is tinged with the same colour. 


In the Indian Museum are nineteen specimens registered as follows :— 


a Off the S. Coast of Arabia; 110 fms., ‘Investigator.’ 108,89,173 84 mn. 
LORS ON Bae? Gay 307 98), IVPES: 
= Persian Gulf. F. H. Townsend. BO 7m 


30, Squilla supplex, Wood-Mason. 
JDE WIL, de: CO), 
1875. Squilla supplex, Wood-Mason, Proc. As. Soc. Bengal, p. 232, reprinted in Ann. Mag. Nat. 
Hist. (4), XVII, p. 263 (1876). 
1880. Squilla supplex, Miers, Ann. Mag. Nat. Hist. (5), V, p. 20. 
1894. Squilla supplex, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 511. 
1895. Squalla supplex, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 4, pl. ii, fig. 2; pl. iii, 
fig. 2. 


This and the two succeeding forms may easily be recognized from all other species 
of the genus by the increased number of carinae (more than eight) on the abdominal 
somites. 

In Squilla supplex the carapace and abdomen show no trace of punctuation. The 
carapace, measured behind the antero-lateral angles, is about half its median length 
excluding the rostrum. The median carina is sharp and distinct except for its anterior 
bifurcation’ which is obsolete. ‘The intermediate and lateral carinae are indistinct in 
the small type specimen but quite clear in the larger examples. The lateral margins 
are not angled in front of the posterior corners, but antero-laterally are produced as 
sharp spines, which, however, fail to reach as far forwards as the level of the rostral 
base. The breadth of the rostrum is a little greater than its median length and its 
upturned lateral margins converge gradually to the broadly-rounded anterior margin. 
Dorsally the rostrum is smooth except for an indistinct median carina in its anterior 
half. 

The eyes are short. The breadth of the cornea is about equal to the total length 
and the corneal and peduncular axes are set at right angles to one another. The an- 
tennular peduncle is shorter than the carapace. There is a three-segmented mandi- 
bular palp. 

The outer inferior margin of the merus of the raptorial claw is bluntly angled 
distally and the carina on the dorsal aspect of the carpus is entire, but terminates 


! Part of a haul of over 500 specimens. 
» In Wood-Mason’s figure it is much too clearly shown. 


1013 | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 83 


abruptly before reaching the anterior edge. The dactylus is provided with five teeth 
including the terminal one and its outer margin is evenly convex and obscurely lobed 
at its proximal end. 

The last three thoracic somites possess the usual submedian and intermediate 
carinae and between these, in the anterior half of the somite, each bears a short and 
rather obscure keel directed outwards and forwards. The fifth somite is bilobed 
laterally ; the anterior lobe consists of a long and sharp antrorse spine, the posterior is 
short and bluntly rounded. The margins of the two following somites are not 
bilobed, but are obliquely truncate (more squarely than in S. /aevis or S. hieroglyphica) 
and are rounded posteriorly. 

The first five abdominal somites, in addition to the usual eight longitudinal carinae, 
possess also a sharp and distinct median keel. This carina never terminates in a 
spine :— 


Carinae. Abdominal somites. 
Submedian a a a 5, 6. 
Intermediate Fe + oe 5, 6. 
Lateral .. a de OO) S545 By Oo 
Marginal ar ze Ar 12391415. 0: 


The telson is rather longer than broad, and is sharply distinguished from the ma- 
jority of the preceding species by the prominent carination of its dorsal surface. The 
median carina is well developed and terminates in a sharp spine overhanging a small 
tubercle. On either side there is a deep longitudinal groove formed by a number of 
confluent pits and beyond this there is a well-marked longitudinal carina continued 
with or without interruption to the apex of the submedian spines. Each of the usual 
oblique rows of pits is replaced by a rather deep groove and the surface between the 
grooves is raised, forming three or four short” but distinct oblique carinae on either 
side of the median ridge. On the edge of the telson the usual six teeth are well deve- 
loped and there is a prominent praelateral lobe. There are three or four submedian 
denticles, six or seven intermediate and one lateral. The post-anal crest is absent. 

In the bifurcate process from the base of the uropod, the outer margin of the longer 
spine bears the usual lobe near the distal end of its proximal half ; the margin anterior 
to this lobe is straight or feebly convex. The inner margin of the process is slightly 
serrate. 

The secondary sexual distinctions appear to be rather pronounced. In the single 
large male the submedian carinae of the fifth abdominal somite, all the carinae of the 
last somite and the median and marginal carinae of the telson, are conspicuously 
swollen (fig. 69) and the propodus of the raptorial claw is strongly dilated at its distal 
end (fig. 69). 


The large male specimen, examined shortly after capture, exhibited a considerable 
amount of lemon yellow colouration on the abdominal somites, telson and raptorial 


1 In the large male specimen only. 
? In the large male these carinae are extremely short, little more than rounded tubercles, and the 
grooves are less conspicuous (fig. 69). 


84 Memoirs of the Indian Museum. Noces 


claws ; but as the specimen had sojourned, though evidently for a brief period only, 
in the stomach of a fish, the pigmentation was perhaps somewhat modified. The black 
pigment, which appears to be fairly permanent in alcohol, defines all the carinae of 
the carapace and of the abdomen as far as the fourth somite. There are also dark 
median transverse patches on the free thoracic and first two abdominal somites, ex- 
tending laterally as far as the impressed line between the submedian and sublateral 
carinae. The inner uropod and spines of the bifurcate process are tipped with black 
and the terminal segment of the outer uropod is suffused with the same colour. 


Only three examples of Squilla supplex are known ; all are preserved in the Indian 
Museum. 


> Bombay. F. Stoliczka. ea NDR, 
7426 i : 

(28 Bombay. Bombay Nat. Hist. Soc. 19,73 mm. 

31 Tuticorin, S. India. J. Hornell. IS: 


10 


The last specimen was found in the stomach of Polynemus tetradactylus, Shaw. 


31. Squilla costata, De Haan. 
Plate VI, figs. 70—72. 

1884? Squilla costata, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. 5. 
1849. Squilla costata, De Haan, vbid., text, p. 223. 
1880. Squilla costata, Miers, Aun. Mag. Nat. Hist. (5), V, p. 21. 
1894. Squilla costata, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 511. 
1903. Chloridella costata, Rathbun, Proc. U. S. Nat. Mus., XXVI, p. 55. 
1910. Squilla costata, Fukuda, Annot. Zool. Tapon., VII, p. 151. 

In this and in the succeeding form the lateral margins of the sixth and seventh 
thoracic somites are bilobed as in the S. guinquedentata and S. nepa groups. By this 
feature they are readily separated from S. supplex, from which, as from all other species, 
they are distinguished at a glance by the profuse carination of the abdominal somites. 

The principal characters of Squlla costata are as follows :— 

The carapace (fig. 70) is narrow and its breadth behind the antero-lateral angles 
is about two-fifths its median length including the rostrum. The median carina is 
distinct and is continuous in front of the well-marked cervical groove except for a gap 
at the base of the short anterior bifurcation. Posterior to the cervical groove the 
median carina is conspicuous and is bifurcate in its anterior two-thirds. The lateral and 
intermediate carinae are distinct. Between the median carina and the gastric groove, 
there are on each side numerous short carinules or tubercles. In the Burmese example 
(fig. 70) they are disposed obliquely on the area in front of the cervical groove, but 
behind it are situated transversely. In the Japanese specimen this feature is less 
evident. Other tubercles occur on the sides of the carapace and are most distinct in 
the posterior part. The antero-lateral angles are produced to strong spinous termina- 
tions that extend as far as or a little beyond the level of the rostral base. The 
rostrum is about as long as wide; its upturned lateral margins are slightly convergent 
and meet in a broadly rounded apex ; on its dorsal surface it is finely rugose or with 
a few obscure tubercles and in the larger specimen there is a faint median carina. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 85 


The corneal portion of the eyes is set very obliquely on the stalk and its breadth, 
in the specimens examined, is about one quarter the median length of the carapace 
excluding the rostrum. The antennular peduncle is as long as the carapace (rostrum 
excluded). The mandibular palp is entirely missing. 

The outer inferior margin of the merus of the raptorial claw is rounded distally 
and the dorsal carina of the carpus terminates bluntly before reaching the distal edge 
(fig. 71). The dactylus bears six or seven' teeth including the terminal one and its 
outer margin is convex and very obscurely notched at the base. 

The lateral margins of the fifth, sixth and seventh thoracic somites are bilobed 
(fig. 70). The anterior lobe on the fifth consists of a sharp antrorse spine ; the other 
lobes are rounded. The sculpture on the somites is shown in fig. 70. In addition to 
the intermediate and submedian carinae there are on the sixth and seventh somites 
four longitudinal carinae between the submedians, while on the eighth somite there are 
three. The whole surface is covered with numerous short ridges that are sometimes 
confluent with one another and so form a coarse mesh-like reticulation. 

On the abdominal somites the usual four pairs of carinae may be distinguished 
and, in addition, between the submedians there are three complete carinae and occa- 
sionally traces of others. Outside the submedians there are one or more additional 
carinae which are sometimes incomplete. 

The whole surface external to the submedian carinae is beset with short irregular 
tubercles and ridges (fig. 72). The lateral carinae of the first five somites are bicarinate ? 
and traces of similar sculpture are sometimes found on the intermediates. None of 
the median carinae end in spines :— 


Carinae. Abdominal somites. 
Submedian Be : ae (4) 5, 6. 
Intermediate ss oe 2 E(E)L2 245 0: 
Lateral Er En (E20 Sy ela Rs ©: 
Marginal Bis sig a T2: 3, A Ge 


The telson bears numerous carinae and tubercles disposed as shown in fig. 72. 
The median carina terminates in a long and slender spine. The marginal teeth are 
longer than is customary and there may or may not be a denticle on the edge in front 
of the lateral. There are three or four submedian denticles, six to eight intermediate 
and one lateral. 

The inner and longer spine of the ventral process of the uropods bears a sharply- 
acute lobe on its edge and the margin anterior to this lobe is concave. The inner 
margin of the process is serrate. 

Of the two specimens examined, the smaller example from the Burmese coast 
differs in several details from the larger individual which was sent me from Japan— 
the type locality. It is possible that the differences are due merely to age or to normal 
variation, but the Indian specimen may represent a distinct race. 


1 The larger specimen bears six teeth—the customary number; the smaller example (fig. 71) 
possesses only one raptorial claw which is furnished with seven. 
* Not shown in fig. 72. 


86 


Memoirs of the Indian Museum. 


Mora 


The principal distinctions between the two specimens are as follow :— 


Burmese Coast. 

1. The median carina of the rostrum is very 
indistinct. 

2. There are seven teeth on the raptorial 
dactylus (one dactylus only examined). 

3. The anterior lateral lobe of the sixth thoracic 
somite is nearly as broad as the posterior. 

4. The submedian carinae of the fourth ab- 
dominal somite terminate in spines. 

5. ‘The submedian carinae of the sixth abdominal 
somite are bicarinate and there are three carinae 
between them. 

6. The carinae which terminate in the submedian 
teeth of the telson edge are proximally bifurcate. 


7. Thbereis no praelateral denticle on the margin 
of the telson. 

8. On the external edge of the basal segment 
of the outer uropod there are ten movable spines. 


Japanese Coast. 
I. The median carina of the rostrum is distinct. 


2. There are six teeth on the raptorial dactylus. 


3. The anterior lobe of the sixth thoracic somite 
is very much narrower than the posterior. 

4. The submedian carinae of the fourth abdo- 
minal somite do not terminate in spines. 

5. The submedian carinae of the sixth abdo- 
minal somite are entire and there is only a single 
carina between them. 

6. The carine which terminate in the sub- 
median teeth of the telson edge are not bifurcated 
proximally. 

7. There is a very conspicuous praelateral den- 
ticle on the margin of the telson. 

8. On the external edge of the basal segment of 
the outer uropod there are seven movable spines. 


The specimens examined do not show any distinctive colouration. 


We are indebted to Mr. Alan Owston for the specimen from the Japanese coast :— 


S044 
10 


Okitsu, Suruga Gulf, Japan. 


A. Owston. Ig, 65 mm. 


For the opportunity of describing and figuring the only known Indian example 
of this scarce species I have to thank Mr. A. Patience who has kindly permitted me to 
examine the specimen obtained on the Burmese coast by Messrs. Simpson and Rudmose 


Brown. 


This individual is a male, 31°5 mm. in length. 


Miss Rathbun records a specimen 87 mm. in length. 

Squilla costata was originally described from Japan (De Haan) and, except for the 
specimen from the Burmese coast which is mentioned above, the few examples that 
have since been found were all obtained on the coasts of that country. The species is 
known from Wakanoura and Nagasaki (Rathbun) and from Misaki (Fukuda). 


32, 


Squilla multicarinata', White. 


Plate VI, figs. 73—76. 


1847. 
1848. 


1849. 
1880. 
1894. 
1903. 


Squilla multicarinata, White, List Crust. Brit. Mus., p. 84 (sine desc.) 
Squilla multicarinata, White, Proc. Zool. Soc., p. 144, pl. vi, figs. I, Ia. 
Squilla multicarinata, White, Ann. Mag. Nat. Hist. (2), IV, p. 381. 
Squilla multicarinata, Miers, Ann. Mag. Nat. Hist. (5), V, p. 20. 
Squilla multicarinata, Bigelow, Proc. U. S. Nat. Mus., XVII, p. 511. 
Squilla multicarinata, Nobili, Boll. Mus. Torino, XVIII, No. 455, p. 38. 


This species is allied to Squilla costata, but in addition to the possession of a three- 
segmented mandibular palp, is still more profusely carinate, the entire dorsal surface 


of the animal being closely ribbed. 


1 See addendum, p. 196. 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 87 


The details of the carination (figs. 73—76) are as follows :— 

On the carapace the median carina is distinct throughout its course and is bifurcated 
in front for half its length anterior to the cervical groove. Behind this groove there 
are 26 to 30 longitudinal carinae, of which 18 are sharper and rather more conspicuous 
than the rest. The carinae are least numerous in the extreme anterior part of the 
carapace, where there are only 16 in a transverse series. The rostrum bears 5 carinae ; 
two of these are quite close to, but distinct from, the lateral margin, and two, which 
only extend for less than half the entire length, occur at the base on either side 
of the well-developed median keel. There are two dorsal carinae on the basal 
segment of the antenna and one on the succeeding segment. On the outer surface of 
the merus of the raptorial claw there is a single long carina running forwards and 
upwards. - 

The fifth thoracic somite bears four #ansverse carinae and is the only part of the 
animal in which the carination is not longitudinal. Only the anterior carina extends 
across the middle line. Prior to reaching the latter point the posterior one turns 
obliquely forwards and, before arriving at the anterior margin, turns abruptly back to 
the hinder edge. The posterior half of this somite therefore bears, near the centre, 
four short carinae which are almost longitudinal. 

The last two thoracic and all the abdominal somites are covered with close-set 
longitudinal carinae, most of which terminate in spines on the posterior margin. In 
well-grown examples there are 28 on the sixth thoracic, 25 on the last thoracic and 41 
to 45 on the somites of the abdomen. The median, submedian, intermediate, lateral 
and marginal carinae are prominent and can easily be distinguished from the others. 
Of these the following end in spines :— 


Carinae. Abdominal somites. 
Median Bs a ae ZI AN 
Submedian Res ab aus i, Dy Bp Aig Bio On 
Intermediate a 53 TAT: 
Lateral AR M, af Bh Cs Sig Oe 
Marginal oe ae 50 234,510: 


There are ten or eleven carinae on either side of the median crest of the telson, a 
few of which are not continuous. With the exception of the two innermost pairs they 
run to the marginal teeth and to alternate denticles of the submedian and intermediate 
series ; their exact arrangement is, however, subject to much irregularity. On the 
underside of the telson there is a sharp post-anal crestand eight or nine other carinae on 
either side. There are also several sharp carinae on both upper and lower surfaces 
of the uropod and on the basal and proximal segments of the exopod. 


Apart from the disposition of the carinae, Sgwilla multicarinata differs from S. 
costata in the following features :— 

The carapace is broader than in S.costata and its anterior breadth is a little less 
than half its median length including the rostrum. The antero-lateral spines are 
considerably shorter. The eyes are more expanded and the cornea is set trans- 
versely on the stalk.. The mandibular palp is present and composed of three 


88 Memoirs of the Indian Museum. NO 18%, 


segments. The dactylus of the raptorial claw usually bears five! teeth including the 
apical one. 

The anterior marginal lobe of the fifth thoracic somite is rather broader at the 
base ; that of the sixth somite is truncate and the posterior lobe of the seventh somite 
is subacute and is more strongly produced than its fellow. The teeth on the margin of 
the telson are shorter and behind the lateral as in the Japanese example of S. costata 
there is a sharp and well-developed praelateral denticle; there are three submedian den- 
ticles, seven or eight (rarely nine or ten) intermediate and one lateral. The bifurcate 
process from the base of the uropods bears a small rounded lobe on the outer edge of 
the inner and longer spine. The inner edge of the process is provided with five to 
eight sharp spines. The basal segment of the uropod is only a little longer than 
the ultimate. 

The colour of well-preserved spirit specimens is characteristic. Dusky patches 
are usually visible on the dorsal surfaces of the second and fifth abdominal somites, and 
the distal end of the inner uropod and basal segment of the outer uropod are suffused with 
black. The ultimate segment of the outer uropod is entirely jet-black and in this respect 
differs conspicuously from that of any other species examined (fig. 76). 

There are nine specimens in the Indian Museum :— 


9806—13 
5 Hongkong. Hongkong Museum. 50,32 ,63--72 mm. 
ne Simpson and | 
re Burma, Xmas I. } 
i Rudmose Brown. 19,46 mm. 


Squilla multicarinata has also been recorded from Nagasaki Bay, Japan, from 
the Philippine Is. (White, Miers) and from Singapore (Nobili). 


33. Squilla raphidea, Fabricius. 
Plate ie 77e 


1758. Squilla arenaria marina, Seba, Thesaurus, III, p. 50, pl. xx, fig. 2. 

1798. Squilla raphidea, Fabricius, Ent. Syst. Suppl., p. 416. 

1818. Squilla mantis var. B. major, Lamarck, Hist. Nat. Anim. sans Vert., V, p. 187. 
1825. Squilla raphidea, Latreille, Encycl. Méthod., X, p. 471: atlas (sub S. mantis), pl. cccxxiv 
1837. Squilla raphidea, H. Milne-Edwards, Hist. Nat. Crust., II, p. 524. 

1844? Squilla harpax, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. ı. 
1845. Squilla raphidea, Berthold, Abhandl. Gess. Wiss. Göttingen, III, p. 29. 

1847. Squilla raphidea, White, List Crust. Brit. Mus., p. 84. 

1849. Squilla harpax, De Haan, in Siebold’s Fauna Japonica, Crust., text, p. 222. 

1880. Squilla raphidea, Miers, Aun. Mag. Nat. Hist. (5), V, p. 27. 

1882. Squilla raphidea, de Borre, C. R. Ent. Soc. Belge, (III), 20, p. cxi. 

1888. Squlla raphidea, de Man, Journ. Linn. Soc., XXII, p. 296. 

1892. Squilla raphidea, Thallwitz, Abhandl. kön. Mus. Dresden, no. 3, p. 55. 

1893. Sgunlla raphidea, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 453. 

1894. Squila raphidea, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 535. 


' Miers (1880 (a), p. 20) found seven teeth on the left-hand dactylus of a specimen from the 
Philippine Is. The additional teeth in this case were perhaps formed subsequent to an injury. 
* I am indebted to Mr. Patience for an opportunity of examining this specimen. 


IgI3.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 89 


1898. Squilla raphidea, de Man, Zool. Jahrb. Syst., X, p. 694. 

1901. Squilla raphidea, Lanchester, Proc. Zool. Soc., II, p. 553. 

1903. Chloridella raphidea, Rathbun, Proc. U.S. Nat. Mus., XXVI, p. 55. 

1903. Squilla raphidea, Nobili, Boll. Mus. ‘Torino, XVIII, No. 455, p. 38. 

1906. Squilla raphidea, Laanchester, Fascic. Malayenses, Zool. III, p. 133. 

1906. Squilla raphidea, Tattersall, Ceylon Pearl Oyster Rep., V, p. 166. 

1910, Squilla vaphidea, and var. africana, Balss, Abh. math.-phys. Klasse K. Bayer. Akad. Wiss., 
Suppl., Bd. II, Abh. 2., p. 8, text figs. 2a—b. 


The dorsal surface of the carapace and abdomen is, as a rule, smooth, polished 
and impunctate, but in very old examples is finely rugose. The breadth of the cara- 
pace, measured behind the antero-lateral angles,is considerably less than half its median 
length. The postero-lateral angles are very narrowly rounded and the lateral margin, 
in the middle of its posterior half, projects as an acute lobe, the posterior edge form- 
ing a rectangle with the adjacent margin of the carapace. The antero-lateral angles 
end in small spines which in large specimens are a trifle upturned ; they fail to reach 
the level of the rostral base. In front of the cervical groove the median carina is 
distinct and is continued forwards beyond the small mid-dorsal pit ; the anterior 
bifurcated portion is however entirely absent. The posterior margin is deeply con- 
cave and the imperfectly calcified third and fourth thoracic somites are exposed. 


The rostrum is very variable in shape. It is smoothly elevated in the middle 
and its upturned lateral margins meet in an acute or narrowly-rounded apex. In some 
examples the rostrum reaches only to three-quarters the length of the antennular 
somite and the lateral margins are evenly concave, while in others it may reach to the 
distal edge of the somite and may have sinuous margins—concave on either side of the 
apex. 

The corneal and peduncular axes of the eye are at right angles. The cornea is 
sausage-shaped and its greatest breadth is fully one and a half times the length of the 
whole organ. The antennular peduncle is about as long as the median length of the 
carapace. 

The mandibular palp is composed of three segments. 

The merus of the raptorial claw is longer than in most species of Squilla ; when 
naturally flexed it projects backwards beyond the postero-lateral angle of the cara- 
pace. The outer inferior margin terminates distally in a blunt angle. The carina on 
the external aspect of the carpus is not sharp and that on its dorsal edge is entire and 
bends strongly inwards before reaching the distal margin. The propodus is charac- 
teristic. The row of fine pectinations which exists in most species along the margin 
opposed to the dactylus is replaced by a series of strong fixed spines ; seven or eight 
of these spines are longer than the others and alternate more or less regularly with them. 
As in allied species there are three strong movable spines on the inner basal edge of this 
margin. The dactylus is provided with eight (rarely nine) teeth including the terminal 
one. The outer margin is evenly convex in females and young males, but, in large 
examples of the latter sex, is excavate at the base, the distal border of the excavation 
being produced outwards as a strong rectangular process. 


90 Memoirs of the Indian Museum. Nor I, 


The last three thoracic somites (fig. 77) bear submedian and intermediate carinae; 
the former are often obscure or quite obsolete, while the latter are more prominent 
and in large specimens of both sexes terminate posteriorly in spines. On either side 
of the inferioı surface of the fifth somite there is a broad but thin downwardly-directed 
lobe, which is often anteriorly acute or spinous; from its base a deep and wide 
groove runs upwards dividing the somite transversely into two unequal portions. The 
lateral margins of the sixth somite are produced to an acute posterior point ; anteriorly 
a very short, broad and ill-defined process represents all that remains of the prominent 
lobe found in the species of the S. nepa group. The margins of the seventh somite 
resemble those of the sixth, but the anterior process is never represented by more than 
a slight angulation. 

There are eight longitudinal carinae on the first five abdominal somites and six on 
the sixth. Those of the submedian pair are often very faintly marked except on the last 
somite where they appear as strong blunt ridges. The following carinae end in spines:— 


Carinae. Abdominal somites. 
Submedian ah fe mie 6.! 
Intermediate ae a on (el), A, 
Wateralen: se er be BDA RO: 
Marginal an #2 Le I By By alg Sc 


The telson is thick and massive. The median carina has the form of a strong blunt 
ridge which, except in very large examples (where it appears to have been worn away), 
terminates posteriorly in a spine. The ridge is continued beyond the spine and, as a 
rule, projects as a median tubercle on the distal margin. On either side of the median 
crest are a few oblique rows of pits. The six marginal teeth are sharp and prominent 
and in old specimens somewhat rugose, each forming the termination of a short rounded 
carina. There is no praelateral denticle. In large specimens of both sexes the entire 
margin of the telson is strongly swollen and tends to obscure the denticles. Of the 
latter there are four to six submedian, seven to thirteen (usually nine or ten) inter- 
mediate and one lateral. The post-anal carina is distinct and in old specimens more 
or less tuberculate. 

In the bifurcate process from the base of the uropod the inner spine is about twice 
the length of the outer and bears a small lobe on its outer margin near the end of its 
distal half. 

Among Indo-pacific species Squilla raphidea is peculiar in that the margins of the 
telson are inflated in large specimens of both sexes. The angled external edge of the 
raptorial dactylus, characteristic of the mature male, is, as a rule, well-marked in all 
examples of that sex measuring 190 mm. or more in length ; but one male of 190 mm. 
is remarkable for the entire absence of this feature. 

Balss has described a variety under the name, var. africana, distinguished by 
the form of the rostrum, the sharpness of the submedian carinae of the post-abdomen 
and the thickened teeth and denticles on the margin of the telson. The rostra of 


! Often worn down and obsolete in very large specimens. 
? In large specimens only. 


S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. OI 


several specimens in the Indian Museum resemble that figured by Balss, others are 
normal or short and with convex margins as in De Haan’s figure of S. harpax. The 
other characters mentioned by Balss are also seen in some specimens, but it appears 
quite impossible to correlate them with any particular type of rostrum. 

In most spirit specimens the hinder margins of all the segments of the post-ab- 
domen except the first and last are defined by a line of black pigment. There are, as 
a rule, two black spots on the superior margin of the raptorial merus near its distal 
end, a spot or streak on the inside of the carpus close to the dorsal carina, a spot on 
either side of the propodus near its distal end and a round and often well-defined spot 
at the base of the telson on each side of the median carina. The distal ends of the 
uropods are also suffused with black pigment ; but in the terminal segment of the 
exopod the suffusion is confined to the inner longitudinal half. 


1913.] 


The following specimens of Squilla raphidea are in the Indian Museum :— 


9834-40 

6 

= | Hongkong. Hongkong Museum. 189,149, 82170 mm. 
1843—9 { G. Dennys. 

9 

= Singapore. Raffles Museum. 19,253 mm. 


Port Blair, Andamans. 
Mergui Archipelago. 


Rangoon. 


Off Irrawaddy Delta; 20 fms., 


15° 20 N., 94° 55° E. 
Akyab, Tenasserim. 


7318—20 


10 
7322 


6 
7324 
10 


M. of R. Hughli, 
Ganges Delta, 


Off Puri, Orissa Coast. 


Off Gopalpur, Ganjam Coast ;: 
25—28 fms. 


Off Madras Coast. 
Ceylon 


| Bombay. 


(purchased.) 
Mus. Colir. 
Mus. Colir. 


‘ Investigator.’ 


I. H. Burkill. 


J. H. Row. 
W. M. Daley. 


J. Butler. | 
A. J. Milner. 


( ‘ Investigato1. | 


‘ Golden Crown.’ 


‘Golden Crown.’ 


‘ Golden Crown.’ 
Colombo Museum. 


\ Bombay Nat, Hist. Soc. 
| J. Caunter. 


| 


I? ,124 mm. 
38,12, IOoI—175 mm. 
Io, 235 mm. 


IS ,19,53, 100 mm. 


Ig, 193 mm. 


93,82, 92—300 mm. 


I3',22, 243—288 mm. 


IG', 100 mm. 


Ig', 152 mm. 


Id, 127 mm. 


83,92, 72—335! mm. 


1 This is, I believe, the largest known Stomatopod. 


92 Memoirs of the Indian Museum. NOrsNEe 


FE Karachi. Karachi Museum. Id, 108 min. 
Sl Persian Gulf; 28°59’ N., ‘ Investigator.’ 1,290 
SOUS eee 5 iss 
nm (no history.) 28,52 ,83—27I mm. 
I have also examined the following specimens from other sources :— 
Suruga Gulf, Japan. A. Owston. 19,188 mm. 
Bombay Harbour. Bombay Nat. Hist. Soc. 19, 208 mm. 


Squilla raphidea has been recorded from the following localities:—Japan (De 
Haan, Rathbun, Balss) ; Hongkong (Bigelow) ; Formosa (Balss) ; Philippine Is. 
(Miers) ; Borneo (Miers) ; Java and Java Sea (de Man, Thallwitz) ; Malay Peninsula 
(Miers, de Man, Nobili, Lanchester) ; Mergui Archipelago (de Man) ; Madras (Hender- 
son) ; Palk Bay and Rameswaram (Tattersall) ; E. Africa (Balss) ; Zanzibar (Miers). 


34. Squilla annandalei, Kemp. 

Plate VII, figs, 78—80. 

IgII. Sguilla annandalei, Kemp, Rec. Ind. Mus., VI, p. 99. 
This species is very closely allied to S. raphidea, but differs in the following parti- 

culars :— 
I. The rostrum (fig. 78) bears some resemblance to that seen in the form of 
S. raphidea figured by De Haan under the name of S. harpax. The apex, 
however, is more bluntly rounded. 

. The dorsal processes of the ophthalmic somite are more sharply-pointed distally. 

3. The antennular peduncle is at least as long as the carapace and rostrum 
combined. 

A. The anterior lateral lobe of the sixth thoracic somite (fig. 80) is prominent and 
acute. Itis a little shorter than the posterior lobe, the latter, however, being 
better developed than in S. raphidea. 

5. The anterior lateral lobe of the seventh thoracic somite is small, Bu distinct 
and acute (fig. 80). 

6. The intermediate carinae of the last three thoracic somites end in spines. These 


D 


are never found in S. vaphidea of a corresponding size. 
7. The submedian carinae of the fifth abdominal somite end in spines. The com- 
plete formula is :— 


Carinae. Abdominal somites. 
Submedian se A Sie 4 Os 
Intermediate ae ais 2 T2 AO: 
Lateral .. Bi + a6 yn Sy Cy Sy Os 
Marginal a = + it, By 32, 5s 


8. The dorsal carina of the telson is sharp and considerably higher than in S. 
vaphidea. It terminates in a sharp spine overhanging a single tubercle, but 
is not continued beyond this to the distal margin. 


The corneal portion of the eyes is a trifle larger than in S. raphidea, specimens 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 93 


of similar size being compared. Submedian carinae are very distinct on the last three 
thoracic and on all the abdominal somites. The last two somites and telson are finely 
rugose and, frequently, a fine pearly sheen is visible on various parts of the body ; 
most conspicuously on the under-surface of the telson. The marginal teeth of the 
telson are rather longer than in S. raphıdea ; there are six to eight submedian denticles, 
seven to eleven intermediate and one lateral. 

In other structural details S. annandalei seems to bear an exceedingly close 
resemblance to the preceding species. There are eight teeth on the raptorial dactylus, 
a row of stiff spines along the upper margin of the propodus and an acute angular lobe 
on the lateral margin of the carapace. The single male example has only one raptorial 
claw and that has obviously been broken off and subsequently regenerated. The dac- 
tylus shows no trace of the external process found in adult S. raphidea of the same sex. 

The pigmentation of the external segment of the outer uropod is quite distinctive 
and is conspicuous in all the specimens. The segment is entirely jet-black except for a 
white midrib (fig. 79). In S. raphidea there is merely a rather indefinite suffusion of 
black along the inner longitudinal half. There is also a dark transverse patch, some- 
times broken up into four separate spots, on the second abdominal somite and various 
short transverse streaks on other somites (fig. 79). For the rest the dark markings 
resemble those of S. raphidea ; the two black spots near the proximal end of the 
telson are distinct and sharply defined. 

The four specimens of Squilla annandalei were all found in comparatively deep 
water in the Gulf of Martaban :— 


T5 Gulf of Martaban; 61 fms., ‘Investigator.’ 192,115 mm. TYPE. 
14° 40’ N., 95° 52’ E. 

u Gulf of Martaban; 67 fms., ‘Investigator.’ Id, 106 mm. 
T4° 26’ N., 96 23, E. 

2 Gulf of Martaban; 55 fms., ‘Investigator.’ 19,101 mm. 
14° 54’ 30” N., 96° 13’ E. . 

ar Gulf of Martaban: 53 fms., ‘ Investigator.’ 19,90 mm. 


142 382.122 N, 06.247302 E. 


* Leptosquilla schmeltzii’’ (A. Milne-Edwards). 
Plate VII, figs. 81—83. 
1873. Squilla schmeltzii, A. Milne-Edwards, Journ. Mus. Godeffroy, I, iv, p. 87, pl. ii, fig. 7. 


1880. Squilla schmeltzi1, Richters, in Mobius’ Meeresfauna Mauritius, p. 168. 
1880. Leptosquilla schmeltzii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 13. 


There can be little doubt that the Stomatopod to which this name has been applied 
is merely a late post-larval example of the genus Squilla. One of its most prominent 
features, that on which Miers founded the genus Leptosquilla, is the peculiar elongated 
form of the antennular somite' and it is clear from comparison with Bigelow’s figure of 
the ‘‘ first stage of the adult form ’’ in Squilla quadridens (1894, p. 548, text-fig. 28) 


! Miers states that it is ‘‘ the ophthalmic segment which is greatly elongated and prolonged beyond 
the rostrum.’’ This is clearly an error. 


94 Memotrs of the Indian Museum. Norte 


that this is characteristic merely of a particular stage in the development of certain 
species. 

There is a single specimen of this form in the Indian Museum which appears to 
belong to the same species as that described by Milne-Edwards. The chief characters 
are sufficiently well shown in the figures (figs. 81-83) to render detailed description 
unnecessary. 

I am inclined to agree with Hansen (1895, p. 69) that the form is in all pro- 
bability related to one of the species of Sguilla to which Miers applied the name 
Chloridella, though the possibility of some affinity with such species as S. /asciata, 
S. lata and S. scorpio cannot be overlooked. 

In the very elongated cylindrical eyes L. schmeltzi bears no resemblance to any 
adult species of Squilla ; it is probable, however, that this feature is subject to con- 
siderable modification during further growth. But the chief difficulty that is en- 
countered in an attempt to discover relations between this form and any species of 
Squilla hitherto known from the Indo-pacific region concerns the structure of the rap- 
torial claw. The dactylus of L. schmeltzi bears seven teeth and, among species of 
Squilla with a single lateral process on the fifth thoracic somite, this number is found 
only in S. armata, a species of isolated position and a peculiar distribution confined to 
southern latitudes. 

_ In the present state of our knowledge it seems impossible to suggest any plausible 
view of the relations of this form ; but it may prove considerably easier when the early 
post-larval history of the adult species is better known. 


The single specimen in the Indian Museum is registered as follows :— 


7 Port Blair, Andamans. J. Wood-Mason. IS, II mm. 


Genus Pseudosquilla, Dana. 


1837. Squilla, Squilles trapues, H. Milne-Edwards, Hist. Nat. Crust., II, pp. 518, 525. 
1837. Gonodactylus, H. Milne-Edwards, ibid. pp. 528, 530 (partim). 

1849. Squilla, sect. Parallele, De Haan, in Siebold’s Fauna Japonica, Crust., p. 221. 
1852. Pseudosquilla (Guérin ined.) Dana, U.S. Explor. Exped., Crust., p. 621. 

1880. Pseudosquilla, Miers, Anu. Mag. Nat. Hist (5), V, p. 108. 

1894. Pseudosquilla, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 408. 

Carapace little narrower in front than behind with conspicuous gastric grooves ; 
cervical groove usually absent, never visible mid-dorsally ; no carinae on carapace ; 
antero-lateral angles not spinous, cornea of eye rarely bilobed. Mandibular palp 
composed of three, rarely of two, segments. Merus of raptorial claw articulating 
terminally with ischium ; ventral surface of merus channelled longitudinally through- 
out its length ; upper margin of propodus finely pectinate externally ; dactylus not 
inflated at base and armed with at most three teeth excluding terminal one. Shorter 
ramus of last three thoracic limbs linear, composed of a single segment. Free 
thoracic and abdominal somites compressed, first five of the latter without sharp 
longitudinal carinae. Telson with median carina and one to five pairs of additional — 
carinae; submedian, intermediate and lateral marginal teeth, well-formed, the first 


T9rT3.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 95 


with at least the tips movable; no submedian denticles, never more than three inter- 
mediate denticles. Ventral process of uropods consisting either of two well-formed 
spines, inner longer than outer, or vice versa and with an additional spine or spine 
and spinules on inner margin near base. 

In structure there do not appear to be any marked secondary sexual differences. 

One species, the very curious and rare P. ferusacci from the Mediterranean, 
disagrees in several important particulars from the diagnosis given above and seems 
in fact to be almost exactly intermediate in character between Pseudosquilla and 
Squilla. The principal features in which alliance with the latter genus is shown are (i) 
the well marked cervical groove, (ii) the presence of the posterior reflected part of 
the marginal carina of the carapace and (iii) the presence of fine submedian, inter- 
mediate and lateral carinae on the abdomen. ‘The form of the telson is, however, 
similar to that found in Pseudosquilla and the characters afforded by the ventral 
process from the basal segment of the uropods seem to indicate a position near 
P. empusa, P. cerisii, P. lessons and their allies. 

Another outstanding form of doubtful position is P. stylifera; this species is 
known from the Australian and Chilian Coasts and there is some evidence to show 
that different races inhabit these widely separated localities. 

Of the twelve species of Pseudosquilla that are known, nine have been found 
within the area with which this memoir is concerned and two of these, P. ciliata and 
P. oculata, have also been taken in the Atlantic. Two forms, P. cerisii and P. ferus- 
saci, appear to be restricted to the Mediterranean and one, P. lessoni, is known only 
from the Pacific Coast of America. 

In Pseudosquilla ciliata and its immediate allies a curious process springs from 
the dorsal aspect of the antennal protopodite. This consists of a flat elongated plate, 
directed forwards, and provided inferiorly with a deep vertical keel. In some cases 
this process seems to afford useful specific indications. 

The species of Pseudosquilla seem to prefer rough ground, inhabiting coral reefs, 
oyster-beds and similar situations. They are invariably found in shallow water, the 
greatest depth from which any species is known being 20-23 fathoms (P. ciliata, 
Bigelow). 

The carinae on the telson, in species which possess the full complement, consist 
of a median and five pairs and for these I have used the terms “submedian,”’ 
inermediaren traiter a, second are and <omarcinal.. This ter 
minology is the same as that employed in the case of Odontodactylus (see text- 
fig. 4, p. 12), but the ‘‘ second submedians ” are never found in the present genus. 


Key to the species of Pseudosquilla. 


I. Dactylus of raptorial claw with teeth on its inner margin. 
A. Basal process of uropods terminating in two large teeth, its inner margin 
smooth. 
I. Telson with only three carinae on either side of median crest, the 
first and second laterals being absent. 


Memoirs of the Indian Museum. 


A. Eyes long and cylindrical ; cornea set very obliquely on stalk 
B. Eyes short and flattened; cornea set transversely on stalk. 
I. Rostrum without a median distal spine; a pair of dark spots on 
carapace, usually surrounded by a white ring 
2. Rostrum with a slender median distal spine; a pair of dark spots 
on carapace not surrounded by a white ring : 
Telson with four pairs of carinae on either side of median crest, the 
first lateral present. 
A. Breadth of cornea scarcely equal to whole length of eye; Six 
posterior spines on last abdominal somite; first lateral carinae 
terminating behind base of intermediate 


II. 


of telson parallel, 

marginal teeth. . 

B. Breadth of cornea greater en whole ans of eye; on He 

rior spines on last abdominalsomite; first lateral carinae of telson 

posteriorly divergent, coterminous with lateral marginal teeth .. 

B. Basal process of uropods with a single large additional tooth or with a 

series on inner margin. 
I. Telson with not more than three carinae on either side of the median 
crest. 

Rostrum less than twice as broad as long; abdomen with nine fine 
longitudinal carinae; telson without submedian carinae. 

Rostrum almost three times as broad as long ; first five abdominal 

somites without carinae; telson with one pair of submedian 


A. 
B. 


carinae ; 58 co 

Telson with full en of carinae, er on either side of median 
crest. 

A. Raptorial dactylus with three teeth! ; 

I. Basal process of uropod bearing three large teeth, one terminal 

and two on inner margin. 
a. Rostrum with sharp median spine and obtuse antero-lateral 
angles 80 © BE 
b. Rostrum with sharp median spine and acute or spinous antero- 


JOG 


telson carinae not spinous. 


lateral angles 50 
2. Basal process of uropod with two lage teeth and numerous 
small spines on inner margin 
B. Raptorial dactylus with four teeth,! no carinae of len 
with two or three sharp spines, intermediate with two distal 
spinules, first lateral with one 
Dactylus of raptorial claw with spinous termination, but rom teeth on 
[Rostrum longer than broad; ophthalmic somite largely 
eyes long with very oblique cornea; telson with intermediate 
carinae; basal process of uropod terminating in a single large tooth} 


II. 
inner marein. 
exposed ; 


1. Pseudosquilla ciliata’ (Fabricius). 


1787. Syuilla ciliata, Fabricius, Mantiss. Insect., I, p. 333. 
1790. Cancer ciliatus, Linnaeus, Syst. Nat., ed. XIII, I, p. 2990. 


! Including the terminal tooth. 
? See addendum, p. 196. 


IVor ee 


ciliata, p. 96. 


ornata, p. 100. 


oxyrhyncha, p. IOI. 


oculata, p. 102. 


megalophthalma, p. 103. 


[ferussaci.] 


empusa, p. 104. 


[cerisir.] 
[lessoni.] 


dofleini, p. 104. 


pilaensis, p. 105. 


stylifera, p. 106. 


1913.] 


1793 
, 1796 
1798 
1818 
1825 


S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 97 


. Squilla ciliata, Fabricius, Ent. Syst., II, p. 512. 

. Cancer (mantis) ciliatus, Herbst, Krabben u. Krebse, II, p. 102. 

. Squilla ciliata, Fabricius, Ent. Syst. Suppl., p. 417. 

. Squilla stylifera, Lamarck, Hist. Nat. Anim. sans Vert., V, p. 180. 
. Squilla stylifera, Latreille, Encycl. Méthod., X, p. 472. 


1829-43. Squilla stylifera, Guérin Méneville, in Cuvier’s Iconogr. Règne Anim., III, Crust., p. 19, 


1832 


1837. 
1838. 


1830. 
1830. 
1841. 


1847. 
1850. 
1852. 
1862. 
1869. 
1372. 
1874. 
1380. 
1380. 
1880. 
1882. 
1886. 
1886. 
1887. 
1892. 
1893. 
1893. 
1894. 
1895. 


1898. 
1898. 
1899. 
1899. 
1899. 


pl. xxiv, fig. Ia, b. 

. Squilla ciliata, Owen, Proc. Zool. Soc., p. 6. 

Squilla stylifera, H. Milne-Edwards, Hist. Nat. Crust., II, p. 526. 

Squilla stylifera, H. Milne-Edwards, in Lamarck’s Hist. Nat. Anim. sans Vert., 2nd ed., V, 
p. 324. 

Squilla ciliata (leach, MS.), Owen, Zool. Voy. ‘Blossom’ Crust., p. 90, pl. xxvii, fig. 5, 5a-d. 

Squilla stylifera, Randall, Journ. Acad. Nat. Sci. Philadelphia (1), VIII, p. 147. 

Squilla quadrispinosa, Eydoux and Souleyet, Voy. de la ‘ Bonite,’ Zool., I, Crust., p. 262, 
pl. v, fig. I. 

Squilla ciliata, White, List Crust. Brit. Mus., p. 84. 

Squilla stylifera, Gibbes, Proc. Americ. Assoc., p. 200. 

Pseudosquilla stylifera, Dana, U.S. Explor. Exped., Crust., I. p. 622, pl. xli, figs. 4, a-e. 

Squilla stylifera, A. Milne-Edwards, in Maillard’s l’île Réunion, ann. F., p. 16. 

Squilla stylifera, Clark, Proc. Zool. Soc., p. 3. 

Pseudosquilla stylifera, v. Martens, Arch. f. Naturgesch., XX XVIII, i, p. 146, 

Squilla stylifera, Hoffmann, Rech. Faune Madagasc., V, ii, p. 36. 

Pseudosquilla ciliata, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 108, 458, pl. iii, figs. 7, 8. 

Squilla quadrispinosa, Richters, in Mobius’ Meeresfauna Mauritius, p. 168. 

Pseudosquilla stylifera, Richters, ibid. p. 169. 

Pseudosquilla ciliata, Haswell, Cat. Australian Crust., p. 209. 

Pseudosquilla ciliata, Muller, Verhandl. naturf. Ges., Basel, VIII, p. 471. 

Pseudosquilla ciliata, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p.53, pl. xv, fig. ro. 

Pseudosquilla ciliata, de Man, Arch. f. Naturgesch., LIII, i, p. 57T. 

Pseudosquilla ciliata, Thallwitz, Abhandl. kön. Mus. Dresden, no. 3, p. 55. 

Pseudosquilla ciliata, Henderson, Trans. Linn. Soc., Zool., (2), V. p. 454. 

Pseudosquilla ciliata, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 474. 

Pseudosquilla ciliata, Bigelow, Proc. U.S. Nat. Mus., XVII. p. 499. 

Pseuderichthus communis, Hansen, Isop. Cumac. u. Stomatop. Plankton. Exped., p. 86, 
pl. viii, figs. 5-50. (larval form). 

Pseudosquilla ciliata, de Man, Zool. Jahrb., Syst., X, p. 694. 

Pseudosquilla ciliata, Borradaile, Proc. Zool. Soc., p. 36. 

Pseudosquilla ciliata, Rankin, Ann. N. Y. Acad. Sci., XII. p. 545. 

Pseudosquilla ciliata, Borradaile, in Willey’s Zool. Results, p. 402 (including var. occidentalis). 

Pseudosquilla ciliata, Nobili, Ann. Mus. civ. Genova (2), XX, p. 275. 


1901-3. Pseudosquilla ciliata, Verrill, Trans. Conn. Acad., XI. p. 20. 


1902. 
1902. 
1903. 
1904. 
1906. 
1906. 
1907. 
IQIO. 
1910. 


Pseudosquilla ciliata, Bigelow, Bull. U.S. Fish Comm. for 1900, XX, 2, p. 154, figs. 3, 4. 
Pseudosquilla ciliata, de Man, Abhandl. senck. Ges. Frankfurt, XXV, p. 911. 
Pseudosquilla ciliata, Lanchester, Faun. and Geog. Maldives and Laccadives, I, p. 457. 
Pseuderichthus communis, Jurich, Stomatop. deutsch. Tiefsee-Exped., p. 395, pl. xxix, fig. 1. 
Pseudosquilla ciliata, Tattersall, Ceylon Pearl Oyster Rep., V, p. 167. 

Pseudosquilla ciliata, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 335. 

Pseudosquilla ciliata, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 213. 

Pseudosgunlla ciliata, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 571. 

Pseudosquilla ciliata, Fukuda, Annot. Zool. Japon., VII, p. 145. 


98 Memoirs of the Indian Museum. [Morne 


The carapace is smooth, without carinae, and is considerably longer than its great- 
est breadth. The gastric grooves converge anteriorly ; of the cervical groove no trace 
remains. The antero-lateral angles are rounded. The rostrum is fully one and a half 
times as broad as long, widest in front of the base, and its strongly rounded, lateral 
margins meet apically in a very broad, obtuse, and almost obsolete angle. 

The eyes are cylindrical and about as long as the greatest breadth of the rostrum. 
The cornea scarcely exceeds the breadth of the stalk and is set very obliquely on it. 
It is traversed laterally, both inside and outside, by a narrow longitudinal band com- 
posed of fine parallel grooves. The antennular peduncle is about half the length of 
the carapace and rostrum combined. ‘The dorsal process of the proximal segment of 
the antennal protopodite is flat above and the margin of the vertical keel on its ventral 
surface is concave in lateral view. 

The carpus of the raptorial claw is bluntly carinate dorsally ; the propodus bears 
three movable spines at the base of its pectinate margin and the dactylusis very slender, 
convex externally, and possesses three long flattened teeth including the apical 
one. 

The exposed thoracic somites are smooth dorsally. The fifth has a deep vertical 
groove on its lateral face. The lateral margins of the sixth somite are truncate ; those 
of the seventh are less sharply truncate and, owing to the oblique antero- and postero- 
lateral edges, are much narrower. The eighth somite is narrowly rounded laterally 
with a small and deeply incised apical notch. 

The first five abdominal somites are smooth except for a deep Y-shaped groove on 
the lateral face of the first and for a pair of conspicuous pits on each of the four succeed- 
ing somites. The postero-lateral angles of the fourth end in a small spine in Atlantic 
specimens and in some from the Indo-pacific; those of the fifth somite are invariably 
spinous. The sixth somite has a pair of strong submedian spines on its posterior 
margin and a similar pair at the postero-lateral angles. Between these two pairs 
there is an additional spine on either side which springs from the middle of the somite 
and does not quite reach to the distal edge. The abdomen is very strongly arched 
from side to side and its greatest breadth is contained rather less than five and a half 
times in the total length. 

The telson bears seven carinae on its dorsal surface, the first and second laterals 
being absent. The median carina is high and sharp and terminates in a strong spine 
which may reach as far as the distal margin. The submedians are low and in young 
examples inconspicuous ; they converge distally to the posterior end of the median 
carina and are absent in the anterior third of the telson. The intermediate carinae 
are strong, posteriorly divergent, and end abruptly in a sub-acute apex. The marginal 
carinae are externally curved and reach the edge of the telson in front of the posterior 
marginal tooth. Inside this carina near the proximal end there is a small tubercle. 
On the margin there are six large spines : a pair of long movable submedians and two 
pairs, the intermediates and laterals, which are fixed. The edge between the sub- 
medians is usually rather deeply incised, rarely entire. On the inner side of the inter- 
mediates there is a minute spinule succeeded by a rounded lobe and the margin 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 99 


between the intermediates and laterals is concave or slightly sinuous with a small 
spinule at the base of the latter. The ventral surface of the telson is smooth. 

The external margin of the peduncular segment of the uropod is rounded; 
dorsally there is a blunt longitudinal carina which is produced as a spine over the 
articulation of the basal segment of the exopod. The ventral process is long, reaching 
almost to the apex of the exopod; distally it is cut into two strong $Spines, the inner 
of which is usually a trifle longer than the outer. There are seven to nine movable 
spines on the outer margin of the proximal segment of the exopod ; the terminal one, 
which is the longest, fails to reach, or only just reaches the distal end of the ultimate 
segment. On the ventral surface there is a single fixed spine at the distalend. The 
paddle or ultimate segment of the exopod is at least three-quarters the length of 
that which precedes it. 


The colour of spirit specimens is not specially distinctive ; the large eye-spot on 
either side of the carapace, which occurs in some of the allied species, is missing. Clark 
(1869, sub Squilla stylifera) has given a most interesting account of the habits of this 
species and has described the colour of the living animal as follows :— 

““ When first hatched the larvae are of a delicate yellowish green, and are very 
active. As they grow they assume a mottled grey, and the swimmerets and legs 
become pea-green. The green gradually increases in brightness, but it is not till they 
have reached a length of three inches that the colours of the adult appear. The male is 
then of a beautiful bluish green, with the jaw-feet, the swimmerets, and the branchiae, 
as well as the antennae and the fimbriae which border the different organs, of a cherry- 
red. The female is clouded with brown and grey, presenting much the appearance of 
tortoise shell, and the red about her is much less vivid than in the male.’’ 

Specimens preserved in formalin and examined shortly after capture were of a 
uniform dull olivaceous yellow, very faintly mottled with white laterally, and tending 
to a slightly redder shade on the telson and to a green tone on the inner and outer 
uropods. The antennal and antennular peduncles, the base of the antennal scale, and 
the raptorial claw were mottled with dull yellow and white and the raptorial dactylus 
was in addition marbled with red. The other thoracic and abdominal appendages 
were olivaceous, sometimes faintly mottled and all the fringes of setae were red. 


According to Brooks’ observations (1886) Atlantic specimens of this species differ 
in certain small details from those found in the Indo-pacific seas, and Borradaile (1899) 
proposed to distinguish the former by the name var. occidentalis. Bigelow, however, 
in 1002, supplied a description and good figures of the Atlantic specimens from which it 
appears that only one character (v.c. the presence or absence of a postero-lateral spine 
on the fourth abdominal somite) remains to distinguish the two forms. Now that 
Tattersall has found by actual comparison that a specimen from Ceylon possesses all 
the characters which Brooks noted as distinctive of his West Indian specimens, there 
seems little to justify Borradaile’s statement in 1907 that ‘‘it is not clear if the true P. 
ciliata is found in the Atlantic.’’ 

Of the sixteen specimens, all from Indo-pacific waters, which are preserved in the 


100 Memoirs of the Indian Museum. Do I, 


Indian Museum, three possess spines at the postero-lateral angle of the fourth abdominal 
somite, while, in all, the inner spine of the bifurcate process of the uropod is, as shown 
in Bigelow’s figure, slightly longer than the outer. 

The specimens examined are registered as follows :— 


— Australia, .. Queensland Museum. .. 19,76 mm. 

= Upolu, Samoa. ..  (purchased.) .. 19,81 mm. 

> Andaman Is. | B. Ford, V. Ball 

“a i Ve Wood-Mason. En, 

TES Andaman Is. .. G.H. Booley. .. 29,61 and 75 mm. 

PT N. Cheval Paar, Ceylon. : .. I. Southwell. .. 2822, 50-55 mm. 
a Persian Gulf. en .. FE. H. Townsend. .. 2629 ,27-51 min. 

305 Mauritius. .. Stege. ra une 


I have also seen three examples, kindly sent me on loan by Mr. Alan Owston. 
These were found at Iriomote, Vayeyama, Liu Kiu Is. 

Pseudosquilla ciliata appears to be a common species, of constant occurrence 
in coral reefs, and has a wide distribution in Indo-pacific and Atlantic waters. In the 
former region it has been recorded from the Ogasawara Is., Japan (Fukuda), Hawaiian 
Is. (Owen, Randall, Miers, Eyd. and Soul., Dana, Brooks, Bigelow), Solomon Is. 
(Miers), Fiji Is. (Miers, Dana), Loyalty Is., New Britain and Ellice Is. (Borradaile), 
Australia (Miers), Arafura Sea (Pocock), New Guinea (Miers, Thallwitz), Amboina (de 
Man), Celebes (de Man), Sooloo Sea and Philippine Is. (Miers), Sumatra (Nobili), Timor 
(30 fms.), Ceylon (Muller, Tattersall), Minikoi (Lanchester), Maldives (Lanchester), 
Madras (Henderson), Red Sea (Nobili), Seychelles (Borradaile), Mauritius (Richters, 
Clark), Reunion (Milne-Edwards, Hoffmann), Madagascar (Lenz) and from Diego Garcia, 
Wazin I. and Zanzibar (Borradaile). 

In the Atlantic the adult is known from Bermuda (Bigelow) Bahama Is. (Rankin, 
Bigelow), S. Carolina (v. Martens), Florida Keys, Porto Rico and Culebra Is. (Bigelow) 
and from St. Thomas (Bigelow, 20-23 fms. ; Brooks, 2 fms.). 

The larva has been recorded from the Indian Equatorial current (Jurich) from 
Ceylon (Tattersall) and from the Chagos Archipelago ; Alphonse Is., Amirante group; 
Providence Is. and between Mauritius and Cargados (Borradaile). In the Atlantic it 
has been recorded from the Sargasso Sea and the N. Equatorial current (Hansen). 


2. Pseudosquilla ornata, Miers. 
1861. Pseudosguilla oculata, Heller, Verhandl. zool-bot. Ges. Wien, XI, p. 497. 
1865. Pseudosquilla oculata, Heller, Reise ‘ Novara’ Exped., Crust., p. 124. 
1880. Pseudosquilla ornata, Miers, Ann. Mag. Nat. Hist. (5), V, p. III, pl. iii, figs. 5, 6. 
1887. Pseudosquilla ornata, de Man, Arch. f. Naturgesch., LIII, i, p. 571. 
1894. Pseudosquilla ornata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 500. 
1894. Pseudosquilla ornata, Ortmann, Denk. med-wiss. Ges. Jena, VIII, p. 60. 
1907. Pseudosquilla ornata, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 213. 


Pseudosquilla ornata is closely allied to P. ciliata but differs from it in the following 
characters :— 


1913.] S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. IOI 


I. The rostrum is shorter, nearly twice as broad as long. 

2. The eyes are much stouter, shorter than the greatest breadth of the rostrum 
and the cornea is dorso-ventrally flattened and is set transversely on the stalk. 

3. The dorsal process of the basal segment of the antennal protopodite is deeply 
chanelled dorsally and the margin of the vertical keel on its ventral surface 
is not concave in lateral view. 

4. The seventh thoracic somite is squarely truncate laterally and is not narrowed, 
and the lateral margin of the eighth shows only obscure indications of the 
deeply-incised notch which occurs in P. ciliata. 

5. The anterior limb of the Y-shaped groove on the lateral wall of the first 
abdominal somite is obsolete. 

6. There is a deep incised groove isolating two separate lobes between the sub- 
median and intermediate spines of the telson, and a similar groove defining 
a similar lobe just inside the lateral spines. These lobes replace the small 
spinules noticed in the preceding species. 

7. The outer spine of the bifurcate process of the uropod is much larger than the 
inner and reaches to the distal end of the exopod. 

8. The ultimate segment of the exopod isvery small, less than half the length of the 
preceding segment and the outermost of the 9-11 movable spines which fringe 
the external margin of the latter, reaches beyond the apex of the former. 

g. The whole animal is narrower, the greatest breadth of the abdomen being 
about one-sixth the total length. 


In most spirit specimens a pair of conspicuous dark eye-spots, which are frequently 
defined by a white ring, are found on the carapace. 


There are only two specimens in the Indian Museum :— 
aS Samoa. .. (purchased.) 5. Lo), 82 mm. 


= Mauritius. ,. Stege. De Oy A Em. 


This species appears to be much scarcer than P. ciliata; it has been recorded from 
Tahiti (Heller) ; Samoa (Ortmann) ; Kagoshima, Japan (Ortmann); Philippines (Miers) ; 
Amboina (Miers, de Man, Ortmann); Salomon Atoll, Chagos Arch. (Borradaile)- 
Coetivy, Seychelles (Borradaile) and Mauritius (Bigelow). 


3. Pseudosquilla oxyrhyncha, Borradaile. 
1898. Pseudosquilla oxyrhyncha, Borradaile, Proc. Zool. Soc., p. 37, pl. vi, figs. 9, ga-d. 
1907. Pseudosquilla oxyrhyncha, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 213. 

This species is known to me only from Borradaile’s description and figures. It is 
evidently extremely closely allied to P. ornata, but differs from it in possessing a delicate 
apical spine on the rostrum. Borradaile (1907) states that, in addition, the last segment 
of the expodite of the uropod is outreached by the inner spine of the bifurcate process, 
but in his figure (1898) the reverse is shown. The dorsal edge of the carpus of the rap- 
torial claw terminates (according to the figure) in a spine and in this respect differs from 
the examples of P. ornata in the Indian Museum. In other details, such as the form of 


102 Memoirs of the Indian Museum. Vor ARE 


the eyes, of the telson, and of the lateral margins of the sixth and seventh thoracic 
somites, there appears to be the closest resemblance between the two forms. 

The colour is as follows :—‘‘ Abdomen olive-green, mottled with paler green ; 
carapace greenish brown, mottled with cream; raptorial claw brown (dactylus purple) 
mottled with cream.’’ There is also a pair of large dark spots on the carapace, not 
surrounded by a white ring. 

The only known specimen, a male, 88 mm. in length, was found at Rotuma, Fiji Is. 


4 Pseudosquilla oculata (Brullé). 

1830-44. Squilla oculata, Brullé, in Webb and Berthelot’s Iles Canaries, Zool., II, ii, Crust., 
p. 18, pl. unique, fig. 3. 

1880. Pseudosquilla oculata, Miers, Ann. Mag. Nat. Hist. (5), V, p. 110, pl. iii, figs. 3, 4. 

1880. Pseudosquilla oculata, Richters, in Mobius’ Meeresfauna Mauritius, p. 160. 

1891. Pseudosquilla oculata, de Man, Notes Leyden Mus., XIII, p. 59. 

1893. Pseudosquilla oculata, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 474. 

1894. Pseudosquilla oculata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 500. 

1895. Pseudosquilla distinguendus, Hansen, Isop. Cumac. u. Stomatop. Plankton. Exped., p. 86. 

1904. Pseuderichthus distinguendus, Jurich, Stomatop. deutsch. Tiefsee-Exped., p. 39, pl. xxvii, 
fig. 5. 

1907. Pseudosquilla oculata, Borradaile, Trans. Linn. Soc. Zool. (2), XII, p. 214. 

1907. Pseuderichthus distinguendus, Borradaile, ibid., p. 215. 

This species is also a close ally of P. ornata, but is readily distinguished from it 
and from both the preceding species by the possession of an additional pair of short 
longitudinal carinae on the telson. These carinae, the first laterals, are parallel 
and terminate behind the base of the intermediate marginal teeth. 

In other respects the points of distinction between P. oculata and P. ornata are 
few :— 

I. The rostrum is provided with an apical spine.! 

2. The eyes are flattened as in P. ornata, but are shorter and more thick-set ; the 
cornea is transverse, very noticeably wider than the stalk, and its greatest 
breadth is almost equal to the length of the whole organ. 

3. The ultimate segment of the exopod is even smaller than in P. ornata, being much 
less than half the length of the preceding segment. It is outreached both by 
the distal external spine of the basal segment and by the outer (and longer) 
spine of the bifurcate process. 


De Man has compared the type specimen of this species, which was found at the 
Cape Verde Is., with two from Samoa, and notes that the only differences he can find 
are that in the Pacific examples the eye-peduncles are a little more dilated at the 
distal end and that the rostrum is slightly more transverse with the apical spine a trifle 
longer. 

According to Miers the colour of this species is described by Webb and Berthelot 
as being of a green colour, with numerous yellow rounded spots; there is a large round 


! This spine in the single specimen examined, is much larger than is indicated in Miers’ figure. 


1913. ] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 103 


green spot on each side bordered with a yellow ring ; penultimate joint of raptorial 
limbs bordered with green and yellow ; dactyli rose-coloured. 
There is only a single small specimen of P. oculata in the Indian Museum : — 


5 


Society Is. Otago Museum. 70927 


Pseudosquilla oculata has been recorded from Samoa {de Man) ; Macclesfield Bank, 
China Seas (Pocock) and from Mauritius (Richters, Bigelow). In the Atlantic it is 
known from the Cape Verde Is. (Brullé) and from Madeira (Miers). 


5. Pseudosquilla megalophthalma, Bigelow. 
1894. Pseudosquilla megalophthalma, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 500. 
1906. Pseudosquilla megalophthalma, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 336. 

P. megalophthalma, of which I have seen no specimens, is related to P. oculata, 
and possesses the same number of carinae on the telson. It may be distinguished by 
the following characters :— 

I. The rostrum is broadly heart-shaped, truncated at the base, and is interme- 

diate in form between that of P. ornata and P. oculata. 

2. The eyes are large and triangular. The conical portion is set transversely on 
the stalk and its breadth is considerably greater than the length of the whole 
organ. 

3. In addition to the usual six spines on the last abdominal somite there is a small 
one on the inner side of each of the intermediates. 

4. On the dorsal surface of the telson the first lateral carinae! are not parallel 
as in P. oculata, but are posteriorly divergent and are continued to the tips 
of the lateral spines. 

5. The distal segment of the exopodite of the uropods is larger than in P. oculata 
or P. ornata. 


The raptorial claw also is longer and more slender, and, when folded, extends from 
the apex of the eyes to the most posterior part of the carapace. 

In the type specimen the postero-lateral angles of the last five abdominal somites 
are spinous; only the last three in the specimens recorded by Nobili. The outer of the 
two spines of the bifurcate process of the uropod is longer than the inner in Bigelow’s 
example, but in Nobili’s specimens the two are subequal. 

The colouring, according to Nobili is characteristic :—‘‘ Sur la carapace il y a 
une grosse tache oculiforme noire, entourée par une ligne jaunâtre. Les angles posté- 
rieurs du cinquième et du sixième segment de l’abdomen sont aussi colorés de noir avec 
une ligne jaunatre. Je dos du doigt des pattes ravisseuses, et les épines et les crêtes 
du telson ont une couleur violette.’’ 

Bigelow’s specimen, which is 68 mm. in length, was found at Mauritius, the five 
smaller examples recorded by Nobili were taken at Obock in the Red Sea and at Dji- 
bouti. 


! It is possible that these carinae are not in reality first laterals, but are homologous with the 
second lateral carinae found in P. dofleini, pilaensis and cerisii. No figure of the telson is extant. 


TO4 Memotrs of the Indian Museum. [Vor. IV, 


6. Pseudosquilla empusa (De Haan). 
1844? Squilla empusa, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. 6. 
1849. Squilla empusa, De Haan, ibid., text, p. 224. 
1880. Pseudosquilla ? empusa, Miers, Ann. Mag. Nat. Hist. (5), V, p. 113. 
1884. Pseudosquilla empusa, Miers, Voy. H.M.S. ‘ Alert,’ p. 567. 
1894. Pseudosquilla ? empusa, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 499. 

The rostrum is broadly truncate in front, almost three times as wide as long and is 
impressed in the middle. The cornea of the eyes, according to De Haan’s description 
is scarcely wider than the middle of the stalk ; in the figure, however, it appears to be 
considerably broader. The raptorial dactyli bear three teeth including the terminal 
one. 

The thorax and first five abdominal segments have the same form as in P. ciliata. 
The sixth somite is furnished with six indistinct carinae ; the submedian and inter- 
mediate carinae do not terminate in spines. On the telson there is a median carina, 
a pair of submedians, one pair of laterals and perhaps also a pair of marginals. The 
six large marginal teeth are broad and flattened ; De Haan does not state that the sub- 
medians are movable, and the feature is not apparent in his figure. The basal process 
of the uropods terminates in two large teeth, the outer longer than the inner. Inaddi- 
tion there is one stout tooth, and possibly a series of smaller teeth also, on the internal 
margin of the process. 

Pseudosquilla empusa has not been rediscovered since it was originally described 
by De Haan. The type specimen, about 83 mm. in length,' is recorded from Japan. 


7. Pseudosquilla dofleini, Balss. 
1910. Pseudosquilla dofleini, Balss, Abh. math.-phys. Klasse k. bayer. Akad. Wiss., Suppl., Bd. IT, 
Abh. 2, p. 7, text-fig. I. 

The principal characters of this species are as follows :— 

The rostrum consists of a short basal part with rounded lateral margins and a 
strong median spine. The cornea of the eyes is greatly expanded, much wider than 
the stalk and is set obliquely. The dactylus of the raptorial claw is furnished with a 
tubercle at the base of its outer edge ; it bears three teeth, the terminal one long, the 
other two much shorter. 

The lateral margins of the last three thoracic somites are rounded. ‘The postero- 
lateral angles of the fifth abdominal somite are spinous, those of the four preceding 
somites, though sharply angled, do not bear spines. The last somite has six longitu- 
dinal ridges, terminating in spines, as in P. ciliata. On the telson the full number of 
carinae are present, five on either side of the median keel. The intermediate carinae 
are continued to the apex of the submedian spines, the first laterals stop close behind 
the base of the intermediate teeth and the second laterals are coterminous with the 
lateral teeth. 

The basal process of the peduncular joint of the uropods bears a rowof small teeth 


! There is reason to believe that De Haan’s figure represents his specimen at its full size. 


1013.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 105 


on its inner margin. It terminates in two larger teeth, the outer about twice the length 
of the inner. 

Pseudosquilla dofleini is allied to P.cerisi and P. lessomi, but is readily distinguished 
from them by the numerous teeth on the inner margin of the uropodal process. 

The above account is derived from Balss’ figure and description of the only known 
specimen. This example a female, 85 mm. in length, was found in Sagami Bay, Misaki, 


Japan. 
8. Pseudosquilla pilaensis, de Man. 


1888. Pseudosquilla pilaensis, de Man, Journ. Linn. Soc., XXII, p. 296. 
1894. Pseudosquilla pilaensis, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 499. 
1906. Pseudosquilla pilaensis, Nobili, Ann. Sci. Nat., Zool. (9), IV, 1906, p. 336. 

The principal features of this well-marked species are as follows :— 

The rostrum terminates in a prominent acute spine reaching a little beyond the 
eyes ; its basal portion is about twice as broad as long with slightly obtuse antero- 
lateral angles. ‘The eyes are short and the cornea is dilated and conspicuously bilobed. 

The dactylus of the raptorial claw possesses four teeth including the terminal one, 
differing in this respect from all other species of the genus; its external margin bears 
a sharply rounded lobe at the proximal end. 

The lateral margins of the sixth and seventh thoracic somites are rounded, 
broadly in front and narrowly behind. The postero-lateral angles of the fourth 
and fifth somites are spinous. The fifth bears on either side a deep longitudinal 
sulcus and a carina close tothe margin. The sixth somite bears six smooth triangular 
elevations ending in sharp spines and between the submedians and intermediates, 
there is a small rounded tubercle. 

The telson is twice as broad as long and, as in P. dofleini, bears the full number 
of carinae. The median terminates in a sharp spine; the submedian consists of 
a carina in the anterior half ending in a spine and succeeded by one, more commonly 
by two, spines; the intermediate is entire and ends in a spine overhanging a small 
tubercle before reaching the margin behind the outer intermediate denticle; the first 
lateral may be interrupted and terminates in a spine overhanging a tubercle at 
the base of the intermediate marginal tooth. The second lateral is continued to the 
apex of the lateral marginal tooth. Between the two laterals there is, in the single 
specimen examined, an additional short carina ending in a spine. On the margin the 
usual three pairs of primary teeth are prominent, the submedians being mobile. 
There are two rounded intermediate denticles and one lateral. The inferior surface 
is smooth except for a pair of spines on either side placed opposite the interspaces 
of the primary marginal teeth. 

The basal process of the uropods consists of a long and sharp spine, bearing 
a small spine on its outer margin and a series of five to seven similar spines on 
its inner margin. 

In colour the single example is pale with dark pigmentation along the sides of the 
carapace and abdomen. 


“~~ Port Maria, Elphinstone I., Mergui Archipelago. ‘ Investigator.’ 2 745 mm. 


106 Memoirs of the Indian Museum. LOG. ION, 


Hitherto known from two specimensonly. Thetype, 88 mm. in length, wasfound 
at Elphinstone I. (de Man), while the second example is recorded from the Red Sea 
(Nobili). 

9. Pseudosquilla stylifera (H. Milne-Edwards). 
Plate VII, figs. 84, 85. 


1837. Gonodactylus styliferus, H.Milne-Edwards, Hist. Nat. Crust., II, p. 330, pl. xxvii, figs. 9-14. 
1839. Gonodactylus styliferus, Randall, Journ. Acad. Sci. Philadelphia (1), VIII, p. 147. 

1840. Gonodactylus styliterus, Nicolet, in Gay’s Hist. fisica de Chile, Zool., III, p. 225, pl. ii, fig. 3. 
1880. Pseudosquilla stylifera, Miers, Ann. Mag. Nat. Hist. (5), V, p. 112. 

1894. Pseudosquilla styhifera, Bigelow. Proc. U.S. Nat. Mus., XVII, p. 502, fig. 3, p. 505. 

1900. Pseudosquilla stylifera, Holmes, Occ. Papers Calif. Acad. Scı., VII, p. 220. 

1900. Pseudosquiila stylifera, Whitelegge, Mem. Australian Mus., IV, p. 198. 

1910. Pseudosquilla bigelowi, Rathbun, Proc. U.S. Nat. Mus., XX XVIII, p. 608. 


The carapace is broad, its posterior breadth being equal to its mid-dorsal length. 
The cervical groove is distinct laterally, but is absent in the middle line. The 
antero-lateral angles are subrectangular, the postero-lateral broadly rounded and 
carinate. ‘The rostrum is smooth above, triangular, longer than broad, with straight 
lateral margins converging to a narrow rounded apex. 

The cornea of the eyes is greatly swollen and is set very obliquely on the stalk. 
In dorso-internal view the peduncle terminates in a narrow triangular area, invested 
on two sides by the cornea. The dorsal processes of the ophthalmic somite consist of 
a pair of large triangular lobes directed straightly outwards on either side and wholly 
exposed in dorsal view. The anterior margin of the somite is produced between the 
bases of the eyestalks and terminates in an acute point. There are no distinct dorsal 
processes on the antennal protopodite (fig. 84). 

The mandibular palp is composed of two or three segments. 

There is a stout tooth on the dorsal edge of the raptorial carpus at its distal end. 
The propodus bear only two mobile spines at the proximal end of its upper margin ; 
the outer of the two edges opposed to the dactylus is furnished with a series of fine 
pectinations on its basal two-thirds. The dactylus consists of a single stout spine, at 
the base almost square in section. Its inner margin is microscopically serrate, but 
is without trace of the large teeth found in most species of the genus. 

The margins of the sixth and seventh thoracic somites are rounded and reflected a 
little outwards. The postero-lateral angles of the abdominal somites are not spinous. 
The first four somites are smooth; the fifth has a shallow longitudinal depression on 
either side ; the sixth has eight blunt longitudinal carinae which do not terminate in 
spines. The two submedian pairs are obscure or obsolete in very large specimens. 

On the telson there are five longitudinal carinae, a median, one pair of parallel 
intermediates and the marginals. The median carina does not terminate in a spine. 
The marginal teeth are large and broad and each is strengthened by a short rounded 
keel. The submedians have small movable tips. There are no submedian or lateral 
denticles; but between the submedian and intermediate teeth there are one or two 
conspicuous rounded lobes (fig. 85). 


1013.| S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 107 


The peduncular segment of the uropods is externally ridged, but does not possess 
a spine at the distalend of the upper margin. The ventral process is strikingly different 
from that of any other known Stomatopod. It terminates in a stout tooth, homologous 
with the inner one of the two found in P. ciliata, and the external margin behind the 
apex of this tooth is convex and is continued evenly backwards to the point of articu- 
lation of the exopod, on this margin near its distal end is found a minute spinule 
representing the outer tooth of P. ciliata and the other species. The internal margin of 
the process is smooth.' The basal segment of the exopodite is extremely short and bears 
four or five mobile spines on its external margin, the outermost much the longest. 
Both the endopod and ultimate segment of the exopod are very large and broadly 
expanded ; the latter is fully two and a half times the length of the proximal seg- 
ment (fig. 85). 

The colouring of spirit specimens is not characteristic; Whitelegge notes that the 
outer laminae of the uropods are brilliant violet in living examples. 


In two large specimens from the Australian coast the mandibular palp is two- 
jointed * and there are two prominent lobes between the submedian and intermediate 


Pseudosquilla stylifera. Yast abdominal somite, telson and uropods of an Australian specimen. 


marginal teeth of the telson (see text-fig.). In a slightly smaller example from Chili 
there is only a single intermediate lobe on the telson (fig. 85), and the mandibular palp 
is composed of three quite distinct joints. 

Both Milne-Edwards and Bigelow, in their figures of S. American specimens, show 
only a single intermediate lobe on the telson ; but Whitelegge does not mention the 
existence of two in the examples which he records from New South Wales, while Milne- 
Edwards figures the mandibular palp of a Chilian example in one case with two seg- 
ments (1837, pl. 27, fig. 10), and in another with only a single segment (ibid., fig. 11). 

There can be little doubt that the Australian and American specimens both belong 
to the same species, for, with the exception of the two characters noticed above, the 
examples from the two localities are, as nearly as possible, identical. ‘The distinction 
in the number of segments in the mandibular palp is consequently the more remarkable, 


' Miers states, perhaps in error, that this margin is denticulated. 
* The two distal segments of the normal three-segmented palp are fused. 


108 Memoirs of the Indian Museum. Worse 


for among other Stomatopoda in the Indian Museum any two species which show 
very close structural affinity invariably agree in this character. It is not improbable 
that distinct races of P. stylifera exist on the coasts of Australia and America, but this 
cannot be established with any certainty without the examination of large numbers 
of specimens from the two localities. 

The three examples in the Indian Museum are registered as follows :— 


7583 


m Disaster Bay, Victoria. .. Australian Museum. 50, BS, US, WHO sana, 


120 : a ; 
=, Coquimbo, Chili. .. Berlin Museum. Lo LG), OB an, 


Pseudosquilla stylifera has been recorded from Chili (Milne-Edwards, Nicolet, 
Miers), from California (Bigelow, Holmes) from the Hawaiian Is. (Randall), and from 
Newcastle Bight, New South Wales (Whitelegge). It is strange that the species 
has not been found on the New Zealand coast. 


Post-larval stages of Pseudosquilla. 


Two small specimens, 19 and 27 mm. in length, belong to what has been termed 
the ‘ monodactyla’ stage of this genus.' Hansen has shown that this form is not a 
distinct species, as was supposed by Milne-Edwards and Miers; but is a ‘ zwischen- 
stadium ’ in the development of members of this genus. In the absence of teeth on the 
raptorial dactylus and of carinae on either side of the median crest of the telson, the 
appearance of these forms is so strikingly dissimilar to that of the adult species that 
the mistake made by the earlier writers is not astonishing. 

Specimens of Pseudosquilla oculata in the ‘ monodactyla ’ stage have been recorded 
of lengths varying from 28 to 34 mm., while those of a similar stage of P. ciliata range 
from 16 to 21°3 mm. 

I am uncertain as to the species to which the two examples in ae Indian Museum 
should be referred. It is probable that they belong to P. ciliata since that is the only 
species of Pseudosquilla known from the localities in which they were found. 

It will be noticed that among the recorded specimens of P. ciliata (p. 100 ) are 
two specimens which also measure only 19 and 27 mm. in length. These examples 
possess all the adult characteristics ; but their existence does not necessarily prove 
that the ‘ monodactyla’ of the same size cannot belong to that species, for it is very 
probable that an actual shrinkage in total length may take place at the close of the post- 
larval stages. 

The examples agree closely with the individual, 20 mm. in Le which de Man 
has recorded from Amboina, and differ in the various points which have been men- 
tioned by that author from the description and figures given by Miers. 

The specimens are registered as follows :— . 

Croate: 02 00% IN, Gx? 10 By PO Investoaton 9 oz 


6 
cn Port Blair, Andaman Is. .. J. Wood-Mason. .. I2,Igmm. 


! For literature dealing with these post-larval stages see A. Milne-Edwards, 1877-8, p. 232; Miers, 
880, p. 110, pl. iii, figs. 1,2; de Man, 1887, p. 571, pl. xxii a, fig 6: Hansen, 1895, p. 85; Bigelow, 
1902, p. 156, and Nobili, 1906, p. 336. 


LORS] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 109 


Genus Lysiosquilla, Dana. 
1825. Coronis, Latreille, Encycl. Method., X, p. 474 (nom. praeocc.). 
1837. Squilla (Squilles fine-tailles, rst sect.), H. Milne-Edwards, Hist, Nat. Crust., II, p. 518. 
1837. Coronis, H. Milne-Edwards, ibid. p. 530. 
1849. Squilla (sect. Maculatae), De Haan, in Siebold’s Fauna Japonica, Crust., p. 220. 
1852. Lysiosquilla, Dana, U.S. Explor. Exped., Crust., p. 616. 
1880. Lysiosquilla, Miers, Ann. Mag. Nat. Hist. (5), V, p. 5. 
1894. Lysiosquilla, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 302. 
1910. Lysiosquilla, Giesbrecht, Faun. Flor. G. v. Neapel, XX XIII, p. 44. 

Carapace littleif at all narrowed anteriorly ; cervical groove wholly absent orscarcely 
traceable, never continuous from side to side; longitudinal carinae absent ; antero- 
lateral angles never spinous. Cornea of eyes small or considerably expanded. Mandi- 
bular palp composed of three segments. Merus of raptorial claw articulating terminally 
with ischium ; ventral surface of merus longitudinally hollowed throughout its length ; 
upper margin of propodus finely pectinate ; dactylus not inflated at base, armed with 
teeth on inner margin. Penultimate segment of third and fourth thoracic limbs close to 
insertion of ultimate segment flattened and transversely ribbed. Shorter. ramus of last 
three thoracic limbs composed of two segments, basal segment short, ultimate segment 
longer, linear or oval. Free thoracic and abdominal somites flat, depressed and with- 
out longitudinal carinae. Telson broader than long, in Indo-pacific forms of two 
distinct types. First type without dorsal spines and with posterior margin unarmed 
or cut into a few large blunt teeth. Second type with a curved transverse tow of spines 
or prominences near distal end, with sharp primary teeth, a number of submedian 
spinules and a few intermediate denticles. One slender movable spine situated beneath 
submedian teeth or wholly replacing them. Ventral process of uropods consisting of 
two large spines, inner longer than outer ; inner spine rarely suppressed. 

Some of the species are transversely barred with deep blue-black or black pigment. 

Secondary sexual distinctions are usually absent ; but the raptorial claw of very 
large female L. maculata and L. glabriuscula is modified. 

In some, perhaps in all, species of the genus soft elongated papillae are found 
on the antennal protopodite. These are well-developed in L. maculata and are three 
in number, one situated antero-dorsally and curved backwards and inwards towards 
the base of the rostrum and two on the ventral surface at the base of the endopod, 
a long one directed forwards and a much shorter one pointing backwards. In 
L. acanthocarpus, L. mulhfasciata, L. spinosa and L. insignis only one of these pro- 
cesses is found, placed on the ventral surface and pointing outwards or backwards. 

One Indo-pacific form, L. maculata, is found in the Atlantic, but appears to be 
very scarce in those waters ; another is so closely allied to L. biminiensis from the 
Bahamas that it has only been accorded subspecific rank. With these exceptions no 
Indo-pacific species of the genus is known to occur in the Mediterranean and Atlantic 
or on the Pacific coasts of America. 

The twelve species found in the Indo-pacific fall readily into two groups distinguished 
at a glance by the presence or absence of a transverse dorsal series of spines near the 
distal end of the telson. The first group, comprising those species without these spines, 


IIO Memoirs of the Indian Museum. [VoL. IV, 


includes L. maculata, the only common Indo-pacific representative of the genus, and 
three species described from unique specimens. One of the latter, L. crassispinosa, 
Fukuda, furnished with a remarkably large number of movable spines on the pen- 
ultimate segment of the raptorial claw, appears to be an isolated form, butmay perhaps 
find allies among the less well-known Atlantic species. The typical representative of 
the second group is L. acanthocarpus. L. insignis and L. latifrons form a small highly 
specialized section characterized by the elaborate ornamentation of the telson, while 
L. spinosa, in which the inner spine of the ventral process of the uropods is suppressed, 
seems to have no close affinity with any other known species. 

Practically nothing is known of the habits of Indo-pacific species of Lysiosguilla. 
Brooks, however, notes that those of L. excavatrıx are very different from those of 
Squilla empusa, and it is probable that his observations afford an explanation of the 
extreme scarcity of many species. L. excavatrıx inhabits much deeper burrows than 
S. empusa ; it lies in wait for its prey at the mouth of the burrow and seldom ventures 
more than a few inches away. Sguwilla empusa on the contrary wanders about and is 
in consequence caught without much trouble. The burrows of L. excavatrıx extend 
vertically downwards for a distance of several feet and Brooks found it almost im- 
possible to obtain them by digging. Under these conditions it will readily be under- 
stood that the difficulty of obtaining specimens is very great and when the species 
inhabits comparatively deep water, as some are known to do, it can only be by the 
merest chance that a specimen is caught at all. 

The greatest depth at which any species of the genus has been found is 235 
fathoms (L. insignis). 

Many undescribed forms doubtless remain to be found and a number of those that 
are already known stand in need of redefinition. 


Key to the Indo-pacific species of Lysiosquilla. 
I, Telson without dorsal spines; limbs of sixth and seventh thoracic somites 
with shorter ramus linear. 
A. Upper margin of raptorial propodus with not more than four movable 
spines; postero-lateral angles of first five abdominal somites not 
spinous. 
I. Raptorial dactylus with eight to eleven teeth.! 
A. Exposed thoracic somites without longitudinal wrinkles; sixth 
abdominal somite not grotesquely sculptured; terminal 
tooth of raptorial claw not dilated at apex. 
I. Rostrum cordiform, at least as wide in front of base as at 
base, not grooved near margin; raptorial dactylus with 
ten or eleven teeth! in male De ale .. maculata, p. III. 
2. Rostrum triangular, widest at base. with a deeply-incised 
antero-lateral groove; raptorial dactylus of male with 
eight teeth! ae a ER .. do. var. sulcirostris, 


{Ds 26160 
B. Exposed thoracic somites irregularly and longitudinally wrink- 


Including the terminal tooth. 


1013.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 
led; sixth abdominal somite grotesquely sculptured; ter- 
minal tooth of raptorial claw dilated at apex 
II.- Raptorial dactylus with fifteen teeth! 


B. Upper margin of raptorial propodus with a series of ten or more movable 
spines; postero-lateral angles of abdominal somites spinous 
II. limbs of 


sixth and seventh thoracic somites with shorter ramus broadly ovate. 


Telson with a transverse row of dorsal spines near distal margin; 
A. Only three dorsal spines in a transverse row on telson in adults; outer 
spine of ventral process of uropods much longer than inner 
B. More than three dorsal spines in a transverse row on telson in pace 
inner spine of ventral process of uropods longer than outer. 
I. Five dorsal spines on telson; median spine simple and acute. 
A. Rostrum with median spine and rounded or subrectangular 
antero-lateral angles. 
I. Raptorial dactylus with six or seven teeth ;! postero-inferior 
margin of last abdominal somite without spines; 
with three pairs of large marginal teeth. 


telson 


a. Penultimate tooth of raptorial dactylus much shorter 
than antepenultimate. 

i. Raptorial dactylus with two lobes of about equal 

size at base of external margin Do 

ii. Raptorial dactylus with a very large distal and a 

small proximal lobe at base of external margin .. 

b. Penultimate tooth of raptorial dactylus longer than 

antepenultimate .. 


2. Raptorial dactylus with eleven teeth; ! postero-inferior 
margin of last abdominal somite with a series of spines; 
telson with four pairs of large marginal teeth 

B. Rostrum with sharp median and antero-lateral spines [telson 
with four pairs of large marginal teeth] se 

II. Five dorsal spines on telson, median one trilobed, or seven distinct 

and separate spines. 

A. Rostrum more than one and a half times as long as broad; eyes 
large: a pair of posterior spines on sixth abdominal somite 
in addition to those at postero-lateral angles 

B. Rostrum broader than long; eyes small; sixth no 


somite with spines at postero-lateral angles only 


1, Lysiosquilla maculata (Fabricius). 


Plate VIII, figs. 86-91. 


. Squilla maculata, Fabricius, Ent. Syst., II, DST 


. Squilla maculata, Fabricius, Ent. Syst. Suppl., p. 415. 
. Squilla maculata, Lamarck, Hist. Nat. Anim. sans Vert., V, p. 188. 
. Squilla maculata, Desmarest, Dict. des Sci. Nat., XXVIII, p. 341. 


! Including the terminal tooth. 


III 


miersi, p. TIO. 
capensis, p. 117. 


crassispinosa, p. 117. 


spinosa, p. 118. 


acanthocarpus, p. 120. 
multifasciata, p. 122 
biminiensis subsp. pact- 

ficus, p. 124. 
tigrina, p. 125. 


vicina, p. 126. 


insignis, p. 126. 


latifrons, p. 128 


Squilla arenaria terrestris or locusta, Rumphius, Amboinsche Rariteitkamer, p. 4, pl. iii, fig. R. 


. Cancer (mantis) arenarius, Herbst, Krabben u. Krebse, II, p. 96, pl. xxviii, fig. 2. 


TI Mémoirs of the Indian Museum. NVorenyz 


1825 Squiila maculata, Latreille, Encycl. Method. X, p. 470. 

1837. Squilla maculata, H. Milne-Edwards, Hist. Nat. Crust., II, p. 518, pl. xxvi, fig. II. 

1838. Squilla maculata, H. Milne-Edwards, in Lamarck’s Hist. Nat. Anim. sans Vert., 2nd ed., V, 
P- 323. 

1839. Squilla arenaria, Randall, Journ. Acad. Sci. Philadelphia (r), VIII, p. 146. 

1847. Squilla maculata, White, List Crust. Brit. Mus., p. 83. 

1849. Squilla maculata, De Haan, in Siebold’s Fauna Japonica, Crust., p. 221. 

1852. Lysiosguilla maculata, Dana, U.S. Explor. Exped., Crust., p. 616. 

1877. Lysiosquilla maculata, Miers, Proc. Zool. Soc., p. 138. 

1879. Lysiosquilla maculata, Miers, Phil. Trans. Roy. Soc., CLXVIII, p. 494. 

1880. Lysiosquilla maculata, Miers, Ann. Mag. Nat. Hist. (5), V, p. 125, pl. i, figs I, 2 and p. 458. 

1881. Lysiosquilla maculata, Lenz and Richters, Abhandl. Senck. Ges., Frankfürt, XII, p. 428. 

1886. Lysiosquilla maculata, Brooks, Voy. H.M.S. ‘Challenger,’ X VI, Stomatop., p. 45, pl.x, figs. 1-7. 

1887. Lysiosquilla maculata, de Man, Arch. f. Naturgesch., LIII, i, p. 571. 

1892. Lysiosquilla maculata, de Man, in Weber’s Zool. Ergebn. Nied. Ost-Ind., II, p. 518. 

1892. Lysiosquilla maculata, Thallwitz, Abhandl. kön. Mus. Dresden, No. 3, p. 55. 

1893. Lysiosquilla maculata, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 452. 

1894. Lysiosquilla maculata, Bigelow, Proc. U. S. Nat. Mus., xvii, p. 508. 

1894. Lysiosquilla maculata, Ortmann, Denk. med.-wiss. Ges. Jena, VIII, p. 60. 

1895. Lysiosquilla maculata, Hansen, Isop. Cumac. u. Stomatop. Plankton. Exped., p. 74. 

1898. Lysiosquilla maculata, Borradaile, Proc. Zool. Soc., p. 37 (habits). 

1898. Lysiosguilla maculata, Whitelegge, Proc. Linn. Soc. N.S.W., XXIII, p. 368. 

1899. Lystosqiilla maculata, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 276. 

1902. Lysiosquilla maculata, Stebbing, S. African Crust., II, p. 46. 

1902. Lysiosquilla maculata, de Man, Abhandl. senck. Ges. Frankfürt, XXV, p. 910. 

1910. Lysiosquilla maculata, Balss, Abh. math.-phys. Klasse k. bayer. Akad. Wiss, Suppl., Bd. II, 
Abh. 2, p. 5. 

1910. Lysiosquilla maculata, Fukuda, Annot. Zool. Japon., VII, p. 146, pl. iv, fig. 3. 

1910. Lysiosquilla maculata, Stebbing, Ann. S. African Mus., VI, p. 406. 


The whole dorsal surface of the animal is smooth and polished. The carapace is 
strongly convex from side to side and is almost as broad anteriorly as posteriorly. 
There are no carinae on the carapace. The gastric grooves are sharp and distinct, 
but the cervical groove is obsolete and is represented merely by a short and shallow 
depression on the outer side of the gastric. The lateral margins are convex, the antero- 
and postero-lateral angles are rounded and the posterior margin is deeply concave. 

The rostrum (fig. 86) is broader than long, cordiform and widest at a point some- 
what in advance of its proximal end. ‘The lateral margins are not upturned and con- 
verge rapidly to an acute, but rounded, apex behind which they are often slightly con- 
cave. ‘The dorsal surface is smooth except for a blunt longitudinal carina in the 
anterior third. . 

The eyes are large. The breadth of the cornea is about equal to the greatest length 
of the cornea and stalk combined, and the corneal and peduncular axes are very oblique. 
On its outer margin the eyestalk bears a small but not inconspicuous tubercle. The 
anterior margin of the ophthalmic somite is sharply pointed, and its dorsal processes 
consist of a pair of upstanding lobes which terminate subacutely on either side of the 
rostral apex. | 


1903.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. TT 


The antennular peduncle is considerably less than half the length of the carapace, 
excluding the rostrum. There are three soft papillae on the antennal protopodite 
(see p. 109). The mandibular palp is composed of three segments. 

The outer inferior margin of the merus of the raptorial claw is rounded distally. 
The dorsal carina of the carpus is blunt and terminates in a sharp spine which may 
overhang the anterior margin. In males and young females the width of the propodus 
is nearly one quarter its greatest length ; the margin in contact with the dactylus bears 
a series of fine pectinations externally, while internally at its proximal end it is pro- 
vided with four movable spines. The dactylus is slender and bears nine to eleven 
teeth (usually ten, very rarely nine), including that at the apex. Thelatter is usually 
more than twice the length of the next of the series. The outer margin of the dactylus 
is straight or slightly concave (fig. 91). 

In very large females, as has been noticed by Miers, the form of the raptorial claw 
is strikingly different (fig. 90). In a specimen of 186 mm. the carpus and basal part 
of the propodus bear tufts of long hairs, and the latter segment is much wider than in the 
male, being more than one-third as broad as long ; it is, moreover, provided with only 
two movable spines at the proximal end of its upper margin. The dactylus also is 
strongly convex externally, and the armature consists only of seven or eight nicks 
along the inner margin in addition to the strong terminal tooth. 

The third and fourth thoracic somites are exposed in dorsal view. The lateral 
margin of the fifth is transversely channelled by a broad and shallow groove, separating 
blunt anterior and posterior carinae which converge as they approach the inferior aspect 
of the somite. The lateral margins of the sixth and seventh somites are rounded in front 
and subrectangular behind. The shorter ramus of the last three thoracic limbs is linear. 

Near the anterior margin of each post-abdominal segment except the first and last 
there is a short and rather faint longitudinal furrow which divides the somite anteriorly 
into three sub-equal portions and, on either side of the middle line, a very shallow 
depression is sometimes visible delimiting a very faint longitudinal elevation. ‘The 
lateral margin of each somite (except the last) is concave, and its postero-lateral angle 
is rounded. The sixth somite has a transverse groove on either side of the middle 
line near its anterior edge, an oblique groove external to this and beyond this again a 
shallow longitudinal furrow which defines a swollen lateral lobe. In addition there are 
sometimes a number of obscure and irregular elevations and depressions on either side 
of the smooth median portion of the somite. On its inferior margin the sixth somite 
bears a single strong tooth at each antero-lateral angle, projecting backwards over the 
- base of the uropod. 

The telson is nearly twice as broad as long, convex above and concave below. 
On both dorsal and ventral surfaces it presents a feeble median triangular elevation 
limited by a pair of very shallow posteriorly-convergent grooves. On either side there 
is a large oval patch of more or less regularly disposed pits which are in some cases 
scarcely visible, but usually are specially conspicuous in the neighbourhood of the 
lateral and antero-lateral margins. Along the anterior and antero-lateral edges of the 
telson there is a well-marked groove, which is interrupted in the middle line and fades 


II4 Memoirs of the Indian Museum. Worle 


away laterally before reaching the external marginal tooth. The ventral surface is 
obscurely pitted on either side. The posterior margin of the telson is indistinctly 
notched in the middle line and bears on each side two, three, or more usually four, 
blunt lobes, the outermost or two outermost being generally sharper and more con- 
spicuous than the rest. 


The inner dorsal edge of the peduncular segment of the uropod terminates in a 
short spine. The inferior bifurcate process consists of two long spines. Each is 
triangular in section, bearing three longitudinal grooves and three carinae ; the inner 
is nearly twice the length of the outer and is considerably longer than the segment 
from which it arises. The proximal segment of the exopod bears a fixed distal spine on 
its inferior aspect and a series of eight or nine, which are movable, on its outer margin. 


In the genus Squilla marked secondary sexual characters, when present, are found 
in the male only and differentiate large examples of that sex from females and young 
males. In L. maculata, however, the reverse is the case, special modifications occurring 
only in large females. Among specimens in the Indian Museum the peculiar form of 
the raptorial claw ' (mentioned above) is found only in a single female which measures 
186 mm. In another scarcely smaller example (182 mm.) the raptorial teeth, though 
somewhat reduced in size, are still well formed and there are, as usual, four movable © 
spines on the propodus. In both these females the eye is small and it seems probably 
that this again is a special feature of large examples of this sex (c/. figs. 87 and 88). 
The following table contains measurements of five females and seven males. It will 
be noticed that, as in all Stomatopoda, the relative size of the eye diminishes with 
growth and that, whereas the proportions in young females are closely comparable with 
those of males, the breadth of the cornea in the two large females is very much less 
than would be expected to occur in males of a similar length. 


Corneal Index.? 
Total length. Sex. | Length of carapace. De of 
F ? 
m: mm, mm. 
283 J 53°5 ae 20 
208 d 39 © 3:9 
186 2 30 ae > an 
182 2 32 DE 
144 d' 27 Te 27 
> À a 6°5 3°7 : 
123 C4 22°5 6a 3-6 à 
112 CA 20°5 "5 3:7 = 
a 5 129 5%: 2e 3°5 
100 2 18-5 5 BR à 
% 2 175 53 33 . 
2 1 145 25 à pe 
5 2 En 45 29 ie 


" De Man’s sugsestion that the teeth are merely worn down in large females cannot be supported, 
for the whole form of the propodus and dactylus is altered 


* The number of times that the breadth of the cornea is contained in the median length of the 
carapace (see p. Q). 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 115 


The colouring of L. maculata is very striking and is usually distinct in spirit speci- 
mens. There are, as a rule, three broad transverse bands of blue-black pigment on the 
carapace; the fifth thoracic and last abdominal somites are blue-black, while the other 
segments of the post-abdomen are broadly bordered both anteriorly and posteriorly with 
the same colour. A typical scheme of colour is shown in fig. 88 ; sometimes the 
pigmentation is better developed (fig. 87) and occasionally it is much less marked, with 
narrow bands on the carapace, the fifth thoracic somite entirely pale, and a large pale 
patch on the last abdominal somite. ‘The telson may show threelarge blue-black spots, 
isolated (as in Herbst’s figure), or partially connected with one another; more usually 
there is a pale anterior transverse band and a pale bilobed distal patch. In one speci- 
men the anterior band is divided by oblique bars of pigment into three separate spots 
(fig. 89). In the uropods the spines of the bifurcate process are blue-black distally as is 
also the inner exopodal segment. The peduncular segment is blue-black proximally 
and the outer segment of the exopod is divided obliquely into dark proximal and pale 
distal areas. The endopod is blue-black except for a narrow transverse band at the base. 


The specimens in the Indian Museum are registered as follows :— 


rs Fiji Is. Australian Museum. 19,182 mm. 
TS New Guinea. Australian Museum. 1,283 mm. 
= Christmas I., Polynesia. D. G. Stead. (Raptorial claws only.') 
ue Nicobars. F. A. de Roepstorff. TOR. 
3031 Andamans. Homfray and ‘Tytler. Id, C4. 93 mm. 
om Andamans. B. Ford. (Fragment.) 
= Madras. Madras Museum. 23, 137, 144 mm. 
HE Coconada, Madras Pdy. G. W. Wicks. 63,32, 69-185 mm. 
3037 (No history.) i ©, Oe onen), 


5 


The following specimens from other sources have also been examined :— 
Christmas I., Polynesia. D. G. Stead. Id, 270 mm. 


Ceylon. Colombo Museum. 24,29 ,09-251 mm. 


Lysiosquilla maculata seems to occur only in shallow water ; though never found 
in abundance, it has a wide Indo-pacific distribution extending from Japan and Oceania 
to South Africa. It has been recorded from the following localities: Japan (De Haan, 
Fukuda), Hawaiian Is. (Randall, Miers), Marquesas Is. (Bigelow), Duke of York Is. 
(Miers) , Samoa (Miers, Thallwitz, Bigelow, Balss), Fiji (Miers), Rotuma (Borradaile), New 
Guinea (Nobili, Thallwitz, Whitelegge), Amboina (Brooks, de Man, Ortmann, Nobili), 
Goram (Miers), Batjan (de Man), Flores (Thallwitz), Celebes (de Man), Philippine Is. 
(Miers, Brooks), Penang (Miers), Tuticorin and Madras (Henderson), Mysore (Thallwitz), 
Rodriguez (Miers) and Durban (Stebbing). 


1 The raptorial dactyli are 80 mm. in length, and must have belonged to a specimen measuring 
over 300 mm. 


II6 Memoirs of the Indian Museum. orne 


In the Atlantic this species has been recorded from Antigua, W. Indies (Stebbing) ' 
and the larva, Lysierichthus duvaucellit, Guérin, has been recorded by Hansen (1895) 
from the vicinity of the Cape of Good Hope (32° 30’ S., 15° 0’ E.). 


var. sulcirostris, nov. 
Plate VIII, figs. 92, 93. 

This variety differs from typical specimens in the following points :— 

1. The rostrum is triangular, widest at the base, and is produced to a much sharper 
point than in typical L. maculata. In the anterior two-thirds there is on each 
side a sharply-defined and deep groove running parallel to the lateral margin 
(fig. 92). 

2. The dactylus of the single male specimen resembles that of typical male ma- 
culata, but has only eight teeth including the terminal one (fig. 93). 

3. The sixth abdominal somite is grooved as in the typical form, but much less 
distinctly, and the irregular wrinkling on either side is absent. 

The colour pattern is closely similar to that of L. maculata, but the dark trans- 

verse bands are much narrower than in typical examples of the species. 

L. sulcirostris is perhaps specifically distinct from L. maculata, but its resemblance 

to the latter form is so great that I hesitate to adopt such a course on the evidence of 
a single specimen. 


a Andaman Is. .. Homfray and Tytler. oC 5 WU) warn, 


Apparently found in company with a typical specimen of L. maculata. 


2. Lysiosquilla miersi, De Vis. 
1883. Lysiosguilla miersii, De Vis, Proc. Linn. Soc. N.S.W., VII, p. 321. 
1894. Lysiosgwilla miersii, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 504. 

De Vis’ description of the only known specimen runs as follows :— 

‘“ Carapace smooth, in the form of a truncate isosceles triangle, rounded at the 
posterior lateral angles. Rostrum elongate, cordiform, acuminate, with a median ridge 
on its produced limb. Exposed segments with irregular longitudinal wrinkles on the 
dorsum, which is slightly depressed, the wrinkling and depression becoming more pro- 
nounced posteriorly. Anirregular semi-lunar depression on each segment near the lateral 
line. Penultimate segment, narrow, deeply sculptured in a grotesque fashion. Telson 
smooth, with a bold sagittate median ridge and four teeth on the lateral margin, of which 
the anterior two are strong. Ten long sharp teeth on the dactyli of the raptorial limbs 
including the terminal one, which near the tip is anteriorly dilated and presents a sharp 
edge. All the teeth finely serrated on the posterior edge. Four long sharp teeth on 
the propus, the posterior one moderately recurved. 


! According to Stebbing’s account the specimen from this locality differs from any Indo-pacific ex- 
ample that I have seen in having the fifth pleon segment denticulate along the hind margin, the sixth 
denticulate in an arched proximal! band and round the distal margin, and the telson with three spines on 


each side and the truncate portion cut into five square teeth on one side and six on the other side of a small 
median emargination. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacıfic Region. Tel 


Colour light horn-brown. Exposed thoracic segments with a broad black basal 
band. Post-abdominal segments with a similar band at each suture. Carapace and 
penultimate joint of raptorial limbs with three broad bands, appearing beneath the sur- 
face, a large black blotch on each side of the telson, and on the rami of the appendages 
of the sixth segment ; a conspicuous white longitudinal band on each side of the 
dorsum.’’ 

As noted by De Vis this species belongs to the maculata section of Lysiosguilla. It 
seems to be easily distinguished by the irregular wrinkling on the exposed thoracic 
somites, by the sculpture of the last segment of the abdomen and by the dilated apex 
of the terminal spine of the raptorial claw. 

L. miersi is known only from a single specimen (sex and length not noted) found on 
sand banks at Moreton Bay, Queensland. 


3. Lysiosquilla capensis, Hansen. 


1895. Lysiosquilla capensis, Hansen, Isop. Cumac. u. Stomatop. Plankton. Exped., p. 74. 
1910. Lysiosquilla capensis, Stebbing, Ann. S. African Mus., VI, p. 406. 


Hansen’s short description of this form runs as follows: “ eine Art, die von allen 
anderen dadurch leicht zu unterscheiden ist, dass sie ausser dem Enddorn 14 Dornen 
an dem Dactylus besitzt, 7 Uropoddornen und Telson wie bei Lys. maculata (F.) geformt 
(x Exemplar von Port Elisabeth im Museum zu Strassburg).’’ A fuller account of 
this specimen is badly needed. 

Lysierichthus pulcher, Hansen, which according to its author is the larva of L. 
capensis, is recorded from the Atlantic, N.W. of the Cape of Good Hope (32° 30’S., 
ROUE) 


4. Lysiosquilla crassispinosa, Fukuda. 
1910. Lysiosquilla crassispinosa, Fukuda, Annot. Zool Japon., VII, p. 146, pl. iv, figs. 4, 4a. 


Fukuda gives the following diagnosis of this species :—‘‘ Eyes with the corneal 
region nearly globular. The manus of raptorial limb bearing, besides the usual serra- 
tion, ten or more acute jointed spines on the inner margin ; dactylus armed with ten 
teeth including the terminal one. Rostrum triangular, terminating in an acute median 
spine. Carapace smooth, with all its angles rounded. Last two thoracic and first two 
abdominal segments provided with a pair of inconspicuous carinae. All abdominal 
segments with their postero-lateral angles ending in spines. The whole dorsal surface 
of the last two segments and a part of that of the fifth abdominal segment covered 
with irregularly shaped granules. Hight stout spines or lobes on the postero-lateral 
margin of the telson, the median crest of which is broad and somewhat obscurely 
defined. ‘The inner spine of the dorsal prolongation of uropod much longer than 
outer.’’ For further details Fukuda’s long and careful description should be consulted. 

The unique example of this curious species, afemale 275 mm. in length, is recorded 
from the Sagami Sea, Japan. 


118 | Memoirs of the Indian Museum. VOLE 


5. Lysiosquilla spinosa (Wood-Mason). 
Pate WIQDL, HS, C4. 
1847. Coronis tricarinata, Gray MS., White, List. Crust. Brit. Mus., p. 85 (sine desc.). 
1875. Coronis spinosa, Wood-Mason, Proc. As. Soc. Bengal, p. 232 ; reprinted in Ann. Mag. Nat. 
Hist. (4) XVII, p. 263 (1876). 
1878. Squilla indefense, Kirk, Ann. Mag. Nat. Hist. (5), II, p. 466. 
1879. Squilla indefensa, Kirk, Trans. N. Zealand Inst., XI; p. 394 (text-fig.) and p. 401. 
1879. Squilla laevis, Hutton (non Hess), Trans. N. Zealand Inst., XI., p. 340. 
1880. Lysiosquilla spinosa, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 12, 125, pl. i, figs. 10-12. 
1882. Squilla tridentata, Thomson, Trans. N. Zealand Inst., XIV, p. 230. ; 
1886. Squilla indefensa, Fihol, Miss. de l’île Campbell, III, 2e, p. 436, pl. lv, fig. 3.1 
1886. Squilla laevis, Filhol, ibid. p. 491. 
1891. Lysiosquilla spinosa, Chilton, Trans. N. Zealand Inst., XXIII, p. 61, pl. x, figs. 1-3. 
1894. Lysiosquilla spinosa, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 503. 
1894. Lysiosquilla (?) tridentata, Bigelow, ibid. p. 503. 
1895. Lysiosquilla spinosa, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 1, pl. i, figs.1-3. 
1901. Lysiosquilla spinosa, Lanchester, Proc. Zool. Soc., II, p. 554. 
1909. Lysiosquilla spinosa, Chilton, in Subantarctic Is. of N. Zealand, II, p. 615. 
1910. Lysiosquilla spinosa, Chilton, Trans. N. Zealand Inst., XLIII, p. 139. 

The carapace and abdomen are smooth and polished dorsally and the former is con- 
siderably wider posteriorly than anteriorly. The gastric grooves are distinct through- 
out their course and the cervical is represented by a short transverse depression on 
either side as in L. maculata. The antero- and postero-lateral angles are broadly 
rounded and the posterior margin is deeply concave. The rostrum is perfectly smooth 
dorsally ; it is a little broader than long and its strongly convex lateral margins con- 
verge to a small but acute apex. 

The corneal portion of the eyes is a trifle wider than the stalk, not bilobed, and the 
corneal and peduncular axes are slightly oblique with regard to one another. ‘The 
anterior margin of the ophthalmic somite is truncate but, in dorsal view, is entirely 
concealed by the rostrum. The dorsal processes of the somite are for the same reason 
quite invisible. The antennular peduncle is less than one and a half times the 
length of the carapace, excluding the rostrum. The dorsal processes of the antennular 
somite are acute and reach to the base of the eyestalks. The mandibular palp is com- 
posed of three segments. 

The outer inferior margin of the merus of the raptorial claw is subrectangular, but 
rounded distally. The dorsal edge of the carpus is not carinate but its upper margin 
ends in a sharp spine overhanging the propodal articulation. There are three movable 
spines at the base of the pectinate edge of the propodus. ‘The dactylus is sinuous ex- 
ternally with a very small outstanding proximal lobe. It bears from nine to fourteen 
teeth * (including the terminal one) which increase regularly in size from the base to the 
apex. 


! A bad reproduction of Kirk’s rough text-figure. ü 

* Of the two specimens examined, the type has ten spines, while the other has ten on one side and 
eleven on theother. The type of Thomson’s S. tridentata has only four teeth on the raptorial dactylus; 
but the specimen is exceedingly small; only '75 inches in length. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. II19 


The lateral margins of the fifth thoracic somite are greatly compressed antero- 
posteriorly and form a thin outstanding blade which, in dorsal view, looks like a spine. 
The sixth and seventh somites are rounded antero-laterally, while their postero-lateral 
angles are almost rectangular. The basal segment of the last three thoracic limbs bears 
a short stout tooth on its external aspect. The first five abdominal somites are dorsally 
smooth and their postero-lateral angles do not project backwards as spines. The sixth 
somite possesses a shallow longitudinal groove on either side and its postero-lateral 
angles are sharply spinous. > 

The telson is semicircular in shape and is more than twice as broad aslong. On its 
dorsal surface near the distal margin it is provided with a transverse series of three 
short but acute spines ; the two lateral form the terminations of carinae, while the 
median is situated at the end of a rather conspicuouslobe. The submedian, intermediate 
‚and lateral marginal spines are well developed. The submedians are movable and the 
edge between them, which is slightly emarginate, bears four to seven minute denticles. 
There are two spines, the outermost of which is the largest, between the submedians 
and intermediates, and one between the intermediate and the lateral. The ventral 
surface of the telson is smooth. 

The basal segment of the uropod bears a single dorsal spine on its distal margin, 
projecting over the articulation of the inner segment of the exopod. The outer spine 
of the bifurcate process (fig. 94), is enormously larger than the inner ; it is broad, very 
greatly expanded laterally, and is deeply keeled ventrally. The inner spine of the 
process is, in the type specimen, represented merely by a small rounded lobe ; it is 
larger and sharply spinous in an example from New Zealand. In addition, there is 
another small ventral spine at the inner distal angle of the basal segment which pro- 
. jects over the base of the inner uropod. The proximal segment of the exopod bears 
a single spine ventrally and distally and a series of five or six, which are movable, along 
its outer margin. Of these spines the two outermost are long and recurved. 


Chilton (1891, p. 66) has given an account of the adult colouration. There is, it 
appears, a very great difference in this respect between the two sexes, for the male is 
greyish in general tone, while the carapace of the female is similar, but darker, and the 
whole of the central part of the abdominal segments is coloured a bright red. Lan- 
chester (1901) notes that the small specimens which he examined were suffused with 
rosy red and sprinkled with red and black spots, the latter forming two conspicuous 
black marks over the submedian marginal spines of thetelson. From these two accounts 
it is clear that the transverse bars of blue pigment that characterize the majority of the 
species of Lysvosguilla are entirely absent in L.spinosa. No trace of the original colour 
remains in the case of the two specimens preserved in the Indian Museum. 


For the elucidation of the synonymy of this species we are indebted to Chilton 
who has been able to examine the type specimens of S. laevis, S. indefensa and S. tri- 
dentata. The latter species was described from a specimen ‘75 inches in length and 
it possesses only four teeth on the dactylus of the raptorial claw. It is probable, as 
Chilton remarks, that it is a young individual, but nevertheless, on the assumption that 
it is to be referred to L. spinosa, it is not very easy to account for the small number of 


120 Memoirs of the Indian Museum. [Vor. IV, 


dactylar teeth, for the specimens which Lanchester records from Penang, and which 
also were only ‘75 inches in length, possess twelve to fourteen teeth. It is, however, 
not impossible that the post-larval development of the species may vary in different 
localities. 

The two examples of L. spinosa in the Indian Museum were found at the following 
localities :— 

3052 Port Blair, Andamans. .. J. Wood-Mason. 50 tO, 06m SINAPIE. 


oe Dunedin, New Zealand. 2) Wee tcon + I 9,74 mm. 


The species has been recorded from numerous localities in New Zealand and in 
the neighbouring islands: Chatham Is., Kapiti and Waikanae, near Wellington 
(Kirk), Waipapa Pt. and Port Pegasus, Stewart I. (Thomson), Port Chalmers; Otago 
Harbour, Resolution I. (Chilton), Napier District (Hamilton) and Auckland Is., in the 
stomach of Notothenia microlepidota (Hutton). In addition it was found by Sir J. C. 
Ross’ Antarctic Expedition (Miers) and is known from Penang (Lanchester). Accord- 
ing to Chilton most of the specimens which have been recorded were taken from the 
stomachs of fish. 

6. Lysiosquilla acanthocarpus', Miers. 
1847. Cornis acanthocarpus, Gray MS., White, List Crus. Brit. Mus., p. 85 (sine desc.). 
1880. Lystosquilla acanthocarpus, Miers, Ann. Mag. Nat. Hist. (5), V, pp. II, 125, pl. i, figs. 7-9. 
1882. Lysiosquilla acanthocarpus, Haswell, Cat. Australian Crust., p. 206. 
1886. Lysiosquilla sarasinorum, Müller, Verhandl. naturf. Ges. Basel, p. 478, pl. iv, fig. 3. 
1894 Lysiosgunlla acanthocarpus, Bigelow, Proc. US. Nat. Mus., XVII, p. 503. 
1894. Lysiosquilla saracinorum, Bigelow, ibid., p. 503. 
1904. Lysiosquilla acanthocarpus, Jurich, Stomatop. Deutsch. Tiefsee-Exped., p. 374, pl. xxvi, 
fig. 3. 

The dorsal surface is smooth and polished. The greatest breadth of the carapace 
is about equal to its mid-dorsal length ; for the rest, it bears a close resemblance to that 
of L. maculata. ‘The rostrum is widest at the base and is usually a little longer than its 
greatest breadth. It terminates in a slender median spine and possesses in addition 
prominent antero-lateral angles on either side. These are rectangular or slightly obtuse, 
and the margin both before and behind them is concave: the former more strongly 
so than the latter. The rostrum is thus composed of an almost square proximal portion 
which bears at its distal end a sharp spine with a very broad base. 

The eyes are composed of two closely-approximated lobes. The greatest breadth 
of the cornea is about equal to the greatest length of the stalk. The anterior edge of the 
ophthalmic somite is rounded, but, when the animal is viewed from above, is entirely 
concealed by the rostral plate, as are also the dorsal processes of the same somite. The 
antennular peduncle is less than half the length of the carapace excluding the rostrum. 
The dorsal processes of the antennular somite consist of two long and sharp spines 
directed obliquely outwards and forwards and reaching to the middle of the eyes. The 
mandibular palp is composed of three segments. 


vx, !See addendum, p. 106. 


OS S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. LA 


The outer inferior margin of the raptorial claw is rounded distally. The carpus is 
not carinate above, but its dorsal margin is provided with a sharp distal spine. Along 
its inner edge the propodus is finely serrate and bears four large movable spines near 
the base. Externally the dactylus is convex and at its proximal end is cut into two 
small subequal lobes, the distal of which is the more obtuse. Internally it is provided 
with five, more commonly with six spines, including the large terminal one. The teeth 
are as a rule microscopically serrate and the penultimate is much shorter and more 
slender than the ante-penultimate. 

The lateral margins of the fifth thoracic somite are vertically grooved ; those of 
the two succeeding somites are truncate with rounded anterior and posterior angles. 
The shorter ramus of the limbs of the last thoracic somite is linear, those of the two 
preceding somites broadly ovate. 

The first five abdominal somites do not possess spines at ‘ihe postero-lateral angles. 
The sixth bears a stout spine at each of these angles and its dorsal surface is very ob- 
scurely furrowed longitudinally on either side. On its ventral surface this somite pos- 
sesses a slender curved spine at its outer anterior angles, projecting backwards over 
the base of the peduncle of the uropods. 


The telson is more than twice as broad as long. Near its base it bears dorsally a 
pair of shallow oblique grooves which run towards the antero-lateral angles, but disappear 
close to a small and obscure tubercle before reaching it. Distally there is a dorsal series 
of five sharp spines arranged in a semicircle. These spines do not form the termina- 
tions of longitudinal carinae as they do in certain other species of Lysiosquilla. On the 
margin three pairs of fixed primary spines may be distinguished ; the submedians are 
the smallest and the laterals much the largest. Close beneath each submedian there is a 
conspicuous movable spine and the margin between the two is occupied by a series of 
five or six (rarely four) pairs of small, sharp, fixed denticles. There are three stout 
. denticles, two of which are usually placed close together, between the submedians and 
intermediates and one between the intermediates and laterals. Beneath the base of 
each lateral spine there is a small, acute, outwardly-directed lobe. The inferior surface 
of the telson is swollen with a large depression behind the anus. 

The peduncular segment of the uropods bears a longitudinal carina along its 
antero-lateral margin, and a small dorsal spine posteriorly near its articulation with 
the exopod. The ventral bifurcate process consists of two long tricarinate spines, the 
outer of which is less than two-thirds the length of the inner. Behind these spines the 
lower surface bears a median carina which runs backwards and terminates abruptly in 
a blunt tooth close to the proximal margin. 


The outer margin of the basal segment of the exopod bears a series of six or seven 
movable spines of which the two last are much longer than the rest. There is also a 
short fixed ventral spine projecting over the articulation of the ultimate segment, and 
inwards of this a small rounded plate bearing a series of long setae. The anterolateral 
angle of the inner uropod is always permanently bent over and rests against the dorsal 
surface. 


122 Memoirs of the Indian Museum. Voralvs 


The only male specimen examined is very small (48 mm.) and does not possess 
any special structural modifications characteristic of its sex. 

The colouration of spirit specimens is very characteristic. The whole of the ros- 
trum except the apical spine and a narrow proximal band is black. There are 
three broad transverse bands of the same colour on the carapace, the posterior of which 
encloses a pale oval spot on either side. The last three thoracic and all the abdominal 
somites have each a broad, median, transverse, black band, leaving pale anterior 
and posterior borders. On the sixth somite the posterior pale border is sometimes 
obsolete. The telson has a pair of large rounded patches of the same black pig- 
ment, involving both the intermediate and lateral dorsal spines and frequently coal- 
escing anteriorly. In some specimens there is a single large patch which is cut into 
four distal lobes. The inner uropod is entirely black, as are also the spines of the 
bifurcate process except at the extreme apex. The peduncular segment is dark proxi- 
mally ; the first segment of the exopod has a round black distal spot, while the 
ultimate segment is obliquely divided into dark anterior and pale posterior portions. 


The collection in the Indian Museum comprises ten specimens :— 


se Port Blair, Andaman Is. G. H. Booley. 19,41 mm. 

a Coconada, Madras Presidency. G. W. Wicks. 19,5 © ,48—74 mm. 
ae Coconada, Madras Presidency. ‘ Investigator.’ 19,86 mm. 
ue Bombay. Bombay Nat. Hist. Soc. 22,65 and 85 mm. 


Lysiosquilla acanthocarpus has been recorded from Port Essington, N. Australia 
(Miers) , from Penang (Miers) and from Trincomali (Muller, sub S. sarasinorum). 


Miers ' has described a small female Lyszosquilla from Goree I., Senegambia, under 
the name L. acanthocarpus var. septemspinosa. ‘This specimen differs from typical 
acanthocarpus in the slightly transverse rostrum, in the less prominent eyes and in the 
possession of seven teeth on the dactylus of the raptorial claw, the penultimate of which 
is not shorter than the antepenultimate. This form should, perhaps, be recognized as 
a separate species. 


7. Lysiosquilla multifasciata, Wood-Mason. 


1895. Lysiosquilla multifasciata, Wood-Mason, Figs. and Desc. of nine Squillidae, p. I,pl.i, 
figs. 4-7. 

? 1903. Lysiosquilla multifasciata, Nobili, Boll. Mus. Torino, XVIII, no. 447, p. 30. 

? 1904. Lysiosquilla valdiviensis, Jurich. Stomatop. Deutsch. Tiefsee-Exped. p. 372, pl. xxvi, 
figs. 2-28. 

? 1906. Lysiosguilla multifasciata, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 337. 

1910. Lysiosquilla multifasciata, Balss, Abh. math.-phys. Klasse k. bayer. Akad. Wiss. Suppl. Bd. 
II, Abh. 2, p. 6. 


This species is very closely allied to the preceding, differing from it only in the 
following particulars :— 
1. The dactylus of the raptorial claw bears five or six teeth of which the penulti- 


! Miers, Ann. Mag. Nat. Hist. (5), VIII, 1881, p. 368, pl. xvi, fig. 7. 


1013.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 123 


mate is short, as in L. acanthocarpus, but the two lobes at the base of its exter- 
nal margin are very unequal, the distal one being greatly expanded and more 
than twice the size of the proximal. 

2. There are four short evenly-spaced spinules between the intermediate marginal 
teeth and the movable submedians. The innermost of these perhaps repre- 
sents the stout fixed submedian tooth which is found overlying the movable 
spine in L. acanthocarpus, but in the present species is not any larger than 
the three external to it. In the type specimen there are four pairs of fixed 
denticles between the two movable submedians. 

It may also be mentioned that the rostrum is a trifle longer proportionally than in 

L. acanthocarpus, its lateral margins are rather more distinctly convergent anteriorly 
and the antero-lateral angles are a little more obtuse. The eyestalks moreover appear 
to be a trifle more slender and the cornea a little narrower. In other structural details, 
notably in the form, carination and spinulation of the segments composing the uropods, 
the two species are in closest accord. 


The colour pattern is, however, different, at any rate in the type specimen. The 
two anterior dark bands on the carapace are indistinct and are composed of dark 
speckles and mottling; the posterior one is very well marked, especially at the sides 
where there is no trace of the pale enclosed spot which occurs in L. acanthocarpus. Each 
segment of the post-abdomen from the sixth thoracic to the fifth abdominal inclusive 
bears two transverse dark bands. ‘The anterior of these! is very narrow and does not 
reachthe lateral margin on either side, while the posterior, which is about as broad as the 
pale band in front of it, is separated from the distal margin of the somite by a very 
narrow pale border. The sixth abdominal somite possesses a single dark transverse 
band close to the posterior edge. The telson bears four indistinctly separated dark 
spots in much the same position as those of L. acanthocarpus, and in the pigmentation 
of the uropods there is a close resemblance between the two species. 


I am not certain that the specimens which Nobili described in 1906 are correctly 
identified. In his account there is no reference to the curious: form of the external 
lobes on the raptorial dactylus (much the most important structural feature possessed 
by L. multifasciata), while with regard to the colour he merely informs us that ‘‘la 
coloration est celle des autres Lysiosquilla, par bandes transversales noires, dont trois 
sur la carapace.’’ From the description supplied it seems possible that the specimens 
may really belong to L. acanthocarpus. 

Lysiosquilla valdiviensis, Jurich, described from a semi-larval specimen 14°6 mm. 
in length, is probably synonymous with this species. The form of the raptorial claw 
bears a very close resemblance to that of L. mulhfasciata; but the rostrum is without 
trace of antero-lateral angles, and there are only three dorsal spines near the distal 
margin of the telson. The eyes are larger; but this is invariably the case in very 
young specimens and there is good reason to believe that the two characters mentioned 
above are also due merely to immaturity. ‘The description of species from semi-larval 


! Balss notes that this band is obsolete in the specimen which he examined. 


124 Memoirs of the Indian Museum. [Vor. IV, 


forms, as Jurich has done both in this case and in the case of Squilla minor, is greatly 
to be deprecated. 


Wood-Mason’s type specimen is the only example of L. multifasciata preserved 
in the Indian Museum :— 
ee Bombay. Bombay Nat. Hist. Soc. 19,44mm. TYPE. 
Balss has recorded the species from Formosa. Nobili’s examples, which are per- 
haps wrongly determined, were found at Samarinda in Borneo and at Obock in the 
Red Sea. The locality of Jurich’s L. valdiviensis is unknown. 


8. Lysiosquilla biminiensis, Bigelow. 
1893. Lysiosquilla biminiensis, Bigelow, John Hopkins Univ. Circ., No. 106, p. 102. 
1894. Lysiosquilla biminiensis, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 504, text figs. 4-7. 
subsp. pacificus, Borradaile. 
1899. Lysiosquilla biminiensis var. pacificus, Borradaile, in Willey’s Zool. Results, p. 403. 

This species, like the preceding, is a close ally of L. acanthocarpus, but differs from 

it structurally in the following features :— 

I. The dactylus of the raptorial claw bears six teeth, including the apical one, and 
the penultimate is longer than the ante-penultimate. The lobes at the base of 
the external margin have much the same form as in acanthocarpus. 

2. There are only three pairs of denticles, which are comparatively large, between 
the mobile submedian spines of the telson. 

3. The rounded lobe fringed with long setae which is found in L. acanthocarpus 
on the ventral aspect of the basal segment of the outer uropod, projecting over 
the articulation with the ultimate segment, does not exist (fide Bigelow, fig. — 
4) in L. biminiensis and the carina on the lower surface of the peduncular 
segment (which ends in a strong anterior tooth in L. acanthocarpus) also 
appears to be missing. 


The colour of the living animal, according to the full description which Bigelow 
gives, is fawn or pink with black, reddish brown and bright lemon yellow markings. 
The black colouring, which alone persists in alcohol, exists as a narrow border round 
the postero-lateral angles of the carapace. ‘here is also a black posterior transverse 
streak on either side of the last thoracic and fifth abdominal somites and a pair of 
small black eyespots on the telson placed side by side in the median line just in front 
of the dorsal spines. 

The specimen which Borradaile names pacificus differs-structurally from those des- 
cribed by Bigelow only in the most insignificant features, but the colouring is different 
for on each somite from the sixth thoracic to the fifth abdominal inclusive, there is a 
narrow deep-black posterior border. 

Bigelow’s two specimens (48 mm. in length) were found in a burrow in the sand 
at the Bimini Is., Bahamas, while the single individual which Borradaile records was 
found at New Britain and is called pacificus in order to emphasize the remarkable 
distribution of the species. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 125 


9. Lysiosquilla tigrina, Nobili. 
Plate VIII, figs. 95-98. 
1903. Lysiosguilla tigrina, Nobili, Boll. Mus. Torino, XVIII, no. 447, p. 28. 


The carapace is dorsally smooth and polished, and its greatest breadth is about 
equal to its mid-dorsal length. The gastric grooves are well-defined and slightly con- 
vergent posteriorly ; but of the cervicals scarcely a trace remains. ‘The antero- and 
postero-lateral angles are rounded. The rostrum is as long as broad; the lateral margins 
are only slightly convergent anteriorly and terminate in well-marked subacute angles. 
The apex is sharply acute and reaches a little beyond the base of the eyestalk (fig. 95). 

The eyes are nearly one-third the mid-dorsal length of the carapace, and the cornea, 
which is not pigmented at the extreme apex, is obscurely bilobed, its breadth being 
about equal to the length of the stalk. The antennular peduncle is about half the 
length of the carapace. The mandibular palp is composed of three segments. 

The outer inferior margin of the raptorial claw (fig. 96) is bluntly angled distally ; 
the carpus is not carinate dorsally but terminates in a sharp spine overhanging the 
carpo-propodal articulation. The propodus bears four mobile spines at the proximal 
end of its pectinate edge. The dactylus is provided with eleven spines, including the 
apical one, the penultimate being very slightly shorter than the next of the series. 
The outer margin is convex and at its proximal end bears two small lobes, the ante- 
rior of which is broad and rounded while the posterior is narrow and subacute. 

The free thoracic and abdominal somites are smooth ; the lateral margins of the 
former are very broadly rounded, almost truncate, and the postero-lateral angles of 
the latter are also rounded except in the case of the last somite, in which they project 
posteriorly as a pair of long and sharp spines. On its inferior margin the sixth somite 
(fig. 98) bears a pair of curved spines on either side at the antero-lateral angles and a 
humber of short spines rather irregularly disposed (nine in the only known specimen) 
along the posterior margin. ‘This series of spines is not known to occur in any other 
species of Lysiosquilla. 

The telson (fig. 97) is smooth and rounded dorsally except for a faint transverse 
depression on either side of the median line ; on its inferior surface there is a central 
depression behind the anus. Dorsally it possesses a series of five spines arranged ina 
curved transverse row near the posterior margin. There are four pairs of large mar- 
ginal teeth, a pair of submedians, two pairs of intermediates and one pair of laterals. 
Between the submedians there are six pairs of small denticles, the outermost of which 
are mobile and longer than the rest, and outside these are three small fixed spines, 
one in each interspace between the primary teeth. These small spines are all 
inferior in position and are situated beneath the level of the larger ones (fig. 98). In 
addition there is a single large spine immediately below the proximal primary tooth. 

On the posterior margin of the peduncle of the uropods there is a small median 
dorsal spine and another at the external angle. The bifurcate process from the 
inferior margin consists of a pair of long curved spines, of which the inner is only very 
slightly longer than the outer, and on the margin inside the process there is also 


126 Memoirs of the Indian Museum. Vor; IV, 


another spine which is stout and much shorter. The proximal segment of the outer 
uropod bears the usual series of movable spines along its external margin. As in L. 
acanthocarpus there is on the ventral side a small fixed spine and a setose lobe project- 
ing over the articulaton of the ultimate segment (fig. 98). 


The type specimen is pale in colour with scattered dark chromatophores on the 
carapace, which are closely aggregated near the posterior margin (fig. 95). The 
rostrum is sprinkled with similar chromatophores which are thickest proximally. On 
each of the free thoracic and abdominal somites there is a broad transverse dark band 
separating narrow anterior and still narrower posterior borders. On the first five 
abdominal somites the dark band shows a tendency to become divided by a narrow 
and obscure, transverse, pale stripe. The colouring of the sixth somite and of the 
telson is shown in fig. 97. The peduncular segment of the uropods is dark as is also 
the inner ramus. There is a dark spot near the distal end of the proximal segment of 
- the exopod and the outer segment is suffused with dark pigment at the base. 


Thanks to the kindness of Mr. J. C. Moulton, I have had an opportunity of ex- 
amining the only known example of this species, and I have been able in consequence 
to supplement Nobili’s account with figures and a few further details. 


The type of Lysiosquilla tigrina is a male, 45 mm. in length. It was found at 
Santubong, Borneo, and is preserved in the Sarawak Museum. 


10. Lysiosquilla vicina, Nobili. 
1904. Lysiosguilla vicina, Nobili, Bull. Mus. Hist. Nat., X, p. 220. 
1906. Lystosquilla vicina, Nobili, Ann. Sci. Nat., Zool. (9), IV, p. 339. 

This species is founded on a single small specimen in which the raptorial claws are 
wanting. It appears to be allied to L. tigrina ; but is distinguished by the following 
characters :— 

1. The rostrum is trispinous, each antero-lateral angle being acutely produced. 

2. The posterior ventral border of the sixth abdominal somite is smooth and with- 

out spines. 

3. As in L. tigrina there are four pairs of large primary teeth on the margin of the 
telson, but the stout additional spine, which occurs in that species below the 
proximal lateral tooth, is missing. 

L. vicina seems to be easily distinguished from all other known species of Lyszo- 
squilla by the trispinous rostrum, and from all except L. tigrina by the four pairs of 
long fixed spines on the margin of the telson. 

The type and only known example, a male only 27 mm. in length, was found at 
Obock in the Red Sea in sand inhabited by Balanoglossus. 


11, Lysiosquilla insignis, Kemp. 
Plate IX, figs. 09-102. 
1011. Lystosquilla insignis, Kemp, Rec. Ind. Mus., VI, p. 94. 
The carapace is smooth, longer than its greatest breadth and is obtusely angled 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 127 


antero-laterally. There is no cervical groove, but the gastric grooves are distinct 
throughout. The rostrum has the form of a narrow isosceles triangle and is more than 
one and a half times as long as broad. There are no antero-lateral angles and the 
lateral margins, which are feebly sinuous, converge to a very fine spinous apex (fig. 00). 

The eyes are very large. The cornea is oblique and obscurely bilobed ; its greatest 
breadth is one-third the mid-dorsal length of the carapace and is about equal to the 
total length of the whole eye. Seen from above, the anterior margin of the ophthalmic 
somite and its dorsal processes are entirely concealed by the rostrum. ‘The antennular 
peduncle is nearly half the length of the carapace (excluding the rostrum), and its ulti- 
mate segment is only half the length of the penultimate; the dorsal processes of the 
somite consist of a pair of sharp divergent spines. The mandibular palp is composed 
of three segments. 

The carpus of the raptorial claw (fig. 100) is not carinate, but its dorsal margin ter- 
minates in a sharp spine ; the propodus bears four movable spines at the base of its 
pectinate margin. ‘The dactylus is provided with seven or eight teeth including the 
terminal one, the penultimate being, as in L. acanthocarpus, very noticeably shorter 
than the ante-penultimate. The external margin is concave and at its base bears two 
lobes, the anterior of which is obsolete, while the posterior has the form of a sharply 
acute tooth. 

The posterior portion of the lateral margin of the fifth somite projects outwards 
as a thin vertical blade. The sixth somite has an angular dorsal elevation on either 
side near the anterior edge ; antero-laterally the margin is deeply excavate and behind 
this projects in the form of a truncate lobe with rounded subrectangular anterior and 
posterior angles. The seventh somite is broadest posteriorly ; its lateral margins are 
obliquely truncate in front and rather sharply rounded behind (fig. 99). The basal 
segment of the last three thoracic appendages terminates posteriorly in a sharp spine. 
The shorter ramus of the limb attached to the sixth somite is rather broadly oval, that 
of the eighth much narrower, though it could scarcely be termed linear, while that of 
the seventh is intermediate in form. 

The first four abdominal somites are smooth dorsally. The postero-lateral angles 
of the first three are rounded; those of the last three are sharply spinous. The fifth 
somite bears an obscure longitudinal carina on either side in its posterior half. The 
sixth somite, in addition to the spines at its postero-lateral angles, possesses a dorsal pair 
situated close to the distal margin (fig. 101). Each of the latter forms the termination 
of an irregular blunt carina, and between this and the outer margin there is also another 
carina which runs obliquely towards the postero-lateral spine. On its ventral surface 
the sixth somite bears a sharp curved spine at its antero-lateral angles situated im- 
mediately in front of a prominent rounded lobe. 

The telson (figs. IOI, 102) is rather less than twice as broad as long. It bears on 
its dorsal surface two pairs of tubercles and eight longitudinal carinae, the outermost 
of which is bifurcated proximally.. The peculiar arrangement of these carinae, which 
are slightly irregular and not precisely similar on either side, is shown in fig. ror and 
does not call for detailed description. In place of the curved series of dorsal spines 


128 Memoirs of the Indian Museum. [Voy. IV, 


which occurs in most of the preceding species, there is in L. ıinsignis a very peculiar 
armature which consists of a trilobed median prominence, a sharp intermediate spine 
and an angular lateral lobe. These are not separated by the smooth declivous dorsum 
of the telson, asin L. acanthocarpus and the other species with a dorsal circlet of spines, 
but are connected by a sharp transverse ridge. On the margin of the telson there are 
two very large teeth, the intermediate and the lateral, on each side and two long 
movable submedian spines. Between the latter are six pairs of small denticles (fig. 102). 
There are four short spines' between the mobile pair and the intermediate tooth, and 
one between the intermediate and the lateral. 

The peduncular segment of the uropods is carinate along its antero-lateral margin 
and a second carina, parallel to the above, runs to the middle of the distal margin where 
it is continued as a long spine extending over the base of the proximal segment of the 
exopod. The bifurcate process consists of a pair of large tricarinate spines, the outer 
of which is scarcely shorter than the inner. In addition there is a sharp inferior spine 
at the point of articulation of the inner uropod. On the external margin of the 
basal segment of the outer uropod there is a series of six mobile spines; the two outer- 
most are curved and many times longer than the next of the series. On the ventral 
surface there is a stout fixed distal spine projecting over the base of the ultimate 
segment. 

No trace of any colour remains in the single specimen examined. 


©® Off N. Andaman I., 235 fathoms, .. ‘ Investigator’. Gy) UG) 210) amen. NDS, 
50° 7’ N., 79° 7' E. 


12. Lysiosquilla latifrons (de Haan). 
1844? Lysiosquilla latifrons, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. 3, 
text, 1840, p. 222. 
1880. Lysiosquilla (Coronis) latifrons, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 10, 25. 
1880. Lysiosguilla brazieri, Miers, ibid., pp. II, 125, pl. 1, figs. 3-6. 
1882. Lysiosguilla brazieri, Haswell, Cat. Australian Crust., p. 206. 
1894. Lysiosquilla latifrons, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 503. 
1894. Lysiosquilla brazieri, Bigelow, ibid., p. 503. 
1903. Lysiosquilla latifrons, Rathbun, Proc. U.S. Nat. Mus., XX VI, p. 54. 
1910. Lysiosquilla brazieri, Chilton, Trans. N. Z. Inst., XLIII, p. 130. 
I have seen no examples of L. latifrons ; the principal characters of the species 
appear to be as follows :— 
The rostrum is broader than long with very strongly rounded lateral margins and 
a sharp median spine. ‘The cornea of the eyes is bilobed, but is very much smaller than 
in the preceding species.” The dactylus of the raptorial claw bears six, rarely seven, 
teeth. The lateral margins of the exposed thoracic somites are truncate. The short 


! The innermost of these is probably homologous with the fixed submedian spine in L. acanthocarpus 
and other species. 

* In De Haan’s figure, which probably represents a small specimen natural size, the breadth of the 
cornea is about one-fifth the length of the carapace. In Mier’s large example the eye, as might be ex- 
pected, is proportionately smaller. 


ERRATUM. 
P. 128, line 22 from top of page, joy © BO No, 7 Be read A 27 N., 
93°50 H.”’ 


1913.| S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 129 


ramus of the appendages of the sixth and seventh somites is broadly oval; that of the 
succeeding somite almost linear. 

The first five abdominal somites are not spinous at the postero-lateral angles ; 
the sixth is obliquely sulcated on either side with sharp postero-lateral spines. The 
telson bears seven dorsal spines in a transverse row near the posterior margin. Accord- 
ing to the figures the two outermost on either side are separated from one another and 
from the three median by a very distinct furrow which extends forwards half way to the 
anterior margin. There are three pairs of large marginal teeth, the submedians being 
mobile. Between the submedians are ten to fourteen small spinules and there are three 
short spines between the submedians and the intermediate and one between the inter- 
' mediate and the lateral. On the inferior surface there is a single spine behind the anus. 

The descriptions published contain no mention of any distinctive colouring. 

There can be little doubt that, as Chilton has suggested, Miers’ L. brazieri is 
synonymous with L. latifrons. De Haan states that there are seven dactylar teeth in 
the latter form, whereas Miers found six only in L. brazieri ; but, as Miss Rathbun 
has pointed out, six are shown on the right raptorial claw in De Haan’s figure. In 
Miss Rathbun’s example, which, like the type, was found in Japanese waters, both 
raptorial dactyli bear six teeth. 

Lysiosquilla latifrons seems to present some affinity with L. insignis, but the two 
forms are readily distinguished by the characters afforded by the rostrum, eye, raptorial 
dactylus, sixth abdominal somite and telson. 

The largest of the known specimens is about 105 mm. in length. 

The species is recorded from Japan (De Haan), Nagasaki, Japan (Rathbun), Port 
Jackson, New South Wales (Miers), and from Otaki, New Zealand (Chilton). 


Genus Coronida, Brooks. 
1886. Coronida, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 79. 


Carapace strongly, or scarcely at all, narrowed anteriorly, without carinae : 
antero-lateral angles rounded; gastric grooves distinct; cervical groove almost 
obsolete, wholly absent mid-dorsally. Mandibular palp three-segmented or entirely 
absent. Merus of raptorial claw articulating terminally with ischium ; ventral sur- 
face of merus longitudinally hollowed throughout its length ; upper margin of pro- 
podus pectinate proximally along outer edge, distally with a pair of smooth blunt 
ridges; dactylus inflated at base, armed with teeth on inner margin. Shorter ramus 
of last three thoracic appendages linear, composed of two segments. Free thoracic 
somites without carinae. Abdominal somites depressed, first four without carinae 
or sculpture. Last one or two abdominal somites and telson with elaborate orna- 
mentation consisting of close set spinules or of carinae and large tubercles, in the latter 
case a pair of submedian carinae on telson; telson semicircular in shape, marginal 
teeth minute. Ventral process of uropods consisting of a single strong spine with a 
much smaller spine on outer margin. 

There are no marked secondary sexual distinctions. 


130 Memoirs of the Indian Museum. More 


The form of the articulation of the ischium and merus of the raptorial claw 

leads me to believe that the affinities of this genus are with Squilla, Pseudosquilla, 
~ and Lysiosquilla rather than with Odontodactylus and Gonodactylus. 
Of the three known species one inhabits the Atlantic and two the Indo-Pacific. 
The Atlantic form, C. bradyi, is very closely allied to C. tvachura and the two may, 
in fact, belong to the same species; C. bradyi is only known from a single example 
described in 1869. C. multituberculata, though very different in general appearance, 
must certainly be accommodated in this genus. 


Key to the species of Coronida. 


I. Mandibular palp present ; external margin of raptorial dactylus with a 
single blunt lobe at base ; dorsal surface of last abdominal somite and 
telson covered with spinules ; form broad, greatest width of abdomen 
more than one-fifth of total length. 


A. Spinules simple, covering last abdominal somite and telson only .. [bradyt.] 
B. Spinules stellate, covering last abdominal somite and telson and, in 
addition, posterior half of fifth somite oe . trvachura, p. 130. 


II. Mandibular palp absent ; external margin of raptorial dactylus with tie 
teeth; fifth abdominal somite smooth and unarmed; sixth with eight 
dorsal ridges ; telson with two median dorsal carinae and numerous 
large tubercles on either side ; form narrow, greatest breadth of abdo- 


men much less than one-fifth of total length .. multituberculata, p. 132. 


1, Coronida trachura (v. Martens). 
1881. Gonodactylus trachurus, v. Martens, Sitz.-ber. Ges. naturf. Freunde, Berlin, p. 93. 
1884. Gonodactylus trachurus, Miers, Proc. Zool. Soc., p. 16, pl. i, figs. 3, 3a. 
1886. Coronida trachura, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., pp. 7 and 97-80. 
1894. Odontodactylus trachurus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 
1906. Coronida trachura, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 334. 

The carapace is smooth without trace of carinae and is widest posteriorly, its great- 
est breadth being about equal to its median length including the rostrum. The gastric 
grooves are well marked, and on either side of them faint traces of the transverse 
cervical groove may be observed. The antero-lateral angles are angular but not pro- 
duced as spines, the postero-lateral are very broadly rounded. ‘The rostrum is sub- 
rectangular, much wider than long; its lateral margins are parallel, the antero-lateral 
angles almost rectangular, and the apex is formed by a blunt obtuse or subrectangular 
prominence. 

The eyes are about one-third the length of the carapace, excluding the rostrum. 
The cornea is prominently bilobed, set transversely on the stalk, and in breadth is 
rather less than the total length of the whole organ. The ophthalmic somite termi- 
nates in asmall acute apex. The antennular peduncle is a little shorter than the 
carapace excluding the rostrum. ‘The mandibular palp is three-segmented. 

The outer inferior margin of the merus of the raptorial claw is broadly rounded 
distally ; the carpus has a dorsal carina which is cut into one or two sharp teeth near its 
distal end, The propodus bears three movable spines at the base of the margin opposed 


1913.] S. Kemp: Crustacea Stomatcpoda of the Indo-Pacific Region. AT 


to the dactylus. This margin as in allied genera is pectinate, but only along its basal 
two-thirds. In the distal third, where there are no pectinations, the inner and outer 
edges are elevated and form a pair of low smooth crests, between which the proximal 
edge of the dactylus fits when the claw is closed. The dactylus bears four! spines 
including the terminal one. Its outer margin is notched at the base, and in front of 
this notch is strongly inflated somewhat after the fashion of Gonodactylus. 

The free thoracic and first four abdominal somites are smooth dorsally. The 
lateral margin of the fifth possesses a vertical groove, which separates a pair of inferiorly 
convergent carinae, the posterior of which is the highest and in dorsal view resembles 
a lateral spine. The sixth and seventh somites are broadly rounded laterally and the 
short inner ramus of the three pairs of thoracic limbs is linear. 

The postero-lateral angles of the first four abdominal somites are not spinous. 
The posterior half of the fifth somite is rather thickly scattered with small irregular 
tubercles, and close to the external margin there is a fine longitudinal carina which runs 
outwards and terminates in the angular and produced postero-lateral angle. The 
sixth somite is densely covered with stellate or spinous tubercles, and its postero- 
lateral angles end in spines. The posterior margins of both the last two abdominal 
somites are fringed with a series of sharp spinules. 

The telson is rather less than twice as broad as long ; it is strongly convex and, 
like the preceding segment, is covered with closely packed spinous tubercles. Seen 
from below a pair of submedian movable spines may be distinguished, and two pairs of 
large fixed lateral spines ; there are also, near the distal margin, two pairs of inferior 
spines situated inwards of the two pairs of laterals. The ventral surface bears numer- 
ous scattered spinules. 

Tubercles similar to those on the dorsal aspect of the telson are found on the upper 
side of all the segments composing the uropods, but on the endopod and ultimate segment 
of the exopod, they exist only in a narrow median crest. The peduncular segment is 
strongly arched and sharply carinate antero-laterally ; distally it is produced as a spine, 
supported by a short carina, which overhangs the articulation of the basal segment 
of the exopod. The ventral bifurcate process consists of a sharp inwardly curved spine 
which is very broad proximally. Its outer margin bears a sharp tooth near the apex, 
while its inner margin is cut into three or four strong teeth. The process is strength- 
ened by a blunt longitudinal ventral carina. The external margin of the basal segment 
of the exopod carries nine or ten movable spines, and there is also an additional one, 
which is fixed, at the extreme distal end of the ventral surface. The internal margin of 
the inner uropod is concave, the segment being more or less crescentic in outline. 


Two of the specimens examined are pale straw colour in spirit sprinkled with very 
small black chromatophores. These are aggregated and tend to form a dark posterior 
band on the carapace, enclosing a small pale area on either side. In addition, there is 
a median and pair of lateral patches on each of the first five abdominal somites. The 


! In one of the specimens examined there are on one side only three spines, but this appears to be 
quite abnormal. 


132 Memoirs of the Indian Museum. [Vor. IV, 


sixth thoracie is thickly sprinkled with black dots, as are also the lateral portions of 
the two succeeding somites, the basal segments of the thoracic appendages and the outer 
uropods. In the third example the distribution of the pigment is precisely similar ; 
but thechromatophores, which appearto be expanded, form conspicuous black patches. 

As Miers has mentioned, Coronida trachura is very closely allied to C. bradyi (A. 
Milne-Edwards, 1869) from St. Vincent, Cape Verde Is. According to Milne-Edwards’ 
description C. bradyi appears to be distinct from the Indo-pacific species, but a fresh 
account of the Atlantic form is badly needed. 

The three specimens of C. trachura in the Indian Museum are registered as 
follows :— 

en Mauritius. Purchased. 24,19, 38-40 mm. 


The species has been previously recorded from Mauritius (v. Martens, Miers) 
from the Red Sea (Nobili) and from the Pelew Is. (Palass Ins.), Caroline group (v. 


Martens). 
2. Coronida multituberculata (Borradaile). 


1898. Squilla multituberculata, Borradaile, Proc. Zool. Soc., p. 38, pl. vi, figs. 7, a-c. 
1899. Squilla multituberculata, Borradaile, in Willey’s Zool. Results, p. 403. 

Through the kindness of Mr. I,. Doncaster I have been able to examine two of the 
type specimens of Borradaile’s Squilla multituberculata, and have thereby been able 
to satisfy myself that the species must be transferred to the genus Coronida. In the 
total absence of the cervical groove in the median part of the carapace, in the absence 
of abdominal carinae, and in the proximal inflation of the raptorial dactylus, C. multi- 
tuberculata agrees very closely with C. trachura, and the ornamentation of the body 
is, as in the latter species, restricted to the last abdominal somite and telson. More- 
over, in the minute and peculiar characters afforded by the raptorial propodus and by 
the basal segment of the uropods, and its spinous process, there is such a close 
resemblance between the two species that no doubt of their affinity can be entertained. 

The principal characters of C. multituberculata are as follows :— 

The carapace is wide posteriorly and is closely similar to that of the preceding 
species ; its greatest breadth, however, is less than its median length, excluding the 
rostrum. The rostrum is longer than wide and has the shape of an inverted triangle, 
its straight lateral margins diverging anteriorly to a broad, squarely truncate apex. 
The eyes are dorso-ventrally flattened ; the bilobed cornea is not wider than the stalk 
and is set obliquely on it. The mandibular palp is entirely absent. 

The dorsal edge of the carpus of the raptorial claw is cut into two large teeth near 
its distal end. The propodus is proportionately broader than in C. trachura ; on its upper 
margin it bears a series of conspicuous pectinations, and in its proximal portion three 
movable spines. Distally the pectinations are replaced by a pair of smooth crests, 
much as in the preceding species. The dactylus bears four teeth including the terminal 
one ; the basal part is inflated, though not very strongly so, and the external edge of 
this inflated part is cut into three blunt teeth, of which the proximal is the largest. 

The fifth thoracic somite resembles that of C. trachura, but the lateral margins of 
the three following are more truncate, and all the segments of the post-abdomen are 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 133 


proportionately longer than in that species. The first five abdominal somites are quite 
smooth above and their postero-lateral angles are rectangular, but slightly rounded. 
The sixth abdominal somite bears four pairs of longitudinal carinae, the outermost of 
which run along the lateral margins and terminate in subacute postero-lateral angles. 
The carinae of the submedian pair are slightly curved and project a trifle beyond the 
posterior margin ; between them, in one specimen, there is a short median carina in the 
posterior part of the somite. The two intermediate carinae on either side are con- 
nected posteriorly and form a \/-shaped figure, and between the innermost of these and 
the submedians there are several smoothly rounded tubercles. 

The telson is semicircular in outline, about one and a half times as broad as long, 
and is strongly convex above. It bears a pair of posteriorly convergent submedian 
carinae, and the surface is otherwise closely covered with smooth rounded tubercles : 
shortest on the vertex and longest at the edges. The ventral surface is smooth and on 
the posterior margin (seen from below) three sharp spines may be observed, as well as 
series of five submedian and intermediate spinules. 

The outer edge of the basal segment of the uropod is strongly arched as in the pre- 
ceding species, and the process arising from its inferior aspect consists of a single sharp 
spine bearing a very small tooth high up on its outer margin. The proximal segment of 
the exopodite is shorter than the ultimate segment and is provided with a series ofnine 
movable spines. The endopod is curved as in C. trachura, but much less distinctly so. 


No secondary sexual distinctions could be detected. 


Only three specimens of Coronida multituberculata are known. Their very small 
size, 12-13 mm., suggests that they are perhaps immature ; but, with the possible 
exception of the shape of the rostrum, it does not seem likely that their characters 
would be much modified by further growth. One male and two females were found at 
Sandal Bay, Lifu, Loyalty Is. (Borradaile). 


Genus Odontodactylus, Bigelow. 
1894. Odontodactylus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 445 (wbr syn.). 


Carapace little, if at all, narrowed anteriorly ; without carinae ; gastric grooves 
distinct, cervical groove wholly absent ; antero-lateral angles rounded, cornea of eye 
not distinctly bilobed. Mandibular palp present and composed of three segments 
or (?) entirely absent. Ischio-meral articulation of raptorial claw situated at a point 
in advance of proximal end of merus, the latter segment consequently extending back- 
wards considerably beyond the joint ; the inferior surface of merus not channelled 
longitudinally throughout its entire length ; upper margin of propodus not pectinate ; 
dactylus inflated ‘at base, armed with two to nine teeth on inner margin. 
Shorter ramus of last three thoracic appendages linear, composed of two segments. 
Free thoracic and abdominal somites compressed ; distinct carinae present only on 
last abdominal somite. Telson with sharp median carina and one or more carinae 
on either side; strong submedian, intermediate and lateral teeth on distal margin, 
the first with movable apices; no submedian denticles (though there may be minute 


134 | Memoirs of the Indian Museum. [Vor.. IV, 


spinules), one or two intermediate denticles and one lateral. Ventral process of 
uropods terminating in two sharp spines, inner usually shorter than outer. 

There are no well-marked secondary sexual distinctions. 

Ten species of this genus are known; nine have been found in Indo-pacific 
waters and one, O. havanensis, in the Atlantic. No species is known to inhabit both 
areas. 

Specimens of Odontodactylus are, unfortunately, not numerous in collections 
and consequently little is known of the variation that occurs within the limits of 
the species and of the alteration in form due to growth. In taxonomic work the 
precise number and disposition of the carinae on the telson is of great importance, 
but this is a character that may perhaps be modified in the course of growth from the 
post-larval form to the adult. Another feature which appears to have considerable 
systematic value is the size of the eye. In Sqwilla the breadth of the cornea is 
known to decrease proportionately during growth and this, if true in Odontodactylus, 
must have much influence on our views of the limits of the various species. Exami- 
nation of seven specimens of O. southwelli, varying from 21 to 37 mm. in length, 
seems, however, to indicate that this proportional difference, due to growth, is not 
one of any considerable magnitude, at any rate in that form. 

The species appear to live on rough ground, but of their habits nothing precise 
is known. The greatest depth from which any member of the genus has been 
obtained is 163 fathoms (O. havanensis, Bigelow). 

For explanation of the terminology used below in designating the carinae of the 
telson, see text-fig. 4, p. 12. 


Key to the species of Odontodactylus. 


I, Dactylus of raptorial claw with only two or three teeth on its inner margin. 
A, Several carinae on sixth abdominal somite, six of them terminating pos- 
teriorly in spines ; several carinae in anterior half of telson on either side 
of median crest. 
I. Raptorial dactylus with two teeth; second lateral carinae of telson 
present. 

A. Breadth of cornea about one-fifth median length of carapace ; raptorial 
dactylus very strongly dilated at base ; median crest of telson not 
remarkably high, with two pairs of submedian carinae .. = 

B. Breadth of cornea about one-third median length of carapace ; rapto- 
rial dactylus not very strongly dilated at base ; median crest of telson 


scyllarus, p. 135. 


remarkably high with a single pair of submedian carinae ae eultrajen, ps 137 
II. Raptorial dactylus with three teeth ; second lateral carinae of telson ab- 
sent (breadth of cornea about one half median length of carapace) .. : carinifev, p. 138. 
B. Sixth abdominal somite without carinae, with only four posterior spines, 
submedians and postero-laterals ; telson with median carina only .. elegans, p. 139. 


I. Dactylus of raptorial claw with at least six teeth on its inner margin. 
A. Second Jateral carina of telson present. 
I. Telson with two pairs of submedian carinae ; second lateral carina run- 
ning to apex of lateral marginal tooth (text-fig. 4, p. 12) .. japonicus, p. 139. 
II. Telson with a single pair of submedian carinae ; second lateral carina 


I913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. : | 135 


parallel with outer margin, running towards outer edge of intermediate 
marginal tooth. 

A. Breadth of cornea not more than one-third median length of carapace ; 
median portion of exposed dorsal process of opthalmic somite an- 
teriorly concave. 

I. Rostrum evenly rounded anteriorly ; basal segment of outer uropod 


with terminal mobile spine reaching to apex of distal segment .. hanseni, p. 140. 
2. Rostrum anteriorly truncate; basal segment of outer uropod with 
terminal mobile spine not reaching to apex of distal segment .. Jatirostris, p. 141. 


B. Breadth of cornea about one half median length of carapace; median 
portion of exposed dorsal process of ophthalmic somite very strongly 
excavate anteriorly, enclosing a deep and almost circular pit .. southwelli, p. 142. 
B. Second lateral carina of telson absent. 
I. Ophthalmic somite largely exposed in dorsal view; raptorial dactylus 
strongly inflated at base and not notched externally Le .. [havanensis.|! 
II. Ophthalmic somite almost entirely covered by rostrum ; raptorial dactylus 
only very slightly inflated at base and notched externally near its proxi- 
mal end ae ss Bs Me ae ..  brevirostris, p. 143. 


1, Odontodactylus scyllarus (Linnaeus). 


1705. Squilla lutaria, Rumphius, Amboinsche Rariteitkamer, p. 6, pl. iii, fig. F. 

1758. Squilla arenaria prona, Seba, Thesaurus, III, p. 51, pl. xx, fig. 3. 

1767. Cancer scyllarus, Linnaeus, Syst. Nat., 12th ed., I, ii, p. 1054. 

1787. Squilla scyllarus, Fabricius, Mantiss. Insect., I, p. 333. 

1796. Cancer (mantis) scyllarus, Herbst, Krabben u. Krebse, II, p. 99, pl. xxxiv, fig. 1. 

1703. Squilla scyllarus, Fabricius, Ent. Syst., II, p. 512. 

1798. Squilla scyllarus, Fabricius, Ent. Syst. Suppl., p. 416. 

1818. Squilla scyllarus, Lamarck, Hist. Nat. Anim. sans Vert., V, p. 188. 

1825. Gonodactylus scyllarus, Latreille, Encycl. Méthod., X, p. 473. 

1829. Gonodactylus scyllarus, Latreille, in Cuvier’s Régne Anim., 2nd ed., IV, p. 100. 

1837. Gonodactylus scyllarus, H. Milne-Edwards, Hist. Nat. Crust., II, p. 520. 

1838. Gonodactylus scyllarus, H. Milne-Edwards, in Lamarck’s Hist. Nat. Anim. sans Vert., 2nd 
ed VE Wo 3224. 

1836-49. Gonodactylus scyllarus, H. Milne-Kdwards, in Cuvier’s Regne Anim., 3rd ed., atlas, pl. ly, 
1S, Be 

1847. Gonodactylus scyllarus, White, List Crust. Brit. Mus., p. 85. 

1862. Gonodactylus scyllarus, A. Milne-Edwards, in Maillard’s l’île Reunion, ann. F., p. 10. 

1868. Gonodactylus bleekeri, A. Milne-Edwards, Nouv. Arch. Mus. Hist. Nat., IV, p. 65, foonote. 

1874. Gonodactylus scyllarus, Hoffmann, Rech. Faune Madagasc., Crust., p. 43. 

1880. Gonodactylus scyllarus, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 115, 459. 

1880. Gonodactylus bleekeri, Miers, 1bid., p. 118. 

1880. Gonodactylus scyllarus, Richters, in Mobius’ Meeresfauna Mauritius, p. 167. 

1886. Gonodactylus scyllarus, Müller, Verhandl. naturf. Ges. Basel, VIII, p, 471. 

1887. Gonodactylus scyllarus, de Man, Arch. f. Naturgesch., LIII, i, p. 572. 

1894. Odontodactylus scyllarus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 

1894. Odontodactylus bleekeri, Bigelow, ibid., p. 496. 

1894. Gonodactylus scyllarus, Ortmann, Denk. med.-wiss. Ges. Jena, VIII, p. 60. 


1 See p. 144. 


136 Memoirs of the Indian Museum. Vor. IV, 


1898. Odontodactylus scyllarus, Borradaile, Proc. Zool. Soc., p. 36, pl. v, fig. 6. 
1809. Odontodactylus scyllarus, Borradaile, in Willey’s Zool. Results, p. 402. 

1899. Odontodactylus scyllarus, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 276. 
1903. Odontodactylus scyllarus, Rathbun, Proc. U.S. Nat. Mus., XXVI, p. 54. 

The carapace is smooth with its antero-lateral and postero-lateral angles rounded : 
the former more broadly than the latter. It bears no carinae and the cervical groove is 
entirely obsolete. The rostrum is cordiform, widest a little in front of the base, and 
is rather more than one and a half times as broad as long ; seen from above it entirely 
conceals the dorsal processes of the ophthalmic somite. The apex is acute, depressed, 
and ventrally thickened. 

The eyestalks are short and broad and the breadth of cornea, which is a little wider 
than the stalk, is in large specimens contained about five times in the median length 
of the carapace, excluding the rostrum. The antennal scale is very large, between 
‘8 and ‘9 times the length of the carapace, and is nearly three times as long as wide. 

The dactylus of the raptorial claw is very strongly and abruptly swollen at the 
base ; distally it is slender and almost straight and the knife-like edge on its inner 
margin is cut into two strong teeth, the posterior of which springs from the mid-ventral 
point of the inflated portion, and is longer and more slender than the other. 

The thoracic somites are smooth above. The lateral margin of the sixth is truncate 
and rounded at either angle ; that of the seventh is similar but rather less sharply 
truncate and a trifle broader. The lateral margin of the eighth somite is produced to a 
rounded point and is notched antero-laterally. 

The median portion of the first five abdominal somites is smooth, but on each 
side there is a well-defined | -shaped groove running parallel to the anterior and lateral 
borders. On the fifth and sixth somites, also, a broad depression well above the lateral © 
margin is confined by two longitudinal ridges the innermost of which is proximally 
notched. Traces of similar depressions are visible on the sides of the two preceding 
somites. The postero-lateral angles become gradually more acute from before back- 
wards, and those of the fourth and fifth somites terminate in spines. The sixth somite 
bears four pairs of longitudinal carinae ; the submedian and lateral endin spines which 
project beyond the posterior margin, while, of the two intermediate pairs, the inner- 
most is entire and the outer ends in spines which fail to reach the distal edge. Between 
the submedian and first intermediate carinae there is a prominent proximal tubercle, 
and another which is elongated and forms a stout keel, is found between the second 
intermediates and the laterals. 

In the single specimen examined the height of the median crest of the telson 
above the general surface is less than one-sixth the greatest breadth of the telson, and 
below its small terminal spine there are two blunt points. In the anterior half there 
are on each side two closely-approximated submedian carinae which meet or almost 
meet at the distal end of the median ridge, an intermediate which is continuous, 
though not very clearly so, with that which runs to the apex of the submedian spine, 
and the second lateral which fuses distally with the marginal to form the lateral mar- 
ginal spine. The first lateral carina, found in O. japonicus (see text-fig. 4, p. 12) 
does not exist in the anterior part of the telson. There is a strong carina in the 


1013.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacijic Region. 137 


posterior half of the telson which supports the intermediate spines and another, shorter 
and far less conspicuous, which runs out on to the intermediate marginal denticle. 
The six principal marginal teeth are well-developed ; the submedians are broad and 
have small movable tips. In the large specimen examined there are only the most 
obscure traces of minute submedian spinules. Normally there is one large intermediate 
and one lateral denticle, which may bear one or two minute apical spinules. The 
ventral surface of the telson is quite smooth. 

The ventral process of the uropods bears two blunt keels on its lower surface and 
is cut distally into two spines, the inner of which is about two-thirds the length of the 
outer. The proximal segment of the exopod is provided with twelve movable spines 
on its outer margin ; the distal segment extends beyond the apex of the last of these 
spines and is only a little more than half the length of the proximal. 

In colour the preserved specimen is olivaceous green with obscure traces of marb- 
ling. The carinae of the sixth somite and telson are quite pale, and the antennal scale 
is pale with a large dark distal patch. According to Milne-Edwards’ coloured figure 
the marbling is conspicuous in fresh specimens and the raptorial dactylus, the spines 
on the outer uropod, and all the fringes of setae are crimson. 

Odontodactylus bleekeri, founded on a specimen from Batavia, is described by 
A. Milne-Edwards as follows: ‘‘ Cette espèce . . . . se distingue du Gonodactylus 
scyllarus par la forme de la plaque rostrale, qui, bien qu’élargie a sa base, se termine 
en pointe, et par l’existence d’une caréne médiane lamelleuse que porte en dessus le 
dernier segment de l’abdomen ; cette carène est notablement moins élevée que celle du 
Gonodactyle cultrifer.’’ A Milne-Edwards seems to have misunderstood the characters 
of O. scyllarus, for, in the features which he notes, his new species does not seem to 
present any difference from that form. 

Of Odontodactylus scyllarus I have seen only a single example :— 


3032 
5 


Mauritius. Purchased. 19,115 mm. 


This species appears to have a wide Indo-pacific distribution extending from 
Samoa and Japan to Madagascar. It has been recorded from Samoa (Miers, Bigelow) ; 
Rotuma (Borradaile); Kii, Japan (Rathbun); New Britain (Borradaile); New 
Guinea and Goram I. (Miers) ; Amboina (Miers, de Man, Ortmann, Nobili) ; Celebes 
(Rumph); Lette I.,Timor(de Man); Batavia(A. Milne-Edwards) ; Trincomali (Müller) : 
Seychelles (Miers) ; Zanzibar (Miers) ; Madagascar (Miers, Richters); Réunion (Milne- 
Edwards, Hoffman); Mauritius (Miers). 


2. Odontodactylus cultrifer (White). 
1850. Gonodactylus cultrifer, White, Proc. Zool. Soc., p. 96, pl. xvi, figs. I, 2. 
1851. Gonodactylus cultrifer, White, Ann. Mag. Nat. Hist. (2), VII, p. 422. 
1880. Gonodactylus cultrifer, Miers, Ann. Mag. Nat. Hist. (5), V, p. 117. 
1894. Odontodactylus cultrifer, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 
1901. Gonodactylus culirifer, Lanchester, Proc. Zool. Soc., II, p. 555. 
Of this species I have seen no specimens. According to the accounts of White, 
Miers and Lanchester and from the figures which the first of these authors has given, 
it may be separated from O. scyllarus by the following characters :— 


138 Memoirs of the Indian Museum. | NO I, 


I. The rostrum is quadrilateral, with the sides distally convergent and the an- 
terior margin straight. 

2. The eyes, according to White’s figure, are much larger, the breadth of the 
cornea being about one-third the length of the carapace excluding the 
rostrum. 

3. The dactylus of the raptorial claw is much less strongly dilated at the base, 
but otherwise resembles that of O. scyllarus. 

4. There are only six carinae on the sixth abdominal somite, the inner interme- 
diate pair found in the preceding species being absent. 

5. The median crest of the telson is very remarkably elevated ; its height above 
the general surface is nearly equal to the breadth between its base and the 
lateral margin. The submedian carinae are entire ; the submedian spines are 
less expanded at the base than in O. scyllarus and their movable tips appear 
to be larger. 

6. The basal segment of the outer uropod is, according to the figure, only about 
two-thirds the length of the distal segment ; there are only eight or nine 
movable spines on its external margin. 


Lanchester notes that in his specimen the rostrum does not nearly reach the base 
of the eyestalks as it appears to do in White’s figure. Even in the latter a greater 
portion of the ophthalmic somite is exposed than in O. scyllarus. 


White remarks that in his dried specimen the greater part of the upper surface is 
tinged with a reddish hue, while along the middle of the back there is a pale line. 


O. cultrifer is known only from two examples, 4 and 44 ins. in length. One is 
recorded from China (White, Miers) and the other from Kelantan in the G. of Siam 
(Lanchester). 


3. Odontodactylus carinifer (Pocock). 
1893. Gonodactylus carinifer, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 478, pl. xx B, figs. 4, 4a, b. 
1894. Odontodactylus carinifer, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 
O. carinifer is known only from a single small specimen which is distinguished 
from the two preceding species by the following characters :— 


1. The rostrum is semicircular, not quadrilateral or cordiform. 

2. The eyes are extremely large, according to Pocock’s figure the breadth of the 
cornea is exactly half the median length of the carapace excluding the 
LOSC NS 

3. The dactylus of the raptorial claw is moderately dilated at the base and is 
armed along its inner margin with three sharp subequal teeth in addition to 
the spinous termination. 

4. The sixth abdeminal somite bears six carinae only, as in O.cultrifer. The 
telson also resembles that of this species, but the median crest is only half 
the height, and the seccrd lateral carina, fourd in the two preceding species 
running to the apex of the lateral marginal teeth, is absent. 


1923.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 139 


Pocock observes that O. carinifer may possibly prove to be a young example of 
O. cultrifer, but this, I believe, is improbable. 


The single specimen, a male, 24 mm. in length, wascaught in 24 fathoms on the 
Holothuria Bank, China Seas. 


4. Odontodactylus elegans (Miers). 
1884. Gonodactylus elegans, Miers, Voy. H.M.S. “ Alert,’ p. 566, pl. Li, figs. B, b. 
1894. Odontodactylus elegans, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 
Odontodactylus elegans may be readily separated from all other species of the genus 
by the sculpture of the sixth abdominal somite and telson. 


1. The rostrum is transverse, subtriangular, with its anterior margin regularly 
arcuated. 

2. According to the figure the breadth of the cornea is about one-third the 
median length of the carapace and the ophthalmic somite, as in O. scyllarus, 
is entirely concealed in dorsal view by the rostrum. 

3. The raptorial dactylus is only feebly dilated at the base and bears two teeth 
on its inner margin. 

4. On the third, fourth and fifth abdominal somites there are five small pits. 

5. On the posterior margin of the sixth abdominal somite there are four spines ; 
the longitudinal carinae found in other species of the genus are absent. 

6. Except for a single median keel the dorsal surface of the telson is smooth and 
without carinae. There are about twenty-six minute submedian spinules, 
and, on each side, two intermediate denticles and one lateral. 


I have seen no specimens of this species. The two individuals on which Miers 
based his description were found in 13-20 fathoms and 24 fathoms at Providence I., S. 
of the Seychelles. The largest example is only 23 mm. in length and is almost certain- 
ly immature. 


5. Odontodactylus japonicus (De Haan). 
1844.? | Gonodactylus japonicus, De Haan, in Siebold’s Fauna Japonica, Crust., atlas, pl. li, fig. 7. 
1845. Gonodactylus edwardsii, Berthold, Abhandl. kön. Ges. wiss. Gottingen, III, p. 30, pl. iii, fig. 6. 
1849. Gonodactylus japonicus, De Haan, in Siebold’s Fauna Japonica, Crust., Text, p. 225. 
1880. Gonodactylus japonicus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 116. 
1894. Odontodactylus japonicus, Bigelow, Proc. U.S. Nat. Mus. XVII, p. 496. 
1907. Odontodactylus japonicus, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 212. 
1910. Odontodactylus japonicus, Fukuda, Annot. Zool. Japon., VII, p. 145. 
1910. Gonodactylus edwardsii, Baiss. Abh. math.-phys. Klasse k. bayer. Akad. Wiss., Suppl. Bd., IT. 
AND), Ay 195 3% 


The principal characters of this species are as follows :— 

The rostrum is subtriangular, about one and a half times as broad as long ; the 
lateral margins are sinuous, and the apex, which is bluntly pointed, is very strongly de- 
flexed. The eyestalks are short and the greatest breadth of the cornea is contained about 


1 See footnote p. 66. 


140 Memoirs of the Indian Museum. [Vora lve 


five times in the length of the carapace excluding the rostrum. ‘The dorsal processes of 
the ophthalmic somite are composed of two wing-like processes which meet and fuse an- 
teriorly ; they are almost wholly exposed in dorsal view. The antennal scale is remark- 
ably large ; its length is at least equal to that of the carapace excluding the rostrum. 

The dactylus of the raptorial claw is only slightly dilated at the base and the knife- 
like edge on its inner margin is cut into from five to seven teeth. 

The exposed thoracic and first five abdominal somite closely resemble those of 
O. scyllarus, but the lateral depressions and ridges are practically obsolete: The 
postero-lateral angles of the fourth and fifth somites are rounded in the two large 
specimens examined. The sixth abdominal somite bears a rather obscure median carina 
and four others on either side. The lateral, outer intermediate and submedian carinae 
end in spines in De Haan’s figure, but appear to have been worn away in the two large 
specimens in the Indian Museum. Between the outer intermediates and the laterals in 
the anterior part of the somite there is a prominent rounded tubercle. 

The arrangement of the carinae on the dorsal surface of the telson resembles that 
shown in text-fig. 4, p.12. The median carina is not high. The submedians are longi- 
tudinally bisected and consist of a conspicuous inner and a less prominent outer ridge, 
the latter being distinctly beaded. The intermediate carina is continued, though with 
a slight interruption in the middle, to the apex of the submedian teeth. The first 
lateral is obscure ; it passes close to the anterior end of the carina of the intermediate 
marginal teeth, and trends inwards towards the base of the intermediate denticle. The 
second lateral carina is sharp and is continued to the apex of the lateral marginal tooth. 
Close to the anterior margin of the telson are two pairs of large tubercles, one placed 
between the submedian and intermediate carinae, and one between the intermediate 
and first lateral carinae ; the latter tubercle is sometimes obsolete. The marginal teeth 
do not differ greatly from those of O. scyllarus. | 

The ventral process of the peduncular segment of the uropod is cut distally into 
two broad teeth, the outer about twice the length of the inner ; each bears on its lower 
surface a strong flat ridge. The basal segment of the exopod is about one and a third 
times the length of the distal segment, and on its outer margin is provided with a series 
of twelve or thirteen movable spines. These spines are flattened and blunt at the 
apex ; the outermost reaches almost to the end of the distal segment. 

Iwo large specimens, pale yellow in spirit, are preserved in the Indian Museum :— 

“= Hongkong. R. Hungerford. I G', 142 mm. 


“5 Misaki, Sagami Sea, Japan. A. Owston. 16,138 mm. 


Odontodactylus japonicus was described by De Haan from Japan, and has since 
been found in several localities in Japanese waters (Fukuda, Balss). It has been re- 
corded from the Chinese Seas (Berthold, Miers) and from 55 fathoms on the Saya de 
Malha Bank, S.E. of the Seychelles (Borradaile). 


6. Odontodactylus hanseni (Pocock). 


1893. Gonodactylus hansenii, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 477, pl. xx, B, figs. 3-30. 
1894. Odontodactylus hanseni, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. I4I 


According to the description and figures which Pocock has given, this species 
differs from O. japonicus in the following features :— 


1. The rostrum is evenly rounded anteriorly from side to side and is twice (or 
according to the figure rather more than twice) as broad as long. 

2. The breadth of the corneal part of the eye is contained only a little more than 
three and a half times in the median length of the carapace excluding the 
rostrum. The ophthalmic somite is exposed in dorsal view, and the dorsa] 
process is deeply excavate anteriorly. 

3. The antennal scale is a little shorter than the median length of the carapace. 

4. The raptorial dactylus is armed with nine distinct teeth (including the ter- 
minal one ?). 

5. There are four pairs of longitudinal carinae on the sixth abdominal somite 
as in O. japonicus, but the feeble median keel found in that species is missing 
and the proximal tubercle, which is found in most other species between the 
lateral and outer intermediate carinae, is omitted, possibly by error, in 
Pocock’s figure. 

6. On the telson there is only a single submedian carina, the first lateral is absent 
and the second lateral instead of fusing with the marginal to form the lateral 
tooth as in O. scyllarus and japonicus is parallel to the edge and runs towards 
the outer margin of the intermediate tooth. 


The number of movable spines on the outer margin of the uropod is not given ; 
the distal spine is, however, said to reach to the apex of the ultimate segment. 


Pocock records a single female, 60 mm. in length, from the Macclesfield Bank, S. 
China Seas, in 35 fathoms. 


7. Odontodactylus latirostris, Borradaile. 


1907. Odontodactylus latirostris, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 212, pl. xxii, 
figs. 3, 34. 

This species is perhaps not distinct from Odontodactylus hanseni. Borradaile 
does not say anything of the affinities of his new form and was perhaps unaware of 
Pocock’s work at the time of writing. In the descriptions and figures which 
the two authors have supplied I am only able to detect the following trifling 
differences. 


I. The rostrum is a little broader in O. latirostris than in O. hanseni, and its anterior 
margin is more definitely truncate. 

2. The raptorial dactylus bears only seven teeth on its inner margin.’ 

3. On the sixth abdominal somite there is the customary tubercle near the an- 


1 According to Borradaile’s description. In the figure, however, there appears to be an additional 
tooth lying close alongside the apex and a series of obscure serrations on the external margin of the 
swollen basal part of the dactylus. The latter feature, which is not mentioned in the text, does not 
seem to occur in any other species of the genus. 


142 Memoirs of the Indian Museum. [Vor. IV, 


terior margin inside the lateral carina (this is perhaps omitted by error in 
Pocock’s figure). 

4. The telson is proportionately broader in O. latirostris, and the movable tips of 
the submedian spines are shorter. 

5. The terminal movable spine on the outer margin of the uropod fails to reach 
the apex of the ultimate segment. 


In such important characters as the breadth of the eye, the form of the exposed 
dorsal processes of the ophthalmic somite and the details of the carination of the telson 
there appears to be the closest resemblance between the two forms. . 

In O. latirostris the lateral margin of the seventh thoracic somite is rounded and is 
distinctly narrower than that of the sixth, which is truncate. There are ten movable 
spines on the outer margin of the exopodite of the uropods. 

Borradaile records two specimens of this species, the largest 55 mm. in length, 
from the Amirante Is., in 25-30 fathoms. 


8. Odontodactylus southwelli, Kemp. 
Plate IX, figs. 103-106. 
1011. Odontodactylus southwelli, Kemp, Rec. Ind. Mus., VI, p. 94. 

This species is allied to the two preceding, but may be distinguished from them 
at a glance by the enormous size of the eyes. Its principal characters are as follows :— 

The rostrum is almost exactly twice as broad as long; its anterior margin is evenly 
rounded from side to side and is not depressed in the centre of its distal margin. The 
eyes (fig. 103) are enormously dilated, the breadth of the cornea being contained only 
from two to two and a quarter times in the median length of the carapace excluding 
the rostrum. A greater portion of the ophthalmic somite is exposed than in any of the 
allied species ; it bears dorsally a pair of strong anteriorly convergent ridges, and in 
the centre is very conspicuously excavated, the cavity so formed taking the shape of a 
deep and almost circular pit. The antennal scale is about four-fifths the median length 
of the carapace. 

The dactylus of the raptorial claw (fig. 104) is moderately dilated at the base and 
bears seven to nine well-defined teeth on its inner margin in addition to the terminal 
one. | 

The lateral margins of the sixth and seventh thoracic somites are rounded, that 
of the sixth being a trifle broader and more broadly rounded than that of the seventh 
(fig. 105). A lateral depression confined by a pair of blunt carinae is visible on the ~ 
fourth and fifth abdominal somites as in O. scyllarus and traces of a similar depression 
may be seen on the two preceding somites. The postero-lateral angles of the fourth 
and fifth somites are spinous. The arrangement of the carinae on the sixth abdominal 
somite is similar to that of the two preceding species (fig. 106). The submedian, second 
intermediate and lateral pairs of carinae terminate in spines; the first intermediates 
are connected proximally with the submedians, and the usual proximal denticle inside 
the laterals is well-marked. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 143 


On the telson (fig. 106) the carinae are similar to those of the two preceding species. 
The intermediates are continued to the apices of the submedian teeth, and the second 
lateral is parallel to the external edge and does not run to the tip of the lateral tooth. 
In the bifurcation between the submedian teeth there are twelve to sixteen pairs of 
minute spinules ; there are two sharp-pointed denticles between the submedians and 
intermediates and one between the intermediates and laterals. 

The outer of the two spines forming the bifurcate process of the uropod is one and a 
half times as long as the inner and reaches to the apex of the endopod. On the basal 
segment of the exopod there are ten or eleven {rarely nine) movable spines, the outer- 
most reaching almost or quite to the apex of the ultimate segment. 

The specimens in the collection vary considerably in length, but the proportional 
size of the eye remains approximately constant. 

There are seven examples of Odontodactylus southwelli in the Indian Museum :— 


== Andamans. “ Investigator.’ 19,37 mm. TYPE. 
se Andamans ; 53 fms. “ Investigator.’ Ig ,22,22—31 mm. 
"2 Off Cinque Is., Andamans ; 20 fms. ‘ Investigator.’ 19,22 mm. 

ue Off Interview I., Andamans. “ Investigator.’ Id, ca. 28 mm. 

55 N. Cheval Paar, Ceylon. T. Southwell. 19,21 mm. 


10 


9. Odontodactylus brevirostris (Miers). 


1884. Gonodactylus brevirostris, Miers, Voy. H.M.S. ‘ Alert,’ p. 567, pl. li, fig. C. 

1894. Odontodactylus brevirostris, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 496. 

1906. Odontodactylus brevirostris, Tattersall, Ceylon Pearl Oyster Rep., V, p. 172, pl. ii, figs. 16-18, 
1907. Odontodactylus brevirostris, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 212. 


Odontodactylus brevirostris is easily distinguished from all other species of the same 
group (?.e. those with more than three teeth on the raptorial dactylus) by the complete 
absence of lateral carinae in the anterior part of the telson. 

The rostrum is more than two and a half times as wide as long, and its lateral and 
anterior margins form an evenly rounded curve. ‘The middle of its distal margin is 
slightly depressed. The dorsal processes of the ophthalmic somite consist of a pair 
of upstanding triangular projections placed close together near the middle line and in 
dorsal view almost concealed by the rostrum. The eyes are large. The breadth of the 
cornea is contained about two and a half times in the median length of the carapace, 
excluding the rostrum. 

The dactylus of the raptorial claw is only very slightly swollen at the base. It is 
sharply notched at the proximal end of its external margin and bears six to nine (usu- 
ally seven or eight) teeth, not reckoning the spinous apex. 

The lateral margins of the sixth and seventh thoracic somites are rounded, not 
truncate, and are of equal width. The fourth and fifth abdominal somites are spinous 
at the postero-lateral angles and show feeble indications of a lateral depression similar 
to that noticed in O. scyllarus. The sixth abdominal somite bears six longitudinal 
carinae all of which terminate in spines. The inner intermediate carina found in the 


I44 Memoirs of the Indian Museum. [Vor. IV, 


allied species is obsolete, and its position is indicated merely by a low smooth eminence. 
The customary proximal tubercle is present between the outer intermediate carinae 
and the laterals. 

The telson differs from that of most of the preceding species in the entire absence 
of lateral carinae. The intermediate carinae run towards, but are not continuous 
with, those on the submedian marginal spines. The margin between the submedian 
spines is provided with a number (about sixteen pairs) of minute spinules; there are 
two sharply acute denticles between the submedians and intermediates and one between 
the intermediates and laterals. 

The outermost of the two spines forming the bifurcate process of the uropod is 
more than one and a half times the length of the inner, and reaches almost to the apex 
of the exopod. There are ten or eleven movable spines on the outer margin of the basal 
segment of the exopodite and the distal one, which is the longest, reaches to or beyond 
the apex of the ultimate segment. 

In one of the specimens examined the rostrum is only slightly depressed in the 
middle line, as figured by Tattersall, but there is a small incision, perhaps due to in- 
jury, in the middle of the distal margin. In the outer example the centre is more 
obviously deflexed and the margin is entire. In both examples the mandibular palp 
appears to be missing. The specimens are perhaps immature ; but in 7. southwelli 
the palp is distinct, though not very clearly segmented, in individuals of a similar size. 


Brown mottling is visible on the dorsal surface of one of the preserved specimens 
and forms decided patches in the middle of the last three thoracic somites. In this 
individual also the basal segment of the exopod of the uropod is very dark brown except 
for a pale postero-lateral triangular area. 


This species is closely allied to Odontodactylus havanensis, Bigelow (1894, p. 479), 
the only known Atlantic representative of the genus. The form described by the 
American author seems, however, to be distinguished by the following characters :— 

The rostrum is longer and has no median depression; the greater part of the 
ophthalmic somite is exposed in dorsal view; the antennal scale is longer and the 
basal part of the raptorial dactylus is strongly swollen and is not notched externally 
at the base. 

O. havanensis is known from a single specimen, 20 mm. in length, found in 163 


fathoms off Havana, Cuba. 


There are only two examples of Odontodactylus brevirostris in the Indian 


Museum :— 
= Andamans. “ Investigator.’ 19,23 mm. 
TT Andamans, 53 fathoms. ‘ Investigator.’ 19 ,2I mm. 
This species has been previously recorded from the Pearl Banks, G. of Manaar 
(Tattersall), from the Seychelles in 34 fathoms (Borradaile), and from Providence I., S. 


of the Seychelles, in 19 fathoms (Miers). 


TOES S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 145 


Genus Gonodactylus, Latreille. 


1825. Gonodactylus, Latreille, Encycl. Méthod., X, p. 473. 

1837. Gonodactylus, H. Milne-Edwards, Hist. Nat. Crust., II, p. 528 (partim). 

1849. Gonodactylus, De Haan, in Siebold’s Fauna Japonica, Crust., p. 220. 

1880. Gonodactylus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 115 (partim). 

1886. Gonodactylus, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 55. 
1886. Protosquila, Brooks, ıbid., p. 65. 

1894. Gonodactylus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 492. 

1894. Protosquilla, Bigelow, tbid., p. 492. | 

1895. Gonodactylus, Hansen, Isop. Cumac. u. Stomatop., d. Plankton-Exped., p. 86. 
1906. Gonodactylus, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 326. 


Carapace little, if at all, narrowed anteriorly ; without carinae; gastric grooves 
distinct, cervical groove wholly absent; antero-lateral angles rounded or acute. 
Cornea of eye sometimes indistinctly bilobed, mandibular palp present and composed 
of two or threesegmentsorabsent. Ischio-meral articulation of raptorial claw situated 
at a point in advance of proximal end of merus, the latter segment consequently 
extending backwards beyond the joint ; inferior surface of merus not channelled longi- 
tudinally throughout its entire length ; upper margin of propodus not pectinate ; 
dactylus inflated at base, without teeth on inner margin. Shorter ramus of last three 
thoracic appendages linear, composed of two segments. Free thoracic and abdominal 
somites compressed, no sharp carinae on first four abdominal somites. Telson very 
variable in form and sculpture, carinate, tuberculate or spinous. Ventral process of 
uropods terminating in two sharp spines, inner usually shorter than outer. 


There are no well-marked secondary sexual distinctions. 


In reducing Brooks’ Protosguilla to a synonym of Gonodactylus I have followed 
Hansen and Nobili. The character which Brooks employed, the fusion or free articula- 
tion of the last abdominal somite and telson, is, as Hansen has shown, wholly unreliable 
and no other feature sufficiently well-marked to serve as a generic distinction can be 
found. On the contrary it seems that the species here included in Gonodactylus fall 
into four natural groups of approximately equal value, but these do not, I believe, possess 
any greater claims to nomenclatorial distinction than do the groups in Lysiosquilla 
or in other genera. 


The four groups may be distinguished by the following characters :— 


GROUP I.— The anterior margin of the carapace slopes forward on either side of the rostrum, so that 
the antero-lateral angles are well in advance of the rostral base. The rostrum has a sharp median spine and 
its antero-lateral angles are rounded or subacute : in rare cases acute. ‘The mandibular palp is composed 
of three distinct segments. The posterior margin of the sixth abdominal somite is straight or slightly 
convex in dorsal view. The basal segment of the outer uropod projects posteriorly a little beyond the 
articulation of the ultimate segment, and its external margin is armed with a series of straight, or almost 
straight, movable spines. The inner uropod is normal in shape and, asa rule, in the disposition of its setae. 


This group includes G. chiragra, demani, graphurus and their immediate allies. 


Group II.—The anterior margins of the carapace slope forwards as in the preceding group. ‘The 
rostrum is trispinous : the antero-lateral angles are produced to sharp spines often almost as slender as the 


146 Memoirs of the Indian Museum. VorsNVz 


median. ‘The mandibular palp is two or three-segmented.! The posterior margin of the sixth abdominal 
somite is straight or slightly convex in dorsal view. The basal segment of the outer uropod extends far 
beyond the point of articulation of the ultimate segment, and the spines on the external margin of this 
projecting portion are in most cases very distinctly recurved. The inner uropod is always abnormal in 
shape ; it is usually more or less crescentic when seen from above, or may be strongly arched in lateral 
view. ‘The setae are restricted to only a portion of the margin or may invest all the edges and the greater 
part of the upper surface as well. 

This highly-specialized and peculiar group includes six species, all of which seem to be very rare. They 
are G. herdmani, dvepanophorus, brevisquamatus, fimbriatus, spinosocarinatus and furcicaudatus. The first 
two possess a closer affinity with the preceding section than the rest. 


Group III.-—The anterior margins of the carapace do not slope forwards on either side of the rostrum 
and the antero-lateral angles are consequently not in advance of the level of the rostral base. The rostrum, 
typically, is trispinous; the antero-lateral angles are produced to sharp spines similar to the median, but these 
in rare cases are only acute. The mandibular palp comprises two segments only.” The posterior margin of 
the sixth abdominal somite, when seen in dorsal view, is distinctly convex in the middle. The basal seg- 
ment of the outer uropod articulates terminally with the ultimate segment and is armed with straight mov- 
able spines on its external edge. The inner uropod, as in Group I, is normal in shape and in its investment 
of setae. ; 

(This group includes nearly all the species which Brooks referred to Protosquilla. It comprises G. tris- 
spinosus, stoliurus, excavatus, folini, brooksi, spinosissimus and numerous allied forms. 


Group IV.—The anterior margins of the carapace slope forwards on either side of the rostrum, so 
that the antero-lateral angles are well in advance of the rostral base. The rostrum is trispinous, but the 
lateral spines are not so well developed as in most species of the two preceding groups. The mandibular 
palp is wholly absent. The posterior margin of the last abdominal somite is straight in dorsal view. The 
basal segment of the outer uropod projects posteriorly a little beyond the insertion of the ultimate seg- 
ment and bears a series of straight spines on its outer margin. Both the inner uropod and the terminal 
segment of the outer uropod are provided with setae only on the distal half of their outer margins ; the 
inner margin in each case bears sharp spines. 

This group includes a single species only ; G. acanthurus. 


Of the thirty-two species here recognized, twenty-eight are restricted to Indo- 
pacific waters. Three, G. oerstedi, G. festae, and G. folini, are known only from the 
Atlantic, while G. glabrous, a widely distributed Indo-pacific form, is recorded from 
the Mediterranean. . 

Brooks (1893, p. 353) has given a most valuable account of the habits of 
Gonodactylus * in the Bahamas, where he found that the animals made burrows in 
hard stone. He gives most interesting details of their cannibalistic propensities and 
of the manner in which the female tends the egg-masses. Of the habits of the Indo- 
pacific species practically nothing is known ; they seem to prefer rough ground and 
coral reefs and possibly all of them burrow in coral or rock as in the case of the 


! Two species only have been examined; the palp is three-segmented in G. herdmani, two-segmented 
in G. furcicaudatus. 

? Six species examined. 

® Brooks calls the species G. chiragra, but the specimens doubtless belong to G. oerstedi, Hansen. 


DOS] S. KEMP: Crustacea Stomatopoda of the I ndo-Pacific Region. 147 


species observed by Brooks.! Asa rule they are found in shallow water, but G. spin- 
osissimus and G. acanthurus have been recorded from 45-50 fathoms. 


Key to the species of Gonodactylus. 


I. Mandibular palp present ; margins of inner uropod and ultimate segment of 
outer uropod, not bearing spines. 

A. Antero-lateral angles of carapace well in advance of rostral base; posterior 
margin of last abdominal somite straight or slightly convex in dorsal view ; 
basal segment of outer uropod extending at least a little beyond inser- 
tion of ultimate segment. 

I. Rostrum with antero-lateral angles rounded or acute, never sharply spinous ; 
inner uropod normal in shape and in its investment of setae [mandibular 

palp three-segmented]. Group I. 

A. Mid-dorsal portion of telson with three long keels, occasionally with a 
short additional keel on either side at hinder end of median. 
I. No ridge on inner side of keel of intermediate marginal teeth of 
telson. 

a. Dorsal processes of ophthalmic somite large and subtriangular; me- 
dian keel of telson not very strongly arched in lateral view, its 
depth ! less than half its greatest breadth ; no spinules on dorsal 
surface of telson. 

i, Antero-lateral angles of rostrum rounded or subacute ; a pair of 
small round tubercles close to anterior margin of telson; sub- 
median dorsal keels of telson parallel. 

a. Form narrow; abdomen compressed and strongly arched from 
side to side; lateral marginal teeth of telson small but 


distinct, rarely absent .. cs er .. chivagra, p. 155. 
ß. Form broader; abdomen less compressed; lateral marginal 
teeth of telson wholly absent a it .. do.v. platysoma, p. 162. 


ii. Antero-lateral angles of rostrum acute ; no tubercles near anterior 

margin of telson ; submedian dorsal keels of telson posteriorly 
convergent Ber Ei .  acutirostris, p. 163. 

b. Dorsal processes of heine somite consisting a pair os small a 

inconspicuous transverse plates; median keel of telson very 

strongly arched in lateral view, its depth ? fully half its greatest 

breadth ; dorsal surface of telson usually beset with spinules. 
i. Telson broader than long; intermediate marginal teeth well-deve- 


loped ; dorsal surface with spinules a & .. demant, p.164. 
ii. Telson aslong as broad; intermediate marginal teeth rudimentary 
or absent ; dorsal surface with abundant spinules .. .. do.v. spinosus, p. 165. 


iii. Telson broader than long; intermediate marginal teeth rudimen- 
tary; dorsal surface without spinules .. .. do. v. espinosus, p.166. 
2. À ridge on inner side of keel of intermediate marginal fect of telson. 


1 See addendum, p. 197, for further notes on this subject. 
? Measured from the highest point of the median keel to the corresponding point on the lower 
surface. 


148 Memoirs of the Indian Museum. [Moraine 


a. Antero-lateral angles of rostrum rounded; no spinules on dorsal 


surface of telson .. ip = a .. [oerstedr].! 
b. Antero-lateral angles of rostrum acute ; numerousspinules on dorsal 
surface of telson .. 3 { = -..  [festae]. 
B. Mid-dorsal portion of telson with ane long ven 
I. First five abdominal somites smooth os 5 .. glabrous, p. 167. 
2. First five abdominal somites finely grooved transversely es longitu- 
dinally .. a ae 52 de .. graphurus, p. 160. 


II. Rostrum sharply trispinous ; inner uropod abnormal in shape and in its 
investment of setae[mandibular palp two- or three-segmented]. Group 
106 | 
A. Telson normal in shape, its length greater or only a trifle less than its 
length. 
I. Spines on external edge of basal segment of outer uropod forming an 
even series throughout its length, outermost slightly recurved ; 
inner uropod more or less crescenticinshape, apex curved inwards. 
a. Submedian teeth of telson without spinules on inner ınargins ; 
lateral teeth represented by blunt rounded lobes ; dorsal surface 
with three keels and a few rounded tubercles pi .. herdmant, p. 171. 
b. Submedian teeth of telson with spinules on inner margins ; lateral 
teeth sharp and well-formed ; dorsal surface with three keels and 
large sharp spinules es a : .. dvepanophorus, p. 173. 
2. On external edge of basal segment of outer uropod two or three spines 
at distal end form large hooks, proximal part bare or with few 
small straight spines ; apex of inner uropod more or less curved 
outwards. 
a. Intermediate teeth of telson edge large ; lateral teeth absent. 
i. Intermediate ridges of last abdominal somite much narrower 
than submedians ; mid-dorsal portion of telson occupied by 
three large ridges? ; no spinules on inner edges of submedian 
marginal teeth .. 6:8 5 N ..  brevisquamatus, p. 174. 
11. Intermediate and submedian ridges of last abdominal somite of 
equal breadth ; mid-dorsal portion of telson with three keels 
closely packed together in middle, with two fainter keels on 
either side? ; spinules on both edges of submedian marginalteeth fimbriatus, p. 175. 
b. Intermediate teeth of telson edge very small ; lateral teeth minute 
[dorsal surface of telson with nine closely packed keels] .. Spinoso-carinalus, p. 173. 
| B. Telson very abnormal in shape, about four times as broad as long, 
sharply truncate behind ; the truncate portion reniform in shape 
and honeycombed, with a pair of long furci springing from the middle 
of its inferior edge .. sin : ; .. furcicaudatus, p. 176. 
B. Antero-lateral angles of carapace not in advance of rostral bs inedian 
part of posterior margin of last abdominal somite concave in dor view ; 


| Hansen 1895, p. 65 (footnote) and Bigelow, 1902, p. 152, text figs. I, 2. To the synonymy of this 
species as given by the latter author, the following may be added :— 
1890. Gonodactylus chiragra, Pocock, Journ. Linn. Soc. Zool., XX, p. 526 (Fernando Noronha). 
1897. Gonodactylus chiragra, Nobili, Boll. Mus. Torino, XII, no. 280, p. 6 (Darien). 
1901. Gonodactylus falcatus, Moreira, Arch. Mus. Nac. Rio de Janeiro, XI, p. 1 (Brazil). 


* Not reckoning the ridge that runs to the apex of the intermediate marginal teeth. 


1913.] 


basal segment of outer uropod articulating terminally with ultimate 
segment [mandibular palp two-segmented]. Group III. 
I. Large spines on last abdominal somite and telson with a soft fleshy pro- 
cess protruding from apex : ai > : 
Large spines on last abdominal somite and telson, if present, without 
fleshy process at apex. 
A. Distal margin of telson divided into right and left halves by a long and 


very narrow median fissure with its edges partly in contact with one 
another. 


II. 


I. Three round or oval bosses in centre of telson, external boss on each 
side not reaching distal margin. 
a. External boss on each side reaching only to middle of telson. 
i. Median portion of fifth abdominal somite entirely smooth. 
a. Rostrum sharply trispinous ; dorsal processes of ophthalmic 
somite produced, with acute antero-lateral corners : 
ß. Antero-lateral angles of rostrum acute but not spinous ; dorsal 
processes of ophthalmic somite not produced antero-late- 
rally as ae 4 ; 
. Median portion of fifth abdominal somite en, ee) 
a. Rostrum sharply trispinous ; external bosses of telson oval 
in outline : BR Ne 5 
B. Antero-lateral angles af rostrum acute but not spinous ; 
external bosses of telson strictly circular in outline 
b. External boss on each side reaching well beyond middle of telson, but 
not extending to distal margin. 
i. Median portion of fifth abdominal somite smooth ; distal margin 
of telson with three large teeth on either side of median fissure. 
a. Dorsal surface of telson, between bosses, smooth 
B. Dorsal surface of telson, between bosses, covered with well- 
marked irregular depressions 50 ac 
ii. Median portion of fifth abdominal somite furrowed; distal margin 
of telson with four large teeth on either side of median fissure .. 
2. ‘Three bosses in centre of telson, external boss on each side termina- 
ting in a point on distal margin. 
a. Fifth abdominal somite smooth mid-dorsally ; 
ed bosses ; 


sixth with six round- 
bosses of telson smooth and entire st 
5. Fifth and sixth abdominal somites finely grooved ; dorsal bosses of 
telson incised at margins by fine grooves runnings towards summit 
of each boss Sc 30 
B. Distal margin of telson divided into right a left halves by a large, 
wide and deep excavation. 
1. No spines or spinules on dorsal surface of telson. 
a. Distal margin of telson with two large teeth on either side of median 
excavation À sc 
b. Distal margin of telson with tbe large ei blunt teeth on either 
side of median excavation : 
2. Dorsal surface of last abdominal somite ad los u with spines 
or spinules. 
a. With short spinules. 
i. Telson with three pyriform bosses ; spinules straight 


S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 


guerini, p. 192. 


pulchellus, p. 177. 
nefandus, 


pP. 179. 


trispinosus, p. 180. 


tuberosus, p. ISI. 


glaber, p. 182. 
tanensis, p. 183. 


stoliurus, p. 184. 


ectypus, p. 185. 


glyptocercus, p. 186. 


excavatus, p. 187. 


[folini, p. 189.] 


brookst, p. 180. 


150 Memoirs of the Indian Museum. Worms 


ii. Telson with three circular bosses ; spinules with recurved points 
directed backwards oh ase ae USA AD Os 


b. With long spines spinosissimus, p. IQI. 


II, Mandibular palp absent; inner margins of inner uropod and of ultimate 
segment of outer uropod bearing large spines [numerous slender dorsal 
spines at distal end of telson]. Group IV. se as .. acanthurus, p. 193. 


Gonodactylus chivagra and its allies. 


There is considerable difference of opinion respecting the precise relationships 
between Gonodactylus chiragra and its near allies oerstedi, acutirostris, demant, spinosus, 
espinosus, festae, glabrous and graphurus, and Lanchester, whose important paper on the 
subject (1903) deserves careful consideration, holds that all must be regarded as varie- 
ties of a single species. In this view he has not been followed by more recent authors, 
and a careful survey of the large collection in the Indian Museum seems to show that 
he has omitted to lay sufficient emphasis on the more outstanding features of the case. 

The variation shown in these species of Stomatopoda presents problems of rather 
unusual interest and before any final conclusions can be reached much work based on 
living material must be undertaken ; it is, I believe, only by direct observation on the 
post-larval growth, habits and associations that a true estimate of the relationships 
of the various forms can be reached. ‘The variation in these species seems to find no 
parallel among other Stomatopoda, many of which, notably the species of Sqwlla 
belonging to the S. oratoria group, are remarkable for the constancy of small and seem- 
ingly trivial characters. 

The outstanding features of the variation in the chiragra section of the genus 
Gonodactylus appear to me to be as follows :— 

Distributed throughout the Indo-pacific region, occurring on rough ground, in 
rock-pools or on coral reefs, a form which may be called Gonodactylus chiragra, s.s.,' 
is abundant. In the characters afforded by the ornamentation of the last abdominal 
somite and telson a large amount of variation may be found ; the keels with which 
these segments are furnished may be swollen or narrow and may or may not terminate 
in spines ; the median carina on the telson is moreover simple and linear or it may bear 
short lateral branches at its distal end and so take the form of an ‘ anchor.’ Between 
the various forms every shade of intermediate can be found. Intermediate forms in 
any selected character are also commoner than those which exhibit it in an extreme 
development, but in their turn are less abundant than those which conform more or less 
closely with the prevailing type. 

Now in examining large series of examples one feature is predominant in the Indian 
Museum collection : the variation shown among large examples (70 mm. or more in 
length) is very much less than that found in small specimens (60 mm. or less in length), 
and it is further to be noticed that the individuals which in Lanchester’s opinion formed 
connecting links between this form and its allies are also characterized by their ex- 
tremely small size. 


! The characters employed for the separation of this form from its allies are shown in the key 
on p. 147. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. I5I 


From this result various interesting speculations arise. Must we conclude (i) that 
all the small specimens exhibiting extremes of variation die before reaching maturity, 
or (ii) do they represent dwarf races capable of breeding and perpetuating their precise 
type, or (iii) do they at subsequent moults lose their peculiar features and approximate 
to the typical form ?' 

Of these alternatives the second is, I believe, very improbable,” though with our 
present lack of knowledge regarding the size of sexually mature specimens, it cannot be 
wholly disregarded. There remain the first and third ; the former implies a large nu- 
merical preponderance of small individuals, a factor at present unproved, while the latter, 
the theory to which I myself am inclined, can only be properly substantiated by prc- 
longed rearing experiments demanding infinite patience and beset with many technical 
difficulties. This theory nevertheless will acquire much probability if the two others 
can be disproved, a task somewhat easier in performance. 


The question of the nomenclature of this varietal series in G. chivagra is one of less 
interest. Numerous terms such as incipiens, tumidus, anancyrus, etc., have been em- 
ployed ; but if, as I believe, it can be shown that the case is to a very large extent one of 
continuous variation, the adults approaching a standard type while young examples 
exhibit a much greater diversity of form, all the terms must lapse, for the application 
of names to arbitrarily selected points in a perfectly graded series cannot be upheld, 
nor is it clear that any limit can be set to such a process. 

In this connection I would refer to Borradaile’s valuable discussion on ‘ Varieties 
in Decapod Crustaceans’ * and in particular to the following passage in the opening 
paragraph.—‘‘ It must, however, be borne in mind that the intermediate specimens 
between varieties are much less numerous than those which exhibit the peculiarities 
of the varieties in a typical development. Otherwise it is only possible to say that the 
species exhibits considerable range of variation, but that definite varieties are not 
established.’ ’ 

With this statement I am in entire agreement. The author was doubtless un- 
awate of the main features of the present case when, at a later date, he divided the 

species G. chivagra into a considerable number of named varieties and the same time 
added to the named varieties of G. glabrous. That data of value from a distributional 
or from some other point of view are likely to accrue by a precise definition of these 
varietal phases is not improbable, but they should be designated by the letters or 


! In most known instances of variation the distinctions between the varietal and typical forms are more 
readily perceptible in the adult than in the young and instances similar to that found in G. chiragra seem 
to be exceedingly rare. ‘The only case, of which I am aware, that presents features of an analogous nature 
is afforded by Thalassochelys caretta. Gadow notes that in this species of turtle the number of dorsal 
scutes varies from 16 to 24 ; in adults, however, there are as a rule only 16 and abnormalities are from four 
to seven times as common in new-born as in mature specimens. ‘The explanation, according to Gadow, lies 
in the partial or complete correction of the abnormalities in the course of growth. [Gadow, Orthogenetic 
variation in the shells of Chelonia. Willey’s ‘‘ Zoological Results,’’ p. 207.] 

? There is, however, some evidence that this is the case with the form to which the name ‘ smithi’ 
has been applied. 

3 Borradaile, ‘‘ Fauna and Geog. of Maldives and Laccadives,’’ I, p. 193. 


152 Memoirs of the Indian Museum. Nor IH, 


numbers already in use, the term variety being restricted to such cases as conform with 
Borradaile’s definition. 

Turning to the forms that differ from G. chiragra in characters of a more trenchant 
nature, intermediates being rare or perhaps wholly absent, the first that must be con- 
sidered is Wood-Mason’s Gonodactylus platysoma. No description of this form has ever 
appeared and its claim to recognition rests solely on the figures published in 1805. 
G. platysoma may be distinguished from G. chiragra by its broader form and by the size 
and shape of the terminal part of the raptorial dactylus. The telson, with its narrow 
carinae and complete absence of lateral teeth on the margin differs markedly from that 
of any G. chiragra in the collection ; but in this respect the form bears a close resem- 
blance to certain examples to which Ianchester gave the name of G. chiragra var. acutus, 
and it is possible that these specimens are identical with those determined by Wood- 
Mason. Judging from the material in the Indian Museum G. platysoma seems quite 
sufficiently distinct from G. chivagra to claim recognition as a variety. 

In Atlantic waters G. chiragra is represented by a form known as G. oerstedi, 
Hansen, which differs from its Pacific relative in the possession of a small additional 
keel on the inner face of the intermediate teeth of the telson edge. The character is so 
trivial that for many years it entirely escaped detection, but it seems nevertheless to 
prevail with the most absolute constancy. I have followed the general consensus of 
opinion in regarding this form as a distinct species, but some may consider subspecific 
rank more suitable. 

When describing Gonodactylus acutırostris, Dr. de Man mentions that it is possibly 
only a variety of chiragra ; but, though it is true that it may eventually prove to bea 
very abnormal form of that species, it should remain separate pending further informa- 
tion. In the single known example the antero-lateral angles of the rostrum are much 
more acutely produced than in any G. chivagra,' and on the dorsal surface of the telson 
the two small tubercles, that are situated close to the anterior margin in all the allied 
forms, are entirely absent, while the submedian carinae are posteriorly convergent. 
I have seen no specimen possessing any one of these characters. 

Gonodactylus demani, Henderson, and spinosus, Bigelow, are forms which are dis- 
tinguishable at a glance from all the preceding by the presence of spinules on the dorsal 
surface of the telson. Apart from this feature G. demani bears a close resemblance to 
G. chiragra, from which, however, it may be separated by the shape of the dorsal pro- 
cesses of the ophthalmic somite and by the height and extreme convexity of the mid- 
dorsal lobe of the telson when seen in lateral view. The dorsal aspect of the telson 
presents, moreover, a very distinct and characteristic facies, for the three dorsal keels, 
and those which terminate in the submedian and intermediate marginal teeth, are all 
strongly swollen, and are separated from one another by narrow \/-shaped grooves that 
form a striking contrast to the comparatively wide interspaces found practically with- 
out exception in G. chiragra. Lanchester, strangely enough, does not mention this 
form in his synopsis of the varieties of G. chivagra; he has in fact omitted all reference 
to it, though he cites the paper by Henderson that contains the original description. 


! But there is, indeed, no little variation in G. chiragra in this respect. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 153 


But G. spinosus, which is unquestionably more remote from G. chivagra, he includes 
under the latter name as a variety. 

G. demam is, it seems, a smaller form than G. chiragra and does not exceed 40 mm. 
in length. The spinulation of the telson is most astonishingly variable (as will be seen 
by reference to pl. ix, figs. 109-111) ; but even if, as is not unlikely, an individual 
wholly destitute of spinules should appear, the other characters noted above will, I 
believe, remain amply sufficient to justify specific separation from G. chiragra. The 
variation in the number of spinules is unquestionably continuous. 

Gonodactylus spinosus, Bigelow, described one month later than Henderson’s 
G. demani, differs from that species only in the slightly greater proportional length of 
the telson and in the suppression of the intermediate teeth of the margin. But G. 
demam itself shows no little variation in this respect, and in the Indian specimens 
which I have referred to spinosus, these teeth are slightly better developed than in the 
typical individual figured by Lanchester. I am forced therefore to the conclusion, 
already suggested by Nobili, 1906 (a), p. 331, that G. spinosus can, at most, only be 
retained as a variety of G. demani.! The form appears to be rare ; in the details of its 
spinulation it is perhaps as variable as G. demanz,' but on this point little is known. 

Borradaile’s Gonodactylus espinosus, known only from a single specimen of small 
size, differs from spinosus solely in the greater breadth of the telson and in the total 
absence of spinules on its dorsal surface. In G. demani the variation in the number of 
spinules, as illustrated in the material examined, is so great that should a wholly smooth 
example be added thereto the series would still be almost continuous in its variation. 
On analogy therefore with its ally it may be expected thata complete transition between 
spinosus and espinosus will ultimately be found and, in giving the latter name admission 
as a variety, it must be understood that with the access of fresh material there is a pro- 
bability that the name will cease to possess any greater significance than ‘ inciprens 
“ tumidus ’ and others which have been employed in the case of G. chiragra. 

In his inclusion of spinosus and espinosus as varieties of G. chragra Lanchester 
has not, I believe, been followed by any subsequent writer, and I am unable to find any 
sufficient reasons for such a course either in his work or from an examination of the 
material in the Indian Museum. On the contrary the separation of G. chivagra and its 
variety from G. demani and its variety has been conspicuously easy in performance. 

Gonodactylus festae, Nobili, is an Atlantic species resembling G. oerstedi in the 
possession of a short additional carina on the intermediate marginal spines of the telson, 
but differing in the spinulation of this segment. It is only known from the original 
description based on a small number of specimens. It bears, perhaps, much the same 
relation to G. oerstedi that G. demani does to G. chiragra, though, according to Nobili’s 
account, it differs from all these and resembles de Man’s acutirostris in the sharply pro- 
duced antero-lateral angles of the rostrum. a 

Gonodactylus glabrous, Brooks, and G. graphurus, Miers, are more clearly dis- 
tinct and in the possession of five long keels in the middle of the dorsal surface of the 


! See addendum, p. 198. 


154 Memoirs of the Indian Museum. [Git JOM, 


telson are distinguishable at a glance from G. chivagra or any of the other species already 
mentioned. It is true that one particular telson pattern in G. chiragra, in which the 
flukes of the ‘ anchor’ are separated from the distal end of the median carina and form 
short additional carinae on either side of it, seems to represent a type intermediate be- 
tween that species and G. glabrous. But such specimens are invariably extremely small 
in size, and even if it were impossible to separate the two forms at this early age, which 
is not the case, there would still be no justification for uniting the two as Lanchester 
has done. In addition to the distinctions afforded by the telson, G. glabrous and gra- 
phurus may be distinguished from G. chiragra by the small size of the dorsal processes 
of the ophthalmic somite and by the presence of a small tooth at the base of the outer 
spine of the bifurcate process of the uropods ; but these characters are of little value in 
the case of very small specimens. 

G. graphurus is separated without difficulty from G. glabrous by the presence of the 
fine incised lines or grooves which traverse the first five abdominal somites in the 
former species. The character is an obvious one and, as yet, there is no evidence of the 
existence of intermediate forms. G. glabrous seems to be the most abundant of the two, 
and exhibits a very considerable amount of variation in the development of the keels 
on the telson. The keels may be narrow and well separated or swollen and touching one 
another and the three median keels, as well as those on the last somite of the abdomen, 
may or may not terminate in spines. There can be no doubt that the variation, as in 
the case of G. chiragra, is continuous and the same objections that have already been 
made must be urged in this instance also against the use of special names to designate 
particular varietal phases. G. graphurus seems to exhibit a range of variation closely 
similar to that of G. glabrous. 

The species belonging to the chiragra section of Gonodactylus form a very homo- 
geneous group. It seems probable that they are more primitive than the others in- 
cluded in the genus, and it may be inferred that all the other more highly specialized 
forms were derived from the common ancestor of this section. 

The abundance of species of Gonodactylus in the Indo-pacific region and their 
poverty in the Atlantic may perhaps indicate that they have originated in the former 
area, while the very slight distinctions that exist between G. chiragra and G. oerstedi 
may be taken as evidence that their extension to Atlantic waters is of compara- 
tively recent date, though it is possible that unknown factors, absent in the Atlantic, 
may have induced a more speedy differentiation of species in Indo-pacific waters. 


Of the species with a three-segmented mandibular palp, G. chivagra,in the simplicity 
of its structure appears the most primitive, while G. graphurus is perhaps the most 
highly specialized. 

As has already been pointed out, the vast range of variation in these species is the 
more remarkable when the singular constancy of slight specific distinctions in other Sto- 
matopoda is considered. I am of the opinion that four, or perhaps five forms in the 
Indo-pacific, and two in the Atlantic, are sufficiently isolated to merit specific recogni- 
tion ; but from the numerous varietal phases it is impossible, in the present state of 
our knowledge, to draw any satisfactory conclusions. That the ceaseless production of 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 155 


such forms—even though some seem never to reach maturity—will in the ordinary 
course of natural selection result in races specially adapted to particular environmental 
factors is far from improbable, and that it may already have occurred is still an open 
question. Results of great importance may be expected from direct observations on 
living animals and from a comparative examination of large collections from distant but 
strictly localized areas. 

The scheme of classification adopted for these species differs, though not very 
widely, from that given by most previous writers, but is diametrically opposed to 
that suggested by Lanchester. The latter author was, I believe, misled by the small 
size of the specimens which formed the vast majority of his material and many forms 
that in the opinion of all previous authors were entitled to specific distinction he re- 
garded as varieties of a single species. In this there is perhaps no great cause of com- 
plaint, for the classification of some of the forms must long remain a matter of in- 
dividual opinion. But in Lanchester’s scheme, which comprises a single species, 
fourteen named varieties and a large number of lettered and numbered subheads, the 
main features of the case are to a very great extent lost sight of, and certain forms 
which by reason of their structure, geographical distribution or abundance, stand out far 
ahead of all others are, in his account, placed on the same footing as other forms to 
which not nearly the same significance can be attributed. It is here that I must dissent 
most strongly from this author’s views. 

But, in any endeavour to arrive at a true understanding of these matters, the data sup- 
plied by Lanchester and the excellent figures which he has given will always prove of great 
value and though, in a recent revision of the genus, his work is wholly ignored, such a 
course can hardly tend to a satisfactory solution of the problems that are involved. 


1. Gonodactylus chiragra (Fabricius). 
Hate mee, ICY. 

1781. Squilla chiragra, Fabricius, Species Insectorum, I, p. 515. 
1787. Squilla chiragra, Fabricius, Mantiss. Insect., I, p. 334. 
1790. Cancer chiragricus, Linnaeus, Syst. Nat., ed. XIII, I, p. 2990. 
1793. Squilla chivagra, Fabricius, Ent. Syst. II, p. 513. 
1796. Cancer (mantis) chiragra, Herbst, Krabben u. Krebse, II, p. 100, pl. xxxiv, fig. 2. 
1798. Squilla chiragra, Fabricius, Ent. Syst., Suppl., p. 417. 
1823. Squilla chiragra, Desmarest, Dict. des Sci. Nat., XXVIII, p. 342. 
1825. Squilla chiragra, Desmarest, Consid. Crust., p. 251, pl. xliii. 
1825. Gonodactylus chiragra, Latreille, Encycl. Méthod., X, p. 473, Atlas, pl. ccexxv, fig. 2. 
1829. Gonodactylus chiragra, Latreille, in Cuvier’s Régne Anim., 2nd ed., IV, p. 109. 
1832. Gonodactylus chiragra, Owen, Proc. Zool. Soc., p. 6. 
1837. Gonodactylus chiragra, H. Milne-Edwards, Hist. Nat. Crust., II, p. 528. 
1843. Gonodactylus chiragra, Krauss, Stid-afric. Crust., p. 60. 
1847. Gonodactylus chiragra, White, List. Crust. Brit. Mus., p. 84. 
1852. Gonodactylus chiragra, Dana, U.S. Explor. Exped., Crust., p. 623, pl. xli, figs. 6a, b. 
1861. Gonodactylus chiragra, Heller, Verhandl. zool.-bot. Ges. Wien, XI, p. 29. 
1862. Gonodactylus chiragra, A. Milne-Edwards, in Maillard’s l’île Reunion, Ann. F., p. 16. 
1865. Gonodactylus chiragra, Heller, Reise ‘ Novara’ Exped., Crust., p. 126. 
1866. Gonodactylus chiragra, Annesley Proc. Zool. Soc., p. 338 (habits). 


156 


1869. 
1874. 
1875. 
1879. 
1879. 
1880. 
1880. 
1880. 
1880. 
1881. 
1882. 
1884. 
1886. 
1886. 


1887. 
1887. 
1888. 
1890. 
1892. 
1892. 
1803. 
1803. 
1803. 
1894. 
1804. 
1894. 
1897. 
1808. 
1808. 
1899. 


1890. 
1900. 
1901. 
1901. 
1901. 
1902. 


1902. 
1903. 
1903. 


1904. 
1905. 


1906. 
1906. 


. 


Memoirs of the Indian Museum. VO AE 


Gonodactylus chiragra, Hilgendorf, in von d. Decken’s Reise in Ost-afr., III, i, p. 103. 

Gonodactylus chiragra, Hoffmann, Rech. Faune Madagasc., V, ii, p. 36. 

Gonodactylus chiragra, Paulson, Rech. Crust. Mer Rouge, p. 127. 

Gonodactylus chiragra, Hilgendorf, Monatsber. Ak. Wiss. Berlin, p. 846. 

Gonodactylus chiragra, Miers, Phil. Trans. Roy. Soc., CL XVIII, p. 495. 

Gonodactylus falcatus, Kossmann, Zool. Ergebn. Roth. Meer., III, p. 100. 

Gonodactylus chiragra, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 118 and 459. 

Gonodactylus chiragra, Richters, in Mobius’ Meeresfauna Mauritius, p. 169. 

Gonodactylus chragra, Stossich, Boll. Soc. Adriat. Sci. Nat. Trieste, p. 218. 

Gonodactylus chiragra, Lenz and Richters, Abhandl. Senck. Ges. Frankfürt, XII, p. 428. 

Gonodactylus chiragra, Haswell, Cat. Australian Crust., p. 210. 

Gonodactylus chiragra, Miers, Voy. H.M.S. ‘ Alert,’ pp. 298 and 565. 

Gonodactylus chiragra ?, Müller, Verhandl. naturf. Ges. Basel, VIII, p. 471. 

Gonodactylus chiragra, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop. p. 56, pl. xy, 
fig. 4 (partim). 

Gonodactylus chiragra, de Man, Arch. f. Naturgesch., LIII, i, p. 573. 

Gonodactylus chiragra, Walker, Journ. Linn. Soc., XX, p. 113. 

Gonodactylus chiragra, de Man, Journ, Linn. Soc., XXII, p. 299. 

Gonodactylus chiragra, Pocock, Ann. Mag. Nat. Hist. (6); V, p. 72. 


Gonodactylus chiragra, de Man, in Weber’s Zool. Ergebn. Nied. Ost-Ind., II, p. 518. 


Gonodactylus chiragra, Thallwitz, Abhandl. kön. Mus. Dresden, no. 3, p. 54. 

Gonodactylus chiragra, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 454. 

Gonodactylus chiragra, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 474. 

Gonodactylus smithi(? var. of G. chiragra), Pocock, ibid., p. 475, pl. xx, B, fig. I. 

Gonodactylus chiragra, Ortmann, Denk. med.-wiss. Ges. Jena, VIII, p. 60. 

Gonodactylus chiragra, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 495 (partim). 

Gonodactylus chiragra, Wedenissow, Bull. Soc. Ent. Ital., XXVI, p. 417. 

Gonodactylus chiragra, Whitelegge, Mem. Australian Mus., III, p. 148. é 

Gonodactylus chiragra, de Man, Zool. Jahrb. Syst., X, p. 694, pl. xxxviit, fig. 77. 

Gonodactylus chiragra, Borradaile, Proc. Zool. Soc., p. 34, pl. v, fig. 4: pl. vi, fig. 8. 

Gonodactylus chiragra, Borradaile, in Willey’s Zool. Results, pp. 400, 401 (including vars. 
anancyrus and smith). : 

Gonodactylus chiragra, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 276. 

Gonodactylus chiragra, Nobili, Ann. Mus. Civ. Genova, (2), XX, p. 519. 

Gonodactylus chiragra, Lenz, Zool. Jahrb. Syst., XIV, p. 477. 

Gonodactylus chiragra, Nobili, Ann. Mus. Zool. Naples, I, 3, p. 16. 

Gonodactylus chiragra, Lanchester, Proc. Zool. Soc., II, p. 555. 

Gonodactylus chiragra, de Man, Abhandi. Senck. Ges. Frankfürt, XXV, p. 912 (including vars. 
affinis and confinis, pl. xxvii, fig. 66). 

Gonodactylus chiragra, Schenkel, Verhandl. naturf. Ges. Basel, XIII, p. 582. 

Gonodactylus chiragra, Nobili, Boll. Mus. Torino, XVIII, no. 447, p. 32. 

Gonodactylus chiragra, Lanchester, Faun. Geog. Maldives and Laccadives, I, p. 445 et seq., 
pl. x-xiii, figs. 1-7 and 10-12 (including vars. tumidus, acutus, snuthii, segregatus, incipiens and 
anancyrus. 

Gonodactylus chiragra, Jurich, Stomatop. deutsch. Tiefsee-Exped., p. 375, pl. xxvi, figs. 4-5a. 

Gonodactylus chiragra, Lenz. Abhandl. Senck. naturf. Ges. Frankftirt, XX VII, p. 386 (including 
var. smithi, p. 387). { 

Gonodactylus chiragra, Nobili, Ann. Sci. Nat. .Zool. (9), IV, p. 327 (including var. smithi, p. 329) 

Gonodactylus chiragra, Tattersall, Ceylon Pearl Oyster Rep., V, p. 167 (including vars. smithii 
and incipiens and G. glabrous var. affinis, p. 169). 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 157 


1907. Gonodactylus chivagra, Borradaile, Trans. Linn. Soc. Zool. (2), XII, p. 211 (including vars. 
affınis, smithi, incipiens, anancyrus, confinis and tumidus). 

1910. Gonodactylus chiragra, I,enz, in Voeltzkow’s Reise in Ost-Africa, II, p. 571 (including vars. | 
tumidus, smithüi and incipiens). 

1910. Gonodactylus chiragra, Fukuda, Annot. Zool. Japon., VII, p. I4L. 


The carapace is slightly narrower in front than behind and its greatest breadth is 
about three quarters its median length excluding the rostrum. The posterior margin 
is straight or very slightly concave and the anterior margins on either side of the rostrum 
slope forwards, so that the broadly rounded antero-lateral angles are very considerably 
in advance of the rostral base. The rostrum bears a stout median spine which may 
project beyond the middle of the eyestalks ; the lateral margins are anteriorly divergent 
and the antero-lateral angles are subacutely, or, in rarer instances, rather broadly 
rounded. ~ 

The dorsal processes of the ophthalmic somite (fig. 107) consist of two large sub- 
triangular plates, separated distally by a narrow fissure ; the anterior margin of each is 
oblique and is recedent from the middle line. The eyes in well-grown specimens are 
about one quarter the median length of the carapace and reach almost to the middle of 
the second segment of the antennular peduncle. The cornea is set very obliquely on 
the stalk. 

The mandibular palp is composed of three segments. 

The propodus of the raptorial claw is greatly dilated at its distal end in both sexes 
and its upper margin is deeply channelled for the reception of the dactylus. The outer 
edge of this channel is as a rule very finely spinulose, but not pectinate, in its proximal 
half; distally the edges are flattened and both are transversely striate. At the 
proximal end of the inner edge there is a single mobile spine. The basal portion of 
the dactylus is very strongly inflated ; the slender finger bears microscopic spinules 
on its inner margin and is flexed inwards near the apex. 

The lateral margins of the sixth and seventh thoracic somites are truncate with 
rounded anterior and posterior angles. Those of the eighth somite are produced to a 
bluntly rounded point. ‘The first five abdominal somites are quite smooth except for a 
sharp L-shaped groove running along the lateral and antero-lateral margins. The 
postero-lateral angles of the first four somites are rounded, those of the fifth bluntly 
rectangular. The posterior margin of the sixth somite is straight or slightly convex 
in dorsal view and on the upper surface of this somite there are six longitudinal 
ridges which usually terminate in spines. The latter in young specimens are not in- 
frequently large and stout, while in very old examples they may be wholly obsolete. 
The carinae of the lateral pair terminate in the postero-lateral angles. 


The telson is broader than long and in the middle of its dorsal surface there are 
three smooth longitudinal ridges. Close to the anterior margin, immediately below the 
intermediate carinae of the last abdominal somite, there are two small rounded tubercles. 
In large specimens a small lateral outgrowth is usually visible at the posterior end of the 
median ridge trending forwards on either side, and thus giving to the whole keel the form 
of an anchor. The flukes of this anchor are occasionally wholly absent both in large 


158 Memoirs of the Indian Museum. [Vor. IV, 


and small specimens and, in the latter, they are sometimes quite separate from the 
median keels and so form incomplete additional carinae between the medians and 
‘ laterals. The distal end of the median keel may be rounded, bluntly pointed or sharply 
spinous either with or without the addition of anchor-flukes. The flukes themselves, 
in cases where they are separate from the median ridge, may also terminate in spines. 

The three main keels may take the form of thin narrow crests or may be greatly 
swollen, but in practically every known instance there is a definite interspace between 
them, and between them and the keels on the marginal teeth. The submedian and in- 
termediate marginal teeth are well-developed and each is supported by a prominent 
longitudinal ridge. There are as a rule numerous spinules on the inner edges of the sub- 
medians and a small rounded lobe or denticle in conjunction with one or two spinules 
between the submedians and intermediates. At the tip of each submedian tooth there 
is a small movable spinule. The lateral margin is sharply carinate anteriorly and 
terminates in a sharp notch representing the lateral tooth. This tooth is never well- 
developed, and in some cases is perhaps wholly absent.' 

The basal segment of the uropods is provided with a short dorsal spine projecting 
over the first segment of the exopod. Inferiorly it is produced to a long broad process, 
divided by a distal emargination of less than halfits entire length into two strong teeth. 
The outer of these reaches almost or quite to the apex of the exopod and is usually a 
trifle longer than the inner. The internal margin of the outer spine is unarmed and 
does not bear a small tooth as in large specimens of G. glabrous and graphurus. The 
basal segment of the outer uropod projectsa little beyond the articulation of the ultimate 
segment,and bears, as a rule, ten or eleven movable spines—the outermost sometimes 
slightly recurved at the apex—on its external edge. The inner uropodis oval and 
elongate and, as is customary, bears a single row of setae on its margins. 

In the table on p. 159 I have attempted, with the help of the key given by Borra- 
daile (1907, p. 211), to arrange the 224 specimens that have been examined under differ- 
ent varieties, giving the sex and approximate length in mm. of each specimen. ‘The 
form K (= confinis ’) is not represented in the collection. 

Owing to the fact that the series is perfectly continuous in its variation it has 
proved well nigh impossible to catalogue the various individuals in a satisfactory 
manner, but conclusions of some interest are none the less indicated. 

The table will, I believe, be found to substantiate my statement (p. 150) that 
young examples exhibit a much greater range of variation than those of large size. 
This is well shown in the case of the specimens referred to H (=‘ afinis’), and is also 
noticeable in those catalogued as HA, AD, D, HD and HF, while some corroboration 
is afforded by a study of the data given by Lanchester. Of the three explanations 
which I have suggested the most probable in my view is that the young forms alter 
during subsequent moults and eventually reach or nearly reach the standard adult 
pattern; it is possible, however, that this is not the case with D ( =‘ smithi ’). 

In the next pass it will be seen at a gine that A i LUE ’) is far more 


ite iS ae in every ae of G be dr I eck seen, but is a in all en of the 
var. platysoma. 


S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 


1913.] 159 
aia | |S © là 
S | ‘ à : ' | s <8 2 
wel es 
S à A.=“ incipiens. 3 = S as S ë à 2 à : 2 
3 S ~ 5 & | à > = 
I = il S S | i À 7 1 SE I Sales 
S , 8 | Fi S > Q 
a i 2 een A Il 
< ñ A | & 
qc < jan} 
= | | z = | \ 
Queensland, Australia .. | 902 2 65 an 
Liu Kiu Is., Japan | | 378 | 
En LE | 371 
a = | 1983 3 94 
Borneo 378 3 89 
: | G55 | 
Mergui Archipelago and 2 50 4 2 80-92 9240| 921 | 234 
Tavoy. _ ik | IS 2 2 32-49 : 2 36) 3 9 17\ 27 
Nicobars 3 105 | 
2 | 
À 928 945 58 100-105 | 91 6 96|0 78 19367 940 |9 23 
$22) 239 58 3290-99 | «83 | 927 | 
918 933119 6 9 80-89 278 
917 29 31128109 70-79 | 60 
Andamans Sr 223) Og5 2160-00 38 
2220| 24 1950-59 
3 18 2 49 | 
| 7 BEE 279135. 38 | 
| RO Chae | À 
= | Tea HO Srl ke 3 68] d 80 | 
| 9231 42 7870-79 | 62,034 
| 53 5260-69 | 833 | | 
Coco Is. al Id 1957-58 | 
| 23 1943-49 
2 36 
IG 1929 2 
Preparis I. RES 
| 023 352 250 
9,22 2 48 
218 | 3 9 38 
Ceyl ? 16 | ? 32 
Fos | 22 28 
| | 214 
Br 975 954 272973869859 | 
Karachi 2250,54 d46 | | 
39 ea) | | Ber 
Mine 9 73 2 56 d 70) (353 2 30 
2957,58 | 254849 | 1827| 
Persian Gulf ra 3940-48 de | le aol 
| Is 2930-39 | 26:50 Dal = 
Dr | 350 
se (242 
Mauritius 2 30 
Locality unknown 2 89 2 68 
i 2 66 ne fs 
Number of specimens 12 14 140 | & Bo FF 
Number 50 mm. or more 
in length © x 124 os 2 Salze © 0 
| 


160 Memoirs of the Indian Museum. [Vor IV, 


numerous than any other form and comprises the vast majority of large specimens. 
From the fact that it is connected by numerous intermediates with practically every 
other type, it may be suggested that this is the starting-point from which the others 
have diverged; on the other hand it may be argued that A is a specialized type that has 
become more or less isolated in such localities as the Andamans, Cocos Is., and the 
Persian Gulf—the only places from which any considerable number of specimens have 
been examined. Before any definite conclusions on this point can be reached, data 
supplied by large collections from other localities are necessary. 

As regards evidence of isolation among other varieties, it will be noted that A 
seems to be entirely absent from the Ceylon Coast. Only a few specimens are available 
and most of these were obtained on the Pearl Banks where G. chiragra seems to occur 
sparingly in company with large numbers of G. glabrous. None of the specimens are 
large ; the majority belong to D (= ‘ smithi ’), while some, all very young, are referred 
to H. Further material is necessary before any decided opinion can be formed, but 
there appears to be some indication that D represents a small race that exists in this 
locality to the exclusion of the more generally abundant form A. JLanchester’s work - 
seems also to suggest that this type prevails at Male Atoll in the Maldives, but the 
number of recorded specimens is again very small. 

From the figures given it will be seen that males and females occur in equal pro- 
portions, and that there is no trace of any correlation between sex and varietal form. 

The colouring of this species in life appears to be fully as variable as the detailed 
structure of its telson. Among the limited number of fresh specimens that have been 
examined, the general tone was in some cases yellow, in others dull orange, brick red or 
olivaceous brown. Numerous other shades doubtless occur and occasional reference 
to them is found in the literature ; it may well be that colour changes due to environ- 
ment, and possibly to other factors, still remain to be discovered. Males are in most 
cases very darkly coloured on the ventral surface and in this respect differ strikingly 
from females which are invariably pale beneath. 

That G. chiragra is capable of making a loud snapping noise there can be little 
doubt, but the means by which this is effected has not, so far as I am aware, been 
determined.' It is possibly caused by snapping the fingers as in many Alpheidae, but 
it may perhaps be due merely to striking stones on other surrounding objects with its 
prehensile claws. Annesley (1866, p. 338), who succeeded in keeping a specimen alive 
for more than a month, notes that it made a loud sound by knocking its claws against 
the glass in which it was confined, but the noise was produced quite inadvertently when 
the animal was engaged in attacking an ophiuroid. 

Annesley found that his specimen fed voraciously on actinians. 


The localities from which the specimens examined were obtained’ are indicated 
in the table on p.159. The largest examples are males, 105 mm. in length. The 


1 See addendum, p. 198. 
? The majority of these specimens are from the Indian Museum collection, but others, which were 
kindly sent me for examination by Prof. Kishinouye, Mr. J. C. Moulton and Dr. J. Pearson, are included. 


TOIS.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 161 


majority were found when shore-collecting, but a few, all very small, were taken at 
depths of 20, 30 and even 41 fathoms. 

Gonodactylus chiragra has been recorded from a vast number of localities in the 
Indo-pacific region. It extends from the Gulf of Suez (Miers, Kossmann) and the Red 
Sea (Miers, Nobili) along the African Coast southwards to Natal Point, where, according 
to Krauss, it is rather rare. Kastwards of this it seems to occur in every favourable 
situation, and on the Australian coast reaches south to the Swan River in the west and 
to Port Molle in the east (Miers). It appears to be absent from the S. coast of Australia 
and from New Zealand. In Oceania it is abundant and has been found as far east 
as Tahiti (Heller). Northwards it extends as far as Japan (Fukuda); but it has not, I 
believe, been taken at the Hawaïian Is. 

Miers’ record from Panama, the only one from the Pacific coast of America, re- 
quires confirmation and Nobili (1897, p. 6) and Moreira (1901, p. I, sub G. falcatus) 
appear to have been unaware of Hansen’s separation of the Atlantic form under the 


a? 


NS 
N x NN 


rd 
WE 


SL RR, 


RES NZ AS 


al 


I 
ji 


7 
Op # 


Sy 
% 
TY 


Fig.2. 


Fic. 1. Gonodactylus chiragra var. platysoma. 

Fic. 2. Gonodactylus chiragra, typical form. 
name of G. oerstedi when they recorded specimens from Darien and the Brazilian coast. 
From Hansen’s notes (I895, p. 65, footnote) and from the further study of numerous 
specimens by Bigelow (1902, p. 152), it seems certain that every Atlantic specimen 
exhibits the characters of G. oerstedi and all references of G. chivagra from these 
localities may be placed with confidence in the synonymy of that species. 


162 Memoirs of the Indian Museum. [Worse 


G. chiragra has been recorded by Heller (1863, p. 309), and others, from the Medi- 
terranean ; but these records do not seem to be authentic. Steuer (IQII, p. 734) has 
re-examined specimens from Heller’s collection found in the Adriatic, and has shown 
that they are to be referred to Lanchester’s G. chiragra var. mutatus, a form of 
G. glabrous. This is the only record that can be accepted of the occurrence of the genus 
Gonodactylus in the Mediterranean. 


var. platysoma, Wood-Mason. 


1895. Gonodactvlus platysoma, Wood-Mason, Figs. and Desc. of nine Squillidae, p. 11, pl. iii, figs. 3-9 
(sine desc.). 
? 1903. Gonodactylus chiragra var. acutus, Lanchester, Faun. and Geog. Maldives and Laccadives, I, 


P- 447, pl. xxiii, figs. 2, 3, 3a. 
?1910. Gonodactylus chiragra var. acutus, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 571. 

The specimens to which Wood-Mason gave the name of Gonodactylus platysoma 
seem by their structure to possess better claims to varietal recognition than any of the 
forms listed on p.159. G. platysoma, as its name implies, is much more depressed 
than G. chragra and is proportionately broader. The greatest breadth of the abdomen 
is from 21°5 to 23°5 % of the total length of the animal including the rostrum, whereas 
it is always less than 20 % in typical G. chiragra. The difference does not appear great 
when expressed in figures, but on actual comparison of specimens is a very conspicuous 
feature (see text-fig., p. 161). 

The measurements of six specimens of var. platysoma are given below in con- 
junction with similar measurements of eight G. chiragra, s.s. With respectto the figures 
of the latter form it is to be noted that in normal specimens the percentage of length to 


= = 
3 
: é 5) 
© Qg Ge 
in 5 one 
ae d o 
LOCALITY. Sex. no a5 ae 
| lee pd ‘3 = 
us ga “3 
| 32 | aes a D 
2.5 | oo 0.0 
© | 4 H 
a | © A 
G. chiragra var. platysoma. mm. mm. % 
Andamans 3 80 175 22 
Do. Q 79 18°5 23'5 
Do. 3 77 16°5 21°5 
Society Is. 9 75 160 | 2I'5 
Mauritius 9 70 5 | 21°5 
Do. au 2 48°5 II’O 22°5 
G. chiragra, s.s. | | 
Andamans od 100 19°0 IQ 
Do. ae HIN ic! 92 17'0 | 19°5 
Do. 88 alone | 82 15°0 185 
Do. Q 82 15'5 19 
Do. 3 78 15/0 19 
Do. Q 72 13:0 19'5 
Coco Is. 2 71 14'0 19'5 
Do. 9 57 II‘0 19°5 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 163 


breadth is usually not more than 18:5 ; the majority of the specimens were specially 
selected because they appeared broader than was customary. 


Along with this character is correlated a difference in the dactylus of the raptorial 
claw. In G. chiragra the slender terminal portion is long and its apex is always very 
definitely inturned. In the var. platysoma, however, the finger is much shorter and is 
not, or is only very slightly, bent at the tip. 

In the form of the telson all the specimens of var. platysoma correspond very closely 
with the individuals to which Lanchester gave the name of acutus (1903, pl. xxiii, fig. 3). 
The keels on the dorsal surface are narrow and the median one is not anchor-shaped and 
does not terminate in a spine ; the lateral marginal teeth are entirely absent. Lan- 
chester’s examples are perhaps identical with those named by Wood-Mason ; but the 
point cannot be definitely settled, for the former author does not mention the other 
characters that separate Platysoma from typical chiragra. It is rather remarkable 
that, among normal G. chiragra in the Indian Museum, no specimen occurs with this 
type of telson, whereas all the examples of the var. platysoma from three widely distant 
localities, agree in possessing it. It seems possible, therefore, that this feature is 
correlated with the other characters of the variety; but the variation in the telson is 
so great among typical G. chiragra that it would be astonishing if the form proved 
constant in a large series of specimens. 

The six examples retain but little trace of their original colour, but in four that are 
better preserved than the rest there is a pair of conspicuous round black spots, placed 
close together in the middle of the first abdominal somite. These spots seem only to 
occur very rarely in typical G. chiragra and perhaps constitute a special feature of the 
variety. 

The specimens are registered as follows :— 


sen Port Blair, Andamans. G. H. Booley. 2312,77—80 mm. TYPES. 
spss Society Is. Otago Museum. 19,75 mm. 
“032 Mauritius. Purchased. 22, 48°5 and 70 mm. 


The examples which Lanchester recorded as G. chiragra var. acutus were taken at 
Minikoi, and those which Lenz attributed to the same form were found at Zanzibar and 
Madagascar. 

2. Gonodactylus acutirostris, de Man. 
1898. Gonodactylus chiragra var. acutirostris, de Man, Zool. Jahrb. Syst., X, p. 695, pl. xxxviii, 
figs. 775, c. 
1899. Gonodactylus chiragra var. acutirostris, Borradaile, in Willey’s Zool. Results, p. 400. 
1903. Gonodactylus chiragra var. acutirostris, Lanchester, Faun. and Geog. Maldives and Laccadives, 
I, p. 454. 
1907. Gonodactylus acutirostris, Borradaile, Trans. Linn. Soc. Zool. (2), XII, p. 210. 

This species, known from a single specimen only, may perhaps prove to be merely 
an abnormal example of G. chiragra; it is distinguished by the following features :— 

1. The antero-lateral angles of the rostrum are acute: distinctly sharper than in 

G. chragra. 
2. Ihe customary pair of tubercles near the anterior margin of the telson are 


164 Memoirs of the Indian Museum. [Vor. IV, 


wholly absent, and the submedian ridges in the middle of thedorsal surface are 
strongly convergent posteriorly. 


The only known specimen, a female 56 mm. in length, is recorded by de Man from 
the Mergui Archipelago. Patience has examined an example from the same locality, 
which in the form of the rostrum resembles the type of this species, while the telson 
shows the characters of chiragra (form acutus). 


3. Gonodactylus demani,' Henderson. 
Plate IX, figs. 108-111. 


1887. Gonodactylus, n. sp., de Man, Arch. f. Naturgesch., LIII, 1, p. 574, pl. xxii a, fig. 7. 

1893.” Gonodactylus Demanüi, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 455, pl. xl, figs. 23, 24. 

1905. Gonodactylus Demani, Nobili, Boll. Mus. Torino, XX, no. 506, p. II. 

1905. Gonodactylus spinosus, Lenz, Abhandl. Senck. naturf. Ges. Frankfürt, XXVII, p. 387, pl. xIvii, 
fig. 12. 

1906. Gonodactylus De Mani, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 330. 

1906. Gonodactylus de Mani, Nobili, Bull. Sci. France et Belg., XL, p. 158. 

1907. Gonodactylus demani, Borradaile, Trans. Linn. Soc. Zool. (2), XII, pp. 210, 212. 

1910. Gonodactylus De Mani, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 572. 

This species is very closely allied to G. chiragra and appears to be distinguished 
from it only by the following characters :— 

1. The dorsal processes of the ophthalmic somite are (fig. 108) extremely small and 
inconspicuous and take the form of two very small transverse plates with 
straight or almost straight anterior edges. 

2. The telson (figs. 109-111) has three swollen ridges in the middle of its dorsal 
surface as in G. chiragra, but the median one is very strongly convex in 
lateral view and its depth, measured from the summit to a point vertically 
below on the inferior surface is equal to half its extreme breadth. The three 
median ridges and those running to the apices of the submedian and inter- 
mediate marginal spines occupy practically the whole surface of the telson 
and are separated from one another merely by narrow V-shaped furrows; the 
interspaces found in G. chivagra are absent. This feature is well shown on 
the submedian spines ; the latter are evenly convex from side to side from 
the apex to the base and never bear a median keel distinct from the general 
surface as in G. chiragra. "Ihe telson, in all known specimens except one (var. 
espinosus) is beset with tubercles or spinules that vary greatly in size and 
number. 

In the typical form of G. demani the telson is broader than long, the intermediate 
marginal teeth are well-developed and the lateral teeth, though small, are usually 
quite distinct. The figures on pl. IX (figs. 109-111) will give some idea of the range 
of variation in the number and size of the spinules. In most examples they are absent 


! See addendum, p. 198. 
* Henderson’s G. demani has one month’s priority over G spinosus, Bigelow; the latter is here re- 
garded as a variety of the former. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 165 


from the summits of the median telson ridges, but Nobili has noted that they may 
cover the entire surface as is usually the case with the var. spinosus. It will be noticed 
that the median ridge of the telson is sometimes very much broader than the others, 
and this is the case with the type specimen figured by Henderson. 

Spirit specimens do not show any very characteristic colouring. Most of the 
examples are yellowish brown with a speckling of black chromatophores, which tend to 
form a transverse band in the posterior third of the carapace, lateral patches on the 
sixth thoracic somite, and a pair of ill-defined spots in the middle of the sixth thoracic 
and first abdominal somites. 


G. demani is a small species and does not seem to exceed 40 mm. in length. The 
specimens in the Indian Museum are not numerous:— 


oe Bombay. F. W. Townsend. Id, 28 mm. 

Se Persian Gulf. F. W. Townsend. 234 9 , 19-37 mm. 
ae Henjam Is., Persian Gulf. W. T. Blanford. 19,28 mm. 

Aa Karachi. Karachi Museum. 23,27, 30mm. 

10 

ae ‘ Arabian Sea.’ Karachi Museum. 19,20 mm. 

un Ibo Archipelago, Portuguese E. Africa.! J. F. Simpson. 1929, 27-39 mm. 


An additional specimen lent by the Bombay Natural History Society has also been 
examined :— 


Persian Gulf, Pearl Banks, 5-7 fms. IQ ,22 mm. 


Gonodactylus demani has been recorded from Pulo Edam on the Java Coast (de 
Man), from Rameswaram in the G. of Manaar (Henderson) from the Persian Gulf and 
Red Sea (Nobili) and from Zanzibar (Lenz, Borradaile). 


var. spinosus, Bigelow. 
Pate IDSC, 11, 102. 


1893. Gonodactylus spinosus, Bigelow, John Hopkins Univ. Circ., no. 106, p. IOI. 

1894. Gonodactylus spinosus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 493. 

1903. Gonodactylus chiragra var. spinosus, Lanchester, Faun. and Geog. Maldives and Laccadives, 
Vag [Oh FR 4 Dci te 307. 

1906. Gonodactylus spinosus, Nobili, Ann. Sci. Nat. Zool. (9). IV, p. 330. 

1907. Gonodactylus spinosus, Borradaile, Trans. Linn. Soc. Zool. (2), XII, pp. 210, 212. 

The telson is as long as broad, and the incision between the submedian marginal 
spines is slightly deeper than in typical G. demam. ‘The intermediate teeth of the telson 
edge are very small, and the laterals are minute or wholly absent. The dorsal surface 
is closely covered with small spinules, but in some cases the summit of the median ridge 
is bare. 

The claims of this form to varietal recognition are very slight and appear to rest 
almost solely on the greater length of the telson, and thereduced size of theintermediate 


! IT am indebted to Mr. Patience for the opportunity of examining these specimens. 


166 Memoirs of the Indian Museum. [Vor kv. 


marginal teeth. In other respects its characters are closely paralleled by certain ex- 
treme forms of typical G. demani. In some of the examples in the Indian Museum 
the lateral and intermediate teeth of the telson are slightly better developed than is 
indicated in Lanchester’s figure (see fig. 112). 

Nobili (1906 (a), p. 330), when recording comparatively large specimens that appear 
to a great extent intermediate between G. demani and var. spinosus , considers it probable 
that the lateral and intermediate telson teeth develop with age. 

The colouring of two of the Indian specimens, examined in weak formalin shortly 
after capture, was peculiar. The general tone was straw or lemon yellow marbled with 
a paler shade, and a few isolated jet-black chromatophores were placed in transverse 
rows on the posterior third of the carapace and on most of the abdominal somites, 
each chromatophore being set in the middle of a pale spot. The yellow colouring 
ceased abruptly at a distinct olivaceous brown line on the last abdominal somite, and 
behind this there was a broad band of pure white involving the ends of all the carinae. 
The telson was mottled with white, olivaceous brown and yellow, and the antennal and 
antennular peduncles were faintly banded with dull red. 

Mr. Patience regards G. spinosus merely as a synonym of G. demani, but I am not 
convinced that this is so. The examples of the form sfinosus which I have been able 
to examine were all found on the Ceylon Pearl Banks, on which, so far as I am aware, 
no specimen of the typical G. demanı has yet been obtained. 


The largest specimen known is 33 mm. in length (Nobili). Fight specimens have 
been examined :— 


7497-8 
10 
8038 
10 


N. Cheval Paar, Ceylon. T. Southwell. 23,16, 16°5 mm. 
Pearl Banks, Ceylon (from Spongodes). T. Southwell. 338,32, 18-22 mm. 


G. demani var. spinosus has been recorded from the Maldives (Lanchester), the 
Red Sea (Nobili) and from Mauritius (Bigelow). 


var. espinosus, Borradaile. 
1898. Gonodactylus espinosus, Borradaile, Proc. Zool. Sos Sip. Olle Wy Ite SE De 


1903. Gonodactylus chiragra var. espinosus, Tanchester, Faun. and Geog. Maldives and Lacca- 
dives, I, p. 455. 


The telson is as broad as, or perhaps a trifle broader than long, the intermediate 
and marginal teeth are obsolete, and on the dorsal surface the customary spinules are 
entirely absent. 

This form seems to be nothing more than a variety of G. demani, and it may ulti- 
mately appear that it is merely an extreme form in a series exhibiting continuous 
variation and, as such, does not deserve recognition under a separate name. As far as 
I am aware it differs from G. demani only in the total absence of the spinules on the 
telson, and in the reduction of the intermediate and lateral teeth of the margin and, 
both in the size of these teeth and in the number of dorsal spinules, G. demani is 
extremely variable. From the var. spinosus it is distinguished by the absence of 
spinules and by the slightly broader form of the telson. 


1913.] 


S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 167 


Borradaile has recorded a single male, olive-green in colour and a trifle more than 
IS mm. in length, from Rotuma, a locality far distant from the regions in which typical 
G. demani or var. spinosus have been found. 


1886. 


1887. 
1887. 
1893. 
1804. 
1804. 
1808. 
1890. 
Igol. 
IOOI. 
1902. 


1903. 


1904. 
1905. 


1905. 
1906. 
1906. 
1906. 


1907. 


1910. 
I9I0. 
IQII. 


4. Gonodactylus glabrous,' Brooks. 
Plate IX, fig. 113: 
Gonodactylus glabrous, Brooks, Voy. of H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 62, pl. xiv, 
fig. 5 ; pl. xv, figs. 7, 9. 
Gonodactylus graphurus, F. Müller, Verhandl. naturf. Ges. Basel, VIII, p. 471. 
Gonodactylus graphurus, de Man, Arch. f. Naturgesch., LIII, i, p. 573. 
Gonodactylus glaber, Henderson, Trans. Linn. Soc. Zool. (2), V, p. 454. 
Gonodactylus glabrous, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 493. 


-Gonodactylus glaber, Ortmann, Denk. med.-wiss. Ges. Jena, VII, p. 60. 


Gonodactylus glabrous, Borradaile, Proc. Zool. Soc., p. 36. 

Gonodactylus glabrous, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 276. 

Gonodactylus glaber, Nobili, Ann. Mus. Zool. Napoli, I, no. 3, p. 17. 

Gonodactylus graphurus, Nobili, ibid., p. 16 (fide Nobili, 1906). 

Gonodactylus glabrous, de Man, Abhandl. Senck. ges. Frankfürt, XXV, p. 913, pl. xxvii, 
fig. 67 (including var. ternatensis, p. 914). 

Gonodactylus chivagra vats. glabrous and mutatus, Lanchester, Faun. and Geog. Maldives 
and Laccadives, I, pp. 448, 451, pl. xxiii, figs. 8, 9, 15. 

Gonodactylus glabrous, Jurich, Stomatop. deutsch. Tiefsee-Exped., p. 370. 

Gonodactylus graphurus, Lenz, Abhandl. Senck. naturf. Ges. Frankfürt, XXVII, p. 387 
(partum, fide Lenz, 1910). 

Gonodactylus glaber, Nobili, Boll. Mus. Torino, XX, no. 506, p. 11. 

Gonodactylus glabrous, Tattersall, Ceylon Pearl Oyster Rep., V, p. 167. 

Gonodactylus graphurus, Nobili, Bull. Sci. France et Belg., XL, p. 159. 

Gonodactylus glaber, Nobili, Ann. Sci. Nat. Zool. (9), iv, p. 333. 

Gonodactylus glaber, Borradaile, Trans. Linn. Soc. Zool. (2), XII, pp. 211, 212 (including 
vars. ternatensis and rotundus). 

Gonodactylus glaber, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 572. 

Gonodactylus glabrous, Fukuda, Annot. Zool. Japon., VII, p. 141. 

Gonodactylus chiragra var. mutatus, Steuer, Sitz.-ber. math -naturwiss. Klasse d. k. Akad. 
Wiss., Wien, CXX, p. 734, text.-fig. 


Gonodactylus glabrous differs from G. chiragra only in the following characters :— 

1. The dorsal processes of the ophthalmic somite (fig. 113) consist of two small 
plates, rounded anteriorly ; they are longer than in G. demani and much nar- 
rower than in G. chiragra. 

2. At the base of the upper margin of the raptorial propodus the small movable 
spine, almost invariably present in G. chivagra, is entirely absent. 

3. On the sides of the 2nd, 3rd, 4th and 5th abdominal somites near the anterior 
margin there is a rather conspicuous pit. This in G. chivagra is very faint or 
wholly absent. 


4. There are five long keels in the middle of the dorsal surface of the telson and 


' The inclusion of Lenz’s Protosquilla glabra in the genus Gonodactylus renders any change in the 


original spelling of this name inconvenient. For further notes on this species see addendum, p. 197. 


168 Memoirs of the Indian Museum. [Vor. IV, 


immediately behind the median there is a pair of small tubercles. The lateral 
marginal teeth are frequently rather more distinct than in G. chiragra. 

5. On the inner side of the external spine of the basal process of the uropods there 
is a small tooth or lobe placed quite near the base. 


The keels on the sixth somite like those of G. chiragra may be swollen or narrow, 
and may or may not terminate in spines. Between the submedians an additional 
narrow longitudinal keel is occasionally found. Except for the usual L-shaped ridge 
along the lateral and antero-lateral margins and for the small pit mentioned above, the 
first five abdominal somites are quite smooth. The five keels in the middle of the 
dorsal surface of the telson may be swollen without distinct interspaces, or narrow and 
well separated ; the three median frequently end in spines. The central keel never, 
or very rarely, assumes the shape of an anchor, but is sometimes subdivided into 
three very short branches at its distal end (see Lanchester, pl. xxiii, fig. 8). In respect 
of the above modifications the species unquestionably exhibits continuous variation, 
but among the Ceylon specimens, which form the vast majority of those examined, a 
spineless form with moderately or greatly swollen keels seems to predominate. 

The Ceylon examples were preserved in formalin, and when examined, not long 
after capture, showed a most remarkable range of colour variation. The commonest 
shade was a uniform olivaceous yellow, while other examples were dull purple, deep 
purplish green profusely mottled, dark olive, olive brown with paler marbling, brick 
red or dull orange. On the last abdominal segment a pair of very small black spots 
were constantly found close to the anterior margin, situated one on either side between 
the submedian and intermediate keels. The keels on this somite and on the telson were 
in most cases defined by a darker tone. 


The specimens of G. glabrous in the Indian Museum are registered as follows :— 


5 Mergui Archipelago. ‘ Investigator.’ Id, 2I mm. 

= Nicobars: F. Stoliczka. 19,35 mm. 

ue Port Blair, Andamans. F. A. de Roepstorff. 19,08 mm. 

= Andamans. “ Investigator.’ Id, 23 mm. 

%% Galle, Ceylon. J. Anderson. 22, 34, 38 mm. 

= Cheval Paar, Ceylon. T. Southwell. 228,569 , 14:5-01 mm. 
a Pearl Banks, Ceylon (from Spongodes). T. Southwell. 19,39, 41-64 mm. 
“© Bombay. Bombay Nat. Hist. Soc. 14,33 mm. 

525 Tho Archipelago, Portuguese E. Africa.! J. J. Simpson. Id',22,50-68 mm. 


In addition the following specimens have been lent me for examination :— 


Ceylon. Colombo Museum. Tess T 9533,85 mm. 
Bombay. Bombay Nat. Hist. Soc. 2,33, 40 mm. 


Gonodactylus glabrous appears to be a common species, but perhaps frequents 


1 For these examples I am indebted to Mr. Patience. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 169 


ground of a different nature to that in which G. chivagra flourishes. Although both 
species occur together on the Ceylon pearl banks, G. glabrous is vastly in the majority. 

G. glabrous is distributed throughout the Indo-pacific region and has been recorded 
from the following localities : Rotuma (Borradaile) ; Liu Kiu Is., Japan (Fukuda) ; 
British New Guinea (Nobili); Sir C. Hardy’s I., N.E. Australia (Henderson) ; Ternate 
(de Man); Samboanga (Brooks); Sulu Sea (Henderson) ; Pulo Edam {de Man) ; 
Ceylon, Tuticorin and Rameswaram (Henderson, Tattersall) ; Maldives and Laccadives 
(Lanchester) ; Persian Gulf (Nobili) ; Aden (Henderson) ; Red Sea (Nobili) ; Seychelles 
(Jurich, Borradaile) ; Cargados Carajos (Borradaile) ; British East Africa (Borradaile) ; 
Zanzibat (Lenz, etc) ; Dar-es-Salaam (Ortmann). Müller’s record of G. graphurus from 
Trincomali doubtless refers to this species. 

Steuer (IOII, p. 734) has examined specimens from Heller’s collection which were 
obtained at Lesina in the Adriatic. This is the only authentic record of any species 
of Gonodactylus in the Mediterranean. 


5. Gonodactylus graphurus, Miers. 

1847. Gonodactylus graphurus, White, List Crust. Brit. Mus., p. 85 (sine desc.). 

1875. Gonodactylus graphurus, Miers, Ann. Mag. Nat. Hist. (4), XVI, p. 344. 

1880. Gonodactylus graphurus, Miers, Ann. Mag. Nat. Hist. (5), V, pp. 120, 459, pl. iii, fig. 9 
(partim). 

1882. Gonodactylus graphurus, Haswell, Cat. Australian Crust., p. 211. 

1884. Gonodactylus graphurus, Miers, Voy. H.M.S. ‘ Alert,’ pp. 298, 566 (partim). 

1886. Gonodactylus graphurus, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 58, pl. xiv, 
figs. I, 4,6: pl. xv, figs. 3, 8. 

1893. Gonodactylus graphurus, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 474. 

1894. Gonodactylus graphurus, Zehntner, Rev. Suisse Zool., II, p. 213. 

1894. Gonodactylus graphurus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 493. 

1894. Gonodactylus graphurus, Ortmann, Denk. med.-wiss. Ges. Jena, VIII, p. 60. 

1901. Gonodactylus graphurus, Lenz, Zool. Jahrb. Syst., XIV, p. 478. 

1903. Gonodactylus chiragra var. graphurus, Lanchester, Faun. and Geog. Maldives and Laccadives, 
I, p. 450. 

1905. Gonodactylus graphurus, Nobili, Boll. Mus. Torino, XX, no. 506, p. II. 

1905. Gonodactylus graphurus, Lenz, Abhandl. Senck. naturf. Ges. Frankfürt, XXVII, p. 387 
(partim, fide I,enz, 1910). h 

1906. Gonodactylus graphurus, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 333. 

1910. Gonodactylus graphurus, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 571. 


Gonodactylus graphurus is closely allied to G. glabrous and may be separated from 
G. chiragra by the same characters that have been given under that species. It may 
be distinguished at a glance from G. glabrous (and from all the other species of the 
same section as well) by the fine transverse and longitudinal grooves on the first five 
abdominal somites. 

The most conspicuous of these grooves starts from the small pit mentioned in the 
case of the preceding species and, after running backwards a short distance, curves 
upwards, and is continued almost to the mid-dorsal line as a transverse groove in the 
posterior third of each somite. The L-shaped antero-lateral groove, found in 
all the species of this group, is distinct and there is also on each somite an addi- 


OU) au) 


170 Memoirs of the Indian Museum. [Vora vs 


tional groove, running almost longitudinally and situated midway between the lateral 
margin, and the commencement of the transverse groove. 


The presence of these grooves (see text-fig. 1), though the only constant charac- 
ter by which G. graphurus may be distinguished from its ally, is, I believe, a feature 
of sufficient importance to justify specific separation ; intermediate specimens appear 
to be wholly unknown. Pocock (1893, p. 475) notes that in a semi-larval form, 10 
mm. in length, the grooves are found only on the fifth abdominal somite, but that 
in all the other numerous examples which he examined, they were well-marked without 
sign of failing. 


Fig, 1: 


Frg 2. 


Fic. I. Gonodactylus graphurus 


5 ee 
Fics an ler First five abdominal somites viewed laterally 


A fine median carina is usually—perhaps invariably—found on the sixth abdo- 
minal somite, but this character is shared by some examples of G. glabrous. 

The other keels on the last abdominal somite and those on the telson appear to 
vary in precisely the same way as in the preceding species. In the two examples from 
Port Molle they are slender, but moderately swollen in the third specimen ; in all three 
the median and submedian keels of the telson terminate in spines. 


The colouring of living G. graphurus is doubtless as variable as that of its allies. 


There are only three specimens in the Indian Museum :— 
7 Australia. Queensland Museum. Id , 60 mm. 


ee Port Molle, Queensland. Australian Museum. 1,19 ,35, 40 mm. 


Both in Henderson’s experience and in my own, G. glabrous occurs on the coasts of 
India to the complete exclusion of G. graphurus, and it would naturally be expected 
that, in the case of the latter species, further indications of a geographical range more 
limited than that of G. glabrous would be found in the published records. It is not 
altogether clear, however, that this is so. The head-quarters of G. graphurus appear 
to lie in an area comprising Oceania, the N. coast of Australia, Amboina and the China 
Seas, but from certain isolated records from the E. African coast, it would seem that the 
species is in reality distributed over the whole Indo-pacific region. 

As regards the African records it will be noticed that Nobili, writing towards the 
end of 1906, includes in his synonymy of G. glabrous, his record of G. graphurus from 


1013.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. A 


Zanzibar made in 1901, and remarks that the specimens collected by MM. Bonnier and 
Ferez in the Persian Gulf—evidently those that he recorded as G. graphurus in April, 
1906—are also examples of G. glabrous. It is therefore apparent that little confidence 
can be placed in Nobili’s other record from Zanzibar made in 1905. 

But, in addition to Nobili, Lenz on two occasions notes the occurrence of G. gra- 
phurus at Zanzibar. In his latest paper (1910), he refers some of the specimens recorded 
in his previous account to G. glabrous, but it seems hardly possible that there can be 
any mistake regarding the remainder, for the author notes the presence of grooves on 
the abdominal somites. 

The type specimens of G. graphurus were obtained at Samoa (Miers) and since 
1886, when the features of G. glabrous were first recognized, the following records, 
believed to be authentic, have been made: Samoa (Lenz) ; Thursday I. (Ortmann) ; 
Baleine Bank, N.W. Australia, 20 fms. ; Baudin I., 8-15 fms. ; Arafura Sea ; Holo- 
thuria Bank, China Seas, 15-24 fms. (Pocock) ; Amboina (Zehntner) ; Zanzibar ! 
(Lenz). 

The confusion between this and the preceding species hasin a large measure arisen 
from the use of characters derived from the carination of the last somite and telson, 
while the more valuable feature of the presence or absence of grooves on the first five 
segments of the abdomen seems to some extent to have been overlooked. 


6. Gonodactylus herdmani, Tattersall. 
JA ana IC, ee ol or 
1906. Gonodactylus herdmani, Tattersall, Ceylon Pearl Oyster Rep., V, p. 169, pl. i, figs. 8-10. 
1907. Gonodactylus herdmani, Borradaile, Trans. Linn. Soc. Zool. (2), XII, p. 210. 

The carapace is about one and a third times as long as broad and is not narrower 
in front than behind. The anterior margins on either side of the rostrum slope for- 
wards, and the rounded antero-lateral angles are well in advance of the rostral base. 
The rostrum is sharply trispinous ; the median spine may extend beyond the middle 
of the eyestalks or may reach only to their base. 

The dorsal processes of the ophthalmic somite consist of a pair of small transverse 
plates with broadly-rounded anterior edges. The cornea is not wider than the stalk 
and is set very obliquely on it. 

The mandibular palp is three-segmented, but the division between the two proxi- 
mal segments is rather indistinct. 

The form of the raptorial claw is almost exactly the same as that of G. chivagra; 
there is a single large movable spine on the dorsal surface of the propodus close to the 
proximal end. 

The lateral margins of the eighth thoracic somite are rounded ; those of the sixth 
and seventh somites are more or less truncate with rounded anterior and posterior 
angles. 

The first five abdominal somites, except for the customary L-shaped carina along 
the antero-lateral margins, are entirely smooth, though in large specimens one or two 
minute pits may be discerned in the middle of the dorsal surface. The postero-lateral 


172 Memotrs of the Indian Museum. orme 


angles are not spinous. The posterior margin of the sixth somite is straight or slightly 
convex in dorsal view, and on its dorsal surface there are six rounded longitudinal 
ridges, the outermost of which are coterminous with the acute postero-lateral angles. 
The four median ridges are distally rounded in the two type specimens and in two 
which I have examined ; but in another example—a small one—each of the subme- 
dians terminates in a minute spinule. 

The telson is much broader than long and bears on either side near the anterior 
margin, the small rounded tubercle which occurs in most of the preceding species. In 
the middle of the dorsal surface are three ridges that are smooth anteriorly, but poste- 
riorly tend to break up into a few large tubercles. ‘The median ridge is much wider 
than the other two and, in the specimens examined, bears three tubercles at its distal 
end and sometimes one other, much less distinct, on either side near the middle. Each 
submedian ridge is composed of an elongate anterior portion, and a large rounded 
posterior tubercle, the latter being sometimes entirely separate from the former.' The 
tubercles in the type specimen are rather more numerous, as will be seen by comparing 
pl. X, fig. 114 with the figure given by Tattersall. On the margin of the telson 
the submedian and intermediate teeth are large and well-developed ; the former have 
movable tips but bear no spinules on their inner margins ; the intermediate denticle is 
obsolete. Hach tooth is strengthened by a dorsal carina broken up into a small number 
of large tubercles. The lateral marginal teeth take the form of lobes ; they are com- 
paratively prominent in the type, but in the specimens examined are very broadly 
rounded. 

The peduncular segment of the uropod is carinate dorsally and terminates in a 
stout spine projecting over a similar carina on the proximal segment of the exopod ; 
the ventral process is composed of two spines of equal length, the outer a little broader 
than the inner. The first segment of the exopod projects far beyond the articulation 
of the second segment, and on its outer margin bears an even series of ten or eleven 
spines. The outermost spines are slightly recurved : more so than is ever the case with 
G. chiragra. ‘Theterminal segment is comparatively long; it is strongly carinate above 
and bears setae only in the distal half of its outer margin. The endopod is a com- 
paratively solid structure, not laminar as in the preceding species. It is more or less 
crescentic in shape with a concave inner margin and ends anteriorly in a subacute 
angle well in advance of its point of articulation with the peduncular segment. It 
bears a strong carina near its external edge and inwards of this carina is feebly ridged 
longitudinally both above and below. Setae are present only on the outer half of its 
distal margin. 


Living specimens are quite pale in colour with dull yellow marbling, darkest at the 
antero-lateral corners of the abdominal somites and tending to a more reddish tone on 
the sixth somite and telson. The ridges and tubercles of these last two segments are 
pure white. The propodus of the raptorial claw has a red-brown patch at the extreme 
distal end, and near the apex of each of the rostral spines, there is a transverse red band. 


! In an abnormal specimen from the Colombo Museum the left submedian ridge is absent; but its 
distal tubercle persists. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 173 


The black spots mentioned by Tattersall are distinct, but are decidedly irregular in 
size and distribution. 
The larger of the type specimens measured 28 mm. ; the three examples which 
I have been able to examine are rather smaller. 
en N. Cheval Paar, Ceylon. T. Southwell. 2B Gg 13, AS sia, 
Ceylon Pearl Banks. Colombo Museum. I 9,22 mm. 


The types and only other known examples are recorded by Tattersall from coral 
reefs in the G. of Manaar. ; Ä 


7. Gonodactylus drepanophorus, de Man. 
1902. Gonodactylus drepanophorus, de Man, Abhandl. Senck. Ges. Frankfürt, XXV, p. 914, 
pl. xxvii, fig 68. 
1907. Gonodactylus drepanophorus, Borradaile, Trans. Linn. Soc., Zool. (2), XII, p. 210. 
In the long description given by de Man I am only able to detect the following 
differences between this species and G. herdmani :— 
I. The carinae of the last abdominal somite are rather narrower and all of them 
terminate posteriorly in spines. 
2. The submedian and intermediate teeth of the telson are rather longer and more 
slender, and the inner edges of the former are beset with spinules. 
3. The lateral teeth of the telson are large, well-formed and sharply acute. 
4. The tubercles on the dorsal surface are rather more numerous and each, in place 
of being rounded, is sharply pointed. 
5. The inner uropod has the same form, but is much more strongly bent inwards 
cieulerapex 


The type specimen was speckled with black chromatophores comparable with 
those of G. herdmanı. 

A single specimen, a female 18 mm. in length, is recorded by de Man from Ternate, 
one of the Molucca Islands. No other examples are known. 


8. Gonodactylus spinoso-carinatus, Fukuda. 


1910. Gonodactylus spinoso-carinatus, Fukuda, Annot. Zool. Japon., VII, p. 143, pl. iv, figs. 2, 2a. 


The telson, which has formed the subject of a longand careful description by Fukuda, 
readily distinguishes G. spinoso-carinatus from all other species of Gonodactylus. In 
brief, the whole surface except for the base of the submedian spines is completely covered 
by nine longitudinal keels. These keels are smooth dorsally, with minute prickles on 
the lateral borders, and a few spinules directed obliquely backwards at their distal ends. 
The median keel is very broad anteriorly. The submedian teeth are large and are 
separated by a deep incision with spinulose margins ; the intermediate and lateral 
teeth are exceedingly small. 

The peduncular segment of the uropods is furnished with three or four spines on 
its dorsal surface in addition to the terminal one. The proximal segment of the exopod 
bears five or six spines on its outer margin ; the first two or three are slender and 


174 Memoirs of the Indian Museum. [VoL. IV, 


straight, the others, situated at the distal end behind the articulation of the ultimate 
segment are stout and very strongly recurved. The endopod is curved outwards. 
Fukuda does not give any details of the disposition of setae on the inner uropod and on 
the last segment of the outer uropod, but from his figure it appears that they exist only 
on the distal half of the outer margins as in G. herdmani and drepanophorus. 

In other respects G. spinoso-carinatus appears to resemble the two preceding species 
closely ; but it is much narrower in form, the greatest breadth being less than one- 
seventh the total length. 

Only two examples are known, the largest being 28°5 mm. in length. They are 
recorded by Fukuda from Jogashima, Sagami Prov., Japan. 


9. Gonodactylus brevisquamatus, Paulson. 
Plate X, figs. 115, 116. 
1875. Gonodactylus brevisquamatus, Paulson, Rech. Crust. Mer Rouge, p. 127, pl. xxi, figs. 3-3g. 
1906. Gonodactylus brevisquamatus, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 331. 

The rostrum is sharply trispinous as in the three preceding species ; the antero- 
lateral angles of the carapace are acute. 

In the middle of the sixth somite there are two smooth and broad elevations, 
placed parallel to one another in the male, but posteriorly divergent in the female. 
The lateral margins are swollen, but scarcely carinate, and between them and the sub- 
median elevations there is on each side a narrow intermediate keel. ‘There are no 
spines on the posterior margin. 

The telson (figs. 115, 116) is broader than long, and at its distal end bears two pairs 
of large teeth, the submedians and intermediates. The former are separated by a 
large rounded emargination and are about twice the length of the latter; an angular 
incision, reaching to the same level as the median emargination, divides the submedians 
from the intermediates. There are no spinules on the inner edges of the submedian 
teeth. The lateral margins of the telson are evenly curved and there is no trace what- 
ever of lateral teeth. In the middle of the dorsal surface there is a large, smoothly- 
rounded, oval elevation placed between a pair of narrower and less prominent keels. 
The base of the submedian spines is somewhat swollen and a ridge extends from near 
the anterior margin to the apex of the intermediate spines. 

The peduncular segment of the uropods appears to resemble that of G. herdmani. 
The basal segment of the exopod projects far beyond the articulation of the ultimate 
segment. The proximal part of the outer margin is smooth (fide Nobili) or with a few 
small spines (fide Paulson’s fig.), while at the distal end there are two or three stout 
spines which are very strongly recurved. The ultimate segment of the exopod is 
covered with setae over its entire surface; the endopod is similarly clothed! and is 
strongly curved outwards. 

Through the help of my friend Dr. Calman I am able to reproduce two figures of 
this species from Paulson’s very rare work. 


1 The setae on the surface of these segments are not shown in Paulson’s figure. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 175 


G. brevisquamatus is known to reach a length of 23 mm. It appears to be exceed- 
ingly scarce and has as yet only been found in the Red Sea. 


10, Gonodactylus fimbriatus, Lenz. 
1905. Gonodactylus fimbriatus, Lenz, Abhandl. Senck. naturf. Ges. Frankfürt, XXVII, p. 388, 
pl. xlvii, fig. rr. 
1907. Gonodactylus fimbriatus, Borradaile, Trans. Linn. Soc. Zool. (2), XII, p. 211. 
1910. Gonodactylus fimbriatus, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 572. 


Gonodactylus fimbriatus is very closely allied to G. brevisguamatus, and appears to be 
distinguished from it only by the following characters :— 


1. Of the three pairs of longitudinal ridges on the last abdominal somite the sub- 
medians and intermediates are about equal in width, while those of the lateral 
pair are slightly narrower. 

2. The marginal teeth of the telson are closely similar to those of G. brevisquamatus, 
but the intermediates are rather larger and there are spinules on the inner edge 
of the latter and on both edges of the submedian teeth. 

3. In the middle of the dorsal surface of the telson there is a large oval prominence 
with a narrower lenticular keel pressed close against it on either side and 
between these, and the thickened lateral margin there are two other feeble 
longitudinal ridges. At the hinder end of the median boss there are three 
small tubercles and two other tubercles, homologous perhaps with those of 
G. chiragra, are found near the anterior margin. All the median dorsal ridges 
stop before reaching the posterior third of the telson, terminating well behind 
the base of the submedian emargination. 


The uropods seem closely similar to those of G. brevisguamatus, but in the ventral 
process of the peduncular segment the inner spine is less than half the length of the 
outer, and is perhaps shorter than in that species. The basal segment of the exopod 
bears no spines on the anterior half of its external margin, but is furnished distally with 
three large recurved teeth. The endopod isflexed outwards and the margins, both of 
this segment and of the distalsegment of the exopodite, appear to be fringed with setae 
as in G. brevisquamatus; their surfaces, however, according to Lenz’s figure, are not 
clothed with hairs. 


Gonodactylus fimbriatus reaches a length of 28 mm. and is recorded from Zanzibar 
(Lenz) and from Coetivy, Seychelles (Borradaile). 

Since the above was written I have, thanks to the courtésy of the author, had an 
opportunity of seeing the manuscript of Mr. Patience’s valuable account of the Stomato- 
poda collected by Messrs J. J. Simpson and R. N. Rudmose Brown in the Mergui Archi- 
pelago and in Ibo Archipelago in Portuguese East Africa. In this paper Mr. Patience 
gives, under the name G. brevisquamatus, a detailed account and figures of a specimen 
obtained at Mergui. G. fimbriatus is cited as a synonym and strong reasons for the 
adoption of this course are adduced. Inasmuch, however, as I have not been able to 
examine any specimens of this scarce species, I have preferred to let my notes stand in 


176 Memoirs of the Indian Museum. [Vor.. IV, 


their original form referring those interested in the question to Mr. Patience’s full dis- 
cussion. 


11. Gonodactylus furcicaudatus (Miers). 
1880. Gonodactylus furcicaudatus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 124, pl. iii, figs. 13-16. 


1886. Protosguilla (?) furcicaudatus, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 78. 
1907. Gonodactylus furcicaudatus, Nobili, Mem. Acc. Sci. Torino (2), LVII, p. 410, pl. ui, fig. 6. 


The carapace is fully one and a half times as long as wide, and the anterior margins 
on either side of the rostrum are straight and trend a little forwards, so that the narrow- 
ly-rounded antero-lateral angles are in advance of the rostral base. The rostrum is 
trispinous, but the lateral spines are less slender than in G. herdmani, and the basal 
undivided part is considerably longer. 


The mandibular palp is composed of two segments. There is no movable spine at 
the base of the raptorial propodus. 


The sixth abdominal somite is somewhat similar to that of G. brevisquamatus. In 
the middle there are two large smooth prominences of no great height and nearly as 
broad as long, and on either side there isa short intermediate ridge and a smooth lateral 
swelling which is coterminous with the rectangular antero-lateral angles. 


The form of the telson is extremely peculiar and consists of a broad and very thick 
anterior portion, abruptly truncate behind and with two long furci projecting from the 
middle of the lower edge of the truncate portion. The dorsal part is nearly four times 
as broad as long, strongly convex from side to side with its ridged lateral margins pos- 
teriorly divergent. An obscure tubercle is found on either side near the anterior end, 
and three feeble and partially confluent swellings may be detected near the hinder edge. 
The latter edge is nearly straight from side to side, but obscure median, interme 
diate and lateral lobes, separated by four very shallow emarginations, are shown. 
There are numerous stout spinules on this edge, disposed in an irregular fashion. 
Behind this margin the telson, which is of very considerable thickness is abruptly 
and vertically truncate and the truncate portion, seen from behind, presents a kidney- 
shaped outline and is hollowed and honey-combed in an extremely curious manner. 
From the inferior edge of this portion, and from its middle point, two stout furci arise 
which are longer than the rest of the telson. They are closely approximate at the base, 
but near the tips they diverge outwards ; a few very fine spinules are visible on their 
anterior margins. 


The peduncular segment of the uropods terminates in a dorsal spine projecting 
over the base of the exopod and, of the two spines that form its ventral process, the inner 
is more slender than the outer and only about half its length. The ultimate segment of 
the exopod articulates with the basal segment at about the middle point of the latter. 
On the external margin of the basal segment there are five small straight spines in the 
anterior part, while at the distal end there are three stout recurved hooks. The ulti- 
mate segment of the exopod is clothed all over its dorsal surface with setae of two 
kinds. The central part, more particularly near the base, is beset with short club- 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 177 


shaped hairs ; the others, which occupy the rest of the surface, are long and slender. 
The endopod rises straight upwards from its articulation and having reached the 
upper level of the telson bends abruptly backwards in a right angle. It is a solid 
structure and is beset with setae like the last segment of the exopod. The setae on 
its upper surface form a dense brush. 


Miers states that his largest specimen was about 29} mm. in length. 


The only example in the collection is a co-type presented by the British Museum :— 
73 (Locality ?) British Museum (H.M.S. ‘ Herald’). If ,24 mm. 


10 


The locality at which the type specimens were obtained is unfortunately unknown, 
and since the species was originally described it has only once been found : Nobili 
(1907) records a single small example from Matakea in Polynesia, N.E. of Tahiti. 


ı2. Gonodactylus pulchellus, Miers. 
DlateX nes 007 TLS: 

1880. Gonodactylus trispinosus var. pulchellus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 122. 
1893. Protosquilla trispinosa, Henderson, Trans. Linn. Soc., Zool. (2), V, p. 455. 
1894. Gonodactylus trispinosus var. pulchella, Ortmann, Denk. med.-wiss. Ges. Jena, VIII, p. 61. 
1902. Protosquilla trispinosa var. pulchella, de Man, Abhandl. Senck. ges. Frankfürt, XXV, p. 920. 
1906. Gonodactylus trispinosus var. pulchellus, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 326. 
1906. Protosquilla trispinosa var. pulchella, Tattersall, Ceylon Pearl Oyster Rep., V, p. 173. 

The carapace is a little longer than broad and its posterior margin is concave. 
The anterior margins on either side of the rostrum are also concave and do not protrude 
forwards ; the subacute antero-lateral angles are consequently not in advance of the 
rostral base. The rostrum is trispinous and the sharp lateral spines are only a trifle 
stouter and shorter than the slender median one. The undivided basal part of the 
rostrum is short and wide, its length being much less than that of the median spine. 

The dorsal processes of the ophthalmic somite can be clearly seen between the rostral 
spines. They consist of a plate, straight in front except for a narrow median incision, 
with sharp antero-lateral angles that project transversely outwards almost as far as the 
external edge of the basal part of the eyestalk. The eyes themselves are cylindrical 
and in length about one-third the median length of the carapace (a trifle longer in small 
specimens) ; the cornea is scarcely wider than the stalk and is set very obliquely on it. 
The eyestalks extend to a little beyond the second segment of the antennular peduncle. 

The mandibular palp is composed of only two segments. 

The basal part of the dactylus of the raptorial claw is strongly swollen, the distal 
portion is slender and slightly curved. 

The lateral margins of the sixth thoracic somite are broadly rounded, those of the 
seventh truncate with rounded angles, those of the eighth subacute. 

The median portions of the first five abdominal somites are entirely smooth except 
that each bears four minute and very obscure pits in a transverse line near the anterior 
margin, while three others, a central and a pair of lateral, may be found in the middle 
of each somite except the first. In all these five somites there is a well-defined ridge 


178 Memoirs of the Indian Museum. [Vor. IV, 


on either side defining the lateral and antero-lateral margins, and on the second, third 
and fourth there are two broad and shallow lateral depressions between the thickened 
lateral edge and the feeble ridge that limits the smooth central portion of the somite. 
These two depressions are separated by a longitudinal elevation which is itself furrowed 
This sculpture is inconspicuous, but may be seen without much difficulty in speci- 
mens of good size, especially after the removal of all surface moisture. The fifth somite 
is strongly puckered and wrinkled laterally. The postero-lateral angles of all four 
somites are subacute, but not spinous. 

The sixth abdominal somite is almost immovably articulated with the telson and 
the middle of its posterior margin is concave in dorsal view. It bears six oval bosses 
in a transverse row and the outermost, which is the longest (more a ridge than a boss), 
trends outwards to meet the external margin in a small postero-lateral spine. In most 
perfect specimens these elevations are clothed with short setae; the interspaces be- 
tween them are coarsely and irregularly punctate. 

The general form of the telson is shown in fig. 118. The median of the three dorsal 
bosses or elevations is somewhat triangular in shape while the submedians, which ‚though 
longer, reach only to about the middle of the telson, are more or less pyriform. The 
lateral margins of the telson and the surfaces of all three elevations are usually beset 
with short setae similar to those on the last abdominal somite, elsewhere the surface 
is coarsely reticulate. The distal margin is divided by seven fissures, cut right through 
the telson, into eight lobes. These fissures ! are exceedingly narrow ; they are widest 
proximally, but distally their margins are usually in contact. The median incision 
extends almost to the base of the central elevation ; those on either side reach about 
half way to the lateral elevations. Each distal lobe of the telson comprises an outer 
subacute portion and an inner movable denticle. In addition there are numerous 
denticles on the margins of the median incision, and on the convex inner edges of the 
submedian lobes. 

The process from the base of the uropods consists of two sharp teeth: the inner 
at least half the length of the outer. The two segments forming the exopod articulate 
normally with one another; the innermost bears nine movable spines on its external 
margin. The margins of the distal segment and of the endopod are uniformly beset 
with setae. 

Miers regarded this form merely as a variety of G. trispinosus, and in this he has 
been followed by most subsequent writers ; but, although I have not been able to com- 
pare the two, there is, I believe, little doubt that G. pulchellus is entitled to rank as a 
distinct species. 


The colouring of fresh specimens is rather striking. The general tone is pale oliva- 
ceous brown due to a profuse marbling of warm olive brown on a pale ground. The 
whole surface is in addition sprinkled with very small black chromatophores. On the 
carapace there are faint indications of anterior and post-median dark transverse bands, 
while, on the post-abdomen, two pairs of dark spots on the sixth thoracic, and a dark 
median patch on the first abdominal somite are generally conspicuous. Median patches 


1 The fissures are best seen if the telson is examined from below. 


FOL) S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 179 


are also sometimes found on the last two thoracic, and on the fourth and fifth abdominal 
somites. The marbling is on the whole very uniform in all the fresh specimens ex- 
amined, but the distribution of the various patches of pigment is too complicated to 
admit of description. In some specimens the general tone of the colouring tends to a 
dull purple on the sixth somite and telson. ‘The merus of the raptorial claw may have 
a vertical crimson streak near its distal end and the anterior parts of the propodus and 
swollen base of the dactylus are always white. The ventral surface is pale throughout 
except that the posterior half of the telson is suffused with brown. 


There are in the Indian Museum specimens of Gonodactylus pulchellus, registered as 
follows :— 


5 N. Cheval Paar, Ceylon. T. Southwell. 23,22, 37-43 mm. 
37 Pearl Banks, Ceylon (from Spongodes). T. Southwell. 33, 22, 26-37 mm. 
a Ban Cie. F. H. Townsend. 19,25 mm. 
=! Portuguese E. Africa.! J. J. Simpson. 2, JE 


The species has been previously reported from Ceylon (Miers, Tattersall), Aden 
(Nobili) and Dar-es-Salaam (Ortmann). It is probable, too, that the examples which 
Henderson has recorded from Ceylon under the name of Protosquilla trispinosa are to be 
referred to this species for there is reason to believe that G. trispinosus does not extend 
as far west as India. 


13. Gonodactylus nefandus, Kemp. 
IEEE, WES, 10), 720. 
1011. Gonodactylus nefandus, Kemp, Rec. Ind. Mus., VI, p. 93. 


This species is very closely allied to the preceding, but may be separated from it 
without difficulty by the form of the rostrum, by the shape of the dorsal processes of the 
ophthalmic somite, and by the shallow incisions separating the distal lobes of thetelson. 

The distinctions between the two species are as follows :— 

1. The antero-lateral angles of the carapace are rectangular and slightly rounded, 
not subacute, and the anterior margin on either side of the rostral base is 
straight. 

2. The lateral spines of the rostrum are acute but very broad, forming a striking 
contrast with the very slender median spine; the undivided basal part is much 
longer than in G. pulchellus, its length being equal to that of the median spine. 

3. The antero-lateral angles of the plate forming the dorsal process of the ophthal- 
‘mic somite are rectangular and do not project outwards as in the preceding 
species. ‘The eyes, moreover, are noticeably stouter and their length, in full- 
grown specimens, is considerably more than one-third the median length of 
the carapace. 

4. The base of the raptorial claw is less strongly swollen and the distal part is 
rather stouter and a little less curved. 


! I am indebted to Mr. Patience for this specimen. 


180 Memoirs of the Indian Museum. - [Vor. IV, 


5. The lateral margin of the sixth thoracic somite is more sharply rounded and 
that of the seventh is less clearly truncate. 

6. The dorsal pits and lateral sculpture on the first five abdominal somites are 
wholly obsolete. 

7. The boss-like elevations on the sixth somite and telson exhibit no trace of the 
clothing of setae found in the preceding species, and the rest of the surface 
shows only the feeblest indication of pitting and reticulation. 

8. The three bosses on the telson are less elevated and are separated only by a 
shallow groove ; in consequence, they appear to be partially connected with 
one another anteriorly. The median fissure in the distal part is similar to 
that in G. pulchellus and is cut right through the telson ; but the margins 
touch one another throughout all, or almost all, their length. The four lobes 
on either side are separated by incisions that are much shallower than in the 
preceding species. On the dorsal surface they are continued forwards as 
grooves almost halfway to the base of the lateral bosses, but they are not cut 
right through the telson for this distance and are not widened proximally as 
in G. pulchellus. 

9. The inner spine of the process from basal segment of the uropods is not more 
than half the length of the outer. The inner uropod is broader in proportion 
to its length, and the outer margin of the proximal segment of the exopod 
bears from ten to twelve movable spines. 


No characteristic colouration can be found in spirit specimens. 


The forty-seven specimens in the Indian Museum were found at the following 
localities :— 


ue Andamans. J. Wood-Mason. I5d, 192, 11-39 mm. 
| TYPES. 
2% Andamans. J. Wood-Mason. 19,32 mm. 
> Andamans, 20 fathoms. “ Investigator.’ 19,16 mm. 
85% Port Blair, Andamans. G. H. Booley. 24, 19 and 27 mm. 
238 Cheduba, Arakan Coast, 7 fathoms. “ Investigator.’ 43,42, 16-40 mm. 
Southern portion of Malacca Strs. “ Sherard Osburn.’ 19,20 mm. 


14. Gonodactylus trispinosus, Dana. 


1847. Gonodactylus trispinosus, White, List Crust. Brit. Mus., p. 85 (partim, nom. nud.). 
1852. Gonodactylus trispinosus, Dana,! U.S. Explor. Exped., Crust., p. 623. 


1 In this work Dana attributes the species to White, and cites as his authority pl. V of the Voyage of 
the ‘Erebus’ and ‘ Terror,’ a work that, in its completed form, was only issued twenty-four years sub- 
sequently (1874). In this volume, moreover, the text is written by Miers ; there are only four plates, and 
there is no account or figure of G. trispinosus. It seems, however, from Miers’ introductory statement that 
the plates were issued under White’s supervision many years previously, and copies must have been in Dana’s 


possession when writing in 1852. Of the absence of plate V in the completed account, no explanation is 
forthcoming. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 181 


1865. Gonodactylus trispinosus, Heller, Reise ‘ Novara’ Exped., Crust., p. 126. 

1876. Gonodactylus trispinosus, Miers, Cat. Crust. N. Zealand, p. go. 

1880. Gonodactylus trispinosus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 121, pl. iii, fig. 10 (Partim). 
1882. Gonodactylus trispinosus, Haswell, Cat. Australian Crust., p. 211. 

1886. Gonodactylus trispinosus, Filhol, Miss., de l’île Campbell, III, 2e, p. 436. 

1886. Protosguilla trispinosa, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 71. 

1891. Protosquilla trispinosa, Chilton, Trans. N. Zealand Inst., XXIII, p. 61. 

1893. Gonodactylus trispinosus, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 476. 

1898. Protosquilla trispinosa, Borradaile, Proc. Zool. Soc., p. 33, pl. v, figs. I, Ia. 

1899. Protosquilla trispinosa, Borradaile, in Willey’s Zool. Results, p. 400. 

This species and G. tuberosus may be distinguished at a glance from the two pre- 
ceding forms by the close and conspicuous corrugations in the middle of the fifth abdo- 
minal somite. In other respects G. trispinosus is very similar to G. pulchellus, from 
which, according to Borradaile’s description and figures, it would seem to differ only in 
the following features :— 

1. The antero-lateral angles of the carapace are not subacute, scarcely even 

rectangular. 

2. The lateral margin of the last thoracic somite is broadly rounded. 

3. The bosses on the last abdominal somite and telson are clothed with setae as in 

G. pulchellus, but the lateral pair on the telson are rather more broadly oval in 
outline. The telson is also rather more rounded at the apex than in the allied 
species, and the surface behind the elevations, in place of being reticulate, is 
marked with parallel impressed lines. The fissures in the distal margin do not 
appear to be widened proximally. 


I have seen no examples of Gonodactylus trispinosus. It has been recorded from 
the Fiji Is. (Dana) ; Loyalty Is. (Borradaile) ; Auckland Is. (Heller) ; Swan R., W. 
Australia (Miers); and Baleine Bank, N.W. Australia (Pocock). Chilton considers 
Heller’s record doubtful. 

Hoffmann (1874, p. 43) includes G. trispinosus in a list of Stomatopods from Mauri- 
tius ; but Miers and Brooks both regard this record as untrustworthy and a consideration 
of the known geographical range of the species supports their view. Pocock notes, on 
Hansen’s authority, that the two specimens from Sharks Bay which Miers referred to 
this species are in reality examples of Muller’s G. stolvurus. 


15. Gonodactylus tuberosus, Pocock. 
1893. Gonodactylus tuberosus (? var. of G. trispinosus) Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 476, 
(Olly nox Brie 
This species ! seems to bear much the same relation to G. trispinosus that G. 


nefandus does to G. pulchellus. 
The rostrum is very similar to that of G. nefandus, but the lateral spines are even 


1 Pocock does not mention the median corrugations on the fifth abdominal somite, but from his re- 
marks concerning the close resemblance of this species to G. trispinosus (specimens of which were before 
him at the time) the presence of this important character may be inferred. 


182 Memoirs of the Indian Museum. YO LOY, 


stouter than in that species; the undivided basal part is as long as the median spine 
thus agreeing with nefandus and differing from trispinosus and pulchellus. 

The form of the telson is somewhat different from that of any of the preceding 
species. In shape it is more or less square, but the submedian lobes project distally 
beyond the next pair, and so break the evenness of the posterior margin. ‘The fissures 
separating the lobes are well-marked, and the outermost on each side is specially broad 
and deeply cut, widening distally. The dorsal bosses are higher than in the other 
species, more particularly the median one, and the laterals are round (not oval) in shape. 

I have seen no examples of Gonodactylus tuberosus. The two known specimens 
were obtained on the Macclesfield Bank, China Seas, 37 fathoms (Pocock). 


16, Gonodactylus glaber (Lenz). 
Plate X, fig. 121. 


1905. Protosgunlla glabra, Lenz, Abhandl. Senck. naturf. Ges. Frankfürt, p. 388, pl. xlvii, fig. 13. 
1010. Protosquilla glabra, Lenz, in Voeltzkow’s Reise in Ost-Afrika, II, p. 572. 

The carapace is fully one and a half times as long as wide, and is slightly broader 
behind than in front. The posterior margin is almost straight, and the anterior margins 
on either side of the rostral base slope slightly backwards and are feebly concave. 
The antero-lateral angles are rectangular in the small specimen, rather more acute in the 
larger individual. The rostrum is sharply trispinous. The median spine is slender 
and very long ; it reaches almost to or beyond the middle of the eyestalks, and is fully 
twice as long as the undivided basal part. The lateral spines are only about half the 
length of the median, stouter and curved forwards. 

The dorsal processes of the ophthalmic somite consist of a pair of plates which ex- 
tend outwards almost to the external edge of the proximal part of the eyestalks ; their 
anterior and postero-lateral margins are slightly convex and meet in a subacute point. 
The eyestalks are two-fifths the length of the carapace in the larger example, longer in 
the smaller individual ; they reach to the end of the second segment of the antennular 
peduncle. The cornea is not noticeably wider than the stalk and is set obliquely on it. 
The mandibular palp is composed of two segments. 

The dactylus of the raptorial claw is strongly swollen at the base and its external 
margin is notched posteriorly ; the slender terminal part is curved inwards at the apex. 

The lateral margins of the sixth and seventh thoracic somites are truncate with 
broadly rounded anterior and posterior angles. The first five abdominal somites are 
smooth dorsally , but show obscure traces of pitting similar to that noticed in G. pulchellus. 
Laterally there is a well-defined L-shaped ridge on each somite except the first, running 
parallel to the anterior and external edges. ‘The fifth somite is obscurely furrowed 
longitudinally on either side of the smooth median portion. The sixth somite bear six 
eroded tubercles on its dorsal aspect, and the outermost of these, the longest and narrow- 
est, meet the external margins in a blunt postero-lateral point. 

The telson is a trifle wider than long and appears to be immovably connected with 
the preceding segment. Of the three mid-dorsal bosses the two outer ones are much 
longer than in any of the preceding species ; they are pyriform in shape and terminate 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo- Pacific Region. 183 


in an evenly rounded apex before reaching the distal margin. The three lobes are 
connected with one another in the extreme anterior part of the telson and, in the larger 
specimen, the margins of the lateral pair are feebly corrugated. Along the lateral 
edge of the telson there is a prominent rounded ridge ; this ridge is continued to the 
outermost tooth on the distal margin, and is obliquely grooved in its posterior third. 
In the centre of the distal margin there is a deep and very narrow fissure extending 
half-way to the median boss, and on either side of this the edge is inflated and so forms 
a pair of obscure ridges. Three stout teeth, separated by triangular notches are found 
in each half of the posterior margin. There are numerous submedian denticles and one 
on the inner aspect of each of the lateral teeth. 

The inner spine of the bifurcate process of the uropod is extremely small, not more 
than one quarter the length of the large and broad outer one. There are eight movable 
spines on the external aspect of the basal segment of the outer uropod. 

There can be little doubt that the two specimens on which the above description 
is based are correctly referred to Gonodactvlus glaber. They differ, however, from Lenz’s 
account in having the carapace and median rostral spine slightly longer proportionally. 
The keels on the lateral margins of thetelson are not clearly shown in the originalfigure, 
and there is no mention’of the proximal notch on the external margin of the raptorial 
dactylus. 

Spirit specimens are pale or brownish in colour with scattered black chromatophores 
which tend to form bands or patches in the posterior part of the carapace, on the sixth 
and seventh thoracic and on the first, fourth and fifth abdominal somites. 


The two examples of Gonodactylus glaber in the Indian Museum are registered 
thus :— 


# Gt. Coco I., N. Andamans. “ Investigator. ’ 19,26 mm. 


=e Gt. Coco I., N. Andamans. “ Investigator.’ ur, DR 


Previously known from Ceylon and Zanzibar (Lenz). The largest recorded speci- 
men is 30 mm. in length. 


17. Gonodactylus tanensis (Fukuda). 
1911. Protosquilla tanensis, Fukuda, Annot. Zool. Japon., VII, p. 285, pl. xi, figs. 1, 2. 


Of this species I have seen no specimens. It is, as Fukuda has remarked, closely 
allied to G. glaber, but may readily be distinguished from it by the sculpture of the 
last abdominal somite and telson. 

In addition to the usual prominences, the last abdominal somite is ““ completely 
covered with irregularly-marked depressions.’’ The telson bears three bosses, 
situated much as in G. glaber, but with larger interspaces. These interspaces are 
covered with irregular depressions and in Fukuda’s figure look almost as if honey- 
combed. The median distal fissure and the lateral marginal lobes are nearly as in 
G. glaber and are provided with similar spinulation. 


184 Memoirs of the Indian Museum. [Vor. IV, 


The inner spine of the bifurcate process of the uropods, though very much 
shorter than the outer, appears to be noticeably larger than in the allied species. 
There are only seven movable spines on the outer margin of the proximal segment of 
the exopodite. _ 

Fukuda records a single male, 30°5 mm. in length, from Tanegashima, Osumi 
Province, Japan. 


18, Gonodactylus stoliurus (Müller). 


1880. Gonodactylus trispinosus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 121 (partum) and p. 460 (fide 
Pocock, 1893, p. 476). 

1886. Protosguilla stoliura, Müller, Verhandl. naturf. Ges. Basal, VIII, p. 477, pl. iv, fig. 2. 

1887. Protosguilla stoliura, de Man, Arch. f. Naturgesch., LIII, i, p. 576. 

1894. Protosquilla stoliura, Zehntner, Rev. Suisse Zool., II, p. 213. 

1894. Gonodactylus stoliurus, Ortmann, Denk. med.-wiss. Ges. Jena, VIII, p. 61. 

1902. Protosquilla stoliura, Schenkel, Verhandl. naturf. Ges. Basel, XIII, p. 582. 


The principal characters of this species seem to be as follows :— 

The carapace is only very slightly longer than wide, with a concave posterior 
margin and rectangular, but rounded, antero-lateral angles. The rostrum bears three 
sharp spines ; the median one reaches only to the base of the eyestalks, while those of 
the lateral pair are slightly shorter. 

The eyes are cylindrical and the cornea is not wider than the stalk. The inner 
edge of the raptorial dactylus is finely serrate, and the basal part is greatly inflated. 

The first four abdominal somites are smooth dorsally, with blunt postero-lateral 
angles. The upper surface of the fifth is wrinkled and grooved in a symmetrical 
fashion. On the sixth somite there are six dorsal prominences, those of the median 
pair are rounded, the others eroded at the margins. The surface of the sixth somite 
and hinder part of the fifth is beset with numerous very small spinules. Bs 

The telson is distinctly broader than long and bears, as usual, three prominent 
rounded elevations on its dorsal surface. The median boss is triangular in shape, while 
those of the lateral pair are pyriform, rounded posteriorly, and reach to the distal third 
of the telson. On either side, between the lateral elevations and the thickened external 
margin, there is a well-marked rounded longitudinal ridge. On the distal margin there 
are eight broad triangular teeth ; between the two submedians there is a very narrow 
fissure extending forwards almost to the base of the median boss. The teeth on either 
side are separated by short notches and bear small spinules on their edges. In addition 
the whole surface of the telson is beset with spinules similar to those found on the two 
preceding somites. 

The outer spine of the process from the base of the uropods is twice the length of 
the inner. The basal segment of the exopod bears ten or eleven spines on its external 
margin. 

By the length of the lateral elevations on the telson, and by the very narrow median 
fissure in the middle of the distal margin, this species is easily distinguished from all 
species except G. glaber ; from the latter it may be separated at a glance by the wrink- 
ling in the median part of the fifth abdominal somite, by the presence of four pairs of 


HOT] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 185 


teeth in place of three on the distal margin of the telson, and by the numerous spinules 
on the telson and last abdominal somite. 


Schenkel (1902, p. 581) gives an account of the colouration. 


I have seen no examples of Gonodactylus stoliurus. It has been recorded from 
Amboina (Muller, Ortmann, Zehntner and de Man) and from Makassar (Schenkel). 
Pocock (1893, p. 476) notes, on Hansen’s authority, that the specimens from Sharks 
Bay, W. Australia, which Miers referred to G. trispinosus, are in reality examples of 
this species, and it is clear that the individual from Amboina, which Miers subsequently 
recorded under the same name (1880 (4), p. 640) also belongs to G. stoliurus. The 
largest known specimen is a male 61 mm. in length (Schenkel). 


19. Gonodactylus ectypus (Müller). 
1880. Gonodactylus folinii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 123 (fide Hansen, 1895, p. 90). 
1886. Protosquilla ectypus, Müller, Verhandl. naturf. Ges. Basel, VIII, p. 476, pl. iv, fig. I. 

Gonodactylus ectypus, along with the succeeding species, G. glyptocercus, may be 
distinguished from all the preceding forms by the elongate shape of the two submedian 
bosses of the telson which extend throughout the whole length of the segment and ter- 
minate in a tooth on the distal margin. 

In G. ectypus the antero-lateral angles of the carapace are subacute, and the median 
of the three spines on the rostrum reaches (according to Müller’s figure) to about the 
middle of the eyes. 

The first four abdominal segments are smooth dorsally, but on the sides of the 
fifth there are three pairs of oblique furrows which are posteriorly convergent. The 
sixth somite bears six dorsal tubercles, the outermost of which are flattened and thorny 
while the other four are rounded and convex. 

The telson is a little longer than wide. ‘The median of the three dorsal bosses is 
oval and smooth and is produced to a posterior point, while those of the lateral pair 
are also smooth, and in the posterior half of the carapace are regularly narrowed to a 
sharp tooth-like apex which extends beyond the distal margin. On either side of these 
eminences there are two ridges running parallel to the external margin, both of which 
terminate distally in teeth. The outer of these is better developed than the inner. 
In the middle of the distal margin two broadly-rounded submedian lobes are visible, 
separated by an incision that seems to be rather wider than in the preceding species ; 
on either side are three teeth forming the terminations of the submedian bosses and 
of the lateral carinae already mentioned. The spinules usually found on the distal 
margin appear to be missing. 

The outer spine of the ventral process of the uropod is much longer and broader 
than the inner. 

Miiller notes that large examples are dark brown in colour, while younger individuals 
are light yellowish grey with brown speckles. 

There are no examples of Gonodactylus ectypus in the Indian Museum. Müller 
described the species from six specimens, the largest 3 cms. in length, from Trincomali 
Bay, Ceylon, and the only other recorded individual was found at Mauritius (Miers, 
sub G. folinii, fide Hansen). 


186 Memotrs of the Indian Museum. Worle 


20. Gonodactylus glyptocercus, Wood-Mason. 

1875. Gonodactylus glyptocercus, Wood-Mason, Proc. As. Soc. Bengal, p. 232; reprinted in Ann 
Mag. Nat. Hist. (4), XVII, p. 263 (1876). 

1880. Gonodactylus glyptocercus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 122. 

1886. Protosquilla cerebralis, Brooks, Voy. H.M.S. ‘Challenger,’ XVI, Stomatop., p. 72, pl. xiv, 
iS, 2, 35 06 soul, was, 2, 2 

1886. Protosquilla glyptocerca, Brooks, ibid., p. 75. 

1887. Protosguilla cerebralis, de Man, Arch. f. Naturgesch., LIII, i, p. 575. 

1898. Protosquilla cerebralis, Borradaile, Proc. Zool. Soc., p. 33, pl. v, fig. 6a. 

1899. Protosquilla cerebralis, Borradaile, in Willey’s Zool, Results, p. 399. 

1902. Protosquilla cerebralis, de Man, Abhandl. Senck. Ges. Frankfürt, XXV, p. 921. 

1910. Protosquilla cerebralis, Fukuda, Annot. Zool. Japon., VII, p. 130. 


Gonodactylus glyptocercus is readily distinguished from its ally G. ectypus by the 
peculiar sculpture on the telson and last two abdominal somites. 

The length of the carapace in the median line is more than one and a half times its 
breadth at the subacute, but rounded, antero-lateral angles. The posterior margin is 
slightly concave as are also the anterior edges on either side of the rostral base. The 
rostrum is sharply trispinous. The median spine reaches to half the length of the 
eyestalks, is fully three times as long as the basal undivided part, and is longer and more 
slender than the forwardly curved lateral spines. 

The dorsal processes of the ophthalmic somite are similar in shape to those of 
G. glaber. ‘The eyes are cylindrical ; in large examples they are about one third the 
median length of the carapace and reach a little beyond the middle of the second seg- 
ment of the antennular peduncle. In dorso-lateral view the cornea is set almost trans_ 
versely on the stalk. 

The mandibular palp is composed of two segments. 

The basal part of the dactylus of the raptorial claw is strongly swollen; the ex- 
ternal margin is notched (sometimes very obscurely) in its proximal portion and the 
inner edge is microscopically serrate. The lateral margins of the sixth and seventh 
thoracic somites are rounded, the latter rather more broadly than the former. 

Except for the customary lateral L-shaped marginal ridge the first four abdominal 
somites are entirely smooth. The postero-lateral angles of the first two are rounded, 
those of the third slightly more pointed, those of the fourth subacute. The dorsal 
surface of the fifth somite is characteristically grooved. The grooves are very fine and 
separated by broad and flat longitudinal ridges; in large examples there are about 
twenty-four in a transverse series, but a few of these bifurcate before reaching the 
posterior margin. On either side of the middle line the grooves are parallel, but 
laterally they run obliquely inwards and backwards. The postero-lateral angles are 
bluntly pointed. 

On the sixth somite the customary elevations are obscure, and the whole surface is 
finely grooved in a complicated, but symmetrical pattern, the details of which are well 
shown in the figure given by Brooks (sub P. cerebralis). A peculiar (-shaped ridge 
on either side near the anterior margin is perhaps worthy of special mention. The 
postero-lateral angles are rounded. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 187 


The telson is considerably broader than long. The median elevation is low, some- 
what triangular in shape, and is continued to the posterior margin as a pair of blunt 
tidges. The submedian elevations are oval and extend posteriorly to the extreme 
distal edge where each terminates in a small submedian tooth. These three elevations 
are bounded by furrows and their margins are characteristically incised by numerous 
grooves running towards the centre of each. Along the lateral edges of the telson 
there are four blunt ridges ; the innermost projects as an intermediate tooth on the 
distal margin, while the three others are united posteriorly to form a lateral tooth. 
The median fissure is well-marked ; it is cut right through the telson for nearly half 
its distal length and is continued forwards on the dorsal surface as a deep groove. 
The convex margins between the submedian teeth bear a series of spinules, the outer- 
most of which is larger than the rest, and there is in addition a small spinule on the 
inner aspect of each of the two lateral teeth. 


The inner spine of the process from the base of the uropods is more than half the 
length of the outer, and the proximal segment of the exopod bears eight to ten movable 
spines on its external edge. 

In small specimens, up to 12 mm. in length, the grooves on the last two abdominal 
somites are imperfectly developed, but the characteristic sculpture is found on the 
telson. 

Comparison of Brooks’ figures of Protosquilla cerebralis with Wood-Mason’s type 
of G. glyptocercus leaves no doubt that the two forms are synonymous, as de Man has 
already suggested. The Indian specimens, however, differ from the figures in the 
“ Challenger ’ report in the particulars mentioned by Fukuda. 

The dorsal surface of many preserved specimens is closely mottled with dark 
pigment except on the last abdominal somite and telson. Other specimens are pale 
with four dark spots on the sixth thoracic somite and dark median patches on the 
seventh thoracic and first and fourth abdominal somites. 


There are sixteen specimens in the Indian Museum, registered as follows :— 


3038 : 4 r 
5; Nicobars. S. Kurz. 1 O AR sooo. VDE, 
7 à 
5s06-17 ¢ Gt. Coco I., N. Andamans. ‘ Investigator.’ 63,82, I0'5-29 mm. 
9 
T Kabusa Is., Mergui Archipelago. ‘ Investigator.’ 19,21 mm. 


Gonodactylus glyptocercus has also been recorded from the Fiji Is. (Brooks) ; 


Rotuma, the Loyalty Is. and New Britain (Borradaile) ; the Ogasawara Is. (Fukuda) ; 
Ternate (de Man) and Pulo Edam, near Java (de Man). 


21. Gonodactylus excavatus, Miers. 


Plate ere 12), 107485, 


1880. Gonodactylus excavatus, Miers, Ann. Mag. Nat. Hist. (5), V, p. 123, pl. iti, figs. IT, 12. 
1886. Protosquilla excavata, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 78. 
1893. Gonodactylus excavatus, Pocock, Ann. Mag. Nat. Hist. (6), XI, p. 476. 


188 Memoirs of the Indian Museum. Von AVE 


This species along with the four following is easily distinguished from the others 
in the section (7.e. species 12-20) by the very wide median excavation in the distal 
margin of the telson. 

The carapace is broader behind than in front and, at its narrowest, is about two- 
thirds the median length. The posterior margin and the anterior margins on either side 
of the rostral base are slightly concave ; the antero-lateral angles are produced and 
subacute. The rostrum is sharply trispinous; the median spine reaches as far as the 
corneal portion of the eyes; those of the lateral pair are much shorter and rather 
stouter. The basal part of the rostrum is extremely short, little if at all longer than 
one quarter the length of the median spine. 

The dorsal processes of the ophthalmic somite have much the same form as in 
G. glaber. The eyes reach to about the middle of the second segment of the antennular 
peduncle ; they measure one-third the median length of the carapace in large specimens, 
but are longer in small examples. They differ from those of the preceding species in 
being distinctly flattened and the cornea, seen in dorso-lateral view, is set transversely 
on the stalk. The mandibular palp is composed of two segments. 

The basal part of the dactylus of the raptorial claw is greatly inflated and its ex- 
ternal margin is notched proximally, as in G. glaber. 

The margins of the sixth and seventh thoracic somites are truncate with rounded 
anterior and posterior angles. The median portions of the first five abdominal somites 
are entirely smooth, but each possesses laterally the L-shaped ridge noticed in the 
preceding species; the postero-lateral angles of the first three are rounded, those of 
the fourth subacute, while those of the fifth are strongly produced and spinous. In 
the fifth there is a rather deep furrow on either side which runs to the extreme distal 
edge of the somite, external to a small rounded lobe on the posterior margin. ‘The sixth 
somite bears three pairs of smooth elevations ; the innermost are the shortest, those of 
the intermediate pair are constricted in the middle, while the outermost, which are the 
longest, meet the external margin in an obtuse postero-lateral angle. 

The telson is about as long as wide. The three mid-dorsal bosses are similar to 
those of G. glaber, but the median one is longer and is entirely separated by grooves 
from the two laterals. The outline of these prominences is not even: the edges of each 
are furrowed and eroded. As in the preceding species there is a strong rounded keel run- 
ning along the lateral margins. Distally there is a wide and deep triangular excavation, 
which bears on its edges numerous (about twelve) small denticles. This excavation 
forms a sharp contrast with the very narrow median fissure found in the preceding 
species. On either side of this gap there are two small tooth-like lobes, the inner more 
acute than the outer, and between each pair of lobes there are one or two small 
denticles. 

The inner spine of the bifurcate process of the uropod is more than half the length 
of the outer, and on the external margin of the basal segment of the exopod there is a 
series of nine to eleven movable spines. 

A few of the specimens are, in spirit, densely mottled with dark pigment on the 
carapace and first five abdominal segments ; the other examples are faintly speckled 
or colourless. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 189 


The fourteen examples of this species in the Indian Museum were obtained at 
the following localities :— 


u Off Andaman Is., 20 fathoms. ‘ Investigator.’ 23, 49, 1I'5-I8 mm. 
meet "3 Off Little Andaman I., 104 fathoms. “ Investigator.’ 18, 3 ©) 5 1S) weaver, 
ne, Off Table I., Coco group, Andamans, 94 fms. “ Investigator.’ 13,32 ,13-I6'5 mm. 


The locality of Miers’ type specimen of Gonodactylus excavatus is unknown. ‘The 
only other example which has hitherto been recorded was found on the Macclesfield 


Bank, China Seas, in 24 fathoms (Pocock). 


[Gonodactylus folini, A. Milne-Edwards.] 
1868. Gonodactylus folinii, A. Milne-Edwards, Nouv. Arch. Mus. Hist. Nat., IV, p. 65, pl. xviii, 
figs. 8-11. 

1880. Protosquilla folinii, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 70. 

1886. Protosquilla elongata, Brooks, ibid., p. 67, pl. xv, figs. 2, 12; pl. xvi, fig. 4. 

1895. Gonodactylus fohinii, Hansen, Isop., Cumac. u. Stomatop. Plankton-Exped., p. 86. 

Gonodactylus folim is known only from the Atlantic, in the vicinity of the Cape 
Verde Is. Hansen has re-examined the specimen which Miers (1880 (a), p. 123) recorded 
from Mauritius and remarks that it is in reality an example of Gonodactylus ectypus, 
Müller. The species has in consequence no place in a list of Indo-pacific species. 

G. folim is allied to G. excavatus, but the distal excavation of the telson is not so 
deep and bears setae, but no denticles, on its edges. Moreover, the two apical teeth, 
which are found in the preceding species on either side of the excavation, are replaced 
in the Atlantic form by three blunt lobes. 


22. Gonodactylus brooksi (de Man). 
1887. Protosquilla brooksii, de Man, Arch. f. Naturgesch., LIII, i, p. 579, pl. xxii a, fig. 8. 
1910. Protosquilla brooksii, Fukuda, Annot. Zool. Japon., VII, p. 140, pl. iv, figs. I, 1a. 

In the possession of a wide and deep median excavation in the distal margin of 
the telson G. brooksi and the three succeeding species resemble G. excavatus and folini ; 
they are, however, readily distinguished by the presence of stout spinules or long spines 
on the dorsal aspect of the telson. 

The carapace, in G. brooksi, is a little longer than broad ; it is rather wider behind 
than in front, and the antero-lateral angles are rectangularly rounded. The median 
of the three rostral spines is very long, reaching to or beyond the anterior extremity 
of the eyes, while those of the lateral pair are much shorter, and extend obliquely for- 
wards to the base of the eyestalks. The proximal undivided portion of the rostrum 
is exceedingly stout. ‘The eyes are slightly flattened dorso-ventrally ; the cornea is a 
little wider than the stalk, and is set transversely on it. 

The first four abdominal somites are laterally carinate as in the preceding species, 
but are quite smooth dorsally. The postero-lateral angles of the first three segments 
are rectangularly rounded; those of the fourth are pointed, while those of the fifth are 
sharply spinous. The median portion of the fifth somite is smooth in front, but poste- 
riorly bears four longitudinal depressions, often constricted in the middle, and evidently 


190 Memoirs of the Indian Museum. [Gr JOY, 


very variable inform. ‘The lateral portions are grooved longitudinally throughout the 
length of the somite. On the sixth somite there are four large tubercles placed in a 
transverse series. In de Man’s type specimen those of the submedian pair bear each 
two stout spinules, but ten were observed by Fukuda in a larger example from the 
Japanese coast. The lateral tubercles, which may also bear a varying number of 
spinules, are sometimes broken up into a number of smaller eminences. Numerous 
stout spinules also occur between these and the lateral edges, and in the Japanese ex- 
ample several small spinules are found on the anterior margin. 

The telson is about as long as wide and the deep median excavation extends to al- 
most half its entire length. It bears three median bosses: the median being triangular 
and pointed behind, the lateral pyriform, posteriorly rounded and not reaching the distal 
margin. The surface of the telson, like that of the sixth somite, is beset with numerous — 
spinules, but their precise number seems subject to much variation. The median boss 
bears nine in the type specimen, while on those of the lateral pair there are ten. In 
Fukuda’s specimen there are on the contrary about thirty on each of the bosses. The 
lateral margin is armed with six to fourteen spinules, and there may be as many as 
twenty others between this edge and the lateral dorsal bosses. On the margins of the 
large distal excavation there are six to twelve pairs of spinules. 

There is a stout spinule at the proximal end of the basal segment of the uropods and 
the inner spine of the broad ventral bifurcate process is about half the length of the outer. 
The inner segment of the exopod bears ten movable spines on its external margin. 

The variation in the number of spinules in this species is very remarkable. It 
may be shown eventually that Fukuda’s example deserves varietal recognition ; but, 
inasmuch as only three specimens are yet known, it is impossible at present to reach 
any definite conclusion. 

De Man notes the presence of bright green colouration on the tubercles of the last 
two segments, but Fukuda did not find this in the specimen which he records, and ex- 
amined when living. 

There are no examples of Gonodactylus brooksi in the Indian Museum. The two 
type specimens, the largest 28 mm. in length, were found at Pulo Kdam, Java (de Man), 
while Fukuda’s specimen, 39°5 mm., was found in Tateyama Bay, Awa, Japan. 


23. Gonodactylus hystrix (Nobili). 
1899. Protosquilla hystrix, Nobili, Ann. Mus. Civ. Genova (2), XX, p. 276. 


This species, which has not been figured, is evidently very closely allied to G. 
brookst. According to the published account it differs from the latter in having fifteen 
carinae on the fifth abdominal somite (including the two marginal), the seven median 
being limited to its posterior half. The sixth somite is covered with small spinules 
with recurved points, directed backwards, and arranged in three regular series. The 
telson is beset with spinules of a similar character, and the three dorsal bosses are of 
a circular shape. 

Nobili records a single specimen, 15 mm. in length, from Beagle Bay, British New 
Guinea. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. IOI 


24. Gonodactylus spinosissimus, Pfeffer. 
Plate X, figs. 124, 125. 


1888. Gonodactylus spinosissimus, Pfeffer, Mitth. naturh. Mus. Hamburg, VI, 4, p. 35. 

1894. Gonodactylus spinosissimus, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 493. 

1906. Gonodactylus spinosissimus, Nobili, Ann. Sci. Nat. Zool. (9), IV, p. 326. 

1906. Protosquilla spinosissima, Tattersall, Ceylon Pearl Oyster Rep., V, p. 173, pl. ii, fig. 19. 


The carapace is widest posteriorly and is a little longer than its greatest breadth. 
Its posterior margin and the anterior edges on either side of the rostral base are con- 
cave. The antero-lateral angles are subacute. The rostrum is sharply trispinous ; 
the median spine is more than three times as long as the proximal undivided part and 
extends to the cornea of the eyes ; the outer spines are much shorter and reach a little 
beyond the base of the stalk. 

The dorsal processes of the ophthalmic somite are clearly visible between the 
rostral spines and resemble those of G. pulchellus, their acute antero-lateral angles ex- 
tending outwards almost to the external margin of the eyestalk. The eyes are flat- 
tened ; the cornea is wider than the stalk, distinctly bilobed, and the corneal and 
peduncular axes are a little oblique. 

The mandibular palp is composed of two segments. The basal part of the raptorial 
dactylus is very strongly swollen and its external margin is feebly notched proximally. 

The lateral margins of the sixth and seventh thoracic somites are truncately 
rounded, the former more broadly than the latter. Except for the customary antero- 
lateral carinae, the first four abdominal somites are smooth. The postero-lateral angles 
of the first three are rounded, those of the fourth rounded or subacute, those of the fifth 
sharply spinous. The fifth somite is smooth in the middle, but bears three or four longi- 
tudinal carinae on either side separated by furrows. 

The sixth somite is fused with the telson.' It bears four large rounded tubercles, 
a pair of submedians and a pair of laterals, the former situated well in advance of the 
latter. All four tubercles are beset with long and slender spines and in addition there 
is a row of similar spines along the anterior and lateral margins and numerous others 
in the vicinity of the lateral tubercles. 

The telson excluding the spines on the edges is almost as broad as long. ‘The 
lateral margins are only very slightly convex. The wide and deep median distal 
emargination is conspicuous, and the apices on either side areıcomposed of two subequal 
lobes separated by a wide notch. The inner of these lobes is narrower than the outer 
and terminates in a stout spine with a movable spinule on its inner aspect. On the 
dorsal surface there are three rounded bosses. The median is nearly circular, while 
the two forming the lateral pair are slightly oval and are situated much further back- 
wards. All three tubercles bear long and slender spines similar to those on the preced- 
ing segment, and in addition, others are found investing all the margins and the surface 
near thetubercles. Those situated between the tubercles and the spinous lateral margin 
are arranged on either side in two regular longitudinal rows. 


! More completely so than in any other species examined. 


192 Memoirs of the Indian Museum. Worle 


The basal segment of the uropods bears one or two sharp spines on its dorsal surface 
at the proximal end, and the inner spine of its ventral process is fully two-thirds the 
length of the outer. There are nine to eleven movable spines on the external edge of 
the basal segment of the exopodite. 


The colouring of living specimens is rather constant. The whole dorsal surface 
of the carapace and abdomen is densely maculated with dark brown tending to red or 
purplish red at the anterior margin of each post-abdominal segment and especially well- 
defined on the fourth and fifth abdominal somites. There is a pale transverse band a 
little behind the middle of the carapace and, frequently, a large subquadrate pale patch 
on the last thoracic and third abdominal somites close to the posterior margin. The 
sixth abdominal somite and telson, with their investing spines, are red or dull purplish 
red, the boss-like prominences usually paler. The antennular flagella are orange yellow; 
the eyes black with mottled brown stalks. The merus of the raptorial claw is pale or 
mottled with brown and the dactylus is suffused with red. 

The specimens in the Indian Museum are registered as follows :— 


Ei N. Cheval Paar, Cevlon. T. Southwell. 63,109, 12-34 mm. 
nm Ceylon. ex coll. Herdman. 19,28 mm. 
= Pearl Banks, Ceylon (from Spongodes). T. Southwell. If, 25 mm. 


Gonodactylus spinosissimus has been recorded from several localities in the neigh- 
bourhood of Ceylon (Tattersall), from the Red Sea (Nobili), and from Zanzibar (Pfeffer). 
It is usually found in shallow water, but Tattersall records it from 45-50 fms. on Muttu- 
varatu Paar. 


25. Gonodactylus guerini, White. 
1861. Gonodactylus guerinit, White, Proc. Zool. Soc., p. 43, pl. vii, and Ann. Mag. Nat. Hist. (3), 
VII, p. 480. 
1880. Gonodactylus guerinii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 121. 
1886. Protosquilla guerinii, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 75, pl. xvi, 
figs. I, 6. 

This peculiar species, which is not represented in the Indian Museum, is perhaps 
distantly related to G. spinosissimus. According to Brooks’ excellent description it 
may be distinguished by the following characters :— 

1. The antero-lateral angles of the carapace are broadly rounded. 

2. The median spine of the rostrum is a little shorter, and the basal undivided 

part is, according to Brooks’ figure, considerably longer. 

3. The first four abdominal somites have a small sharply defined indentation on 
each side at about one-third the distance from the lateral edge to the middle. 

4. The posterior part of the fifth abdominal somite bears six transverse rows of 
short spines, and is separated from the smooth anterior part by a curved trans- 
verse line. 

5. The sixth abdominal somite is armed with very numerous long cylindrical 
spines each of which ends in a blunt rounded tip from which a soft tubular 
fleshy process protrudes. The customary tubercles appear to be absent. 


1010. S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 193 


6. The telson is covered with regularly arranged spines with fleshy appendages 
similar to those on the preceding segment, twenty-two of the largest occupy- 
ing the median area. The customary tubercles are again absent. On the 
distal margin there are four long spine-like processes which bear secondary 
spines on their edges. These processes are separated by three deep subequal 
notches which, though much wider than the narrow fissures found in 
G. pulchellus and its allies, are nevertheless parallel-sided : the median fissure 
does not take the form of a wide gap or emargination as in the five preceding 
species. 

7. The inner uropod bears five immovable spines placed in a longitudinal row on 
its dorsal surface. 


The function of the curious fleshy processes on the tips of the spines of the last two 
segments is wholly unknown, and in their possession the species is unique among Stoma- 
topoda. 

Brooks notes that his spirit specimen was brown with a pale transverse band on the 
carapace. 


Only two examples of Gonodactylus guerim are known. The type, a male 57 mm. 
in length, was found by H.M.S. ‘ Herald’ at the Fiji Is. (White, Miers) ; the second 
example, a female 28°5 mm. in length, was obtained by H.M.S. ‘ Challenger ’ at Hono- 
lulu (Brooks). 

26, Gonodactylus acanthurus, Tattersall. 
1906. Gonodactylus acanthurus, Tattersall, Ceylon Pearl Oyster Rep., V, p. 171, pl. 1, figs. 11-15. 

"1907. Gonodactylus acanthurus, Borradaile, Trans. Linn. Soc. Zool. (2), XII; p. 210. 

The carapace is about one and a third times as long as broad ; its anterior margin 
trends forwards on either side of the rostrum, so that the subacute antero-lateral angles 
are in advance of the rostral base. The rostrum is trispinous ; the median spine is 
long and reaches almost or quite to the middle of the eyestalks, while the laterals are 
decidedly shorter than in most species of the preceding group, and fail to reach the base 
of the eyestalks. The cornea of the eye is narrower than the stalk and is set very 
obliquely on it. The antennular peduncle is extremely short and is exceeded in length 
by the eyestalks. 

The mandibular palp is entirely absent. 

There is a long movable spine at the proximal end of the raptorial propodus on its 
upper margin, and the external edge of the dactylus is feebly notched at the base. 

The last three thoracic somites have the usual form, and the first five segments of 
the abdomen are entirely smooth except for the customary ridge along the lateral and 
antero-lateral margins and for a few microscopic pits on the dorsal surface. The postero- 
lateral angles of the first four are rounded, those of the fifth subacute. 

The posterior margin of the sixth somite is almost straight in dorsal view and 
bears six smooth longitudinal ridges; the submedians are swollen, both higher and 
broader than the intermediates and the laterals are co-terminous with the spinous 


postero-lateral angles. The submedian and intermediate ridges do not terminate in 
spines. 


104 Memoirs of the Indian Museum. Oma IO, 11633. 


The length of the telson, including the terminal spines, is less than its greatest 
breadth. The submedian, intermediate and marginal teeth are represented by long 
and slender spines placed almost in a transverse row on the distal edge. The sub- 
medians have movable tips and a series of slender denticles on their inner margins ; 
there are two similar denticles between the submedians and intermediates and one 
between the intermediates and laterals. In the middle of the dorsal surface are three 
oval bosses, the median slightly the broadest. They are smooth anteriorly ; but 
posteriorly bear a varying number of spinules and short spines. There are also between 
these bosses and the posterior margin two transverse rows of five spines. The spines at 
the ends of the bosses are the shortest and those on the distal margin the longest ; 
those of the intervening rows are intermediate in length. In the largest specimen in 
the Indian Museum the spines are rather more numerous than in the type specimen 
figured by Tattersall. 

The ventral process of the peduncular segment of the uropods is split into two 
long spines, the inner a little shorter and more slender than the outer. The proximal 
segment of the exopod projects a little beyond the insertion of the ultimate segment 
and bears eight or nine movable spines on its external edge. The endopod and small 
ultimate segment of the exopod bear setae on their outer margins only. The inner 
margin of the former segment bears from four to six flat and acute spines, while three or 
four similar spines are found on the same margin of the latter segment. 


The colouring of fresh specimens is variable. Two examples were quite pale in 
general tone, while a third was bright brick red, faintly mottled with pale dull yellow. 
The patches of black chromatophores which Tattersall has mentioned on the sixth 
thoracic and first abdominal somites were conspicuous, and on the former segment 
there were in addition a pair of lateral patches. ‘The last somite and telson were beauti- 
fully marbled with olive green in the pale specimens, and with dark red in the red 
example, but the two median ridges on the last abdominal somite were pure white. 
On the merus of each raptorial claw there was a dorsal black spot. 


This peculiar species appears to be one of the smallest known Stomatopoda. The 


largest example in the Indian Museum is a trifle longer than either of Tattersall’s type 
specimens. 


7502-4 


ıo N. Cheval Paar, Ceylon, 5 fms. T. Southwell. 32 ,1o-II’5 mm. 


G. acanthurus was hitherto known only from two specimens found in 45-50 
fathoms at Muttuvaratu Paar, Ceylon (Tattersall). 


IL NID NI IHN NN N INN N 


ADDENDUM. 


Since the preceding report has been in press I have had an opportunity of visit- 
ing Kilakarai and Pamban in the Ramnad District, S. India. During the first half 
of the tour I was fortunate in being accompanied by Dr. J. R. Henderson, Superin- 
tendent of the Madras Museum: it was in localities at no great distance from those 
that we visited that Dr. Henderson, some twenty-four years ago, made a great part 
of the extensive collections of Crustacea which he subsequently described in the 
Transactions of the Linnean Society (1893) under the title ‘‘ A Contribution to Indian 
Carcinology.’’ 

The fauna of the district seems to be one of almost inexhaustible richness 
and during the comparatively short period in which we were collecting, we succeeded 
in obtaining nine species of Stomatopoda, including several scarce forms; we were 
also able to make a few interesting observations on the habits of Gonodactylus. 


Squilla quinquedentata, Brooks. [p. 52.] 

Judging by the number of specimens hitherto found in Indian waters this species 
appears to be very scarce, but three large individuals, two males and one female, 
ranging from 117 to 125 mm. in length, were obtained from the fishermen at 
Kilakarai. 

The specimens differ in a few unimportant details from those previously 
described. In two examples the anterior bifurcated portion of the median carina of 
the carapace is distinct, but interrupted and non-existent at its proximal end as in 
Squilla interrupta and in S. oratoria var. perpensa. In all the specimens the anterior 
margin of the ophthalmic somite is feebly emarginate and the posterior lobe of the 
fifth thoracic somite is acute. The lateral carinae of all the abdominal somites end 
in spines, the intermediate teeth of the telson are only very slightly inturned and 
there are four submedian denticles, eight or nine intermediate and one lateral. The 
colouration of the living specimens was not distinctive, the dark patches noticed in 
the previously recorded examples at either end of the median carina of the telson 
being indistinguishable. 

Squilla nepa, Latreille. [p. 60.] 

Found in abundance by the fishermen at Kilakarai. The largest specimen seen 
was a female 134 mm. in length; small individuals, including one only 21 mm. in 
length, were caught among weeds close to the shore. 


Squilla holoschista, Wood-Mason, MS. [p. 64.] 
Not uncommon in fishermen’s nets. 


196 Memoirs of the Indian Museum. NVorenz 


Squilla oratoria var. perpensa, Kemp. [p. 70.] 

A few specimens of no great size were brought in by the fishermen at Kilakarai, 
and a very youngle example, 2I mm. in length, was found among weeds near the 
shore. 

Squilla multicarinata, White. [p. 86.] 

A magnificent example of this scarce and finely sculptured species, a male 
measuring 82 mm., was brought in by a fisherman during our stay at Kilakarai. 
The dusky patches noticed on the second and fifth abdominal somites are not visible 
in this specimen, but in every other respect it agrees precisely with the examples in 
the Indian Museum from Hongkong. 

Squilla multicarinata has not hitherto been recorded west of Burma. 


Pseudosquilla ciliata (Fabricius). [p. 96.] 
A large male, 82 mm. in length, was brought to us by a fisherman at Kilakarai. 
The specimen was bright olive green in life with the fringes of the antennal scale and 
uropods crimson. 


Lysiosquilla acanthocarpus, Miers. [p. 120.] 


A living specimen of this uncommon species, a female 56 mm. in length, was 
discovered at low water on the beach at Kilakarai. The spot where it was found was 
one that we habitually passed, but though careful watch was kept no further speci- 
mens were obtained. A dredge was shot several times a short distance from shore 
and dragged to land, but this too proved unsuccessful. 

Placed in a shallow dish with a little water the specimen gave an interesting 
demonstration of the ease with which the species of this genus can reverse their posi- 
tion in a narrow burrow. When laid on its back it would rapidly assume a normal 
posture by turning its anterior extremity upwards and backwards and creeping along 
its own body. Such an action would be impossible in the case of more firmly seg- 
mented species such as those of the genera Squilla, Pseudosquilla and Gonodactylus 
and there can be little doubt that it is among the species of Lysiosquilla that the 
highest degree of specialization for life in burrows is to be found. 

When picked up, the specimen attacked the fingers of its captor with both ends 
of its body simultaneously, striking out vigorously with its raptorial claws and at the 
same time endeavouring to make wounds with the spinous armature on its telson and 
uropods. 

Its colour did not differ very much from that of specimens preserved for many 
years in alcohol. The rostrum, eye-peduncles, and the basal segments of the anten- 
nules and antennae were mottled with russet brown, black and pure white. The 
proximal part of the antennal scale was white. The dark bands on the free thoracic 
somites and abdomen were blackish grey,separated by ivory white. On the telson 
the black colouring was much more intense than elsewhere and the three pale streaks 
that invade the black patch proximally were pale lemon yellow. Seen from below 
the animal was semi-transparent. 


1913.] S. Kempe: Crustacea Stomatopoda of the Indo-Pacific Region. 197 


The fishermen spoke of a large species with similar colouring, doubtless Lysio- 
squilla maculata ; no specimens, however, could be obtained. 


Gonodactylus glabrous, Brooks. [p. 167.| 


The species occurred in profusion on coral reefs both at Kilakarai and Pamban, 
and at the latter place was also obtained on reefs exposed at low tide. The speci- 
mens were, for the most part, found in crevices and holes in coral blocks and in these 
situations examples of all sizes (from 7°5 mm. in length and upwards) were obtained. 
Very small individuals were also caught by dragging a fine-meshed net through the 
Zostera and other weeds which flourished inside the reef in water not more than one 
or two feet deep. 

In specimens 7°5 mm. in length only the median carina of the telson is distinguish- 
able, but in examples only 1 mm. longer all five are developed. Among the various 
forms which the telson may assume in Gonodactylus chiragra, form A (=incipiens) most 
nearly resembles G. glabrous, and according to Lanchester this form constitutes a true 
connecting link between the two species. It is therefore noteworthy that no speci- 
men was found on this occasion that can be referred to the variety and that even at 
a very early age (specimens 8-9 mm. in length) the distinctions between G. glabrous 
and G. chiragra form H are well-marked. In all G. glabrous exceeding 7°5 mm. in 
length five carinae are developed, and of these five the submedians are invariably 
longer than those of the pair external to them. 

Many females with egg-masses were found and this observation is one of some 
interest seeing that it does not appear to have been made hitherto in Indo-pacific 
waters. The females were found in the coral-masses in holes that appeared to have 
been made originally by Lithodomus or some other rock-boring mollusc, the eggs, which 
‚were of a bright greenish-yellow colour, lying at the bottom of the burrow in a loosely- 
compacted discoidal mass. When disturbed the female rapidly gathered the eggs 
into a ball and, holding them with her thoracic appendages, rushed wildly to and fro 
“ seeking concealment or escape. Even the shock of sudden immersion in alcohol was 
not always sufficient to induce the mother to lay aside her parental responsibilities. 
The egg-masses varied, according to the size of the female, from 8 to 14 mm. in 
diameter, individual ova measuring from ‘7 to ‘85 mm. 

The habits of the species appear to be closely similar to those of G. oerstedi, 
observed by Brooks in the Bahamas; but whereas the latter species is said to con- 
struct burrows for itself in solid coral, G. glabrous seems merely to inhabit those made 
by other animals.' In order to tunnel in hard rock an animal must needs be highly 
specialized and consideration of the very close uniformity in structure in G. glabrous 
and G. oerstedi leads one to believe that adaptation for such a purpose, if present 
in one form, would certainly be present in the other. Although Brooks never saw 
specimens engaged in tunnelling he had no doubt that the species actually made its 


1 Specimens were never found in new and clean-cut burrows, but only in those eroded and over- 
grown with encrusting organisms. 


198 Memoirs of the Indian Museum. [Vor. IV, 


burrows itself and his observations are, as a rule, so accurate that one hesitates to 
suggest that he may in this instance have been misled. 

The source of the noise made by the species of Gonodactylus seems never to have 
been adequately described. In the case of G. glabrous and G. demani, and doubtless 
with other species of the genus also, it is caused by the raptorial claws when these 
limbs are used for offensive or defensive purposes. When a living specimen is seized 
the distal end of the raptorial propodus is fitted into the socket hollowed in the 
merus for its reception, and while still lodged in this socket muscular effort is brought 
to bear. Then by a slight alteration in the relative positions of the mero-carpal 
and carpo-propodal joints the propodus is suddenly released and flies straight out in 
front. The dactylus, which before the release of the propodus is flexed close against 
that segment, also flies forwards so that all the segments of the limb are momentarily 
extended as nearly as possible in a straight line. The action takes place with great 
rapidity and is in consequence somewhat difficult to observe ; it is roughly com- 
parable to a rapier-thrust and a well-grown specimen might, I believe, under favour- 
able circumstances succeed in piercing the skin of the hand. 

The sound appears to be produced by the release of the propodus from the 
socket in the merus. In all probability it is merely a subsidiary result of the mecha- 
nical process involved ; it is not so loud as that made by the Alpheid Macrura, and is 
no doubt less effectual in frightening an opponent. 


Gonodactylus demani, Henderson. [p. 164.] 


This species was not uncommon both at Kilakarai and Pamban, occurring in the 
same situations as G. glabrous. It was in an adjacent locality that Dr. J. R. Hen- 
derson in 1889 obtained the specimens from which he drew up the original descrip- 
tion of the species. 

A curious feature of allthe specimens collected is that the setae on the inner 
margin of the inner uropod are absent. In all other examples of G. demani in the 
Museum collection, with the exception of one from the Henjam Is., Persian Gulf, 
the entire margin is invested with setae. The absence of these setae is clearly shown 
in Henderson’s figure of the type specimen, and on careful examination of the speci- 
mens recently obtained, it is evident that their absence is not traumatic, as no trace 
of the pits from which the setae arise can be detected. 

The character no doubt appears somewhat insignificant and the evidence at 
present available is not sufficient to permit of any suggestions regarding the possible 
existence of local races: the specimens in the Indian Museum prove that both forms 
occur in the Persian Gulf. Considering, however, the differences of opinion that 
prevail regarding the status of the species in the chiragra section of the genus, it is 
well to place on record any definite character such as that noted above; and in the 
present instance a considerable amount of interest attaches to the disposition of the 
setae on the uropod in view of the fact that the form of this segment and its setal 


! It may be mentioned here that in the specimens from Ceylon which I have referred to G. demani 
var. Spinosus the setae invest all the margins of the inner uropod. 


1913.] S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 199 


investment are curiously modified in the small section of the genus that comprises 
G. herdmani, brevisquamatus, furcicaudatus and other species. The S. Indian exam- 
ples of G. demani are, in my opinion, more closely related to the members of this 
section (in particular to G. herdmani, the least specialized form) than any other 
known species of the genus not included therein. 

In the majority of the specimens the spinulation of the telson consists of a few 
sharp prickles at the distal ends of the carinae and at the bases of the submedian 
and intermediate teeth. The median carina, as in the type specimen, is often con- 
siderably wider than those on either side. 

The colour of living specimens was very variable, but usually dark in tone. As 
a rule three characteristic pale patches were distinct : (i) a narrow transverse band 
at the junction of the middle and posterior thirds of the carapace, (ii) a lateral 
streak on either side of the first abdominal somite, and (iii) a narrow transverse band 
at the end of the last abdominal somite which is continued laterally as a patch at 
the distal end of the peduncle of the uropods. In addition, living specimens in- 
variably possessed a number of small brilliant blue spots surrounded by a black ring. 
The spots were always arranged in transverse series on the thorax and abdomen, and 
at the base of the telson, their number and arrangement being subject to much 
variation. The spots immediately turn black when the specimen is placed in 
alcohol. 

Several females of this species were found guarding their eggs. The egg-mass is 
usually about 5 or 6 mm. in diameter and individual ova measure about ‘8 mm. 

_ No examples of Gonodactylus chiragra were obtained and I am inclined to 
believe that the species does not occur on the reefs which we examined. The 
experience which I obtained in the course of this visit has confirmed my belief that 
the species of the chivagra section of the genus Gonodactylus are well-defined , though 
variable within certain limits and showing a tendency towards the formation of local 
races. That G. glabrous and G. demani are specifically distinct, could, I venture to 
believe, hardly be doubted by any biologist who had had the opportunity of visiting 
the reefs at Kilakarai and Pamban, and had observed large numbers of individuals 
of both forms in their natural environment. 

In concluding this note I take the opportunity to express my indebtedness to 
Dr. J. R. Henderson, whose long experience and hearty co-operation in making the 
collection on which the note is based proved of the greatest possible assistance. 


LIST OF THE SPECIES OF STOMATOPODA. KNOWN FROM THE ATLANTIC AND THE 
MEDITERRANEAN AND FROM THE PACIFIC COAST OF AMERICA, WITH 
A FEW OF THE MORE IMPORTANT REFERENCES AND SYNONYMS. 


SQUILLA. 


Squilla aculeata, Bigelow, 1893. 
1894. Squilla aculeata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 523, text. figs. 15, 16. 
Panama ; Chili. 
Squilla alba, Bigelow, 1893. 
1894. Squilla alba, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 539, pl. xxii. 
Bahama Is. 
Squilla armata, H. Milne-Edwards, 1837. 


Sas D A, 
Squilla bifor mis, Bigelow, 1893. 
1894. Squilla biformis, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 532, pl. xxi, text-fig. 20. 
Bay of Panama, G. of California ; 85-259 fms. 


Squilla desmaresti, Risso, 1816. 


1816. Squilla desmarestii, Risso, Crust. Nice, p. II4, pl. ii, fig. 8. 

1863. Squilla desmarestii, Heller, Crust. südlich. Europ., p. 307. - 

1880. Squilla desmarestii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 28. 

1910. Squilla desmarestii, Giesbrecht, Faun. Flor. G. Neapel, XXXIII, p. 25, pl. i, figs. 6, 7, pl. iit. 
Mediterranean ; British Coasts. 


Squilla dubia, Milne-Edwards, 1837. 


1837. Squilla dubia, H. Milne-Edwards, Hist. Nat. Crust., II, p. 522. 
? 1852. Squilla rubrolineata, Dana, U.S. Explor. Exped., Crust., p. 618, pl. xli, fig. 2a, b. 
1880. Squilla dubia, Miers, Ann. Mag. Nat. Hist. (5), V, p. 24. 
1894. Squilla dubia, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 518. 
1910. Chloridella dubia, Rathbun, Proc. U.S. Nat. Mus., p. 565, pl. liv, fig. 3. 
E. Coasts of America ; West Indies; Ecuador; Peru. 


Squilla empusa, Say, 1818. 


1817. Squilla empusa, Say, Journ. Ac. Sci. Philad., I, p. 250. 
1880. Squilla empusa, Miers, Ann. Mag. Nat. Hist. (5), V, p. 23, pl. ii, fig. 12. 
1894. Squilla empusa, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 525. 

E. Coast of N. America ; West Indies ; W. Africa. 


Squilla gracilipes, Miers, 1881. 
1881. Squilla gracilipes, Miers, Proc. Zool. Soc., p. 75, pl. vii, fig. 8. 
1890. Ptervgosquilla laticauda, Hidgendorf, Sitz. Ges. naturf. Fr. Berlin, p. 172, text-fig. 
1895. Pterygosguilla laticauda = Squilla gracilipes, Hansen, Isop. Cumac. u. Stomatop. Plankton-Exped., 
p- 09. 
W. Coast of Patagonia. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 201 


Squilla intermedia, Bigelow, 1893. 
1894. Squilla intermedia, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 530, text-fig. IQ. 
1902. Squilla intermedia, Bigelow, Bull. U.S. Fish Comm. for 1900, XX, 2, p. 130. 
G. of Mexico ; Little Bahama Bank ; Porto Rico. 


Squilla mantis, Latreille, 1802. 
1825. Squilla mantis, Latreille, Encycl. Method., X, p. 471. 
1863. Squilla mantis, Heller, Crust. südlich. Europ., p. 306, pl. x, figs. 15-10. 
1880. Squilla mantis, Miers, Ann. Mag. Nat. Hist. (5), V, p. 21, pl. ii, fig. 11. 
1910. Squilla mantis, Giesbrecht, Faun. Flor. G. Neapel, XX XIII, p. 25, pl. i, figs. 1, 2, 9, pl. ii, pl. v, 
figs. 43-60. 
Mediterranean, Portugal, English Channel, North Sea. 


: Squilla neglecta, Gibbes, 1850. 
1850. Squilla neglecta, Gibbes, Proc. Americ. Assoc., p. 200. 
1880. Squilla neglecta, Miers, Ann. Mag. Nat. Hist. (5), V, p. 23. 


South Carolina. 
Squilla pallida, Giesbrecht, 1910. 


1910. Squilla pallida, Giesbrecht, Faun. Flor. G. Neapel, XX XIII, p. 25, pl. i, fig. 5, pl. ii. 


Mediterranean. 
Squilla panamensis, Bigelow, 1893. 


1894. Squnlla panamensis, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 526, text-figs. 17, 18. 
Bay of Panama ; W. Coast of Mexico ; California ; Brazil (?). 


Squilla parva, Bigelow, 1893. 
1894. Squilla parva, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 518, text-figs. II, 12. 


Bay of Panama ; W. Coast of Mexico. 


Squilla polita, Bigelow, 1893. 
1894. Squilla polita, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 513, text-fig. 8. 
California. 
Squilla prasinolineata, Dana (Ives), 1852. 
1852. Squilla prasinolineata, Dana, U.S. Explor. Exped., Crust., p. 620, pl. xli, figs 3, a-c. 
1880. Squilla dufresnii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 18, pl. ii, figs. 8, 9. 
1891. Squilla prasinolineata, Ives, Proc. Acad. Sci. Philadelphia, p. 184. 
1901, Squilla prasinolineata, Moreira, Arch. Mus. Nac., Rio Janeiro, XI, p. 5 (partim ?). 


Brazil, Yucatan, Cuba.! 


Squilla prasinolineata, Miers (nec Dana). 


1880. Squilla prasinolineata ?, Miers. Ann. Mag. Nat. Hist. (5), V, p. 19, pl. ii, fig. Io. 
1891. Squilla prasinolineata (Miers nec Dana), Ives, Proc. Acad. Sci. Philadelphia, p. 185. 
1894. Squilla prasinolineata (Dana ?, Miers), Bigelow, Proc. U.S. Nat. Mus., XVII, p. 520. 
? 1901. Squilla prasinolineata (Dana), Moreira, Arch. Mus. Nac., Rio Janeiro, XI, p. 5 (partim). 
Brazil. 

Moreira considers that this form is at most a variety of the real S. prasinolineata of Dana; but it 
seems evident that the examples which Miers described under the names S. prasinolineata ? and S. 
dufresnit are specifically distinct and Ives’ subsequent remarks on the synonymy appear reasonable. It 


' Specimens from this locality, received in exchange from the Berlin Museum, have been examined. 


202 Memoirs of the Indian Museum. [Vor. IV, 


is probable, therefore, that a new name will be wanted for the present form when further specimens have 
been examined and the question has been finally settled. 


Squilla quadridens, Bigelow, 1893. 
1894. Squilla quadridens, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 511. 
Florida. 
Squilla rugosa, Bigelow, 1893. 
1894. Squilla rugosa, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 541, text-figs. 23, 24. 
G. of Mexico. 
PSEUDOSQUILLA. 
Pseudosquilla cerisii (Roux), 1828. (See p. 96.) 
1828. Squilla cerisii, Roux, Crust. Méditerran., p. innum., pl. v. 
1863. Squilla cerisii, Heller, Crust. südlich. Europ., p. 308. 
1880. Pseudosquilla cerisii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 114. . 
1910. Pseudosquilla cerisii, Giesbrecht, Faun. Flor. G. Neapel, XX XIII, p. 34, pl. i, fig. 3; pl. iv, figs. 1-36. 
Mediterranean. 
Pseudosquilla ciliata (Fabricius), 1787. 
See p. 96. 
Pseudosquilla ferussaci (Roux), 1828. (See p. 96.) 
1828. Squilla ferussacii, Roux, Crust. Mediterran., p. innnm., pl. xxviii. 
1863. Squilla ferussacii, Heller, Crust. südlich. Europ., p. 308. 
1880. Squilla ferussacii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 16. 
1910. Pseudosquilla ferussaci, Giesbrecht, Faun. Flor. G. Neapel, XX XIII, p. 34, pl. i, fig. 8; pl. iv, figs. 
37-48. 
Mediterranean. 
Pseudosquilla lessoni (Guérin), 1830. (See p. 96.) 
1830. Squilla lessonii !, Guérin, Voy. ‘ Coquille,’ Crust., pl. iv, fig. 1. 
1832. Squilla spinifrons, Owen, Proc. Zool. Soc., p. 6. 
1837. Squilla lessonit, H. Milne-Edwards, Hist. Nat. Crust., II, p. 527. 
1837. Squilla monoceros, H. Milne-Edwards, zbid., p. 526. 
1877. Pseudosquilla marmorata, Lockington, Proc. Acad. Sci. California, VII, p. 33. 
1880. Pseudosquilla lessonii, Miers, Ann. Mag. Nat. Hist. (5), V, p. 113. 
1900. Pseudosquilla lessonii, Holmes, Occ. Papers. Calif. Acad. Sci., VII, p. 220. 
1910. Pseudosquilla lesson, Rathbun, Proc. U.S. Nat. Mus.. XX XVIII, p. 565, pl. lüi, fig. 3. 
W. Coast of America from California to Chili. 
Pseudosquilla oculata (Brullé), 1836-44. 
See p. 102. 
Pseudosquilla stylifera (H. Milne-Edwards), 1837. 
See p. 106. 


LYSIOSQUILLA. 
Lysiosquilla armata, Smith, 1881. 


1881. Lysiosquilla armata, Smith, Proc. U.S. Nat. Mus., III, p. 446. 
1894. Lysiosquilla armata, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 507. 
N.E. Coast of America, 17-120 fms. 


' Squilla lessonii on plate, S, cerisii in text, p. 40. 


1913.] S. KEMP: Crustacea Stomatopoda of the Indo-Pacific Region. 203 


Lysiosquilla biminiensis, Bigelow, 1893. 
See p. 124. 


Lysiosquilla decemspinosa, Rathbun, 1910. 
1910. Lysiosquilla decemspinosa, Rathbun, Proc. U.S. Nat. Mus., XX XVIII, p. 566, pl. liii, fig. 3. 


Peru. 
Lysiosquilla desaussurei, Stimpson, 1857. 


1853. Squilla scabricauda, De Saussure, Rev. Mag. Zool., (2), V, p. 367. 
1857. Squilla desaussurei, Stimpson, Boston Journ. Nat. Hist., VI, p. 503. 
1880. Lysiosquilla desaussurei, Miers, Ann. Mag. Nat. Hist. (5), V, p. 8. 
W. Coast of Mexico. 
Lysiosquilla digueti, Coutière, 1905. 
1905. Lystosquilla digueti, Coutière, Bull. Soc. Philom., (9), VII, p. 174, text figs. 1-7. 
G. of California. 
Lysiosquilla eusebia (Risso), 1816. 


1826. Squilla eusebia, Risso, Hist. Nat. Europe Mérid, V, p. 87, pl. iv, fig. 15. 
1869. Squilla eusebia, Nardo, Annot. Crost. ; Mem. Inst. Venetio, XIV, p. 112, pl. iii, fig. 7. 
1880. Lysiosquilla eusebia, Miers, Ann. Mag. Nat. Hist. (5), V, p. 30. 
1910. Lysiosquilla eusebia, Giesbrecht, Faun. Flor. G. Neapel, X XXIII, p. 40, pl. i, figs. 4, 10; pl. v, 
figs. I-42. 
Mediterranean. 
Lysiosquilla excavatrix, Brooks, 1886. 
1886. Lysiosquilla excavatrix, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 48, pl. x, figs. 8-16. 
North Carolina. 
Lysiosquilla glabriuscula (Lamarck), 1818. 


1818. Squilla glabriuscula, Lamarck, Hist. Nat. Anim. sans Vert., V, p. 188. 
1837. Squilla vittata, H. Milne-Edwards, Hist. Nat. Crust., II, p. 519. 
1880. Squilla glabriuscula, Miers, Ann. Mag. Nat. Hist. (5), V, p. 7. 


West Indies ; Florida. 


Lysiosquilla maiaguesensis, Bigelow, 1902. 
1902. Lysiosquilla maiaguesensis, Bigelow, Bull. U.S. Fish Comm. for 1900, XX, 2, p. 158, figs. 10-13. 
Porto Rico ; 161-172 fms. 


Lysiosquilla occulta, Giesbrecht, 1910. 
1910. Lysiosquilla occulta juv., Giesbrecht, Faun. Flor. G. Neapel, XX XIII, p. 131, pl. vi. 
Mediterranean. 
Lysiosquilla plumata, Bigelow, 1902. 
1902. Lysiosguilla plumata, Bigelow, Bull. U.S. Fish Comm. for 1900, XX, 2, 156, figs. 5-9. 
Porto Rico, 25-30 fms. 
Lysiosquilla polydactyla, v. Martens, 1881. 


1881. Lysiosquilla polydactvla, v. Martens, Sitz.-ber. Ges. naturf. Fr. Berlin, p. 92. 
1912. Lystosquilla polydactyla, Doflein und Balss, Mitt. Nat. Hist. Mus. Hamburg, XXIX, p. 40. 
Chili? Tierra del Fuego. 


Lysiosquilla maculata (Fabricius), 1793. 


See p. III. 


1825 
1837 


Memoirs of the Indian Museum. [Vor. IV, 


Lysiosquilla scabricauda (Lamarck), 1818. 
. Squilla scabricauda, Lamarck, Hist. Nat. Anim. sans Vert., V, p. 188. 
. Squilla scabricauda, Latreille, Encycl. Méthod., X, p. 470; Atlas pl. ccexxv, fig. I. 
. Squilla scabricauda, H. Milne-Edwards, Hist. Nat. Crust., II, p. 519. 
. Squilla hoeveni, Herklots, Addit. Faun. Carcin. Afric. occident., p. 17, pl. i, fig. II. 
. Lysiosquilla inornata, Dana, U.S. Explor. Exped., Crust., p. 616, pl. xli, fig. I a-e. 
. Lysiosquilla scabricauda, Miers, Ann. Mag. Nat. Hist. (5), V, p. 7. 
Texas : Florida; S. Carolina; W. Indies; Brazil; Cayenne; W. Africa. 


Lysiosquilla scolopendra (Latreille), 1825. 


. Coronis scolopendra, Latreille, Encycl. Method., X, p. 474. 
. Coronis scolopendra, H. Milne-Edwards, Hist. Nat. Crust., II, p. 531. 


1836-49. Cornis scolopendra, H. Milne-Edwards, in Cuvier’s Regne Anim., Crust., Atlas, pl. lv, fig. 3. 


1880 


1881. 


1869. 
1880. 


1884. 
1886. 


1894. 


1901. 
1903. 


1890 
1895 
1900 
IOOT 


. Lysiosquilla scolopendra, Miers, Ann. Mag. Nat. Hist. (5), V, p. 9. 


Brazil ? . 
Lysiosquilla septemspinosa, Miers, 1881, (See p. 122). 
Lysiosquilla acanthocarpus var. septemspinosa, Miers, Ann. Mag. Nat. Hist. (5), VIII, p. 368. 


W. Africa. 
CORONIDA. 


Coronida bradyi (A. Milne-Edwards), 1869. (See p. 130). 


Squilla bradyi, A. Milne-Edwards, in De Folin and Périer’s Fonds de la Mer. I., Crust., p. 137, pl. 
xvii, fig. II. 

Gonodactylus ? bradyi, Miers. Ann. Mag. Nat. Hist. (5), V, p. 117. 

Gonodactylus bradyi, Miers, Proc. Zool. Soc., p. 16. 

Coronida bradyi, Brooks, Voy. H.M.S. ‘ Challenger,’ XVI, Stomatop., p. 80. 


Bay of St. Vincent. 
ODONTODACTYLUS. 


Odontodactylus havanensis, Bigelow, 1893. (See pp. 135, 144.) 
Odontodactylus havenensis, Bigelow, Proc. U.S. Nat. Mus., XVII, p. 497, pl. xx, text figs. 1, 2. 
Cuba. : 
GONODACTYLUS. 


Gonodactylus festae, Nobili, 1901. (See pp. 147, 153.) 
Gonodactylus festae, Nobili, Boll. Mus. Torino, XVI, no. 415, p. 53. 
Gonodactylus chiragra var. festae, Lanchester, Faun. and Geog. Maldives and Laccadives, p. 456. 


Ecuador, Colombia. 
Gonodactylus folini, A. Milne-Edwards, 1868. 
See p. 189. 


Gonodactylus oerstedi, Hansen, 1895. (See footnote p. 147.) 
. Gonodactylus chiragra, Pocock, Journ. Linn. Soc., XX, p. 526. 
. Gonodactylus oerstedi, Hansen, Isop. Cumac. u. Stomatop. Plankton-Exped., p. 65, footnote. 
. Gonodactylus oerstedi, Bigelow, Bull. U.S. Fish. Comm. for 1900, XX, ii, p. 152, text figs. I, 2. 
. Gonodactylus falcatus, Moreira, Arch. Mus. Nac., Rio de Janeiro, XI, p. 1. 
West Indies to Brazil ; Fernando Noronha. 


1913. | S. Kemp: Crustacea Stomatopoda of the Indo-Pacific Region. 205 


1708. 


1798. 


1798. 


1832. 


1841. 
I841. 


1869. 


DOUBTFUL SPECIES. 
Squilla phalangium, Fabricius, Ent. Syst. Suppl., p. 416. 
““S. pollice falcato quinquedentato : dente tertio quintoque longioribus, corpore laevi. 
Habitat in India orientali Dom. Daldorff. 
Corpus S. Mantis paullo minor, laeve, glabrum, ultime segmento spinoso serratum.’’ 
Perhaps belonging to the acanthocarpus section of the genus Lysiosquilla. 


Squilla ichneumon, Fabricius, Ent. Syst. Suppl., p. 416. 
‘"S. pollice falcato quadridentata cauda margine nodoso spinoso. 
Habitat in India orientali D. Daldorff. 
Praecedente (S. phalangium) adhuc minor. Carpus utrinque lineis tribus elevatis. Cauda 
rotundata spinis crassis nodosis ciliata.’’ 
Possibly a species of Squilla of the ‘ Chloridella’ group. 
Squilla vitrea, Fabricius, Ent. Syst. Suppl., p. 417. 
““ S. thorace laevi carinato : angulis subulatis, pollice falcato sublato inermi. 
Habitat in Oceano atlantico Mus. D. Banks.’’ 
Gonodactylus ensiger, Owen, Proc. Zool. Soc., p. 6. 
‘‘ Gon. pollice edentato, ensato, intus acuto ; vostro 3-spinoso, spina intermedia obsoleta. 
Long 6 unc. Hab. Valparaiso. 
Color flavus ; chelarum caeruleus. 
Praecedenti (i.e. G. chivagra) valde affinis, praesertim sculptura armuturaque segmentorum ulti- 
morum ; sed differt pollice ad basin non ventricoso, ad marginem internum non crenato, 
spina rostri media non producta ; necnon magnitudine.’’ 


Pseudosquilla cylindrica, nom. nud? ) Attributed to Guérin by Eydoux and Souleyet, Voy. de la 
Pseudosquilla lebtodactyla, nom. nud ? Ponte Zoola Crust ls ps 263" 


Squilla mantis, Rondel., Bianconi, Mem. Accad. Sci. Bologna (2), IX, p. 210. 
Probably one of the species included in the oratoria section of the genus Squilla. The 
reference is quoted by Hilgendorf, Monatsber, d. kon. Akad. Wiss. Berlin, 1879, p. 846. 


ug RE EO 


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. Dictionnaire des Sciences Naturelles, Vol. XXVIII, pp. 337-342 (Strassburg 


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. Consid. Gén. des Crustacés, p. 251, pl. xlüüi (Paris). 


Proc ILO, SOC. 
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‘© Die Dekapoden und Stomatopoden der Hamburger Magalhaensischen 
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Proc. American Assoc., 3rd meet- 


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1913.] 
Heller, C., 1865 


Henderson, J. R., 1893 
Herbst, J. F. W., 1796 
Herklots, J. A., 1851 
Hess, W., 1865 
Hilgendorf, F., 1869 
Hilgendorf, F., 1879 


Hilgendorf, F., 1890 


Hoffmann, C. K., 1874 (a) .. 


Hoffmann, C. K., 1874 (b) .. 


Holmes, S. J., 1900 
Hutton, F. W., 1879 


Ives, J. E., 1891 
Jurich, B., 1904 
IK GAO 9, TORTE: 


Kirk, T. W., 1878 
Kirk, T. W., 1879 
Kossmann, R., 1880 
Krauss, F., 1843 
Kunth, A., 1870 


Lamarck, J. B., 1818 
Lanchester, W. F., I90I 


Lanchester, W. F., 1903 


Lanchester, W. F., 1906 


S. KEMP: 


. “A Contribution to Indian Carcinology.’’ 


.. ““ Beiträge zur Kenntniss der Decapoden-Krebse Ost-Australiens.’’ 


.. ‘‘ Synopsis of Californian Stalk-Eyed Crustacea.’’ 


. ‘‘ Die Stomatopoden der deutschen Tiefsee-Expedition.’’ 


. ““ Additions to the Crustacean Fauna of New Zealand.’’ 


. ‘‘ Notes on some New Zealand Crustaceans.’’ 


. ““ Marine Crustaceans. 


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Crustacea Stomatopoda of the Indo-Pacific Region. 209 


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Trans. Linn. Soc. Zool., Ser. 
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. Versuch einer Naturges. d. Krabben und Krebse, Vol. II, pp. 93-103, pl. 


xxxiii-xxxiv (Berlin and Stralsund). 


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Arch. 


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Récherches sur 


Acad. Sci., Vol. VII, pp. 219-221 (San Francisco). 


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Wiss. Ergeb- 
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. “ Preliminary descriptions of new species and varieties of Crustacea Sto- 
y pP 


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100 (Calcutta). 
Ann. Mag. Nat. 


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.. Histoire Naturelle des Animaux sans Vertebres, Vol. V, pp. I86-189 (Paris). 
. ‘‘ On the Crustacea collected during the ‘ Skeat ’ Expedition to the Malay 


Proc. Zool. Soc., Vol. II, pp. 553-555 (London). 

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Fasciculi Malayenses, Zool., Pt. III, p. 133 


Peninsula.’’ 


(London). 


2IO 


Latreille, P. A., 1825 
Latreille, P. A., 1829 
Lenz, H., 1901 


Lenz, H., 1905 


Lenz, H., 1910 


Lenz, H. 
1881 


Linnaeus, C., 1767 
Linnaeus, C. A., 1790 
Lloyd, R. E., 1907 
Lloyd, R. E., 1908 


Lockington, W. N., 1877 


Martens, E. von., 1872 
Martens, E. von., 1881 


Miers, FE. J., 1875 


Miers, E. J., 1876 
Miers, E. J., 1877 
Miers, E. J., 1879 
Miers, E. J., 1880 (a) 
Miers, E. J., 1880 (b) 
Miers, E. J., 1881 (a) 


Miers, E. J., 1881 (b) 
Miers, E. 


J., 1884 (a) 


Miers, E. J., 1884 (0) 


. Encyclopédie Methodique. 


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und Richters, F., 


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. ‘* Remarkable Cases of Variation.’’ 


Memoirs of the Indian Museum. [Vor. IV, 


Entomologie, ou Histoire Naturelle des Crus- 
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ecexxiii—ceexxv (Paris). 


. Cuvier’s Régne Animal, 2nd Edit., Vol. IV, p. 109 (Paris). 
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Reise nach dem Pacific (Schaunisland 1896-1897). 


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.. systema Naturae, I2th Edit., Vol. I, Pt. II, p. 1054 (Holmiae). 
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Rec. Ind. Mus., Vol. II, pp. 29-35, 


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. ““ Remarks on the Crustacea of the Pacific Coast, with descriptions of some 


. ““ Ueber Cubanische Crustaceen.’’ 


. “ Squilliden aus dem Zoologischen Museum in Berlin.’’ 


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Arch. für Naturgesch., Vol. XXXVIII, 
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Sitz.-ber. Ges. 
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. ‘“On some new or undescribed species of Crustacea from the Samoa 


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. Catalogue of the Stalk- and Sessile-Eyed Crustacea of New Zealand, pp. 


88-90 (London). 


. “ On a collection of Crustacea made by the Rev. G. Brown on Duke of 


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. ““ Zoology of Rodriguez, Crustacea.’’ Phil. Trans. Roy. Soc., Vol. CLXVIII, 


PP. 494-495 (London). 

‘“ On the Squillidae’’ Ann. Mag. Nat. Hist., Ser. 5, Vol. V, pp. 1-30 
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““ On a Collection of Crustacea from the Malaysian Region, Stomatopoda.’’ 
Ann. Mag. Nat. Hist., Ser. 5, Vol. V, pp. 458-460 (London). 


. ‘* Account of the Crustacea collected during the survey of H.M.S. ‘ Alert ’ 


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. Report on the Zoological Collections made in the Indo-Pacific Ocean during 


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(London). 

Proc. Zool. Soc., pp. 16-17, 
pl. 1, figs. 3, 3a (London). 


1013.| S. KEMP : 
Milne-Edwards, A., 1862 
Milne-Edwards, A., 1868 
Milne-Edwards, A., 1869 


Milne-Edwards, A., 1873 


Milne-Edwards, A., 1877-78.. 


Milne-Edwards, A., 1891 


Milne-Edwards, H., 1836-40... 
. Histoire Naturelle des Crustacés, Vol. II, pp. 518-531 (Paris). 


Milne-Edwards, H., 1837 
Milne-Edwards, H., 1838 


Moreira, C., 1901 
Müller, F., 1886 
Nardo, G. D., 1869 
Nicolet, H., 1849 


Nobili, G., 1897 
Nobili, G., 1899 


Nobili, G., 1900 

Nobili, G., 1901 (a) 
Nobili, G., 1901 (b) 
Nobili, G., 1901 (c) 
Nobili, G., 1903 (a) 
Nobili, G., 1903 (b) 
Nobili, G., 1903 (c) 


Nobili, G., 1904 


Nobili, G., 1905 


. ‘‘ Faune Carcinologique de l’île de la Réunion.’’ 


. ““ Observations sur la Faune Carcinologique des Iles du Cap-Vert.’’ 


.. Mission Scientifique du Cap Horn. 1882-1883. 


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. ““ Decapodi e Stomatopodi Indo-Malesi.’’ 


. ““ Decapodi e Stomatopodi.’’ 


. ‘‘ Crostacei di Singapore.’’ 


. ““ Contributo all fauna Carcinologia di Borneo.’’ 


« ‘‘ Crostacei di Pondichéry, Mahé, Bombay, etc.’’ 


. ‘* Crostacei di Zanzibar.’’ 


Crustacea Stomatopoda of the Indo-Pacific Region. 2 


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Journ. Mus. Godefroy, Vol. I, 


Zool., Vol. VI, Crustaces, 
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Arch. Mus. Rio de Janeiro, Vol. XI, pp. 1-5 

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Mem. 

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. In Gay’s Historia fisica y politica de Chile, Zoologia, Vol. III, pp. 221-226, 


pl. ii (Paris). 


. ““ Decapodi e Stomatopodi raccolti dal Dr. Enrico Festa nel Darien, a 


Curacäo, La Guayra, Porto Cabello, Colon, Panama, ecc.’’ Boll. Mus. 


Torino, Vol. XII, No. 280, p. 6 (Turin). 


. ““ Contributioni alla conoscenza della fauna Carcinologica della Papuasia, 


delle Molucche e dell’ Australia.’’ 
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Ann. Mus. Civ. Genova, Ser. 2, Vol. 


Ann. Mus. Civ. Genova, Ser. 2, 
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Mus. Torino, Vol. XVI, No. 397, p. 14 (Turin). 
Boll. Mus. Torino, Vol. XVI, No. 415, 
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. “ Decapodi e Stomatopodi Eritrei del Museo Zoologico dell’ Universita di 


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Boll. Mus. Torino, Vol. XVIII, No. 455, pp. 
38-39 (Turin). 

Boll. Mus. Torino, Vol. 
XVIII, No. 447, pp. 28-32, text-fig. 3 (Turin). 

Boll. Mus. Torino, Vol. 
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212 


Nobili, G., 1906 (a) 


Nobili, G., 1906 (b) 


Nobili, G., 1907 


Ortmann, A., 1894 


Owen, R., 1832 
Owen, R., 1839 


Paulson, W., 1875 


Pfeffer, G., 1888 


Pocock, R. I., 1890 (a) 


Pocock, R. I., 1890 (b) 


Pocock, R. I., 1893 


Randall, J. W., 1839 


Rankin, W. M., 1899 
Rathbun, M. J., 1903 


Rathbun, M. J., 1910 


Richters, F., 1880 


Risso, A., I8I6 
Risso, A., 1826 


Roux, P., 1828 


Rumphius, G. E., 1705 
Sens dben TO 


Seba, A., 1758 


M emoirs of the Indian Museum. Wore 


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. “ Übersicht d. v. Herrn Dr. F. Stuhlmann in Egypten, auf Sansibar u. d. 


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.. In Mobius’ Beiträge zur Meeresfauna der Insel Mauritius und der Sey- 


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.. Histoire Naturelle des Crustacés, pp. 112-116, pl. ii, fig. 8 (Paris). 
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. Crustacés de la Méditerranée et de son littoral (not paged), pls. v, xxviii 


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1913.| S. KEMP: 
Schenkel, E., 1902 


Smith, S. I., 1881 


Stead, D. G., 1898 

Stebbing, T. R. R., 1902 
Stebbing, T. R. R., 1908 
Stebbing, T. R. R., 1910 


Steuer, A., IQII 


Stimpson, W., 1857 
*Stossich, M., 1880 


Tattersall, W. M., 1900 


Thallwitz, J., 1892 


Thomson, G. M., 1882 


Verrill, A. E., 1901-3 


Walker, A. O., 1887 
Wedenissow, T., 1894 


White, A., 1847 


Crustacea Stomatopoda of the Indo-Pacific Region. 213 


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LE] 


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White, A., 1849 


White, A., 1850 2 


White, A., 1851 an 


White, A., 1861 


Zool., Vol. XX, p. 113 (London). 


. ““ Di Alcuni Crostacei raccolti nel paese dei Somali dall’lug. I,. Bricchetti- 


Robecchi.’’ Bull. Soc. Ent. Ital., Vol. XXVI, p. 417 (Florence). 

List of the specimens of Crustacea in the collection of the British Museum, 
pp- 83-85 (London). 

‘‘ Description of two new species of Crustacea.’ 
pl. vi, figs. 1, 1 a (London). 


’ 


Proc. Zool. Soc., p. 144, 


. ““ Description of two new species of Crustacea.” Ann. Mag. Nat. Hist., 


Ser. 2, Vol. IV, pp. 381-382 (London). 

‘“ Descriptions of two species of Crustacea in the British Museum.’’ Proc. 
Zool. Soc., pp. 96-97, pl. xvi (London). 

‘ Descriptions of two species of Crustacea in the British Museum.’’ Ann. 
Mag. Nat. Hist., Ser. 2, Vol. VII, pp. 422-423 (London). 


. ‘‘ Descriptions of two species of Crustacea belonging to the Families Cal- 


lianassidae and Squillidae.’’ Proc. Zool. Soc., pp. 43-44, pl. vii; and 
Ann. Mag. Nat. Hist., Ser. 3, Vol. VII, pp. 480-481 (London). 


214 


Whitelegge, T., 1897 
Whitelegge, T., 1898 


Whitelegge, T., 1900 


Wood-Mason, J., 1875 


Wood-Mason, J., 1895 


Wood-Mason, J., and 
Alcock, A., 1891 


Woodward, H., 1879 


Zehntner, L., 1894 


. On some new species of Stomatepod Crustacea. 


Memoirs of the Indian Museum. Vor. INZ 19135] 


. ‘“The Crustacea of Funafuti’ Mem. Australian Mus., Vol. III, p. 148 


(Sydney). 


. ‘‘ Contributions to a knowledge of the Fauna of British New Guinea, 


Crustacea.’’ Proc. Linn. Soc. N.S.W., Vol. XXIII, p. 368 (Sydney). 


. “Scientific Results of the Trawling Expedition of H.M.C.S. ‘ Thetis,’ off the 


coast of New South Wales, in February and March, 1898. Crustacea, 
Pt. 1.” Mem. Australian Mus., Vol. IV, pp. 198-199 (Sydney). 

2” Proc. As. Soc. Bengal, pp. 
231-232 (Calcutta); reprintedin Ann. Mag. Nat. Hist., Ser. 4, Vol. XVII, 
p. 263, 1876 (London). 


. Figures and descriptions of nine species of Squillidae from the coilection 


in the Indian Museum, pp. 1-11, pl. i-iv (Calcutta). 


. ‘‘ Natural History Notes from H.M. Indian Marine Survey Steamer ‘ In- 


vestigator,’ No. 21. Note on the Results of the last season’s Deep-Sea 
Dredging.’’ Ann. Mag. Nat. Hist., Ser. 6, Vol. VII, pp. 271-272 
(London). 


. “ Contributions to the Knowledge of Fossil Crustacea.’’ Quart. Journ. 


Geol. Soc., Vol. XXXV, pp. 549-556, pl. xxvi (London). 


. “ Crustacés de l’Archipel Malais.” Rev. Swisse Zool., Vol. II, pp. 212-213 


(Geneva). 


~ Ne NN Nw er 


acanthocarpus 
acanthurus 

aculeata 

acutirostris 

aculus .. 00 
affınis (Gonodactylus) 
afınis (Squilla) 
africana 

alba 

anancyrus 
annandalei 

avenaria marina 
arenaria prona 
avenaria terrestris 
ayenarius ae 
armata (Lysiosquilla) 
armata (Squilla) 


biformis 
bigelowi 
biminiensis 
boops .. 
bleekeri 

bradyi 

brazieri 
brevirostris 
brevisquamatus 
brooksi 


Cancer 
capensis 
carinifer 
cerebralis 
cerisii .. 
chiragra 
chiragra 
chlorida 
Chloridella 


ciliata 


INDEX. 


Synonyms are printed in italics. 


Page 
4, II, 109, 120, 106. 
4, II, 146, 193. 
3, 6, 200. 
4, II, 150, 152, 163. 
152, 156, 162. 
156, 159. 
54, 57, 66, 70, 72, 76. 
89. 
3, 18, 200. 
151, 156, 150. 
3, 10, 19, 92. 
88, 90. 
135. 
TR 
TEE 
4, 202. 
3, 6, 8, 10, 18, 19, 41. 


3,6, 200. 

106. 

4, 8, II, 124. 

3, 10, 55. 

IB, 187 

4, 130, 204. 

128. 

4, IT, 143. 

4, II, 146, 174, 175. 
4, IT, 146, 189. 


61, 96, 135, 155. 

4, 10, 117. 

4, II, 138. 

186. 

3, 95, 202. 

4, 8,11, 145, 150, 155, 199. 

167, 169, 204. 

3, IO, IQ, 33. 

17, 24, 27, 28, 31, 33, 34. 
66, 84, 88, 200. 

3, 8, 10, 95, 96, 196. 


Clorida 
communis 
confinis 
Coronida 
Coronis 
costata 
crassispinosa 
cultrifer 
cylindrica 


decemspinosa 
decorata 
demani 


dentata 
depressa 
desaussurei 
desmaresti 
digitalis 
digueti 
distinguendus 
dofleini 
drepanophorus 
dubia 
dufresmii 


ectypus 


edwardsii (Gonodac 
edwardsvi (Squilla) .. 


elegans.. 
elongata 


tylu 


Page 
10 24 2737 
97: 
156, 158. 
4, 6, II, 16, 129, 204. 
109, 118, 120, 128, 204. 
3, 10, 18, IQ, 84. 
4, 10, 117. 
ile ait, NEY 
205. 


17208: 

A. Os, Bf 

4, II, 145, 150, 152, 164, 
198. 

46, 47. 

31, 32. 

4, 203. 

3, 6, 200. 

61. 

4, 203. 

102. 

3, 10, 104. 

A, Tit, 144), NZS}. 

3, 200. 

201. 


4, II, 185. 


s) 139. 


60. 
4, II, 139. 
180. 


empusa (Pseudosquilla) 3, 10, 95, 104. 


empusa (Squilla) 
ensiger 
espinosus 
eusebia 
excavatrix 
excavatus 


falcatus 
fasciata 


10h 200 

205. 

4, II, 153, 166. 
4, 203. 

4, IIO, 203. 

4, II, 146, 187. 


156, 204. 
35 7, 10, 17; IQ, 34. 


216 


ferussaci 
festae .. 
fimbriatus 
folini 

folinii 
foveolata 
furcicaudatus 


gibba 

gilesi 

glaber .. 

glaber .. 
glabriuscula 
glabrous .. 


glyptocercus 
Gonodactylus 


Gonodactylus 


gonypetes 
_ gracilipes 
graphurus 
graphurus 
guerini 


hanseni 
harpax 
havanensis 
hemischista 
herdmani 
hieroglyphica 
hoeveni 
holoschista 
hystrix 


ichneumon 
immaculata 
incipiens 
indefensa 
inornata 
insignis 
intermedia 
intermedia 
interrupta 
interrupta 
investigatoris 30 


japonicus 


laevis .. ae 
NGS oo xe 


Index. 
Page 
3, 16, 95, 202. EE 0 50 
4, 146, I49, 150, 153, 204. laticauda 
4, II, 146, 175. latifrons 
4, 146, 189. latirostris 
185. latreillei 
3, 10, 57, 58. leptodactyla 
4, II, 146, 176. leptosquilla 
Leptosquilla 
3. 10, 18, 28. p 7 
lessoni 
3, 6, 10, 19, 39. Ber 
lewisi 
4, II, 182. 
locusta 
167. 3 
tool 203 lutaria 06 
À 5 ; Lysierichthus 
4, II, I46, 150, 153, 167, Lysi ‘ll 
siosquilla 
197. ysiosq 
4, II, 186. 
4, 6,7, 9, 11, 16, 145, 197, maculata 
204, 205. ; ‘ 
Rue 3% maiaguesensis 
San MO e839 ES UI) WA major .. 
as) AG mantis 
3, 10, 54. 
en mantıs 
3 - 4 i ae mantoidea 
a DU) Mas) SO SEE : marmorata 
167. 2 
massavensis 
4, II, 192. ay. 
mauritiana 50 
4, II, 140. megalophthalma .. 
88. microphthalma .. 
4, 6, 134, 144. miersi 
61. | miles 
dy Tt, SG, HY minor: 
Be 16h Bille “monodactyla ? 
204. MONOCCYOS 
3, 7,10, 61, 64, 195. multicarinata 
4, II, 190. multifasciata 
multituberculata 
205. 
3, 6, 10, 46. mutatus 
151, 156, 159. Necroscilla Br 
118, 119. nefandus 00 
AO neglecta 
A, 6, 11,109, 126. nepa 
Dy PO nepa 
58. 
3, 10, 58, 66, 70, 72. occidentalis 
70. occulta © 
By) Oh Fy Oo 180) 80: oculata 
4, ax, 139) oculata 
Odontodactylus 
3, 10, 18, IQ, 49. Odontodactylus es 
118, 119. oerstedi 30 


Page 
3, 10, 17, IQ, 37. 
17, 200. 
A, 36 ts WOT, 
4, II, 141. 
3, 10, 24. 
205. 
3, 6, 10, 18, IQ, 46. 
17, 93- 
3, 95, 202. 
15. 
III. 
135. 
II6. 
4, 6, 7, 10, 16, 109, 196, 
202. 


4, 8, 10, 109, 111. 
4, 6, 203. 

88. 

3, 6, 19, 88, 201. 
61,90, TEL, 135; 155: 
3, 10, 18, IQ, 45. 
202. 

3, 6, 10, 57, 58, 66, 76. 
66, 68. 

3, 10, 103. 

3, 10, 31. 

4, 10, 116. 

3, 10, 36. 

40. 

108, 

202. 

3, 10, 19, 86, 196. 

Jil, Wit SCO), 122; 

4, II, 130, 132. 

167. 


16. 
4, II, 179. 
3, 201. 


3, 10, 19, 57, 60, 76, 195. 
64, 66, 70. 


97; 99. 

4, 203. 

3, 8, 10, 95, 102. 

100. 

4, 6, 7, II, 16, 133, 204. 
130. 

4, 8, 146, 152, 161, 204. 


oratoria > 
oratoria 

ornata 

ovatoria 

oxyrhyncha 


pacificus 

pallida 

panamensis 

parva 

pennata 

perpensa 
phalangium 
pilaensis — 
platysoma 
plumata 

polita 

polita 

polydactyla 
prasinolineata, Dana 
prasinolineata, Ives 
Protosquilla 
Pseuderichthus 
Pseudosquilla 


Pierygosquilla 
pulchellus 


quadraticauda 
quadridens 
quadrispinosa 
quinquedentata 


raphidea 
rotundicauda 
votundus 
rubrolineata 

rugosa I“ 


sarasinorum 00 
scabricauda a0 
scabricauda 
“schmeltzii ’ 
scolopendra 

scorpio 

Sculda 


Page 
3, 10, 66, 196. 
72. 
2 10, 100% 
66. 
3, Io, 101. 


fly Gp Ui, HAA. 
Sn AON. 

35 AO 

3, 201. 

15. 


3, 10, 58, 66, 70, 196. 


204. 

3, IO 105. 

4, II, 152, 161, 162. 
4, 203. 

3, 201. 

76. 

4,203, 

3, 201. 

8 AOL 


5, 145, 176, 177, 180-192. 


97, 102. 


3 10 7 TO TOs G44 OS, 


196, 202, 205. 
17, 200. 
Jy, Son type 


55» 57: 

3, HOW 

97- 

3, 10, IQ, 52, 195. 


3, 10, 19, 88. 
3. 10,133. 


93. 

4, 204. 

Bin UO, TO, WO), PAS 
5, I5. 


Index. 


Sculdidae 

scyllarus 

segregatus 

septemspinosa 

smithi 

southwelli 

spinifrons 

spinosa (Lysiosquilla) 

spinosissimus 

spinosocarinatus 

spinosus (Gonodacty- 
lus) 

spinosus (Gonodacty- 
lus) 

Squilla 


Squilla 


Squillidae 
stoliurus 
stridulans 
stylifera 
stylifera 
sulcirostris 
supplex 


tanensis 
tenuispinis 
ternatensis 
tigrina 
trachura 
tricarinata 
tridentata 
trispinosus 
trispinosus 
tuberosus 
tumidus 


valdiviensis 
vicina 
vitrea 
vittata 


wilsoni 
wood-masoni 


Page 
15. 
4, TT, 135. 
156. 
4, 122, 204. 
156, 159, 160. 
Bb ib, EVA 142: 
202. 
4, II, 109, 118. 
5, II, 146, 191. 
Bo eit, IMO, WB, 


164. 


4, II, 150, 152, 165. 


217 


3 Da Wo 9, IO, 15, 16, 195, 


200-205. 


94, 96, 102, IO4, 109, 135, 


155, 202. 

15. 

5, II, 146, 184. 

3, 6, 10, 57, 66, 78. 
3, 6, 8, I0, 95, 106. 
97- 

4, 10, 116. 

3, 10, 19, 82. 


5, II, 183. 

3p Oy Wey Bish, 10), ZU 
167. 

4, II, 125. 

4, II, 130. 

118. 

118, IIQ. 

5, LL, £460, 180. 
177, 184. 

5, II, 181. 

I5I, I56, 159. 


122, 123: 
4, II, 126. 
205. 

203. 

16. 


3, 10, 58, 66, 76. 


er 


he A N 

en ‘+ > 
ne > 

2 e 


EXPLANATION OF PLATE I. 


i Squilla latreillei (Eydoux and Souleyet). 
Fic. 1.—Jast two abdominal somites, telson and right uropods of a female specimen from the Persian 
Gulf : dorsal view. 
5, 2.—Last two abdominal somites, telson and left uropods of a male specimen from the Persian 
Gulf: dorsal view. 
»» 3.—Right eye in dorsal view. 
»» 4-—Left half of last four thoracic somites in dorsal view. 


Squilla gibba, Nobili. 
»» 5.—Anterior half of the type specimen, a male, in dorsal view. 
», 6.—Right eye in dorsal view. 
»»  7-—Antero-lateral angle of carapace in lateral view. 
,, 8.—Raptorial claw of male. 
»»  9.—Raptorial claw of female. 
,, 10.—Fifth, sixth and seventh thoracic somites in lateral view. 
», 11I.—lLast two abdominal somites, telson and uropods of the type specimen : dorsal view. 
», 12.—I,ast abdominal somite and telson of female in dorsal view. 


Squilla decorata (Wood-Mason). 


>, 13.—Right eye in dorsal view. 

», 14.—Left half of last four thoracic somites in dorsal view. 

», 15.—Last two abdominal somites, telson and right uropods of a small female from the Vé River en- 
trance : dorsal view. 

;, 16.—The same in ventral view. 


Squilla microphthalma, H. Milne-Edwards. 
,, 17.—Right eye in dorsal view. 
,, 18.—Leit half of last four thoracic somites in dorsal view. 


», 19.—Last two abdominal somites, telson and uropods of a female from the Madras Coast. 
,, 20.—The same parts of a female from Karachi. 


Squilla fasciata, De Haan. 
,, 21.—Anterior half of the female specimen from the Andamans : dorsal view. 
5, 22.—Raptorial claw of the same specimen. 


», 23.— last two abdominal somites and telson of the same specimen : dorsal view. 


ZOOLOGY OF THE R. I. M. 8. ‘INVESTIGATOR.’ ee Te 


Photo. Engraved & printed at the Offices of the Survey of India, Calcutta, 1912 


Figs. 1-4 Squilla latreillei. Figs. 5-12 Squilla eibba. Figs. 13-16 Squilla decorata, 
Figs. 17-20 Squilla mierophthalma. Figs. 21-23 Squilla fasciata. 


=. 


Rx 


wears 
ALERTE 


x 


x 


& 


EXPLANATION OF PLATE II. 


Squilla lata, Brooks. 


. 24.—Basal process of right uropod of a specimen from the Gulf of Martaban : ventral view. 


Squilla gilest, Wood-Mason MS. 
25.—Basal process of left uropod in ventral view. 
26.—Anterior half of one of the type specimens in dorsal view. 
27.—Last abdominal somite, telson and left uropods of the male type specimen : dorsal view. 
Squilla armata, H. Milne-Edwards. 

28.—Anterior half of a female from New Zealand : dorsal view. 
29.—Raptorial claw of the same specimen. . 

Squilla scorpio, Latreille. 


30.—Rostrum, carapace and fifth abdominal somite in dorsal view. 


Squilla scorpio var. immaculata, Wood-Mason, MS. 


31.—Anterior half of one of the type specimens in dorsal view. 


ZOOLOGYZORZENEIBER ZTESMESIS: ‘INVESTIGATOR,’ _ CRUSTACEA STOMATOPODA, PLATE II. 


Tr 
Lo wae 
Q eS Ne 
CR —— as LTT 2 
GS 2 > 
TE PA G 4 f Z > GS IR > 
z S : 6 / ee OS LA 


STERN ern 


24x4. 254. 


Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta, 1912. 
S. C. Mondul, del. 
Fig. 24 Squilla lata. Figs. 25-27 Squilla gilesi. Figs. 28-29 Squilla armata. 
Fig. 30 Squilla scorpio. Fig. 31 Squilla scorpio var. immaculata. 


EXPLANATION OF PLATE III. 


Squilla tenwispinis, Wood-Mason. 
Fic. 32.—-Anterior half of a male from the Ganjam Coast in dorsal view. 
„» 33. —Last two abdominal somites, telson and left uropods of the same specimen: dorsal view. The 
gure is erroneous in showing intermediate carinae on the carapace on either side of the gastric 
grooves. 
34.—Telson of the same specimen in lateral view. 


Squilla laevis, Hess. 


»> 35.—Anterior half of a male in dorsal view. 
36.—Raptorial claw of the same specimen. 
37-—Last four thoracic somites in lateral view. 


Squilla hieroglyphica, Kemp. 
38.—Anterior half of the type specimen in dorsal view. 
„» 39.—Raptorial claw. 
,, 40.—Fifth, sixth and seventh thoracic somites in lateral view. 
41.—Last two abdominal somites and telson in dorsal view. 


ZOOLOGY OF THE R. I. M. S. ‘INVESTIGATOR,’ Crustacea STOMATOPODA, PLATE III. 


ERIEITTIRTE 
rer 


0x5. 


34 x5. 


37x14. 


Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta. 1912. 
S. C. Mondul, del. 
Figs. 32-34 Squilla tenuispinis. Figs. 35-37 Squilla laevis. Figs. 38-41 Squilla hieroglyphica. 


EXPLANATION OF PLATE IV. 


Squilla gonypetes, Wood-Mason, MS. 
Fic. 42.—Anterior half of the male type specimen in dorsal view. 
»» 43.—Raptorial claw of the same specimen. 
44.—Last two abdominal somites, telson and uropods of the same specimen: dorsal view. 
Squilla boops, Kemp. 


45.—Anterior half of the type specimen in dorsal view. 
46.—Raptorial claw of the same specimen. 
47.—Last two abdominal somites, telson and uropods of the same specimen : dorsal view. 


Squilla foveolata, Wood-Mason. 


48.—Right half of the last four thoracic somites in dorsal view. 


Squilla nepa, Latreille. 


», 49.—Median carina of carapace anterior to cervical groove. 


Squilla holoschista, Wood-Mason, MS. 


,» 50.—Anterior half of one of the type specimens in dorsal view. 
», 51.—Median carina of carapace anterior to cervical groove. 

1, 52-—Raptorial claw of male. 

»> 53-—Raptorial claw of female. 


ZOOLOGY OF THE R. I. M. S ‘INVESTIGATOR.’ Crusracea Stomatopopa, PLATE IV. 


£ 


s Hr! A 
44-3. HARAS" 
Photo -Engraved & printed at the Offices of the Survey of India, Calcutta, 1912 


Figs. 42-44 Squilla gonypetes. Figs. 45-47 Squilla boops. Fig. 48 Squilla foveolata. 
Fig. 49 Squilla nepa. Figs. 50-53 Squilla holoschista. 


EXPLANATION.OF PLATE V. 


Squilla oratoria, De Haan. 


. 54.—Anterior half in dorsal view. 
55.—Carpus of raptorial claw ; the dorsal tubercles are not quite sharp enough. 


56.—Left half of last four thoracic somites in a specimen from Amoy. 


Squilla oratoria var. perpensa, Kemp. 


57.—Anterior half of one of the type specimens in dorsal view. 
58.—Carpus of raptorial claw. 
59.—Basal process of uropod, left side in ventral view. 


Squilla interrupta, Wood-Mason, MS. 


60.—Anterior half of one of the type specimens in dorsal view. 
61.—Carpus of raptorial claw. 
62.—Basal process of uropod, left side in ventral view. 


Squilla wood-masoni, Kemp. 


63.—Anterior half of one of the type specimens in dorsal view. 
64.—Raptorial claw. 
65.—Basal process of uropod, left side in ventral view. 


Squilla stridulans, Wood-Mason. 


66.—Right half of fifth, sixth and seventh thoracic somites in dorsal view. 


u ZOOLOGY OF THE R. I. M. S, ‘INVESTIGATOR.’ CRUSTACEA STOMATOPODA, PLATE V. 
+ 


63 xR. 


Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta, 1912 


S. €. Mondul, del. 


Figs. 54-56 Squilla oratoria. Figs. 57-59 Squilla oratoria var. perpensa. 


a 


Figs. 60-62 Squilla interrupta. 


2 


EXPLANATION OF PLATE VI. 
Squilla investigatoris, Lloyd. 


. 67.—One of the type specimens, natural size. 


68.—Propodus and dactylus of raptorial claw. 


Squilla supplex, Wood-Mason. 


69.—Last two abdominal somites, telson and left uropods of the male from Tuticorin : dorsal view. 


Squilla costata, De Haan. 
79. —Anterior half of the specimen from Burma in dorsal view. 
71.—Raptorial claw of the same specimen. 
72.—Last three abdominal somites, telson and left uropods of the same specimen : dorsal view. 


Squilla multicarinata, White. 
73.—Anterior half of a specimen from Hongkong in dorsal view. 
74.—Raptorial claw. 
75.—Fifth, sixth and seventh thoracic somites in lateral view. 
76.—Last two abdominal somites, telson and uropods in dorsal view. 


ZOOLOGY OF THE R. I. M. S. ‘INVESTIGATOR.’ Crustacea STOMATOPODA, PLATE VI. 


S. C. Mondul, del. Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta, 1912. 


Figs. 67-68 Squilla investigatoris. Fig. 69 Squilla supplex. Figs. 70-72 Squilla costata. 


Fic. 


EXPLANATION OF PLATE VII. 


Squilla raphidea, Fabricius. 


77.—Left half of fifth, sixth and seventh thoracic somites in dorsal view. 


Squilla annandalei, Kemp. 
78.—Anterior part of carapace, rostrum, eystalks, etc., in dorsal view. 
79.—Outline of a specimen showing the characteristic pigmentation of the abdomen, telson and ulti- 
mate segment of outer uropods. 
80.—Left half of fifth, sixth and seventh thoracic somites : dorsal view. 


““ Leptosquilla schmeltzii ’’ (A. Milne-Edwards). 


81.—Anterior half of the specimen from the Andamans : dorsal view. 
82.—Raptorial claw of the same specimen. 
83.—Last two abdominal somites, telson and right uropods of the same specimen : dorsal view. 


Pseudosquilla stylifera (H. Milne-Edwards). 


84.—Anterior part of carapace, rostrum, eyes, antenna! scales, etc., of a specimen from Chili. 
85.—Last abdominal somite, telson and right uropods of the same specimen. 


ZOOLOGY OF THE R. I. M. S. ‘INVESTIGATOR.’ CRUSTACBA STOMATOPODA, PLATE WALL, 


pes 


19X4. 


| ü 83 x16. 


Photo.-Engraved & printed at the Offices of the Survey of India, Caleutta, 1912 


S.C, Mondul, del. 


Fig. 77 Squilla raphidea. Figs. 78-80 Squilla annandalei. Figs. 81-83 ‘Leptosquilla schmeltzii.’ 
Figs. 84-85 Pseudosquilla stylifera. 


EXPLANATION OF PLATE VIII. 


Lysiosquilla maculata (Fabricius). 


(G. 86.—Rostrum of a typical specimen. 


87.—Sketch of a large female, showing distribution of blue-black pigment. 
88.—Sketch of a male, showing distribution of pigment. 

89.—Telson of another male showing different pigmentation. 
90.—Raptorial claw of large female. 

91.—Raptorial claw of large male. 


Lysiosquilla maculata var. sulcirostris, Kemp. 
92.—Rostrum. 
93.—Raptorial claw. 
Lysiosquilla spinosa, Wood-Mason. 


94.—Left uropod of the type specimen, seen from beneath. 


Lysiosquilla tigrina, Nobili. 


95.—Carapace and anterior appendages of the type specimen. 
96.—Raptorial claw of the same specimen. 
97.—Last abdominal somite and telson of the same specimen : dorsal view. 


08. 


Last two abdominal somites, telson and uropods of the same specimen : 


ventral view. 


ZOOLOGY OF THE R. I. M. S. ‘INVESTIGATOR.’ Crustacea Sromaropopa, Pcare VIII. 


I6xS. 


98x5. 


Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta, 1912 


| 8. C. Mondul, del. 
| Figs. 86-91 Lysiosquilla maculata. Figs. 92-93 Lysiosquilla maculata var. sulcirostris. 


Fig. 94 Lysiosquilla spinosa. Figs. 95-98 Lysiosquilla tigrina. 


ee = 


2 
Spe 
es 
N 
; = 
x 
5 + 
‘ S 
v . 
\ 
t 
1 3 
# 
De 
v \ 
N ’ 1 
"LE 
Ÿ 
à = by id I * 


EXPLANATION OF PLATE IX. 


Lysiosquilla insignis, Kemp. 
Fic. 99.—Anterior half of the type specimen in dorsal view. 
,, 100.—Raptorial claw of the same specimen. 


,, 101.— Last two abdominal somites, telson and uropods of the same specimen : dorsal view. 
,, 102.—Telson viewed from beneath a little obliquely. 


Odontodactylus southwelli, Kemp. 


,, 103.—Carapace and anterior appendages of the type specimen in dorsal view. 

,, 104.—Raptorial claw of the same example. 

,, 105.—Fifth, sixth and seventh thoracic somites in lateral view. 

,, 106.—Last two abdominal somites, telson and uropods of the same specimen : dorsal view. 


Gonodactylus chiragra (Fabricius). 


», 107.—Anterior part of a specimen with rostrum removed to show the form of the dorsal processes of 
the ophthalmic somite. 


Gonodactylus demani, Henderson. 


,, 108.—Anterior part of a specimen with rostrum removed to show the form of the dorsal processes of the 
ophthalmic somite. 


aoe | Tast abdominal somite and telson of three individuals showing the variation in spinulation and 
III. ) in the form of the three median carinae : dorsal views. 


Gonodactylus demani var. spinosus, Bigelow. 


,, 112.—last abdominal somite and telson of a specimen from Ceylon which is somewhat intermediate 
between the variety and the typical form: dorsal view. 


Gonodactylus glabrous, Brooks. 


,, 113.—Anterior part of a specimen with rostrum removed to show the form of the dorsal processes of the 
ophthalmic somite. 


ZOOLOGY OF THE R, I, M. S. ‘INVESTIGATOR,’ Crustacea STOMATOPODA, PLATE IX, 


Photo.-Enéraved & printed at the Offices of the Survey of India, Calcutta, 1912. 


, Mondul, del. 
Figs. 99-102 Lysiosquilla insignis. Figs, 103-106 Odontodactylus southwelli, Fig. 107 Gonodactylus chiragra. 


Figs. 108-111 Gonodactylus demani, Fig. 112 Gonodactylus demani var. spinosus. 
Fig. 113 Gonodactylus glabrous. 


EXPLANATION OF PLATE X. 


Gonodactylus herdmani, ‘Tattersall. 
Fic. 114.—Last abdominal somite, telson and right uropods in dorsal view. 
,, 114a.—Right inner uropod in dorsal view. 
Gonodactylus brevisquamatus, Paulson. 


», 115.—Last two abdominal somites, telson and uropods in dorsal view (after Paulson). 
», 116.—Last abdominal somite, telson and left uropods viewed obliquely (after Paulson). 


Gonodactylus pulchellus, Miers. 
», 117.—Anterior part of carapace, rostrum, eyes, etc., in dorsal view. 
», 118.—Last two abdominal somites, telson and left uropods in dorsal view. 
Gonodactylus nefandus, Kemp. 
119.—Anterior part of carapace, rostrum, eyes, etc., in dorsal view. 
120.—Last two abdominal somites, telson and right uropods in dorsal view. 
Gonodactylus glaber (Lenz). 


», 121.—Last two abdominal somites, telson and right uropods in dorsal view. 


Gonodactylus excavatus, Miers. 
122.—Anterior half of a specimen in dorsal view. 
123.—Last two abdominal somites, telson and left uropods in dorsal view. 
Gonodactylus spinosissimus, Pfeffer. 


124.—Anterior half of a specimen in dorsal view. 
125.—l,ast two abdominal somites, telson and left uropods in dorsal view. 


ZOOLOGY OF THE R. I.M. S. ‘INVESTIGATOR.’ | | si Sion Pie 


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Photo.-Engraved & printed at the Offices of the Survey of India, Calcutta, 1912. 


Ss. C. Mondul, del. 
Fig. 114 Gonodactylus herdmani. Figs, 115-116 Gonodactylus brevisquamatus, Figs. 117-118 Gonodactylus pulchellus. 
Figs. 119-120 Gonodactylus nefandus. Fig. 121 Gonodactylus glaber. Figs. 122-123 Gonodactylus excavatus. 


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RECORDS 
of tbe 


INDIAN MUSEUM 


(A JOURNAL OF INDIAN ZOOLOGY) 


Vol. I, 1907. 


Part I.—Contributions to the Fauna of the Arabian Sea. Records of Hemiptera and Hymenoptera from the Himalayas. 
Further notes on Indian Freshwater Entomostraca. The Fauna of Brackish Ponds at Port Canning, Lower 
Bengal, I—III. A Sporozoon from the Heart of a Cow. Miscellanea. 


Part II.—Revision of the Oriental Stratiomyidæ. Description of an Oligochæte Worm allied to Chætogaster. The 
Fauna of Brackish Ponds at Port Canning, Lower Bengal, IV. Further Note on a Polyzoon from the Himalayas. 
Reports on a collection of Batrachia, Reptiles and Fish from Nepal and the Western Himalayas. The Fauna of 
Brackish Ponds at Port Canning, Lower Bengal, V. Notes on Oriental Diptera, I and II. Miscellanea. 


Part III.—Report on the Marine Polyzoa in the collection of the Indian Museum. The Fauna of Brackish Ponds at 
Port Canning, Lower Bengal, VI. A third note on Earwigs (Dermaptera) in the Indian Museum, with the descrip- 
tion of a new species. Notes on Oriental Diptera, III. Description of a new snake from Nepal. Notes on a col- 
lection of marketable fish from Akyab, with a description of a new species of Laciarius. Description of two fresh- 
water Oligochæte Worms from the Punjab. Notes on Phosphorescence in Marine Animals. Notes on the rats of 
Dacca, Eastern Bengal. Notes on Freshwater Sponges, I—V. Miscellanea. 


Part IV.—Nudiclava monocanthi, the type of a new genus of Hydroids parasitic on Fish. Preliminary descriptions 
of three new Nycteribiide from India. Annotated Catalogue of Oriental Culicidæ. Notes on Oriental Diptera. 
Notes on Freshwater Sponges, VI, VII. Description of a new Cyprinid Fish of the genu Danio from Upper 
Burma. Miscellanea. 


Vol. II, 1908-1909. 


Part I.—The retirement of Lieut.-Col. Alcock, with a list of his papers on Indian Zoology. The Fauna of Brackish 
Ponds at Port Canning, Lower Bengal, VII. Description of a New Dictyonine Sponge from the Indian Ocean. 
Notes on Freshwater Sponges, VIII. Remarkable cases of variation, I. Description of a new species of Lizard 
of the genus Salea, from Assam. The Fauna of Brackish Ponds at Port Canning, Lower Bengal, VIII. Des- 
cription of a new Cavernicolous Phasgonurid from Lower Siam. Descriptions of new species of Marine and 
Freshwater Shells in the collection of the Indian Museum. Notes on Oriental Syrphide, I. Description of a new 
variety of Spongilla loricata. Notes on Oriental Diptera, V. Miscellanea. 


Part II.—Gordiens du Musée Indien. The Fauna of Brackish Ponds at Port Canning, Lower Bengal, IX. Description 
of a new species of Danio from Lower Burma. Rhynchota Malayana, I. Cimex votundatus, Signoret. Notes on 
Freshwater Sponges, IX. Fruit Bats of the genus Pieropus inhabiting the Andaman and Nicobar Archipelagos. 
A new species of Sun-Bird obtained near Darjiling. Three Indian Phylactolemata. On two new species of Eagle- 
Rays (Myliobatide). Description of a new species of the genus Sesarma, Say., from the Andaman Islands. Des- 
criptions of new species of Land, Marine, and Freshwater Shells from the Andaman Islands. 


Part III.—The Fauna of Brackish Ponds at Port Canning, Lower Bengal, X, XI. On some Oriental Solifuge with 
descriptions of new forms. The difference between the Takin (Budorcas) from the Mishmi Hills and that from 
Tibet, with notes on variation displayed by the former. On Caridina nilotica (Roux) and its varieties. Description 
of a new species of Charaxes from the Bhutan Frontier. First Report on the Collection of Culicidæ and Corethridæ 
in the Indian Museum, with descriptions of new genera and species. Miscellanea. 


Part IV.—Report on a collection of aquatic animals made in Tibet by Capt. F. H. Stewart in 1907, I. Notes on 
Aculeate Hymenoptera in the Indian Museum, I. Indian Psychodidæ. Description of a new species of mouse from 
the Madura District, Madras. Some Cleridæ of the Indian Museum. The Fauna of Brackish Ponds at Port Can- 
ning, Lower Bengal, XII. Description of a new species of Saw-Fish captured off the Burma Coast. A new Sting 
Ray of the genus Trygon from the Bay of Bengal. New Micro-lepidoptera from India and Burma. Notes on some 
Chrysomelid Beetles in the collection of the Indian Museum. Six new Cicindelinæ from the Oriental Region. Des- 
cription of a new slug from Tibet. 


Part V.—Revision of the Oriental Leptide. Revised and annotated Catalogue of Oriental Bombylidæ, with descriptions 
of new species. 


Vol. III, 1909. 
Part I.—The Races of Indian Rats. 


Part II.—Notes on Freshwater Sponges, X. Report on a collection of aquatic animals made in Tibet by Capt. F. H. 
Stewart in 1907, II. Note on some amphibious Cockroaches. Description de quelques nouvelles Cécidomyies 
des Indes. Description of new land and marine shells from Ceylon and S. India. Description of two new species 
of Caranx irom the Bay of Bengal. Remarks on some little known Indian Ophidia. Remarks on some forms of 
Dipsadomorphus. A pelagic Sea-Anemone without tentacles. Rhynchota Malayana, II. 


Part III.—Notes on the Neuroptera in the collection of the Indian Museum. New Indian Leptide and Bombylidæ, 
with a note on Comastes, Os. Sac., v. Heterostylum, Macq. Notes on the Trichoptera in the collection of the Indian 
Museum. Diagnoses of new species and varieties of Freshwater Crabs. 1—3. Report on a small collection of 
Lizards from Travancore. Descriptions of three new Cicindelinæ from Borneo. The relation between fertility and 
normality in Rats. Descriptions of a Barnacle of the genus Scalpellum from Malaysia. The Hemipterous family 
Polyctenide. Notes on Freshwater Sponges, XI. Descriptions of two new shells from S. India. Preliminary note 
on a new genus of Phylactolematous Polyzoa. Miscellanea. 


Part IV.—Description of a minute Hymenopterous insect from Calcutta. The Insect Fauna of Tirhut, No. 1. Descrip- 
tions of new species of Botia and Nemachilus. New Oriental Sepsinæ. A new species of Fredericella from Indian 


lakes. Diagnoses of new species and varieties of freshwater crabs, No. 4. On some new or little-known Mygalo- 
morph spiders from the Oriental region and Australasia. 


Vol. IV, 1910. 


No. I.—Second report on the collection of Culicidæ in the Indian Museum, with descriptions of new genera and species. 

Nos. II and III.—The Indian species of Papataci Fly (Phlebotomus). Taxonomic values in Culicidæ. 

Nos. IV.—Revision of the Oriental blood-sucking Muscide (Stomoxine, Philematomyia, Aust., and Pristirhynchömyia, 
gen. nov.). 

No. V.—A new arrangement of the Indian Anopheline. 

No. VI.—A revision of the species of Tabanus from the Oriental Region, including notes on species from surrounding 
countries. 

No. VII.—New Oriental Nemocera. Miscellanea :—Synonymy in Corethrinae. Indian Phlebotomi. 


Nos. VIII and IX.—A revision of the Oriental species of the genera of the family Tabanidae other than Tabanus. Con- 
tributions to the fauna of Yunnan, Part VII. 


No. X.—Annotated Catalogue of Oriental Culicidae— Supplement. 


Vol. V, 1910. 


Part I.—The Hydroids of the Indian Museum, I. Notes on Freshwater Sponges, XII. Descriptions of new Shells in 
the collection of the Indian Museum from Burma, Siam and the Bay of Bengal. Materials for a revision of the 
Phylactolematous Polyzoa of India. Studies on the aquatic Oligochæta of the Punjab. An undescribed Burmese 
Frog allied to Rana tigrina. Miscellanea. 

Part II.—Description d’Ophiures nouvelles provenant des dernières campagnes de ‘‘ I’Investigator ’’ dans l'Océan 
Indien. Description d’Holothuries nouvelles appartenant au Musee Indien. The races of Indian rats, II. Descrip- 
tion of a new species of Scalpellum from the Andaman Sea. Descriptions of five new species of marine shells from 
the Bay of Bengal. Notes on fish from India and Persia, with descriptions of new species. 


Part III.—A new genus of Psychodid Diptera from the Himalayas and Travancore. The Indian barnacles of the sub- 
genus Smilium, with remarks on the classification of the genus Scalpellum. On a subspecies of Scutigerella unguicu- 
lata, Hansen, found in Calcutta. The distribution of the Oriental Scolopendride. Notes on Decapoda in the Indian 
Museum, I. Description of a new species of Nemachilus from Northern India. Notes on the larvæ of Toxorhynchites 
immisericors, Wlk. Description of a South India frog allied to Rana corrugata of Ceylon. Contributions to the 
fauna of Yunnan, Introduction and Part I. Miscellanea. 

Part IV.—Notes and descriptions of Indian Microlepidoptera. On some aquatic oligochæte worms commensal in 
Spongilla carteri. On Bothrioneurum wis, Beddard. Notes on nudibranchs from the Indian Museum. On the 
classification of the Potamonidæ (Telphusidae). Catalogue of the pheasants, pea fowl, jungle fowl and spur fowl 
in the Indian Museum. On certain species of Palaemon from South India. Alluaudella himalayensis, a new species 
of degenerate (7) cockroach, with an account of the venation found in the genera Cardax and Alluaudella.- 
Rhynchota Malayana, III. 


Vol. VI, 1911. 


Part I.—A Rhizocephalous Crustacean from fresh water and on some specimens of the order from Indian seas. Deca- 
poda in the Indian Museum, II. Contributions to the fauna of Yunnan, Parts II to V. Pedipalpi in the Indian 
Museum, I and II. Six new species of shells from Bengal and Madras. Miscellanea :—Fleas from India and China. 
Flies found associated with cattle in the neighbourhood of Calcutta. Mosquito sucked by a midge. Large egg laid 
by a beetle. 

Part II.—Some sponges associated with gregarious molluscs of the family Vermetidae. Aquatic animals from Tibet, 
III. Cyprinidae from Tibet and the Chumbi Valiey, with a description of a new species of Gymnocypris. New 
species and varieties of Crustacea Stomatopoda in the Indian Museum. The development of some Indian Ascala- 
phidae and Myrmeleonidae. Miscellanea :—The occurrence of Dactylopius citri, Risso, in the Himalayas. Note on 
Aquatic Rhynchota. 

Part III.—Nouveaux Chironomides de 1’ Indian Museum de Calcutta. 

Part IV.—Indian Isopods. Systematic notes on the Ctenostomatous Polyzoa of fresh water. Some aquatic Oligochaeta. 
in the Indian Museum. Contributions to the fauna of Yunnan, Part VI. Asiatic species of Crustacea Anostraca 
in the Indian Museum. Freshwater sponges, XIII. Miscellanea :—Synonymy in Corethrinae. The distribution 
of the different forms of the genus /bla. 

Payi V.—Revision of the Oriental Tipulidae. Five new Indian species of Clavicorn Coleoptera. Les Chironomides 
(Tendipedidae) de l’Himalaya et d’Assam. The occurrence of Apus in Eastern Asia. 


Vol. VII, 1912. 


Part I.—Deep-sea Fish obtained by the R.I.M.S.S. ‘‘Investigator’’ during 1910-11. Decapoda in the Indian Museum, 
III. Fauna of Paresnath Hill. Observations on the shallow-water Fauna of the Bay of Bengal made on the 
“ Golden Crown,’’ 1908-09. Coccidae in the Indian Museum, I On the nervous system of Ampullaria globosa. 
Description of a species of Phoridae that causes Myasis in man. Mzscellanea :—Mimicry of a Mutiliid by a spider. 
Capture of Limulus on the surface. Development of the larva of Lingula. Distribution of some Indian and 
Burmese Lizards. 

Part II.—The Mosquitoes of Calcutta. Freshwater sponges, XIV. Pedipalpi in the Indian Museum collection, III and 
IV. New Freshwater Crab from S. India. Decapoda in the Indian Museum, IV. Fauna Symbiotica Indica, I 
and II. Inverteb.ate Fauna oi the Kumaon Lakes. Fauna Symbiotica Indica, III. The Indian Mud-Turtles. 
Anatomy of Atopos sanguinolenia. Cestodes from Indian Fishes. Miscellanea :—Entoprocta in Indian Waters. 
Rearing of Leeches in bara Banki District. Habits of Tiger-Beetles. Schizodactylus monsirosus as bait for birds. 
Macrones menoda var. trachacanthus. Aquatic Tortoises of the Ganges and Brahmaputra. 

Part III.—Gordiens du Musée Indien. A new Tortoise from South India. New species of Branchiodrilus and other 
aquatic Oligochaeta, with remarks on cephalization in the Naididae. Fauna Symbiotica Indica, IV. Freshwater 
Medusa from the Bombay Presidency. A new species of Indian Thrips. Aquatic Chelonia of the Mahanaddi. 
Recent Crinoids from the Indian Ocean. Fauna of Yunnan, VII{. Earthworms. Asiatic Naiades in the Indian 
Museum. Miscellanea:—Malatia Mortality in the Fringe Area of Calcutta. Larval Habits of Toxorhynchites 
immisericors. East Asiatic species of Apus. 

Part IV.—Notes on the surface-living Copepoda of the Bay of Benga) Freshwater sponges of the Malabar Zone Notes 
on the Habits and Distribution of Limnocnida indica. The Anatomy of melo indicus. Two new species of Scoo- 
pendridae. 

Part V.—Indian Psyllidae. Descriptions of some new species of Freshwater Fish from North India New Oriental 
Diptera, I. 


Set Tee eee = 


Vol. VIII, 1912-1913. 


Pari I. ope Results of the Abor Expedition, 1011-12. Introduction. Batrachia. Reptilia. Lepidoptera. 


Porifera. Scolopendridae. Hymenoptera Anthophila. 


Part IT —Mammals. Coleoptera, I-III. Arachnida, I. Dermaptera. Diptera. 


Vol. IX, 1913. 


Part T.—Notes on the collection of Coccidae in the Indian Museum, II. New Indian Empidae. 


MEMOIRS 
of the 


INDIAN MUSEUM 


Vol. I. 
I.—An account of the Rats of Calcutta. By W.C. Hossack. Rs. 5-8. 


No. 

No. 2.—An account of the Internal Anatomy of Bathynomus giganteus. By R. E. 
Eroyp. Rs:2 

No. 3, A and B.— Ihe Oligochæta of India, Nepal, Ceylon, Burma and the Andaman 
Islands, with an account of the anatomy of certain aquatic forms. By W. 
MICHAELSEN and J. STEPHENSON. Rs. 4-8. 

No. 4.—Investigator sicarius, a Gephyrean Worm hitherto undescribed, the type of 
a new order. By F.H. STEWART. Rs. 2. . 

Vol. II. 

No. 1.—Report on the Fishes taken by the Bengal Fisheries Steamer ‘‘ Golden 
Crown.” Part I.—Batoidei. By N. ANNANDALE. Rs. 2 

No. 2.—An account of the Indian Cirripedia Pedunculata. Part Le tem Lepa- 
didz (sensu stricto). By N. ANNANDALE. Res. 2. 

No. 3.—A description of the deep-sea fish caught by the R.I.M.S. Ship ‘‘ Investiga- 
tor.’ since the year 1900, with supposed evidence of mutation in Malthopsis, 
and Illustrations of the Zoology of the R.I.M.S. Ship ‘‘ Investigator,’ Fishes, 
Plates XLIV—L, 1909. By R. E. Lnoyp. Rs. 4-8. 

No. 4.—Etude sur les Chironomides des Indes Orientales, avec description de quelques 
nouvelles espèces d’Egypte. Par J.J. KIEFFER. Rs. 2 

Vol. II. 

NO. I: — Report on the Fishes taken by the Bengal an Steamer “Golden 
Crown.’ Part IJ.—Additional notes on the Batoidei. By N. ANNANDALE. 
Part III.—Plectognathi and Pediculati. By N. ANNANDALE and J. T. JENKINS. 
Part IV:—Pleuronectidae.. By J. T. JENKINS. Rs. 3. 

No. 2.— Studies in post-larval development and minute anatomy in the genera Scal- 
pellum and Ibla. F. H. STEWART, Rs. 4. 

No. 3.—Indian Trypaneids (Fruit-Flies) in the collection of the Indian Museum, 
Calcutta. M. BEZZI. Rs. 6. 

No. 4.—An account of the Oriental Passalidae (Coleoptera) based chiefly on the 


‚collection in the Indian Museum. F. H. GRAVELY. (In the Press). 


Es 


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SMITHSONIAN INSTITUTION LIBRARIES 


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