Further Observations on

Oratosquilla, with Accounts of Two

New Genera and Nine New Species

Crustacea: Stomatopoda: Squillidae

RAYMOND 3. MANNING

m

wtu.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 272
SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of "diffusing knowledge" was expressed

by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph

Henry outlined a program that included the following statement: "It is proposed to

publish a series of reports, giving an account of the new discoveries in science, and

of the changes made from year to year in all branches of knowledge." This theme

of basic research has been adhered to through the years by thousands of titles issued

in series publications under the Smithsonian imprint, commencing with Smithsonian

Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to the Marine Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoo/ogy

Smithsonian Studies in Air and Space

Smithsonian Studies in History and Technology

In these series, the Institution publishes small papers and full-scale monographs

that report the research and collections of its various museums and bureaux or of

professional colleagues in the world cf science and scholarship. The publications are

distributed by mailing lists to libraries, universities, and similar institutions throughout

the world.

Papers or monographs submitted for series publication are received by the

Smithsonian Institution Press, subject to its own review for format and style, only

through departments of the various Smithsonian museums or bureaux, where the

manuscripts are given substantive review. Press requirements for manuscript and art

preparation are outlined on the inside back cover.

S. Dillon Ripley

Secretary

Smithsonian Institution
SMITHSONIA N CONTRIBUTION S T O ZOOLOGY • NUMBER 272

Further Observations on

Oratosquilla, with Accounts of Two

New Genera and Nine New Species

(Crustacea: Stomatopoda: Squillidae)

Raymond B. Manning

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978
ABSTRACT

Manning, Raymond B. Further Observations on Oratosquilla, with Accounts

of Two New Genera and Nine New Species (Crustacea: Stomatopoda: Squillidae). Smithsonian Contributions to Zoology, number 272, 44 pages, 25 figures,

1978.—Six species of the Indo-West-Pacific stomatopod genus Oratosquilla Manning are redescribed from type-specimens: O. anomala (Tweedie, 1935), O.

gonypetes (Kemp, 1911), O. inornata (Tate, 1883), O. perpensa (Kemp, 1911),

0. quinquedentata (Brooks, 1886), and O. -woodmasoni (Kemp, 1911). Oratosquilla tweediei Manning, 1971 is synonymized with O. woodmasoni and O.

jakartensis Moosa, 1975 also is shown to be identifiable with that species. Syntypes of Squilla affinis var. intermedia Nobili, 1903, are identified with four

different species. Nine species are described as new: O. asiatica, O. gravieri, O.

hindustamca, O. pentadactyla, O. solicitans, O. stephensoni, O. striata, O. subtilis, and O. turbata. Species groups in Oratosquilla are reexamined; two are

redefined and five are recognized. Two new genera are proposed for species

previously assigned to Oratosquilla, which now includes 31 nominal species.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Manning, Raymond B 1934-

Further observations on Oratosquilla, with accounts of two new genera and nine new species

(Crustacea: Stomatopoda: Squillidae)

(Smithsonian contributions to zoology ; no. 272)

Bibliography: p.

Supt. of Docs, no.: SI 127

I. Oratosquilla. 2. Crustacea—Classification. I. Title: Further observations on Oratosquilla...

II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no 272.

QL1.S54 no. 272 [QL444.M375] 591'.08s [595'.382] 77-608255
Contents

Page

Introduction *•

Explanation of Terms 1

Acknowledgments 2

Repositories *

Characters in Oratosquilla 3

Species Groups in Oratosquilla 4

Relationships of Species Groups 4

Revised Key to Species Groups in Oratosquilla 5

The gonypetes and perpensa Species Groups of Oratosquilla 5

Discussion 5

List of Nominal Species 7

Systematic Accounts 7

1. Oratosquilla anomala (Tweedie, 1935) 7

2. Oratosquilla asiatica, new species 10

3. Oratosquilla gonypetes (Kemp, 1911) 12

4. Oratosquilla gravieri, new species 14

5. Oratosquilla hindustanica, new species 15

6. Oratosquilla inornata (Tate, 1883) 17

7. Oratosquilla pentadactyla, new species 19

8. Oratosquilla perpensa (Kemp, 1911) 21

9. Oratosquilla quinquedentata (Brooks, 1886) 23

10. Oratosquilla solicitans, new species 25

11. Oratosquilla stephensoni, new species 29

12. Oratosquilla striata, new species 30

13. Oratosquilla subtilis, new species 33

14. Oratosquilla turbata, new species 35

15. Oratosquilla woodmasoni (Kemp, 1911) 36

Two New Genera for Species Previously Referred to Oratosquilla 39

Kempina, new genus 39

Natosquilla, new genus 40

Literature Cited 43

in

Further Observations on

Oratosquilla, with Accounts of Two

New Genera and Nine New Species

(Crustacea: Stomatopoda: Squillidae)

Raymond B. Manning

Introduction

This report was initiated after examination of

material during visits to several European museums

in 1971 revealed the possibility that each of several

supposedly common species of the Indo-WestPacific genus Oratosquilla Manning, 1968 might

actually be made up of more than one species. This

possibility also was supported by examination of

material from a variety of sources.

In 1972 I visited the Zoological Survey of India,

Calcutta, with the primary aim of redescribing

type-specimens of several species originally named

by S. W. Kemp (1911, 1913); most of Kemp's material, which formed the basis for the first and only

monograph of the Indo-West-Pacific stomatopods,

had been deposited in the Zoological Survey. Inasmuch as Kemp's accounts and illustrations are

still in use as the primary source of identification

of Indo-West-Pacific stomatopods today, reexamination of his material was particularly important.

That examination revealed that each of the accounts of three of the common species of Oratosquilla reported by Kemp, O. gonypetes (Kemp,

1911), O. quinquedentata (Brooks, 1886), and O.

perpensa (Kemp, 1911), were based on more than

Raymond B. Manning, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560

one species. The confusion has been compounded

by later authors, including Chopra (1939), Holthuis

(1941), and Manning (1966), among others. These

three species are redescribed herein, and new species, based at least in part on material identified

with these species by Kemp, are newly described.

Some material of these species from other collections, especially that incorrectly identified in the

literature, also is reported here. A fourth species

described by Kemp (1911), O. woodmasoni, is redescribed, and O. tweediei Manning, 1971, and

O. jakartensis Moosa, 1975, are shown to be its

synonyms.

In addition, I have been able to examine typespecimens of three other species: Squilla affinis

var. intermedia Nobili, 1903; Squilla inornata Tate,

1883; and Squilla anomala Tweedie, 1935. Nobili's

variety intermedia is preoccupied by Squilla intermedia Bigelow, 1893, and was considered by Kemp

(1913:58) to be unrecognizable; apparently material

of four different species was included by Nobili

under this name. Squilla inornata and S. anomala,

both of which have been confused in the past with

Squilla oratoria var. perpensa Kemp, are redescribed herein; they appear to be distinct species.

In all, six species are redescribed below and nine

species are newly described.

EXPLANATION OF TERMS.—All of the species of

the gonypetes and perpensa groups described below

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

share certain features which have been omitted

from the descriptions to avoid repetition. These

include: the anterior margin of the ophthalmic

somite is flattened or medially emarginate, unarmed; the ocular scales are subquadrate or truncate in shape, are inclined laterally, and are separate medially; the mandibular palp and four epipods are present; there are eight carinae on each

of the anterior five abdominal somites; there is a

spinule or tubercle ventrolaterally on each side

of the sixth abdominal somite, anterior to the articulation of the uropod; there are three pairs of

marginal teeth, the submedians with fixed apices, as

well as prelateral lobes on the telson.

Oratosquilla woodmasoni differs from the other

species described below in having the anterior margin of the ophthalmic somite broadly rounded and

usually armed with a median spinule. It resembles

the other species described herein in the other

features listed above and these features are not

repeated in its description.

The spine formula for the abdominal somites,

indicating on which somite the carinae are armed

posteriorly, is shown as follows: submedian 5-6,

intermediate 4-6, lateral (1-2)3-6, marginal 1-5.

This formula shows that posterior spines are found

on the submedian carinae of the fifth and sixth

somites, the intermediate carinae of the fourth

through sixth somites, the lateral carinae of the

third through the sixth and occasionally on the

first and second also, and the marginal carinae

of the anterior five somites.

A denticular formula of 3-4, 7-9, 1, for marginal

denticles of the telson, indicates that on each side

of the midline there are three to four submedian,

seven to nine intermediate and one lateral denticles.

Size descriptors used in the text, such as moderate or large, are based on the following scale developed for descriptive accounts of adult stomatopods: very small, total length less than 30 mm;

small, total length 31 to 50 mm; moderate, total

length 51 to 150 mm; large, total length greater

than 150 mm.

Size descriptors for eyes, based on observed range

of corneal indices (carapace length divided by cornea width X100), are as follows: very small, corneal

index greater than 500; small, index 400-499; moderate, index 300-399; large, index less than 299.

In any given species, the indices tend to be smaller

in juveniles and larger in adults.

Range of total lengths given in the section on

measurements is based on material reported herein

and does not necessarily reflect the overall size

range of the species.

Unless otherwise noted, color notes are based

on preserved specimens.

Total length (TL) is measured on the midline,

from the anterior margin of the rostral plate to

a line between the apices of the submedian teeth

of the telson. Carapace length (CL) is measured on

the midline; it does not include the length of the

rostral plate. The prelateral lobe is measured from

the anterolateral angle of the telson to the apex

of the lobe. The segments of the uropodal exopod

are measured dorsally. All measurements are in

millimeters (mm).

Wherever possible, localities have been verified

by reference to U.S. Board of Geographic Names'

Gazetteers. Alternate spellings and coordinates obtained from those gazetteers are entered in brackets when used. If this information is given under

"Material," it is not duplicated under "Distribution."

Original depth data are given throughout. Fathoms are converted to meters by multiplying by

1.83, and given in parentheses.

ACKNOWLEDGMENTS.—The following individuals

provided access to materials in their collections

during visits and/or made available material on

loan, for which I thank them: H. Fechter, Zoologische Staatsammlung, Munich; J. Forest and M.

de Saint Laurent, Museum National d'Histoire

Naturelle, Paris; H.-E. Gruner and G. Hartwich,

Zoologisches Museum, Berlin; L. B. Holthuis,

Rijksmuseum van Natuurlijke Historic Leiden;

R. W. Ingle, British Museum (Natural History),

London; David C. Lee, South Australian Museum,

Adelaide; C. Leonardi, Museo Civico di Storia

Naturale, Milan; and J. Stock, Zoologisch Museum,

Amsterdam. A special word of thanks is due K. K.

Tiwari and H. C. Ghosh, Zoological Survey of

India, Calcutta, for their cooperation and hospitality during a visit there in 1972; without their help

this work could not have been completed.

Part of this work was supported by the Smithsonian Institution through its Research Awards

and Foreign Currency Programs. The original illustrations were prepared by my wife Lilly.

Fenner A. Chace, Jr., Smithsonian Insitution,

took time to go through a draft of the manuscript
NUMBER 272

in his welcome critical manner, for which I thank

him.

REPOSITORIES.—Institutions in which the material

reported herein is deposited are identified by the

following abbreviations:

AM

BMNH

MCSN

MNHNP

QM

RMNH

SAM

USNM

ZMA

ZMB

ZMBS

ZSI

Australian Museum, Sydney

British Museum (Natural History), London

Museo Civico di Storia Naturale, Milan

Museum National d'Histoire Naturelle, Paris

Queensland Museum, Brisbane

Rijksmuseum van Natuurlijke Historic Leiden

South Australian Museum, Adelaide

National Museum of Natural History, Smithsonian Institution, Washington (abbreviation refers to former United States National Museum)

Zoologisch Museum, Amsterdam

Zoologisches Museum, Berlin

Zoologische Staatssammlung, Munich

Zoological Survey of India, Calcutta

Characters in Oratosquilla

Like most squillids, the species of Oratosquilla

are relatively rich in differentiating characters, and

many different features are available for species

discrimination. In spite of the numbers of characters, however, there is a rather remarkable uniformity in a series of specimens of any given species from one locality, as in those of O. perpensa

and O. hindustanica examined during this study.

The major features such as eye size and shape,

shape of rostral plate, shape of the lateral processes

of the exposed thoracic somites, spination of the

abdominal carinae, and shape and size of the lobe

between the spines of the basal prolongation of

the uropod, show no significant variation. In general, there is not enough material of any species

to demonstrate the extent of variation in different

parts of the range of each species.

Characters that should be examined in any study

of Oratosquilla include the following.

General: Nature of texture of body surface,

whether smooth, as in O. woodmasoni, or variously

pitted and eroded, as in several species. This is

best determined when the surface is dried.

Eyes: Shape and size, particularly as reflected

in corneal indices; eyes tend to be relatively larger

in small specimens of any species.

Anterior Margin of Ophthalmic Somite: Truncate or medially emarginate in most species, but

anteriorly rounded, usually with a median point,

in Woodmasoni group.

Antennular Processes: Overall shape as seen in

dorsal view; they are usually unarmed (although

they may appear sharp in dorsal view) but are

produced into distinct spines in O. gravieri, described below.

Rostral Plate: Length/width proportions; shape,

whether trapezoidal, triangular, or rectangular;

shape of apex, rounded or flattened; and presence

or absence of a median carina.

Carapace: Condition of median carina, whether

entire or interrupted, and of anterior bifurcation

of carina, whether present, indistinct (position indicated by pigment), or absent; relative length of

anterolateral spines, especially whether these overreach the base of the rostral plate, as in O. anomala; relationship of anterolateral width to median

length.

Claw: Number of teeth on dactylus; shape of

outer margin of dactylus, whether sinuate, flattened, or evenly curved; presence of a distal spine

or projection on the propodus, as in O. anomala;

condition of the carpal crest, whether uninterrupted or tuberculate; presence of spine at inferodistal angle on outer face of merus.

Lateral Processes of Thoracic Somites: Shape

and relative size of lobes; shape of anterior lobe

of process of sixth somite (triangular in O. perpensa, rectangular in O. solicitans) is particularly

important.

Spination of Abdominal Carinae: Patterns of

spination often are specific characters; submedian

carinae of fourth somite rarely spined (variable in

O. woodmasoni); extent of spination of lateral

carinae also important (unarmed on anterior three

somites in some species, e.g. O. quinquedentatd).

Telson: Presence of dorsal carinae in addition

to median carina and carinae of marginal teeth, as

in O. striata; length of prelateral lobe in relation

to length of lateral tooth; relative length of some

carinae of marginal teeth, as in O. striata versus

O. gonypetes; number and shape of marginal denticles.

Uropod: Relative length and slenderness of

exopod segments, as in O. subtilis versus O. gonypetes; shape and size of lobe between spines of

basal prolongation of uropod; this latter feature

is variable in at least O. hesperia (Manning, 1968b)

and O. woodmasoni, in which it tends to be a

spine in young specimens, a bluntly angled or
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rounded lobe in adults and may vary geographically.

Other information, which probably is extremely

important but which is generally unavailable, is

the color in life and the habitat, including depth

range. Series of all of the species recorded below

accompanied by observations on color and habitat

will be required before some of the systematic

problems can be worked out.

Species Groups in Oratosquilla

Oratosquilla Manning, 1968(a), was recognized

for a group of comparatively large squillids from

the Indo-West-Pacific region, the equivalent there

of the Atlantic-East-Pacific genus Squilla Fabricius,

1787. In 1971 I distinguished eight species groups

in Oratosquilla, primarily as an aid to the recognition of the 23 species then assigned to the genus,

secondarily to emphasize the diversity of the species

then included in it. Since 1971, two new genera

containing species previously assigned to Oratosquilla have been named: Areosquilla Manning,

1976, for species of the indica group, and Busquilla

Manning, 1978, for Oratosquilla quadraticauda

(Fukuda, 1911) and a related new species from

Madagascar. Two new genera are recognized herein,

one for the species assigned to the investigatoris

group and one for those assigned to the mikado

group by me in 1971.

At the present time, five species groups can be

recognized in Oratosquilla: the nepa group (two

nominal species), the oratorio group( four nominal

species), the woodmasoni group (three nominal species), the gonypetes group (nine nominal species),

and the perpensa group (13 nominal species). This

report is concerned with representatives of the latter three groups, and the gonypetes and perpensa

groups are redefined.

Since 1971 the following species of Oratosquilla

have been described: O. arabica Ahmed, 1971 {perpensa group), O. birsteini Makarov, 1971 and O.

vietnamica Blumstein, 1974 (gonypetes group), and

O. jakartensis Moosa, 1975 (woodmasoni group).

The latter species appears to be a synonym of O.

woodmasoni, and it is discussed below under the

account of that species.

RELATIONSHIPS OF SPECIES GROUPS.—Although

there are no clear indications of relationships of

the species groups within the genus, for we can

only guess about whether or not a character is

primitive or derived, some very limited generalizations can be made. One of the characteristics of

the squillids in general is the well developed body

carination, which in Oratosquilla appears to have

been retained in its most "complete" condition

only in the oratorio and nepa groups. In those

groups the median carina of the carapace is entire

throughout its length anterior to the cervical groove

and it terminates anteriorly in a distinct median

bifurcation.

Of these two groups, the Oratoria possibly reflects the oldest stem of Oratosquilla development.

The carination of the body is well developed, the

eyes are relatively large and a distinct meral spine

is present on the claw. The species of the oratoria

group exhibit a relict distribution pattern within

the Indo-West-Pacific region, with two species in

China and Japan, one in Hawaii, and one in the

western Indian Ocean.

The nepa group, comprising two species, possibly

represents an early offshoot of the main stem in the

genus. As in the oratoria group, the median carina

of the carapace terminates anteriorly in a well developed bifurcation, but the bifurcataion differs

from that found in any other squillid in that it

opens anterior to the dorsal pit. The eyes are much

smaller in members of this group than those of

any other group and the meral spine is present.

The woodmasoni group possibly also represents

another early offshoot of the main stem. Representatives are distinguished by the broadly rounded,

usually armed anterior margin of the ophthalmic

somite, which is flattened or medially emarginate

in the other species of the genus, and the broad,

smooth carapace on which the median carina usually lacks a distinct bifurcation; the strong meral

spine also is characteristic of members of this

group, which occurs from Indo-Malaya westward

through the Indian Ocean; two of the species are

found only in the western Indian Ocean.

The remaining two species groups, each of which

appear to represent a different line of development,

differ from those discussed above in that the structure of the median carina of the carapace has been

modified so that it is interrupted at the base of

the anterior bifurcation, and, in some species, the

branches of the anterior bifurcation are poorly

developed or absent. Members of the perpensa
NUMBER 272

group have the meral spine whereas those of gonypetes group lack it. It seems likely that it has been

suppressed in members of this latter group.

Relationships of Oratosquilla to the other squillids will be explored in detail in a review of higher

taxa of stomatopods now in preparation.

Revised Key to Species Groups in Oratosquilla

(Modified from Manning, 1971)

1. Anterior bifurcation of median carina of carapace opening posterior to dorsal pit. Cornea

very small, corneal index 600 or more in adults (TL 100 mm or more), set obliquely on

stalk nepa group

Anterior bifurcation of median carina of carapace, when present, opening anterior to dorsal

pit. Cornea small to large, corneal index less than 600 in adults, set obliquely on stalk 2

2. Median carina of carapace entire, with well formed anterior bifurcation [Anterior margin

of ophthalmic somite medially flattened or emarginate, unarmed. Dactylus of claw with 6

teeth. Surface of body punctate] oratorio group

Median carina of carapace interrupted at base of bifurcation, or branches of bifurcation indisinct or absent 3

3. Surface of body smooth, appearing highly polished. Anterior margin of ophthalmic somite

rounded, often with median spinule [Anterior width of carapace greater than half median

length of carapace. Dactylus of claw with 6 teeth] woodmasoni group

Surface of body minutely or coarsely punctate. Anterior margin of ophthalmic somite medially flattened or emarginate, unarmed 4

4. Inferodistal angle on outer face of merus of claw unarmed [Abdomen usually with dark

dorsal patch(es) on fifth abdominal somite] gonypetes group

Inferodistal angle on outer face of merus of claw produced into distinct spine [Abdomen

lacking dorsal pigmented area on fifth somite] perpensa group

The gonypetes and perpensa Species Groups

of Oratosquilla

DISCUSSION

In 1971 I distinguished the gonypetes, perpensa,

and woodmasoni species groups primarily by the

condition of the anterior bifurcation of the median

carina of the carapace and the number of teeth

on the dactylus of the claw. A comment by Holthuis (1967:12), that material of O. simulans without chelae would be difficult to distinguish from

O. gonypetes also lacking chelae, led me to reexamine characteristics of these three groups. In 1971

I had placed O. gonypetes, which has five teeth on

the dactylus of the claw, in the gonypetes group

and O. simulans, which has six, in the perpensa

group. Further, I had later examined material of

a species near O. gonypetes in which the carapace

was relatively broad, the anterior width being

more than half the median length (but less than

half the length of the carapace and rostral plate

combined), and on that feature would key out in

my earlier key to species groups (Manning, 1971:2)

to the woodmasoni group, in spite of its obvious

affinities with O. gonypetes.

Although the broad carapace is one of the characteristics of the woodmasoni group, there are species in other groups, such as O. striata, described

below, in which the carapace also is relatively

broad. Members of the woodmasoni group are unusually smooth species, with a broad carapace on

which the carinae are poorly developed, and in all

of the species assigned to the group, the anterior

margin of the ophthalmic somite is obtusely

pointed or rounded, almost always with a median

spinule; it is not medially emarginate or flattened

as in the members of the gonypetes and perpensa

groups.

Although I stressed the importance of the number of teeth on the claw as a characteristic of the

species groups in 1971, the present study has convinced me that other features are more important.

The gonypetes group as redefined herein now comprises two sections, one with species that have five

teeth on the claw, the gonypetes complex, and one

with six, the imperalis complex. Members of both

of these complexes share relatively small size and

a roughened surface, which appears pitted and corroded under magnification. In both the inferodis-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tal angle of the merus of the claw is unarmed, and,

at least in the species for which color pattern has

been described, the fifth abdominal somite is ornamented dorsally with a median patch or submedian

patches of dark pigment. As noted above, the anterior margin of the ophthalmic somite is medially

emarginate or flattened and unarmed.

The perpensa group, like the gonypetes group,

includes species with five or six teeth on the claw,

a coarse surface on the body, and a medially emarginate anterior margin on the ophthalmic somite.

Members of this group all have a strong spine on

the inferodistal angle of the merus of the claw and

all lack distinctive dark spots or patches on the

fifth abdominal somite. Four species complexes are

recognized in the perpensa group: the quinquedentata complex, including two species with five teeth

on the claw; the interrupta complex, those species

in which the ridge on the carpus of the claw is

tuberculate, not smooth; and two complexes of

species in which the carpus of the claw is undivided: the perpensa complex including species with

a relatively short, broad rostral plate, and the inornata complex, species with a relatively long, slender

rostral plate.

Oratosquilla imperialis (Manning, 1965) and O.

ornata Manning, 1971 are here transferred from

the perpensa group to the gonypetes group. Both

lack the spine on the merus of the claw, and O.

imperialis, at least, has the dark dorsal patch on

the fifth abdominal somite.

In 1971 I suggested that O. simulans (Holthuis,

1967) from the Red Sea was conspecific with O.

imperialis (Manning, 1965) from Japan. Although

I suspect that they are conspecific, until material

from the two areas can be compared I believe that

both species should be recognized. A recentlydescribed species of the gonypetes group, O. vietnamica Blumstein, 1974, from the Gulf of Tonkin,

is very similar to and may prove to be identical

with O. ornata Manning, 1971, from off Hong

Kong. It also seems certain that another nominal

species of the perpensa group, O. arabica Ahmed,

1971, from the Arabian Gulf, is conspecific with

O. interrupta (Kemp, 1911), a widely distributed

speices which also occurs in the Arabian Gulf. Like

O. interrupta, Ahmed's species has a convex lobe

on the outer margin of the inner spine of the basal

prolongation of the uropod. The other features

used by Ahmed (1971:251) to distinguish O. arabica

from O. interrupta include (1) the first segment

of the antennular peduncle armed with six spines,

(2) a dorsal spine on the antennal peduncle, (3)

the presence of an oval pit in the articulation between the carapace and the rostral plate, (4) the

anterior width of the carapace is more than half

its median length, (5) the ocular scales are cordiform, (6) the outer margin of the intermediate

carina on the sixth abdominal somite with a small

pointed lobe, (7) the presence of a transverse lobe

anterolaterally on the telson, and (8) a denticular

formula of 2-3, 6-8, 1. The first two of these, based

on ornamentation of the antennular and antennal

peduncles, have not been studied in any detail in

stomatopods; they may prove to have value as specific characters but are not now in use as such. There

is always a gap of varying size between the articulation of the rostral plate and the carapace. The

anterior width of the carapace is shown to be much

less than half the median length in Ahmed's illustration (fig. 1), and the ocular scales are more or

less subquadrate and inclined laterally as in all

species of the genus. The interrupted lobe on the

intermediate carina of the sixth abdominal somite

and the short, transverse ridges on the telson are

found in most if not all species of the genus.

Finally, the denticular formula falls within the

range of that reported for O. interrupta in the literature, 2-4, 6-9, 1. Ahmed's species probably

should be considered as a synonym of O. interrupta.

Another lesson from this study is that in the

stomatopods, at least, we have not progressed beyond that stage in systematics in which type-specimens are of primary importance. The types of

older species all should be restudied. Most of the

species characterized below could not have been

recognized without studying Kemp's types, and

Kemp's accounts were, even by present standards,

quite good. Hansen (1926:1) noted that "Kemp's

book is in reality one of the best systematic works

ever used by me. It is difficult to find any useful

specific or generic character not used by him."

However, it seems apparent that Kemp's species

concept was very conservative.

Nominal species in the gonypetes and perpensa

species groups are listed below. Authors and dates

for each species will not be repeated in discussions

of individual species.
NUMBER 272

LIST OF NOMINAL SPECIES

(numbers preceding taxa are those assigned to species

described herein)

I. Inferodistal angle on outer face of merus of claw unarmed (gonypetes group)

A. Dactylus of claw with 5 teeth {gonypetes complex)

3. O. gonypetes (Kemp, 1911)

13. O. sublilis, new species

14. O. turbata, new species

B. Dactylus of claw with 6 teeth (imperialis complex)

O. birsteini Makarov, 1971

O. imperialis (Manning, 1965)

O. ornata Manning, 1971

O. simulans (Holthuis, 1967) {—O. imperialist)

12. O. striata, new species

O. vietnamica Blumstein, 1974 (=O. ornata?)

II. Inferodistal angle on outer face of merus of claw armed

(perpensa group)

A. Dactylus of claw with 5 teeth (quinquedentata complex)

7. O. pentadactyla, new species

9. O. quinquedentata (Brooks, 1886)

B. Dactylus of claw with 6 teeth

i. Ridge on carpus of claw tuberculate (interrupta

complex)

O. arabica Ahmed, 1971 (=O. interrupta})

2. O. asiatica, new species

O. fabricii (Holthuis, 1941)

O. interrupta (Kemp, 1911)

ii. Ridge on carpus of claw smooth, undivided

a. Rostral plate short (perpensa complex)

1. O. anomala (Tweedie, 1935)

8. O. perpensa (Kemp, 1911)

11. O. stephensoni, new species

b. Rostral plate long (inornata complex)

4. O. gravieri, new species

5. O. hindustanica, new species

6. O. inornata (Tate, 1883)

10. O. solicitans, new species

Systematic Accounts

1. Oratosquilla anomala (Tweedie, 1935)

FIGURES 1-3

Squilla affinis var. intermedia Nobili, 1903:39 [part; not

specimens from Nias = O. pentadactyla, new species; preoccupied by Squilla intermedia Bigelow, 1893].

Squilla oratorio var. perpensa.—Parisi, 1922:98 [part; specimen from Singapore only; not Squilla oratoria var. perpensa Kemp, 1911].

Chloridella oratoria.—Schmitt, 1931:147 [part; not Squilla

oratoria de Haan, 1844].

Squilla anomala Tweedie, 1935:45.—Moosa, 1973:147.

MATERIAL.—Tsimei [Chi-Mei, Fukien Province; 24°34'N,

118°06'E], China; 25 Jun 1923: 3$ , TL 83-87 mm (USNM

62188). Siglap [01°19'N, 103°56'E], Singapore, Malaysia;

littoral; M.W.F. Tweedie, leg.; Jun 1934; holotype of Squilla

anomala Tweedie: lj , TL 85 mm (BMNH 1935.12.16.7).

Data same; paratypes of Squilla anomala Tweedie: 1 $, TL

69 mm; 1 $ , TL 82 mm (RMNH 248S). Singapore [01°17'N,

103°51'E], Malaysia; syntype of Squilla affinis var. intermedia

Nobili: 1 $ , TL 66 mm (MCSN).

DESCRIPTION.—Size moderate, total length of

adults less than 100 mm. Body appearing rough,

surface pitted and irregular.

Eye (Figures \b, 2b, 3a) very small to small, cornea bilobed, set obliquely on stalk. Eyes extending

almost to end of first segment of antennular peduncle. Corneal indices 399-527.

Antennular peduncle shorter than carapace. Dorsal processes of antennular somite rounded, unarmed anteriorly.

Rostral plate (Figures la, 2a, 3a) broader than

long, trapezoidal, apex truncate. Median carina

absent.

Anterior width of carapace about half median

length, less than half length carapace and rostral

plate combined. Anterolateral spines (Figures la,

2a, 3a) strong, often extending beyond base of rostral plate. Median carina interrupted at base of

anterior bifurcation, branches of bifurcation distinct. Intermediate carinae turning toward but not

meeting laterals, not extending to anterior margin.

Dactylus of claw with 6 teeth, outer margin

sinuate. Dorsal ridge of carpus (Figure lc) undivided. Inferodistal angle of outer face of merus

with sharp spine.

Exposed thoracic somites (Figures Id, 2c, $b)

each with unarmed submedian and intermediate

carinae, intermediates of fifth somite short, irregular. Lateral process of fifth somite bilobed, anterior

lobe a slender spine, directed anterolaterally or

almost anteriorly, posterior lobe slender, triangular, apex rounded, directed laterally. Lateral process of sixth somite bilobed, anterior lobe large but

smaller than posterior, slender subrectangular, apex

rounded or subtruncate, posterior lobe larger, triangular, apex acute but not sharp. Lateral process

of seventh somite strongly bilobed, anterior lobe

slender, more triangular than that of sixth somite,-

posterior lobe broader, posterior margin sinuous,

apex acute but not sharp.

Submedian carinae slightly divergent on fifth

abdominal somite. Abdominal carinae spined as
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 1.—Oratosquilla anomala (Tweedie, 1935), female, TL 83 mm, China: a, anterior part

of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic

somites; e, submedian teeth and denticles of telson, ventral view; f, basal prolongation of uropod,

ventral view.

a

FIGURE 2.—Oratosquilla anomala (Tweedie, 1935), male holotype, TL 85 mm, Singapore:

a, anterior part of body; b, eye; c, lateral processes of fifth, sixth, and seventh thoracic

somites; d, basal prolongation of uropod, ventral view.

follows: submedian 5-6, intermediae 4-6, lateral

3-6, marginal 1-5.

Telson flattened, slightly broader than long. Prelateral lobe longer than margin of lateral tooth.

Denticles (Figures \e) rounded or subquadrate,

3-4, 7-8, 1. Ventral surface of telson with postanal

keel.

Uropod rather broad, proximal segment of exo-
NUMBER 272

a

FIGURE 3.—Oratosquilla anomala (Tweedie, 1935), male syntype of Squilla affinis var. intermedia

Nobili, 1903, TL 66 mm, Singapore: a, anterior part of body; b, lateral processes of fifth, sixth,

and seventh thoracic somites; c, propodus of claw; d, basal prolongation of uropod, ventral

view.

pod shorter than distal, with 8-9 movable spines

on outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin

of inner spine of basal prolongation (Figures If,

Id, 3d) small, narrower than adjacent spine, margin concave, apex rounded.

COLOR.—Faded in the material reported here.

Tweedie (1935:48) noted: "Distal segment of the

exopodite of the uropod similar to that of S. interrupta, but with a little more black suffusion; the

inner half is never entirely blackish, as in S.

oratorio inornata."

MEASUREMENTS.—Males, TL 66-85 mm; females,

TL 82-87 mm. Tweedie (1935) reported four males

and five females ranging from 58 to 91 mm long.

Other measurements of a female, TL 83 mm: CL

18.3, anterior width 9.3; cornea width 3.5; antennular peduncle length 15.6; rostral plate length

2.2, width 3.4; telson length 15.3, width 16.4.

REMARKS.—Examination of type-series of O.

anomala in the collection of the British Museum

(Natural History) and the Rijksmuseum van

Natuurlijke Historie clearly demonstrated that I

was in error in synonymizing Tweedie's species

with O. perpensa in 1966. These species, two of the

three representatives of the perpensa complex in

the perpensa group, share a roughened body, a

short rostral plate, a broad carapace with relatively

long anterolateral spines, and a claw with an undivided carpus and six teeth on the dactylus, features which characterize the perpensa complex and

serve to distinguish these species from all others

in the genus. Oratosquilla anomala closely resembles O. perpensa, but differs in having a shorter

rostral plate, a broader, more rectangular anterior

lobe on the lateral process of the sixth thoracic

somite, and a smaller lobe between the spines of

the basal prolongation of the uropod. The best

distinguishing feature is the broader anterior lobe

on the lateral process of the sixth thoracic somite;

that lobe is distinctly more slender and triangular

in O. perpensa. Oratosquilla anomala differs from

O. stephensoni, the third representative of the perpensa complex, in having a distinctly concave lobe

between the spines of the basal prolongation of the

uropod; O. anomala also is a smaller species.

It is somewhat surprising to learn that part of

the material assigned by Nobili (1903) to Squilla

affinis var. intermedia and considered by some

earlier authors (Kemp, 1913:58) to be unidentifi-
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

able actually can be identified with O. anomala,

a species also unrecognizable until the types were

examined. One of Nobili's specimens, that from

the collection of the Museo Civico di Storia Naturale, Milan, identified by Parisi (1922) with O.

perpensa, is shown in Figure 3 for comparison with

one of the types of O. anomala, shown in Figure 2.

Nobili (1903) recorded 23 specimens of 5. affinis

var. intermedia from Singapore and two from Nias.

Those from Nias are assigned below to O. pentadactyla, new species. The collection of the Museo

ed Istituto di Sistematica della Universita di Torino included only two specimens of the type-series

from Singapore. One is clearly identifiable with

O. interrupta and the other with O. woodmasoni

(Kemp). I was not able to locate the remainder of

Nobili's type-series.

Another feature of O. anomala which may prove

to be diagnostic is the presence of a distal spine

or angled projection on the extensor (outer) margin of the propodus of the claw (Figure 3c). This

projection is present in Nobili's specimen from

Singapore as well as in the three females from

China (USNM 62188). Not all of the material of

this species was examined at the same time, so the

occurrence of this projection was not determined

in all specimens. This character could prove to be

invaluable for quick recognition of this species,

but much more material will have to be studied

before its worth can be established. The projection

has not been observed in any of the material of

other species of Oratosquilla that I have seen, but

it may have been overlooked.

The specimens from China listed above had been

identified with Oratosquilla oratorio (de Haan)

by Schmitt (1931) (as Chloridella oratorio).

Paratypes of Squilla anomala also are in the collection of the University of Singapore, where they

were transferred from the National Museum of

Singapore (Moosa, 1973).

Apparently the ranges of O. anomala and O.

perpensa overlap in the Indo-Malayan area. So far

as is known, they both are more northern in range

than is the related O. stephensoni, described below.

DISTRIBUTION.—Known only from Tsimei, Fukien

Province, China (Schmitt, 1931), and from Siglap,

Singapore and Morib [02°45'N, 101°26'E], Selangor, Malaysia (Tweedie, 1935; Moosa, 1973); Singapore, Malaysia (Nobili, 1903). Apparently it occurs

in shallow water; some of the types were taken in

the littoral zone and Tweedie (1935) noted that

some specimens came from fishermen's nets.

2. Oratosquilla asiatica, new species

FIGURE 4

MATERIAL.—Keeling [Chilung] River, N Formosa [Taiwan];

Haberer, leg.; May 1903; paratypes: 1$, CL 17.3 mm, 1$ ,

CL 20.5 mm (ZMBS). Anping [An-P'ing or Ampin, 23°00'N,

120°09'E], S Formosa; Haberer, leg.; Jun 1903; paratypes:

1$, TL 81.5 mm, 1 $ , TL 85.5 mm (ZMBS). Pandan,

Panay Island, Antique Province, Philippine Islands; from

trap at mouth of tidal stream; D. G. Frey, leg.; 7 Apr 1946;

holotype: 1 $, TL 93 mm (USNM 125064). Humboldt Bay

[Teluk Jos Sudarso, 02°30'S, 140°50'E], near the mouth of

the Tami River, West New Guinea [West Irian], Indonesia;

25 Jun 1955; paratype: 1 $ , TL 97 mm (RMNH 342S).

DESCRIPTION.—Size moderate, total length of

adults less than 100 mm. Body appearing smooth,

coarsely pitted under magnification.

Eye (Figure 46) moderate to small, cornea bilobed, set obliquely on stalk. Eyes extending almost

to end of first segment of antennular peduncle.

Corneal indices 376-422.

Antennular peduncle about %0 as long as carapace. Dorsal processes of antennular somite produced into unarmed triangular lobes, directed

anterolaterally.

Rostral plate (Figure 4a) slightly broader than

long, lateral margins converging on flattened apex.

Short median carina present.

Anterior width of carapace slightly more than

half median length, less than half length carapace

and rostral plate combined. Anterolateral spine

(Figure 4a) strong but not extending to base of

rostral plate. Median carina scarcely interrupted

(holotype) or distinctly interrupted (other specimens) at base of anterior bifurcation, branches of

bifurcation distinct. Intermediate carinae not extending to anterior margin.

Dactylus of claw with 6 teeth, outer margin sinuate. Dorsal ridge of carpus (Figure 4c) irregular,

with 2-3 rounded tubercles distally. Inferodistal

angle of outer face of merus with sharp spine.

Exposed thoracic somites (Figure 4d) each with

unarmed submedian and intermediate carinae, intermediates irregular, short on fifth somite. Lateral

process of fifth somite bilobed, anterior lobe a

slender spine, directed anterolaterally, posterior lobe

slender, almost spiniform, directed laterally. Lateral
NUMBER 272 11

a

FIGURE 4.—Oratosquilla asiatica, new species, male holotype, TL 93 mm, Philippine Islands: a,

anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

process of sixth somite bilobed, anterior lobe very

small, slender, posterior lobe much broader, apex

obtusely angled, unarmed. Lateral process of seventh somite scarcely bilobed, posterior lobe similar

to that of sixth somite, with low, obtuse prominence on anterior edge.

Submedian carinae slightly divergent on fifth

abdominal somite. Abdominal carinae spined as

follows: submedian 4-6, intermediate 3-6, lateral

(1) 2-6, usually 1-6, marginal 1-5.

Telson flattened, slightly broader than long. Prelateral lobe shorter than or slightly longer than

margin of lateral tooth. Denticles (Figure 4e)

rounded or sub triangular, 3-4, 9, 1. Ventral surface

of telson with postanal keel.

Uropod rather broad, proximal segment of exopod slightly shorter than distal, with 8-9 short

movable spines on outer margin, distalmost not

extending to midlength of distal segment. Lobe on

outer margin of inner spine of basal prolongation

(Figure 4/) low, narrower than adjacent spine,

margin concave, apex rounded.

COLOR.—Carapace and each body segment with

dark posterior line. Dark spot present on articulation of segments of uropodal exopod, color diffuse

on distal segment, not covering inner half. Distal

half of uropodal endopod dark.

MEASUREMENTS.—Males, TL 81.5-93 mm; only

intact female, TL 97 mm. Other measurements of

male holotype, TL 93 mm: CL 20.2, anterior width

10.7; cornea width 5.1; antennular peduncle length

16.7; rostral plate length 3.0, width 3.5; telson

length 17.8, width 18.7.

REMARKS.—Oratosquilla asiatica is a member of

the interrupt a complex of the gony petes group,

those species of the genus in which the claw has

a spined inferodistal angle on the merus, a tuberculate ridge on the carpus, and six teeth on the

dactylus. It differs from O. interrupta and O.

arabica (if the latter species proves to be distinct)

and resembles O. fabricii in having spined submedian carinae on the fourth abdominal somite

and a concave lobe between the spines of the basal

prolongation of the uropod. Oratosquilla asiatica

differs from O. fabricii in having a median carina

on the rostral plate, smaller anterolateral spines

on the carapace, and much slenderer anterior lobes

on the lateral processes of the thoracic somites.

Although little specific ecological information accompanies the few known specimens of this species,

several of the specimens were taken near rivers,

suggesting that the species may be able to tolerate

or may inhabit waters of reduced salinity.

ETYMOLOGY.—The name is from the Latin and

alludes to the distribution of the species.

DISTRIBUTION.—This species is known from local-
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ities in the western Pacific Ocean, including northern and southern Formosa, the Philippine Islands,

and West New Guinea, Indonesia. Its depth range

has not been recorded, but it almost certainly

occurs in shallow water.

3. Oratosquilla gonypetes (Kemp, 1911)

FIGURE 5

Squilla gonypetes Kemp, 1911:96 [part]; 1913:54, pi. 4: figs.

42-44 [part].—Kemp and Chopra, 1921:300.—Chopra,

1939:148 [part?; not illustrated specimen].

MATERIAL.—Off Kabusa Island [Kabosa Island, 12°49'N,

97°53'E], Lower Burma; 25-35 fms (46-64 m); Investigator,

leg.: 19, TL 47 mm (ZSI C320/1). Off Cheduba [Cheduba

Island, 18°48'N, 93°38'E], Arakan coast, Burma; 7 fms (13 m);

Investigator, leg.; lectotype: lj , TL 39 mm (ZSI 3359/7).

Persian Gulf; 26°24'N, 56°02'E; 47 fms (86 m); Investigator,

leg.; paralectotype: 19 , TL 26 mm (ZSI 4421/10). Gulf

of Oman; 25O38'18"N, 56°26'36"E; 73 m; coarse sand and

shells; John Murray Sta 72; 26 Nov 1933: 2^ (1 broken),

TL 55 mm; 1 9 , TL 50 mm (BMNH).

DESCRIPTION.—Size small to moderate, total length

of adults to 55 mm. Body appearing smooth, actually lightly pitted under magnification.

Eye (Figure 5b) moderate to small, cornea bilobed, set obliquely on stalk. Eyes not extending to

end of first segment of antennular peduncle. Corneal indices 358-408.

Antennular peduncle longer than carapace. Dorsal processes of antennular somite produced into

acute lobes directed anterolaterally.

Rostral plate (Figure 5a) slightly broader than

long, appearing elongate, lateral margins converging on rounded apex. Median carina absent.

Anterior width of carapace less than half median

length. Anterolateral spines (Figure 5a) strong, extending almost to, or beyond, base of rostral plate.

Median carina interrupted at base of bifurcation,

branches of bifurcation usually scarcely discernible.

Intermediate carinae slightly turning toward laterals, not extending to anterior margin.

Dactyl us of claw with 5 teeth, outer margin

broadly curved, sinuous proximally, with obtuse

projection basally. Dorsal ridge of carpus (Figure

5c) undivided. Inferodistal angle on outer face of

merus unarmed.

Exposed thoracic somites (Figure 5d) each with

unarmed submedian and intermediate carinae, intermediates of fifth somite irregular, short. Lateral

process of fifth somite bilobed, anterior lobe a

slender spine, directed anteriorly or anterolaterally,

posterior lobe much smaller, slender, apex triangular, almost sharp, directed laterally. Lateral process

of sixth somite strongly bilobed, anterior lobe large,

subrectangular or trapezoidal, apex blunt, posterior

lobe triangular, obtusely pointed, apex blunt. Lateral process of seventh somite strongly bilobed, anterior lobe slightly smaller than that of sixth somite,

triangular, posterior lobe much larger, triangular,

apex obtusely pointed, unarmed.

Submedian carinae divergent on fifth abdominal

somite. Abdominal carinae spined as follows: submedian 5-6, intermediate 3-6, lateral (1) 2-6, marginal 1-5.

Telson flattened, slender, appearing elongate,

length and width subequal. Prelateral lobe subequal in length to margin of lateral tooth. Dorsal

surface of telson occasionally with lines of short

carinae or raised tubercles between curved rows of

pits in large specimens. Denticles (Figure be) usually rounded, 3-4, 6-7, 1, outer submedians and

intermediates largest, inner submedians occasionally

spine tipped.

Uropod slender, proximal segment of exopod

slightly shorter than distal, with 8 movable spines

on outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin of

inner spine of basal prolongation (Figure 5/) small,

narrower than adjacent spine, margin concave, apex

rounded.

COLOR.—Eyes with dark patch laterally on stalk.

Carinae and grooves of carapace lined with dark

pigment; midline of carapace with dark, U-shaped

patch, open anteriorly, anterior to cervical groove.

Claw with dark dorsal and distolateral patches on

merus and distal dark patch on outer face of propodus. Lateral edge of antennal protopod dark.

Carinae and posterior margins of posterior 3 thoracic and all abdominal somites dark, each somite

with longitudinal line of dark pigment between

submedian and intermediate carinae. Lateral processes of sixth and seventh thoracic somites dark

posteromesially. Abdominal somites with short,

curved line of dark pigment above lateral carina,

rectangular middorsal patch on second somite, and

2 square black patches, each lateral to submedian

carina, on fifth somite; posterolateral surface of

sixth somite dark. Marginal carinae of telson dark.

Distal part of proximal segment and inner half of
NUMBER 272 13

a

FIGURE 5.—Oratosquilla gonypetes (Kemp, 1911), male lectotype, TL 39 mm, Burma: a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of uropod, ventral view, enlarged (apices of spines broken).

distal segment of uropodal exopod dark. Distal half

of endopod dark. Basal prolongation of uropod

with dark longitudinal line at level of dark pigment

of endopod.

MEASUREMENTS.—Males, TL 39-55 mm; females,

TL 26-47 mm. Other measurements of female, TL

47 mm: CL 10.6, anterior width 5.0; cornea width

2.6; antennular peduncle length 11.2; rostral plate

length 1.6, width 1.7; telson length 8.6, width 8.4.

REMARKS.—In the original description of this

species, Kemp (1911) mentioned four specimens

from three localities; the same material was reported by him in his more detailed account published in 1913. Because no type designation was

made by Kemp, all four specimens are syntypes.

Two of the specimens, taken off the Andaman Islands, could not be located by me at the Zoological

Survey of India in 1972. Of the two remaining

specimens, a male from off Cheduba, Burma (ZSI

3359/7) is here selected as the lectotype; the small

female from the Persian Gulf (ZSI 4421/10) is a

paralectotype.

In 1913 Kemp identified a female from off

Madras with this species; it is identified with O.

subtilis, described below. Kemp and Chopra (1921)

assigned several specimens from Kabusa Island to

this species; one of those specimens was correctly

identified, but the others are referred to O. subtilis,

below.

Although at least part of the material (the illustrated specimen) identified with this species by

Chopra (1939) from the John Murray collections

may be referable to O. striata, described below, the

three specimens I examined appear to be typical

O. gonypetes.

Material of this species reported by me from

Madagascar (1968) proved to be referable to a third

new species, O. turbata, described below.

Thus material from various parts of the reported

range of O. gonypetes can now be referred to four

different species, all of which occur in the Indian

Ocean. Specimens from the extremes of the range

of O. gonypetes, Japan and South Africa (Manning,

1965, 1971) should be re-examined.

Three of these four species, O. gonypetes, O.

subtilis, and O. turbata comprise the gonypetes
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

complex of the gonypetes group of species in Oratosquilla, those species with an unarmed inferodistal

angle on the merus and only five teeth on the dactylus of the claw. The fourth species, O. striata, is

referable to the imperialis complex of the gonypetes

group, those species with an unarmed inferodistal

angle on the merus of the claw and with six teeth

on the dactylus. As Holthuis (1967:12) pointed out,

specimens of these species which lack claws are

extremely difficult to distinguish. The number of

teeth on the claw may prove to be variable, but

much more material will have to be available before this can be determined. Series of specimens

from a single locality would be most welcome and

might help to solve the problem.

Oratosquilla gonypetes differs from O. subtilis in

having more abdominal carinae armed, in having

a broader uro pod, with the proximal segment of

the exopod shorter than the distal, and in having

a lower, narrower lobe on the outer margin of the

inner spine of the basal prolongation of the uropod; that lobe is broader than the spine in O. subtilis, narrower than the spine in O. gonypetes.

Oratosquilla gonypetes differs from O. turbata in

similar features: the branches of the anterior bifurcation of the median carina of the carapace are

less distinct; the posterior lobe of the lateral process

of the fifth thoracic somite is slender, triangular;

more abdominal carinae are armed; and there are

dark triangles rather than squares on the fifth abdominal somite.

In addition to the difference in the armature of

the claw, O. gonypetes differs from O. striata in

having a much narrower carapace, with the anterior

width less than half the median length, a broader

rostral plate, a broader posterior lobe on the lateral

process of the fifth thoracic somite, and, when present on the telson, fewer, lower dorsal carinae and

tubercles. These dorsal carinae are well developed

in O. striata. Pigment on the fifth abdominal somite is divided into two submedian squares in O.

gonypetes; it is not divided in O. striata.

DISTRIBUTION.—Known with certainty from localities in the Indian Ocean. Records in the literature

have suggested that the range is from Japan (Manning, 1965) to southern Africa (Manning, 1971),

but these records require verification. Indian Ocean

localities include: Kabusa Island, 46-64 m, and off

Cheduba Island, 13 m, both Burma (Kemp, 1911,

1913); Persian Gulf, 86 m (Kemp and Chopra,

1921); and Gulf of Oman, 73 m on coarse sand

and shells (Chopra, 1939).

4. Oratosquilla gravieri, new species

FIGURE 6

Squilla oratorio.—Gravier, 1937:183 [part].—Dawydoff, 1952:

145 [part; not Squilla oratorio de Haan, 1844].

MATERIAL.—Baie d'Along [Yung Ha Long, 20°55'N,

1O7°O5'E], Gulf of Tonkin, North Viet Nam; Dawydoff, leg.;

Oct 1931; holotype: 1 $ , TL 92 mm (MNHNP).

DESCRIPTION.—Size moderate, total length less

than 100 mm. Body appearing smooth, minutely

punctate under magnification.

Eye (Figure 66) of moderate size, cornea bilobed,

set obliquely on stalk. Eyes not extending to end

of first segment of antennular peduncle. Corneal

index 394.

Antennular peduncle almost as long as carapace.

Dorsal processes of antennular somite produced

into anterolaterally directed spines.

Rostral plate (Figure 6a) with length and width

subequal, appearing elongate, lateral margins converging on rounded apex. Median carina absent.

Anterior width of carapace less than half median

length. Anterolateral spines of carapace (Figure 6a)

extending to, but not overreaching, base of rostral

plate. Median carina interrupted at anterior bifurcation, branches of bifurcation distinct. Intermediate carinae turning toward laterals, not extending to anterior margin.

Dactylus of claw with 6 teeth (one claw of holotype regenerating, with 8 teeth), outer margin sinuous, with low, obtuse projection basally. Dorsal

ridge of carpus (Figure 6c) undivided. Inferodistal

angle on outer face of merus produced into slender

spine.

Exposed thoracic somites (Figure 6d) each with

unarmed submedian and intermediate carinae. Lateral process of fifth somite bilobed, anterior lobe a

slender, anteriorly directed spine, posterior lobe

smaller, triangular, apex acute but not sharp, directed laterally. Lateral process of sixth somite

strongly bilobed, anterior lobe acute, slender, triangular, posterior lobe larger, triangular, apex

acute but not spined. Lateral process of seventh

somite bilobed, anterior lobe slightly shorter than

that of sixth somite, triangular, posterior lobe

larger, triangular, apex acute, unarmed.

Submedian carinae subparallel on fifth abdominal somite. Abdominal carinae spined as follows:
NUMBER 272 15

a

FIGURE 6.—Oratosquilla gravieri, new species, male holotype, TL 92 mm, Viet Nam: a, anterior

part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic

somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of uropod, ventral view.

submedian 5-6, intermediate 4-6, lateral 3-6, marginal 1-5.

Telson flattened, longer than broad. Prelateral

lobe longer than margin of lateral tooth. Denticles

(Figure 6e) subquadrate or rounded, unarmed, 4,

5, 1. Ventral surface with short postanal keel.

Uropod broad, proximal segment of exopod subequal in length to distal, with 9 movable spines on

outer margin, distalmost short, not extending to

midlength of distal segment. Lobe on outer margin

of inner spine of basal prolongation (Figure 6/)

small, narrower than adjacent spine, margin concave, apex rounded.

COLOR.—Pattern not distinctive in holotype. Posterior margins of carapace, posterior 3 thoracic and

anterior 5 abdominal somites each with line of dark

pigment: Bases of marginal teeth of telson dark.

Dark spot at articulation of segments of uropodal

exopod, pigment extending onto inner half of distal segment, endopod with distal third dark.

MEASUREMENTS.—Only specimen examined, male

holotype, TL 92 mm. Other measurements of holotype: CL 20.1, anterior width 9.1; cornea width 5.1;

antennular peduncle length 19.6; rostral plate

length 3.0, width 3.1; telson length 17.4, width 15.7.

REMARKS.—This new species belongs to the inornata complex of the perpensa group of species in

Oratosquilla, those species with an elongate rostral

plate. It differs from all species described so far in

having the dorsal processes of the antennular somite produced into sharp spines. The anterior lobe

of the lateral process of the sixth thoracic somite

is slender, as in O. perpensa.

One of the claws of the holotype is regenerating;

it bears eight rather than the normal complement

of six teeth.

ETYMOLOGY.—It seems appropriate to name this

species for the French carcinologist Ch. Gravier,

who was the first to record it from Viet Nam.

DISTRIBUTION.—Known only from the type-locality, the Baie df

 Along, Gulf of Tonkin.

5. Oratosquilla hindustanica, new species

FIGURES 7, 15e

Squilla oratorio var. perpensa Kemp, 1911:98 [part]; 1913:

70 [part; not pi. 5:figs. 57-59].

MATERIAL.—Tuticorin [08°47'N, 78°08'E], Madras, Gulf

of Manaar, India; J. Hornell, leg.; holotype [paralectotype
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 7.—Oratosquilla hindustanica, new species, female paratype, TL 69 mm, India: a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of

uropod, ventral view.

of Squilla oratorio var. perpensa Kemp]: 1 $, TL 66 mm

(ZSI). Data same; paratypes [paralectotypes of Squilla oratorio var. perpensa Kemp]: 12 $, TL 45-68 mm; 17?, TL

40-75 mm (ZSI 7037-41/10). Data same; paratypes [paralectotypes of Squilla oratorio var. perpensa Kemp]: 2 9.

TL 86-95 mm (USNM 125069; ex ZSI 7521-4/10). Data

same; paratypes [paralectotypes of Squilla oratoria var.

perpensa Kemp]: 1$, TL 67 mm; 1 9, TL 69 mm (USNM

143576).

DESCRIPTION.—Size moderate, total length less

than 100 mm. Body appearing smooth, minutely

punctate under magnification.

Eye (Figure 76) moderate to small, cornea bilobed, set obliquely on stalk. Eyes not extending

to end of first segment of antennular peduncle.

Corneal indices 309-412.

Antennular peduncle as long as, or slightly

shorter than, carapace. Dorsal processes of antennular somite produced into acute, slender lobes,

sharp but not spined, directed anterolaterally.

Rostral plate (Figures la, \be) appearing elongate, length and width subequal or width slightly

greater, lateral margins converging on broadly

rounded apex. Median carina absent.

Anterior width of carapace much less than half

median length. Anterolateral spines (Figure la)

strong, extending to or slightly beyond base of

rostral plate. Median carina interrupted at base

of anterior bifurcation, branches of bifurcation

distinct. Intermediate carinae turning toward but

not meeting laterals, not extending to anterior

margin.

Dactylus of claw with 6 teeth, outer margin of

dactylus faintly sinuous. Dorsal ridge of carpus

(Figure 7c) undivided. Inferodistal angle on outer

face of merus with sharp spine.

Exposed thoracic somites (Figure Id) with unarmed submedian and intermediate carinae, intermediates short, irregular, on fifth somite. Lateral

process of fifth somite bilobed, anterior lobe a slender spine, directed almost anteriorly, posterior lobe

triangular, apex acute, directed laterally. Lateral

process of sixth somite bilobed, anterior lobe broad,

apex trapezoidal, truncate or rounded; posterior

lobe larger, triangular, apex obtusely pointed, unarmed. Lateral process of seventh somite bilobed;

anterior lobe smaller than that of sixth somite, triangular; posterior lobe larger, triangular, apex obtusely pointed, unarmed.

Submedian carinae slightly divergent on fifth

abdominal somite. Abdominal carinae spined as

follows: submedian 5-6, intermediate 4-6, lateral

(2)3-6, usually 2-6, marginal 1-5.

Telson flattened, longer than broad. Prelateral

lobe longer than margin of lateral tooth. Denticles

(Figure le) rounded or subquadrate, 4-5, 8-9, 1,

outer submedians and outer intermediates largest.
NUMBER 272 17

Ventral surface with short postanal keel.

Uropod broad, proximal segment of exopod as

long as, or slightly longer than, distal, with 8-9

movable spines on outer margin, distalmost not

extending to midlength of distal segment. Lobe on

outer margin of inner spine of basal prolongation

(Figures 7f, I5e) small, low, narrower than adjacent

spine, margin concave, apex rounded.

COLOR.—Body light, ornamented with lines and

patches of dark chromatophores. Eyestalks with

numerous dark chromatophores, especially on ventral surface. Basal segments of antennae with dorsal and lateral patches of dark chromatophores.

Margins, carinae, and grooves of carapace outlined

with dark chromatophores. Dorsal surface of merus

of claw with longitudinal dark line, outer surface

with dark, distal patch proximal to articulation of

carpus. Propodus of claw with distal dark patch.

Exposed thoracic and all abdominal somites with

dark posterior line, with submedian and intermediate carinae outlined in dark pigment, and with

dark chromatophores middorsally. On abdomen,

area between intermediate and lateral carinae with

curved dark line, convex dorsally. Median area of

telson, curved rows of pits, and bases of marginal

teeth all dark. Midline of proximal segment of uropodal exopod dark, inner half of distal segment

black. Uropodal endopod with margins of distal

third dark, midline clear.

MEASUREMENTS.—Males, TL 45-68 mm; females,

TL 40-95 mm. Other measurements of male holotype, TL 66 mm: CL 14.9, anterior width 6.6; cornea width 4.2; antennular peduncle length 15.6;

rostral plate length 2.4, width 2.5; telson length

13.0, width 11.8.

REMARKS.—This is the common species occurring

on the coasts of "British India" that Kemp named

Squilla oratorio, var. perpensa, and it is one of several species that Kemp included in the type-series

of his variety. Unlike O. perpensa sensu stricto,

however, O. hindustanica has a long, slender rostral plate, a feature that it shares with O. inornata

and O. solicitans, as well as a truncated anterior

lobe on the lateral process of the sixth thoracic

somite. Oratosquilla perpensa, a more eastern species, which was illustrated by Kemp (1913, pi. 5:

figs. 57-59), has a very short rostral plate and a

slender, triangular anterior lobe on the lateral process of the sixth thoracic somite.

Both this species and O. solicitans resemble O.

inornata but differ from that species in having a

much smoother body; O. inornata, like O. perpensa,

has the body surface relatively deeply pitted and

eroded.

The rostral plate and basal prolongation of the

uropod of this species are similar to those observed

in some of the specimens assigned to O. solicitans

below. They are shown in Figure I5e, in contrast

with the same features observed in O. solicitans.

Oratosquilla hindustanica is very similar to O. solicitans, differing in having a more tapering rostral

plate, a smaller lobe between the spines of the basal

prolongation of the uropod, and more divergent

spines on the basal prolongation.

The specimens of this species examined, totaling

more than 30, are remarkably uniform morphologically; the rostral plates, lateral processes of the

exposed thoracic somites, and uropods are basically

the same in all of the specimens examined. For the

present, I believe it should be considered as a separate species from O. solicitans; it may prove to be

the westernmost population of that species or it

may be that both are represented in the IndoMalayan region and that the range of the two species overlaps there. Much more material, documented with observations on habitat and color in

life, will have to be studied before this can be

settled.

The close similarity of the species in the material

assigned by Kemp to O. perpensa as well as the

occurrence in that material of at least two species

with elongate rostral plates, led Chopra (1934:24,

25), who compared some of Kemp's material with

type-specimens of O. inornata, to consider that the

two species then being considered, O. inornata and

O. perpensa, to be identical. They are, in fact,

separate species which themselves are part of a

very confusing species complex.

ETYMOLOGY.—The specific name is derived from

an early name, Hindustan, applied to British India.

DISTRIBUTION.—Known with certainty only from

the type-locality, Tuticorin, Gulf of Manaar, Madras, India. Its range is yet to be determined.

6. Oratosquilla inornata (Tate, 1883)

FIGURES 8, 9

Squilla inornata Tate, 1883:51, pi. 2: figs. $a-c— Manning,

1966:95, fig. 3 [part?].
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 8.—Oratosquilla inornata (Tate, 1883), female lectotype, CL 14.3 mm, South Australia:

a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

MATERIAL.—Gulf of Saint Vincent [35°00'S, 138°05'E],

South Australia; lectotype: 1 $, CL 14.3 mm (SAM). Data

same; paralectotype: 1 9 • TL 39 mm (SAM). 12 mi NNE of

Bowen [20°01'S, 148°14'E], Queensland, Australia; 35-46 m;

Endeavour, leg.: 1 $, TL 91 mm (USNM 111379).

DESCRIPTION.—Size moderate, total length of

adults 100 mm or less. Body appearing rough, surface coarsely pitted, rugose.

Eye (Figures 8&, 96) moderate to small, cornea

bilobed, set obliquely on stalk. Eyes not extending

to end of first segment of antennular peduncle.

Corneal indices 349-414.

Antennular peduncle shorter than carapace. Dorsal processes of antennular somite produced into

acute but blunt lobes directed anterolaterally.

Rostral plate (Figures 8a, 9a) with length and

width subequal or length slightly greater, appearing elongate, lateral margins converging on flattened apex. Median carina absent.

Anterior width of carapace less than half median

length. Anterolateral spines of carapace (Figures 8a,

9a) strong, extending to but not overreaching base

of rostral plate. Median carina interrupted at base

of anterior bifurcation, branches of bifurcation distinct. Intermediate carinae turning toward but not

meeting laterals, not extending to anterior margin.

Dactylus of claw sinuous, outer margin sinuous,

with 6 teeth. Dorsal ridge of carpus (Figures 8c, 9c)

undivided. Inferodistal angle on outer face of merus

with sharp spine.

Exposed thoracic somites (Figures 8d, 9d) each

with unarmed submedian and intermediate carinae,

intermediates short, irregular, on fifth somite. Lateral process of fifth somite bilobed, anterior lobe

a slender spine, directed anteriorly, posterior lobe

smaller, triangular, apex acute but blunt, directed

laterally. Lateral process of sixth somite bilobed,

anterior lobe broad, apex rounded, posterior lobe

much larger, triangular, apex acute, unarmed. Lateral process of seventh somite bilobed, anterior lobe

smaller and more triangular than that of sixth

somite, apex acute, unarmed, posterior lobe larger,

triangular, apex acute, unarmed.

Submedian carinae parallel on fifth abdominal

somite. Abdominal carinae spined as follows: submedian 5-6, intermediate 4-6, lateral 2-6 (3-6 in

young female), marginal 1-5.

Telson flattened, longer than broad. Prelateral

lobe shorter than margin of lateral tooth. Denticles

(Figures Se, 9e) subquadrate or rounded, 4, 7-9, 1.

Ventral surface with short postanal keel.

Uropod broad, proximal segment of exopod subequal in length to distal, with 9 movable spines

on outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin of

inner spine of basal prolongation (Figures 8/, 9/)

of uropod distally erect, narrower than adjacent

spine, margin concave, apex rounded.

COLOR.—Pattern faded in all specimens examined.
NUMBER 272 19

a

FIGURE 9.—Oratosquilla inornata (Tate, 1883), male, TL 91 mm, Queensland: a, anterior part

of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic somites;

e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of uropod,

ventral view.

MEASUREMENTS.—Only male examined, TL 91

mm; only intact female, a juvenile, TL 39 mm.

Other measurements of damaged female lectotype:

CL 14.3, anterior width 6.6; cornea width 4.1; antennular peduncle broken; rostral plate length 2.4,

width 2.0; telson length 12.5, width 11.6.

REMARKS.—Oratosquilla inornata is the most

rugose of the species of the inornata complex. Its

distinctive features include the rugose surface of

the body, the elongate rostral plate with a flattened

apex, the relatively broad anterior lobe on the

lateral process of the sixth thoracic somite, and the

erect, projecting lobe between the spines of the

basal prolongation of the uropod. In an otherwise

very poor figure, Tate (1883, pi. 2: fig. 36) illustrated the relatively broad anterior lobe on the

lateral process of the sixth thoracic somite.

It is likely that most records of O. inornata in

the literature are based on other, similar species,

especially O. hindustanica or O. solicitans. Oratosquilla inornata is known with certainty from localities off southern and eastern Australia; its range

may not extend northward to Indo-Malaya or westward to India.

The larger of the two female syntypes, a damaged

adult female, is here selected as the lectotype. Both

the lectotype and the smaller female paralectotype

are in the collections of the South Australian

Museum.

Also occurring off Queensland is the species identified as O. anomala by Stephenson (1952) and

others and which is named for Stephenson below.

The concave lobe between the spines of the basal

prolongation of the uropod in O. inornata will

serve to distinguish it from O. stephensoni in which

that lobe is straight or almost convex, as in O.

interrupta.

Specimens from other eastern Australian localities from the Endeavour collection (Manning, 1966)

probably are referable to O. inornata; their identity

should be verified.

DISTRIBUTION.—Known only from Bowen, Queensland, in 35-46 m, and from the Gulf of Saint Vincent, South Australia.

7. Oratosquilla pentadactyla, new species

FIGURE 10

Squilla affinis var. intermedia Nobili, 1903:39 [part, specimens from Nias only; preoccupied by Squilla intermedia

Bigelow, 1893].

Squilla quinquedentata.—Kemp, 1913:195 [addendum, not

p. 52].—Holthuis, 1941:244 [not Squilla quinquedentata

Brooks, 1886].
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 10.—Oratosquilla pentadactyla, new species, male paratype, TL 122-5 mm, India: a,

anterior part of body; b, eye; c, carpus of claw, d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

MATERIAL.—Luaha Gundre, Nias, Indonesia; Modigliani,

leg.; syntypes of Squilla affinis var. intermedia; paratypes:

2 9 , TL 56-89 mm (MNHNP). Nias [Pulau Nias, 01°05'N,

97°35'E], Indonesia; Kleiweg de Zwaan, leg.; holotype: 1 $,

TL 124 mm (ZMA). Telok Dalam [PTelukdalem, 00°34'N,

97°49'E], Nias, Indonesia; Kleiweg de Zwaan, leg.; paratype:

1 $, TL 90 mm (ZMA). Data same; paratype: 1 9 , TL 76

mm (USNM 155309). Goenang Sitoli [Gunung Sitoli,

01°17'N, 97°36'E], Nias, Indonesia; Kleiweg de Zwaan, leg.;

paratype: 1 $, TL 138 mm (ZMA). Simaloer [Pulau Simeulue, 02°35'N, 96°05'E], W coast of Sumatra, Indonesia; E.

Jacobson, leg.; Aug 1922; paratype: 1 $, TL 121 mm (ZMA).

Kilakarai [09°14'S, 78°47'E], Ramnad District, S India;

S. W. Kemp, leg.; from fishermen's nets; 17 Feb 1913;

paratype: 1 $, TL 1225 mm (ZSI 8107/10).

DESCRIPTION.—Size moderate, total length less

than 150 mm. Body appearing smooth, lightly

pitted under magnification.

Eye (Figure 106) small to very small, cornea bilobed, set obliquely on stalk. Eyes not extending to

end of first segment of antennular peduncle. Corneal indices 413-521.

Antennular peduncle about y5 as long as carapace. Dorsal processes of antennular somite produced into blunt lobes directed anterolaterally

(occasionally sharp in dorsal view).

Rostral plate (Figure 10a) slightly longer than

broad or with length and width subequal, appearing elongate, lateral margins converging slightly on

flattened apex. Median carina absent.

Anterior width of carapace less than half median

length. Anterolateral spines (Figure 10a) strong,

extending to or beyond base of rostral plate. Median carina interrupted at base of bifurcation,

branches of bifurcation low, scarcely visible in some

specimens. Intermediate carinae turning towards

laterals anteriorly, not extending to anterior margin.

Dactylus of claw with 5 teeth, outer margin sinuous. Dorsal ridge of carpus (Figure 10c) undivided.

Inferodistal angle on outer face of merus with

sharp spine.

Exposed thoracic somites (Figure lOd) each with

unarmed submedian and intermediate carinae.

Lateral process of fifth somite bilobed, anterior

lobe a sharp, anteriorly directed spine, posterior

lobe smaller, triangular, sharp, apex directed laterally or inclined posterolaterally. Lateral process of

sixth somite bilobed, anterior lobe very slender,

posterior lobe broader, apex acute, sharp. Lateral

process of seventh somite distinctly bilobed, ante-
NUMBER 272 21

rior lobe much smaller than obtusely pointed posterior lobe, latter broader than that of sixth somite.

Submedian carinae divergent on fifth abdominal

somite. Abdominal carinae spined as follows: submedian 5-6, intermediate (3)4-6, lateral (1)2-6,

marginal 1-5, usually 5-6, 3-6, 1-6, 1-5.

Telson flattened, slightly broader than long. Prelateral lobe subequal in length to margin of lateral

tooth. Denticles (Figure lOe) rounded or subquadrate, 4, 8-9, 1. Ventral surface with strong postanal

keel.

Uropod broad, proximal segment of exopod

shorter than distal, with 8 short movable spines on

outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin of

inner spine of basal prolongation (Figure 10/) low,

much narrower than adjacent spine, margin concave, apex rounded.

COLOR.—Pattern not distinctive in preservative.

Carapace, posterior 3 thoracic and all abdominal

somites with dark posterior line. Bases of marginal

teeth of telson dark. Inner and distal margin of

proximal segment and inner half of distal segment

of uropodal exopod dark, distal half of uropodal

endopod dark. No trace of dark pigment at each

end of median carina of telson as in O. quinquedentata.

MEASUREMENTS.—Males, TL 90-122.5 mm; females, TL 56-138 mm. Other measurements of

female holotype, TL 124 mm: CL 27.0, anterior

width 12.8; cornea width 5.2; antennular peduncle

length 20.8; rostral plate length 4.0, width 3.7; telson length 23.1, width 24.9.

REMARKS.—This new species closely resembles O.

quinquedentata, and together they comprise the

quinquedentata complex of the perpensa group of

species in Oratosquilla. Specimens of O. pentadactyla were referred to O. quinquedentata by both

Kemp (1913) and Holthuis (1941) and probably by

other authors as well.

Oratosquilla pentadactyla differs from O. quinquedentata in having larger eyes, a narrower carapace, a slenderer anterior lobe on the lateral process of the seventh thoracic somite, and armed

intermediate and lateral carinae on the abdomen

anterior to the fourth somite, and it further differs

in lacking the dark anterior and posterior patches

on the median carina of the telson.

Holthuis (1941) reported on some of the material

listed above and noted some of the differences between the two species, especially those concerned

with the spination of the abdominal carinae. The

specimen from Bombay identified by Kemp (1913)

with O. quinquedentata is correctly identified; his

material from South India is referable to O. pentadactyla.

Apparently the two specimens from Nias which

Nobili (1903) identified with Squilla affinis var.

intermedia were forwarded to the Museum National

d'Histoire Naturelle at Paris in 1906; they are

syntypes of Nobili's species. Another specimen from

Nobili's syntypic series is identifiable with O. anomala, and is discussed above under the account of

that species.

ETYMOLOGY.—The specific name is from the

Greek pente ("five") plus dactylos ("finger"), alluding to the armature of the claw.

DISTRIBUTION.—Known from localities in Indonesia and from southern India. The depth range

for the species has not been recorded, but it apparently lives in relatively shallow water.

8. Oratosquilla perpensa (Kemp, 1911)

FIGURE 11

Squilla oratorio var. perpensa Kemp, 1911:98 [part]; 1913:70,

pi. 5: figs. 57-59 [part].

Squilla oratoria var. inornata.—Holthuis, 1941:248 [part; not

Squilla inornata Tate, 1883].

Squilla perpensa.—Lee and Wu, 1966:49.

MATERIAL.—Tung Rang, S Formosa [Tung Chiang, Taiwan]; trawled; S. Lee nos. 97, 169; 5 Sep 1965: \$, TL

68 mm, 1 $ , TL 82 mm (RMNH 517S). Same loc; Lee and

Wu no. 180; 9 May 1965: 1 9 , TL 66 mm <USNM 125061).

Hong Kong [Hong Kong Island, 22°15'N, 114°H'E]; G. Dennys,

leg.; lectotype: 1 9 , TL 973 mm (ZSI). Data same; paralectotypes: 13 $, TL 73-93 mm, 13 9 . TL 77-97 mm (ZSI

4851-72/9). Data same; paralectotypes: 1 $, TL 92.5 mm,

1 9 , TL 88 mm (USNM 143577). Java, Indonesia; Bleeker,

leg.: 13 , TL 70 mm (RMNH 70S). Off Irrawaddy delta,

Burma; 15°20'N, 94°55'E; 20 fms (37 m); Investigator, leg.;

paralectotypes: 1$, TL 64 mm, 19 , TL 63 mm (USNM

143575).

DESCRIPTION.—Size moderate, total length of

adults less than 100 mm. Body appearing smooth,

actually very rugose when dried or viewed under

magnification.

Eye (Figure 116) moderate to small, cornea bilobed, set obliquely on stalk. Eyes not extending

beyond end of first segment of antennular peduncle.

Corneal indices 363-429.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 11.—Oratosquilla perpensa (Kemp, 1911), male paralectotype, TL 92.5 mm, Hong Kong:

a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

Antennular peduncle almost as long as carapace.

Dorsal processes of antennular somite produced

into acute, occasionally spiniform lobes directed

anteriorly.

Rostral plate (Figure lla) broader than long,

quadrate or trapezoidal in shape, appearing short,

apex flattened. Median carina absent.

Anterior width of carapace about half median

length, not so long as carapace and rostral plate

combined. Anterolateral spines (Figure lla) strong,

extending to or slightly beyond base of rostral plate.

Median carina interrupted at base of anterior bifurcation, branches of bifurcation distinct. Intermediate carinae turning toward but not meeting

laterals anteriorly, not extending to anterior margin.

Dactylus of claw with 6 teeth, outer margin

strongly sinuous. Dorsal ridge of carpus (Figure 1 \c)

undivided. Inferodistal angle on outer face of merus

with sharp spine.

Exposed thoracic somites (Figure lid) each with

submedian and intermediate carinae, those of fifth

somite poorly developed. Lateral process of fifth

somite with anterior lobe produced into long, slender spine, directed anterolaterally, posterior lobe

smaller, triangular. Lateral process of sixth somite

bilobed, anterior lobe slender, triangular, apex

sharp, posterior lobe larger, triangular, apex sharp

but not spiniform. Lateral process of seventh somite bilobed, anterior lobe smaller than that of

sixth somite, triangular, apex sharp, posterior lobe

larger, triangular, apex sharp but not spiniform.

Submedian carinae subparallel on fifth abdominal somite. Abdominal carinae spined as follows:

submedian 5-6, intermediate 4-5, lateral 3-6, marginal 1-5.

Telson flattened, slightly longer than broad. Prelateral lobe subequal in length to margin of lateral

tooth. Denticles (Figure lie) rounded or subquadrate, unarmed, 3^4, 7-9, 1. Ventral surface with

long postanal keel.

Uropod broad, proximal segment of exopod

shorter than distal, with 8-11 (usually 9) movable

spines on outer margin, distalmost not extending

to midlength of distal segment. Lobe on outer margin of inner spine of basal prolongation (Figure

11/) small, erect, prominent, narrower than adja-
NUMBER 272 23

cent spine, margin concave, apex rounded or

angled.

COLOR.—Body appearing dusky, with concentration of pigment dorsally. Median carina, gastric

grooves, and anterior margin of carapace dark,

other carinae outlined by more diffuse lines of dark

pigment. Lateral margin of rostral plate dark.

Merus of claw with diffuse dark band dorsolaterally, propodus with distal patch of dark chromatophores. Exposed thoracic and all abdominal somites

with dark posterior line, submedian and intermediate carinae faintly indicated by lines of dark

chromatophores; lateral carinae flanked mesially by

curved, dark line. Median carina of telson with

anterior and posterior dark patches, bases of marginal teeth dark. Proximal segment of uropodal

exopod with middorsal dark line, inner half of distal segment dark. Distal half of endopod dark. Basal

prolongation of uropod with longitudinal patch of

dark pigment on inner, ventral surface.

MEASUREMENTS.—Males, TL 64-93 mm; females,

TL 63-97.5 mm. Other measurements of female

lectotype, TL 97.5 mm: CL 20.6, anterior width

10.3; cornea width 5.1; antennular peduncle length

19.8; rostral plate length 2.7, width 3.1; telson

length 17.8, width 16.3.

REMARKS.—Examination of syntypes of Squilla

oratoria var. perpensa Kemp in the collection of

the Zoological Survey of India revealed that the

series included representatives of more than one

species. One is a species from southern India with

a long rostral plate which led Chopra (1934) tc

consider O. perpensa to be a synonym of O. inornata (Tate); another is a species with a short rostral

plate, from Burma and Hong Kong, which Kemp

illustrated (1913, pi. 5: fig. 57-59). Although Kemp

(1913:70) intended perpensa to be used for the population occurring off British India, he illustrated

material from Hong Kong. In my opinion less confusion will result from the selection of a specimen

from Hong Kong, a female 97.5 mm long, as the

lectotype of S. oratoria var. perpensa, for Kemp's

monograph is still widely used to identify IndoWest-Pacific stomatopods. The population from

India is named O. hindustanica, above. Kemp's syntypes included material of a third species, named

O. solicitans, below. In all of those new species the

rostral plate is elongate; it is short, subquadrate or

trapezoidal in O. perpensa as restricted herein.

Oratosquilla perpensa, one of the three species

in the perpensa complex of the perpensa group, is

a distinctive species, with a relatively rough surface,

broad carapace, and small eyes. It differs from O.

anomala, a second species of the perpensa complex,

in having shorter anterolateral spines on the carapace and a slender, triangular anterior lobe on the

lateral process of the sixth thoracic somite; that

lobe is distinctly rectangular in O. anomala. Oratosquilla perpensa differs from O. stephensoni, the

third species of the perpensa complex, in being

much smaller and having a concave rather than a

convex or straight lobe between the spines of the

basal prolongation of the uropod.

DISTRIBUTION.—Known with certainty from localities between southern Taiwan and Burma, including Tung Kang, Taiwan, 40-50 fms (73-91.5 m)

(Lee and Wu, 1966); Hong Kong (Kemp, 1911,

1913); Java, Indonesia (Holthuis, 1941); and off the

Irrawaddy delta, Burma, 20 fms (37 m) (Kemp,

1913).

9. Oratosquilla quinquedentata (Brooks, 1886)

FIGURE 12

Squilla quinquedentata Brooks, 1886:21, 26, pi. 1: fig. 3,

pi. 2: fig. 6.—Kemp, 1913:52 [not addendum, p. 195].—

Chhapgar and Sane, 1968:45 [in key].

MATERIAL.—Arafura Sea; 09°59'S, 139°42'E; 28 fms (51

m); green mud; Challenger sta 188; 10 Sep 1874; holotype:

1$, TL 440 mm (BMNH 94.10.16.6). Gulf of Thailand;

P. Naiyanetr, leg.: 19 , TL 119 mm (USNM 125368). Bombay [18°58'N. 72°50'E], India: 1$, TL 103 mm (ZSI 7535/10).

Bombay, India; S. R. Sane, leg.; 1964: 1$, TL 59 mm

(RMNH 511S).

DESCRIPTION.—Size moderate, total length of

adults to *140 mm. Body appearing smooth, with

large, shallow pits visible under magnification.

Eye (Figure 126) small, cornea bilobed, set

obliquely on stalk. Eyes not extending to end of

first segment of antennular peduncle. Corneal indices 506-550.

Antennular peduncle more than % as long as

carapace. Dorsal processes of antennular somite

produced into blunt lobes directed anterolaterally.

Rostral plate (Figure 12a) broader than long,

subquadrate or trapezoidal, apex flattened. Median

carina absent.

Anterior width of carapace slightly less than half

median length. Anterolateral spines (Figure 12a)

strong, extending to or beyond base of rostral plate.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 12.—Oratosquilla quinquedentata (Brooks, 1886), male, TL 103 mm, Bombay: a, anterior

part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic

somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of uropod, ventral view.

Median carina interrupted anteriorly, branches of

bifurcation poorly formed, rarely distinct. Intermediate carinae converging with but not meeting

laterals anteriorly, not extending to anterior

margin.

Dactylus of claw with 5 teeth, outer margin sinuate. Dorsal ridge of carpus (Figure 12c) undivided.

Inferodistal angle of outer face of merus with sharp

spine.

Exposed thoracic somites (Figure 12d) with unarmed submedian and intermediate carinae. Lateral

process of fifth somite bilobed, anterior lobe a sharp

spine directed almost anteriorly, posterior lobe

smaller, broad and triangular, blunt apex directed

laterally. Lateral process of sixth somite bilobed,

anterior lobe slender, subtriangular or rectangular,

posterior lobe broader, triangular, apex bluntly

pointed, unarmed. Lateral process of seventh somite

distinctly bilobed, anterior lobe low, rounded or

subquadrate, much smaller than obtusely pointed

posterior lobe.

Submedian carinae divergent on fifth abdominal

somite. Abdominal carin.ie spined as follows: submedian 5-6, intermediate 4-6, lateral 4—6, marginal

1-5.

Telson flattened, longer than broad. Prelateral

lobe longer than margin of lateral tooth. Denticles

(Figure 12e) rounded, elongate, 3, 7-8, 1. Ventral

surface with strong postanal keel.

Uropod broad, proximal segment of exopod

longer than distal, with 8-9 movable spines on

outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin of

inner spine of basal prolongation (Figure 12/) well

developed but low, narrower than adjacent spine,

margin concave, apex rounded.

COLOR.—Overall appearance dusky. Antennules

with distal dark band on each segment. Carapace

with anterior and posterior margins dark, with irregular band across cardiac region, dark pigment in

U-shaped patch medially. Exposed thoracic and abdominal somites with dark posterior line and with

diffuse dark pigment dorsally, outlined laterally

with dark line lateral to submedian carina and just

above lateral carina. Broad band of dark color

across posterior half of median carina of telson,

extending onto bases of marginal teeth. Anterior

part of median carina of telson with subquadrate

black spot. Uropod with dark pigment on proximal

segment at articulation of exopod segments, and

with black pigment on most of distal half of endopod, apex lighter.
NUMBER 272 25

MEASUREMENTS.—Males, TL 59-140 mm; only

female examined, TL 119 mm. Other measurements of the female: CL 24.8, anterior width 11.8;

cornea width 4.9; antennular peduncle length 22.0;

rostral plate length 3.2, width 4.0; telson length

21.3, width 19.8.

REMARKS.—This is a relatively large and distinctive species, one of two in the quinquedentata complex of the perpensa group, those species with a

distinct inferodistal spine on the merus and five

teeth on the dactylus of the claw. It differs from O.

pentadactyla, the other species of the quinquedentata complex, in having a broader rostral plate and

carapace, smaller eyes, a broader anterior lobe on

the lateral process of the seventh thoracic somite,

and fewer armed carinae on the abdomen. The

dark pigment on the telson also is characteristic

for the species.

Kemp (1913) recorded this species from Bombay

and Kilakarai, India; his material from the latter

locality is referable to O. pentadactyla, described

above.

DISTRIBUTION.—Known with certainty from the

Arafura Sea, 51 m (Brooks, 1886), the Gulf of Thailand, and from Bombay, India (Kemp, 1913; Chhapgar and Sane, 1968). Other records in the literature,

including those of Sunier (1918), Odhner (1923),

and Tweedie (1934), require verification. Their records could be based on O.pentadactyla, which has

a similar range.

10. Oratosquilla solicitous, new species

FIGURES 13, 14, \ba-d

Squilla oratoria var. perpensa Kemp, 1911:98 [part]; 1913:70

[part, not pi. 5: figs. 57-59].

Squilla oratoria var. inornata.—Holthuis, 1941:248 [part;

not Squilla inornata Tate, 1883].

TYPE-MATERIAL.—Sandakan [05°50'N, 118°07'E], Sabah,

Malaysia; A. W. Herre, leg.; 30 Jun 1929; holotype: 1 $, TL

85 mm (USNM 125743). Singapore [01°17'N, 103°51'E],

Malaysia; Stepnani, leg.; paratypes: 6 $, TL 57-68 mm;

2 $ , TL 36.5-70mm (ZMB 12879; 2$ USNM 153951).

Singapore, Malaysia; Capt. Hutcheson, leg.; paratype (paralectotype of Squilla oratoria var. perpensa Kemp): 1 9, TL

78 mm (USNM 143578, ex ZSI 2426/1).

OTHER MATERIAL.—Anping [An-P'ing or Ampin, 23°00'N,

120°09'E], S Formosa; Haberer, leg.; Jun 1905: 3$ , TL 63-69

mm (ZMBS). Gulf of Thailand; P. Naiyanetr, leg.: 1 9, TL

85 mm (USNM 125363). Lena Sing, Mae Nam Chantaburi

[Chanthaburi, 12°29'N, 102°04'E] River, Gulf of Thailand,

Thailand; 7 May 1927: 13 . TL 68 mm (USNM 125065).

Mae Nam Chantaburi River, Tha Chalaep Harbor [Ban Tha

Chalaep, 12°30'N, 102°03'E], Gulf of Thailand, Thailand;

anchorage; R. Rofen, leg., sta 134; 24 Dec 1957: 1 $ , TL 37

mm (USNM 104719). Singapore [01°17'N, 103°51'E], Malaysia; Capt. Hutcheson, leg.; paralectotype of Squilla oratoria

var. perpensa Kemp: 1 $, TL 73 mm (USNM 156256, ex ZSI

2426/1). Sandakan [05°50'N, 118°07'E], Sabah, Malaysia; A.

W. Herre, leg.; 30 Jun 1929: 1 9, TL 96 mm (USNM 168858).

N coast of Java, Indonesia; Buitendijk, leg.; 1904: 1 9, TL

79 mm (RMNH 116S). Tandjoeng Patjinan [Tandjung Patjinan, 07°36'S, 114°02'E], Java, Indonesia; Nov 1938; paratype: 13 , TL 88 mm (RMNH 118S). Makassar [Makasar,

O5°O7'S, 119°24'E], Celebes, Indonesia; Piller, leg.; paratype:

19. TL 82 mm (RMNH 117S). Skroe [Sekru, 02°55'S,

132°14'E], New Guinea, Indonesia; Schadler, leg.; May 1897;

paratype: 1 $, TL 60 mm (RMNH 115S).

DESCRIPTION.—Size small to moderate, total

length of adults less than 100 mm. Body appearing

smooth, minutely punctate under magnification.

Eye (Figures 136, 146) of moderate size, cornea

bilobed, set obliquely on stalk. Eyes not extending

to end of first segment of antennular peduncle.

Corneal indices 290-412.

Antennular peduncle slightly shorter than carapace. Dorsal processes of antennular somite produced into acute lobes directed almost laterally.

Rostral plate (Figures 13a, 14a, 15a-d) with

length and width subequal or width slightly greater,

appearing elongate, lateral margins subparallel,

apex flattened or broadly rounded. Median carina

absent.

Anterior width of carapace less than half median

length. Anterolateral spines (Figures 13a, 14a)

strong, extending to, but usually not beyond, base

of rostral plate. Median carina interrupted at base

of anterior bifurcation, branches of bifurcation distinct. Intermediate carinae turning toward but not

meeting with laterals, not extending to anterior

margin.

Dactylus of claw with 6 teeth, outer margin

straight or slightly sinuous. Dorsal ridge of carpus

(Figures 13c, 14c) undivided. Inferodistal angle on

outer face of merus with sharp spine.

Exposed thoracic somites (Figures 13d, 14d) each

with unarmed submedian and intermediate carinae, intermediates very short on fifth somite. Lateral process of fifth somite bilobed, anterior lobe

a slender spine, directed almost anteriorly, posterior lobe rounded, directed laterally. Lateral process of sixth somite bilobed, anterior lobe large,

broad, apex truncate or slightly trapezoidal, posterior lobe larger, triangular, apex acute, bluntly
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 13.—Oratosquilla solicitous, new species, male paratype, TL 64.5 mm, Singapore: a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

pointed. Lateral process of seventh somite bilobed,

anterior lobe smaller than that of sixth somite,

triangular, posterior lobe larger, triangular, apex

acute but rounded.

Submedian carinae divergent on fifth abdominal

somite. Abdominal carinae spined as follows: submedian 5-6, intermediate 4-6, lateral (1-2) 3-6,

marginal 1-5.

Telson flattened, slightly longer than broad.

Prelateral lobe longer than margin of lateral tooth.

Denticles (Figures l$e, 14e) subquadrate, 3-4, 7-9,

1. Ventral surface with short postanal keel.

Uropod broad, proximal segment as long as or

slightly shorter than distal, with 8-9 movable spines

on outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin of

inner spine of basal prolongation (Figures 13/, 14/,

I5a-d) small to large in size, narrower or broader

than adjacent spine, margin concave, apex rounded.

COLOR.—Grooves and carinae of carapace and

posterior margins of carapace, posterior 3 thoracic,

and all abdominal somites with dark line, narrower

at intermediate carinae of first to fifth abdominal

somites. Dark pigment distally on propodus of claw

and on lateral processes of thoracic somites. Abdominal somites, especially second, each with trace

of dark middorsal patch. Telson with dark spot

on spine of median carina and at bases of submedian and intermediate marginal teeth. Uropod

with dark pigment at articulation of exopod segments, inner third of distal segment of exopod

completely dark, dista.1 third of endopod dark.

MEASUREMENTS.—Males, TL 37-88 mm; females,

TL 36.5-96 mm. Other measurements of male paratype, TL 64.5 mm: CL 14.2, anterior width 6.5;

cornea width 4.1; antennular peduncle length ca.

13.3; rostral plate length 2.2, width 3.3; telson

length 11.7, width U.S.

REMARKS.—If my interpretation of the material

listed above is correct, O. solicitans is not only one
NUMBER 272 27

a

FIGURE 14.—Oratosquilla solicitans, new species, female, TL 96 mm, Sabah: a, anterior part of

body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic somites;

e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of uropod, ventral

view.

of the most common species of the genus in the

Indo-Malayan area, but also is the most variable

in the genus. My original analysis of this material

suggested that two species could be differentiated,

one with a flattened rostral plate and one with a

broadly rounded rostral plate, and, in both, the

lobe between the spines of the basal prolongation

of the uropod was significantly larger than that

found in the very similar O. hindustanica from

India. Indeed, one of the lots examined, from

Sabah, originally contained two females which

could be separated immediately on the shape of

the rostral plate (Figures 13a, 14a, \ba,b) and this

same difference was found in two specimens {syntypes of S. oratorio var. perpensa from the same lot

from Singapore in the collection of the Zoological Survey of India which had been sent to me on

exchange on two different occasions. Further study

indicated variation in these features in other specimens from Indo-Malayan localities. Because of the

distinct possibility that more than one (as many

as four) species are represented in my limited material, I have restricted the type-series to the form

with the distinctly flattened anterior margin on

the rostral plate and the enlarged lobe between the

spines of the basal prolongation of the uropod,

illustrated in Figure IS. The specimen shown in

Figure 14 represents the form with the rounded

anterior margin on the rostral plate and the enlarged lobe on the basal prolongation.

The rostral plate and basal prolongation of the

uropod are shown for each of the four forms of

O. solicitans in Figure \ba-d and for O. hindustanica in Figure \be. As noted above, there appear to

be four distinct forms in O. solicitans, one with a

rounded rostral plate and a large uropod lobe

(Figure 15a), one with a flattened rostral plate and

a large uropod lobe (Figure 156) (solicitans sensu

stricto), one with a rounded rostral plate and a

small uropod lobe (Figure 15c), and one with a

flattened rostral plate and a small uropod lobe

(Figure 15d). These features are shown in Figure

\5e for the very similar O. hindustanica from India.

In each of these forms the anterior lobe of the

lateral process of the sixth thoracic somite is truncate and the spination of die abdominal carinae
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 15.—Rostral plate (upper row) and basal prolongation of uropod (lower row) of Oratosquilla solicitans (a-d) and O. hindustanica (e): a, form with rounded rostral plate and large

lobe between spines of basal prolongation of uropod, female, TL 96 mm, Sabah; b, form with

flattened rostral plate and large lobe between spines of basal prolongation of uropod, female

holotype, TL 85 mm, Sabah; c, form with rounded rostral plate and small lobe between spines of

basal prolongation of uropod, male, TL 68 mm, Thailand; d, form with flattened rostral plate

and small lobe between spines of basal prolongation of uropod, female, TL 85 mm, Thailand;

e, O. hindustanica, female paratype, TL 69 mm, India.

is similar: submedian 5-6, intermediate 4-6, lateral

(1-2) 3-6, and marginal 1-5. Usually the intermediate carinae of the second abdominal somite

are spined, and they are armed on the first somite

in only one specimen. The intermediate carinae

of the first and second somites are unarmed in the

types of O. solicitans, which represent the form

with the flattened rostral plate and the large

uropod lobe. (See Table 1, page 42.)

The available material is not extensive enough

to reveal any differences in eye size which might

be correlated with the observed variation in shape

of rostral plates and uropod lobes, except that the

three specimens from Anping have somewhat larger

eyes than the other material available and they

share a rounded rostral plate as well as a small

uropod lobe. They probably represent a distinct

species. Corneal indices for O. solicitans and O.

hindustanica are summarized below.

In view of the fact that the uropod lobe varies

in size and shape in O. woodmasoni from different

geographic localities between India and eastern

Australia (see account of that species, below), it is

possible that the relative size of the lobe in the

material assigned here to O. solicitans is unimportant, and that this feature can vary geographically

in this species.

If this proves to be true, then it also is possible

that most or all of the specimens with a flattened

rostral plate represent O. solicitans sensu stricto,

a species not now known to range to India, and

that the Indo-Malayan specimens with a rounded

rostral plate are identifiable with O. hindustanica,

now known to occur only off India. The possible

exceptions to this are the three large-eyed specimens from Anping, Formosa, mentioned above,

which may represent a distinct species.

Large series of O. solicitans from throughout its

range will have to be studied before this problem

can be settled. It will be particularly important to

study series from each of several localities in the

Indo-Malayan area.

ETYMOLOGY.—The name is from the Latin for

"vexing," alluding to the problems posed by the

specimens assigned to this species.

DISTRIBUTION.—Western Pacific Ocean, primarily

in the Indo-Malayan region, from Taiwan, the

Gulf of Thailand, and localities in Malaysia and

Indonesia.
NUMBER 272 29

11. Oratosquilla stephensoni, new species

FIGURE 16

Squilla anomala.—Stephenson. 1952:7; 1953:43.—Stephenson

and McNeill, 1955:245, 246 [key].—Stephenson, Williams,

and Cook, 1974:114 [listed] [not Squilla anomala Tweedie,

1935].

MATERIAL.—1.0-1.5 mi E of St. Helena Island, Moreton Bay

[27°15'S, 153°15'E], Queensland, Australia; 4.5 fms (8 m);

trawled; L. Wale, leg.; 15 Feb 1966; holotype: 1 $, TL 134

mm .(USNM 125754). Data same; paratypes: 9 $, TL 108-142

mm; 8$ , TL 109.5-149 mm (USNM 156527; 1$, 1? AM;

1^,19 QM W.5767). Data same; paratypes: 2$, TL 141-144

mm; 5 9, TL 119-147.5 mm (USNM 125752). 2 mi N of the

Pile Light, mouth of the Brisbane River, Moreton Bay,

Queensland, Australia; 6 fms {11 m); muddy; trawl; W. Stephenson and May Rees, leg.; 29 Mar 1966; paratype: 1 $, TL

122 mm (USNM 125067). 2.5 mi E of Woody Point, Moreton

Bay, Queensland, Australia; 6 fms (11 m); muddy; trawl; W.

Stephenson and May Rees, leg.; 29 Mar 1966; paratype: 1 $,

TL 93 mm (USNM 125753).

DESCRIPTION.—Size moderate, total length of

adults 150 mm or less. Body roughened, dorsal

surface coarsely pitted.

Eye (Figure 166) very small, cornea bilobed, set

obliquely on stalk. Eyes not extending to end of

first segment of antennular peduncle. Corneal indices 502-579.

Antennular peduncle shorter than carapace. Dorsal processes of antennular somite produced into

blunt lobes directed anterolaterally.

Rostral plate (Figure 16a) broader than long,

tapering anteriorly, apex truncate. Median carina

absent.

Anterior width of carapace less than half median

length. Anterolateral spines (Figure 16a) strong,

extending to base of rostral plate. Median carina

lacking well marked anterior bifurcation, area of

bifurcation indicated by depression medially. Intermediate carinae slightly turning toward laterals,

falling well short of anterior margin, extending

about to end of median carina.

Dactylus of claw with 6 teeth (7 on 1 side in 1

specimen), outer margin sinuous. Dorsal ridge of

carpus (Figure 16c) undivided. Inferodistal angle

on outer face of merus with sharp spine.

Exposed thoracic somites (Figure 16d) each with

unarmed submedian and intermediate carinae, intermediates of fifth somite reduced. Lateral process of fifth somite bilobed, anterior lobe a slender,

anterolaterally directed spine, posterior lobe broad,

rounded. Lateral process of sixth somite bilobed,

anterior lobe a slender, rounded process, posterior

lobe much larger, triangular, apex bluntly rounded,

unarmed. Lateral process of seventh somite bilobed,

anterior lobe smaller than that of sixth somite,

triangular, apex blunt, posterior lobe much larger,

triangular, apex rounded, unarmed.

Submedian carinae divergent on fifth abdominal

somite. Abdominal carinae spined as follows: submedian 5-6, intermediate 4-6, lateral 3-6, marginal

1-5.

Telson flattened, longer than broad. Prelateral

lobe shorter than margin of lateral tooth. Denticles (Figure 16e) subquadrate or rounded, 2-3,

6-11, 1, usually 3, 8-9, 1. Ventral surface with

short postanal keel.

Uropod broad, proximal segment of exopod as

long as, or slightly longer than, distal, with 7-9,

usually 8, movable spines on outer margin, distalmost not extending to midlength of distal segment.

Lobe on outer margin of inner spine of basal prolongation (Figure 16/) low, narrower than adjacent

spine, margin usually slightly convex or straight,

rarely inconspicuously concave, apex rounded.

COLOR.—Pattern not distinctive in preserved

material. Anterior and posterior margins of carapace and posterior margins of thoracic and abdominal somites dark. Bases of teeth of telson dark.

Uropod with dark spot at articulation of endopod

segments, extending onto inner half of distal segment. Distal third of uropodal endopod dark.

MEASUREMENTS.—Males, TL 93-144 mm; females,

TL 109.5-149 mm. Other measurements of male

paratype, TL 93 mm: CL 20.3, anterior width 9.7;

cornea width 3.9; antennular peduncle length 17.4;

rostral plate length 2.8, width 3.2; telson length

19.4, width 17.6.

REMARKS.—This new species, a relatively large

species of the inornata complex, perpensa group

of Oratosquilla, resembles O. anomala and O. perpensa in having a short, quadrate rostral plate as

well as in the spine formula of the abdomen. Oratosquilla stephensoni, a much larger species than

either, can be distinguished easily by the convex

or straight lobe between the spines of the basal

prolongation of the uropod; in both O. anomala

and O. perpensa that lobe is strongly concave. The

lobe in O. stephensoni resembles that of O. interrupta, but the latter species always has a turberculate rather than an undivided carpus on the claw
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 16.—Oratosquilla stephensoni, new species, male paratype, TL 93 mm, Queensland: a,

anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

as well as distinct branches on the bifurcation of

the median carina of the carapace.

Stephenson's (1952:8) comment on the shape of

the lobe on the basal prolongation of the uropod

led me to the discovery that this species was undescribed.

Oratosquilla stephensoni, which appears to be

common in the waters of Moreton Bay, Queensland, may occur together with O. inornata at localities off Queensland. Oratosquilla stephensoni is

a smoother species, and that feature, as well as its

short rostral plate and convex or straight lobe between the spines of the basal prolongation of the

uropod, should simplify its recognition if the two

species are taken together.

Some of the specimens from one of the lots

(USNM 125752) have commensal bivalves attached

to the body, usually under the carapace.

ETYMOLOGY.—It is a pleasure to name this species for W. Stephenson, University of Queensland,

who first recorded it in the literature and who

donated collections of the species to the National

Museum of Natural History, Smithsonian Institution.

DISTRIBUTION.—Known only from localities in

Queensland, including Townsville, Lindeman Island, and several localities in Moreton Bay, on

muddy bottom in depths between 5.5 and 13

meters.

12. Oratosquilla striata, new species

FIGURES 17-18

tSquilla gonypetes.—Chopn, 1939:140 [listed], 148, fig. 5

[part; not Squilla gonypetes Kemp, 1911].

?SquilIa simulans Holthuis, 1967:7 [part; male paratype only].

MATERIAL.—Persian Gulf; no other data; paratype: 1 $,

TL 34 mm (MNHNP). Gulf of Oman, off Arabian coast;

25°37'N, 56°34'E to 25°39'N, 56°34'E; 79 m; green muddy

sand; Anton Bruun Sta 262A; 1 Dec 1963; holotype: 1 $, TL

47 mm (USNM 168859).

DESCRIPTION.—Size small to moderate, total length

of adults less than 60 mm. Body appearing smooth,

actually minutely pitted under magnification.

Eye (Figure 17&) moderate to small, cornea bilobed, set slightly obliquely on stalk. Eyes not

extending to end of first segment of antennular

peduncle. Corneal indices 386-407.

Antennular peduncle as long as, or slightly

shorter than, carapace. Dorsal processes of antennular somite produced into triangular lobe, apices

acute but not sharp, directed anterolaterally.

Rostral plate (Figure 17a) slightly longer than
NUMBER 272 31

FIGURE 17.—Oratosquilla striala, new species, male holotype, TL 47 mm, Gulf of Oman: a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, telson; /, submedian teeth and denticles of telson, ventral view; g, basal

prolongation of uropod, ventral view.

broad, appearing elongate, lateral margins converging on rounded apex. Median carina absent.

Anterior width of carapace variable, slightly less

than or slightly more than half of median length,

shorter than carapace and rostral plate combined.

Anterolateral spines of carapace (Figure 17a) strong

but not overreaching base of rostral plate. Median

carina interrupted at anterior bifurcation, branches

of bifurcation usually poorly formed but distinct.

Intermediate carinae not sloping toward laterals,

not extending to anterior margin.

Dactylus of claw with 6 teeth, outer margin

faintly sinuous, with obtuse projection basally.

Dorsal ridge of carpus (Figure 17c) undivided.

Inferodistal angle on outer face of merus unarmed.

Exposed thoracic somites (Figure \ld) each with

unarmed submedian and intermediate carinae, submedians divergent, intermediates of fifth somite

short. Lateral process of fifth somite bilobed, anterior lobe a slender, sharp, anteriorly directed

spine, posterior lobe smaller, slender, triangular,

directed anterolaterally. Lateral process of sixth

somite strongly bilobed, anterior lobe large, subrectangular, posterior lobe subtriangular, apex acute

but rounded. Lateral process of seventh somite bilobed, anterior lobe smaller than that of sixth

somite, subtriangular or trapezoidal, posterior lobe

larger, triangular, apex unarmed.

Submedian carinae subparallel on fifth abdominal somite, slightly divergent anteriorly. Abdominal

carinae spined as follows: submedian 5-6; intermediate 3-6; lateral 1-6; marginal 1-5.

Telson (Figures 17e, 18) flattened, slender, slightly

longer than broad. Prelateral lobe slightly shorter

than margin of lateral tooth. Dorsal surface with

numerous longitudinal carinae, number, size, and

arrangement variable. Denticles (Figures \7e,j, 18)

4, 7-8, 1, outer submedians and inner and outer

intermediates enlarged, rounded, remainder spiniform. Ventral surface with short postanal keel.

Uropod slender, proximal segment of exopod

slightly longer than distal, with 8-9 movable spines

on outer margin, distalmost short, not extending to

midlength of distal segment. Lobe on outer margin
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of inner spine of basal prolongation (Figure 17g)

small or moderate in size, narrower than adjacent

spine, margin concave, apex rounded.

COLOR.—Eye with dark lateral patch on stalk.

Carapace with 3 distinct pigmented areas: anterior,

cardiac, and posterior; anteriormost area diffuse;

cardiac patch on midline; posteriormost area

U-shaped with chromatophores extending anteriorly

along gastric grooves to cervical groove. Carinae

and gastric grooves of carapace outlined with dark

chromatophores. Claws with patches of dark chromatophores on basis, distolaterally on merus, and

distally on propodus. Sixth and seventh thoracic

somites with dark submedian carinae, pigment

extending laterally onto posterior margin, and dark

patches lateral to intermediate carinae. Abdomen

with dark posterior line on each somite and with

dark patches medially on second somite (middorsally) and fifth somite (on posterior margin, barely

differentiated into 2 submedian spots). Intermediate

carinae of abdomen lined with dark pigment,

flanked laterally by dark patch between intermediate and lateral carina. Sixth somite dark posterolaterally. Telson pattern faded, but with traces of

curved lines of dark chromatophores on surface,

anterior and posterior dark patches on median

carina, and bases of marginal teeth dark. Uropodal

protopod with dark pigment on dorsal carina, exopod with trace of dark pigment at articulation

FIGURE 18.—?Oratosquilla striata, new species,

telson of female, TL 56 mm, Gulf of Oman

(from Chopra, 1939, fig. 5).

extending onto proximal half of distal segment.

Distal third or half of uropodal endopod dark.

MEASUREMENTS.—Only male examined, TL 47

mm; only female examined, TL 34 mm. Other

measurements of female paratype, TL 34 mm:

CL 8.1, anterior width 4.4; cornea width 2.1; antennular peduncle length 8.1; rostral plate length

1.4, width 1.3; telson length 6.9, width 6.6.

REMARKS.—Oratosquilla striata is the only member of the imperialis complex, gonypetes group, in

which the dorsal surface of the telson is ornamented with numerous, well developed longitudinal carinae. This feature alone will distinguish the

species from the other members of the imperialis

complex.

Material of 0. striata lacking claws might be

difficult to distinguish from O. gonypetes, which

occurs in the same area. In O. striata the dorsal

carinae of the telson are stronger and more numerous than in O. gonypetes (in which carinae usually

are absent). In O. striata the carinae flanking the

median carina are stronger and the carina of the

lateral tooth extends anteriorly beyond the midlength of the prelateral lobe; in O. gonypetes the

carina of the lateral tooth extends almost to the

end of the prelateral lobe. Of the other differences

between O. striata and O. gonypetes mentioned

under the account of the latter species, the difference in color pattern on the fifth abdominal somite

may be the most important; there is a single, dorsal patch of dark pigment, spreading anteriorly on

the submedian carinae, in O. striata, and a pair of

black squares on O. gonypetes.

It seems likely that at least part of the material

from the Gulf of Oman identified with O. gonypetes by Chopra (1939), particularly the illustrated

specimen, a female 56 mm long, is referable to O.

striata. Chopra's figure is reproduced here (Figure

18); the dorsal carinae of the telson are well developed and the carinae of the lateral teeth are quite

long. Only three of the original six specimens reported by Chopra remain in the collection of the

British Museum (Natural History) and that female

is not one of them. The three specimens that I

could examine appear to be referable to O. gonypetes. The John Murray station at which Chopra's

specimens were captured is situated very close to

the type-locality of O. striata.

It seems likely also that the male paratype of

O. simulans (Holthuis, 1967), from the Red Sea,
NUMBER 272 33

may be referable to O. striata. Holthuis pointed

out several differences between his female holotype and the smaller male paratype, including the

presence of dorsal carinae on the telson in the

latter.

The anterior bifurcation of the median carina

of the carapace does not appear to be interrupted

in the female paratype from the Gulf of Oman. It

is interrupted in the male holotype, on which the

dark color of the carina also is interrupted. In

both specimens the branches of the bifurcation are

present and well developed.

ETYMOLOGY.—The name is from the Latin stria

("furrow"), alluding to the ornamentation of the

telson.

DISTRIBUTION.—Known with certainty from the

Persian Gulf and the Gulf of Oman in a depth of

79 m. It may also occur in the Red Sea.

13. Oratosquilla subtilis, new species

FIGURE 19

Squilla gonypetes Kemp, 1911:96 [part]; 1913:54 [part].—

Kemp and Chopra, 1921:300 [part].

Squilla oratoria var. perpensa.—Hansen, 1926:11 [part; not

Squilla oratoria var. perpensa Kemp, 1911].

MATERIAL.—San Andreas Island, Philippine Islands; 13°38'N,

121°58'E; 50 fms (91.5 m); soft grey mud; Albatross sta

5220; 24 Apr 1908; paratype: 1$, TL 52 mm (USNM

77990). Buton Strait, off Tikola Peninsula, S Celebes Island,

Indonesia; 04°31'40"S, 122°49'42"E; 37 fms (68 m); green

mud; Albatross sta 5642; 14 Dec 1909; paratype: 1$, TL 40

mm (USNM 77989). Madura Strait; 07°25'S, 113°16'E; 56 m;

grey mud with some Radiolariae; Siboga sta 2; 8 Mar 1900;

paratype: 1$, TL 30 mm (ZMA). Off Kabusa Island [Kabosa Island, 12°49'N, 97°53'E], Lower Burma; 25-35 fms

(46-64 m); Investigator, leg.; paratypes: 1 $ , TL 39 mm; 1 9 .

TL 43 mm <ZSI, ex C32O/1). Off Yizagapatam [Visakhapatnam,

07°42'N, 83°18'E] coast, Madras, India; 20 fms (37 m); Investigator, leg.; holotype: 1 $, TL 47 mm (ZSI 7536/10).

DESCRIPTION.—Size small to moderate, total length

of adults to 52 mm. Body appearing smooth, minutely pitted under magnification.

Eye (Figure 196) small, cornea bilobed, set

obliquely on stalk. Eyes not extending to end of

first segment of antennular peduncle. Corneal

indices 412-442.

Antennular peduncle longer than carapace. Dorsal processes of antennular somite produced into

blunt lobes directed anterolaterally.

Rostral plate (Figure 19a) with shape variable,

either broader than long or slightly longer than

broad, lateral margins converging on rounded apex.

Median carina absent.

Anterior width of carapace half or slightly less

than half median length. Anterolateral spines (Figure 19a) strong, extending to or almost to base

of rostral plate. Median carina interrupted at base

of bifurcation, branches of bifurcation well developed in some specimens, scarcely visible in others.

Intermediate carinae not sloping toward laterals,

not extending to anterior margin.

Dactylus of claw with 5 teeth, outer margin

evenly curved, with obtuse projection basally. Dorsal ridge of carpus (Figure 19c) undivided. Inferodistal angle on outer face of merus unarmed.

Exposed thoracic somites (Figure I9d) each with

unarmed submedian and intermediate carinae, intermediates of fifth somite irregular, short. Lateral

process of fifth somite bilobed, anterior lobe a

sharp, anteriorly directed spine, posterior lobe

smaller, broad, obtusely subtriangular, rounded,

or subquadrate, directed laterally. Lateral process

of sixth somite strongly bilobed, anterior lobe large,

subrectangular, posterior lobe pentagonal or triangular, obtusely rounded. Lateral process of seventh somite distinctly bilobed, anterior lobe smaller

than that of sixth somite, subtriangular, posterior

lobe much larger, subtriangular, apex acute but

rounded, unarmed.

Submedian carinae subparallel on fifth abdominal somite. Abdominal carinae spined as follows:

submedian 5-6, intermediate 4-6, lateral (3)4-6,

marginal (2-3)4-5.

Telson flattened, slender, slightly longer than

broad. Prelateral lobe subequal in length to margin

of lateral tooth. Denticles (Figure \9e) subtriangular or rounded, usually unarmed, 3-5, 6-7, 1, outer

submedian and outer intermediate larger than remainder, inner submedians occasionally spiniform.

Ventral surface with short postanal keel.

Uropod (Figure 19/, g) slender, proximal segment

longer than distal, with 9-10 movable spines on

outer margin, distalmost not extending to midlength of distal segment. Lobe on outer margin

of inner spine of basal prolongation large, projecting, broader than adjacent spine, margin concave,

apex angled or rounded.

COLOR.—Carinae and grooves of carapace lined
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 19.—Oratosquilla subtilis, new species, female holotype, TL 47 mm, India: a, anterior

part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh thoracic

somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation of uropod, ventral view. Male paratype, TL 52 mm, Philippine Islands: g, uropod.

with dark pigment. Posterior margins and submedian carinae of thoracic somites lined with dark

pigment, sixth to eighth somites each with dark

patch lateral to intermediate carinae. Second and

fifth abdominal somites each with dark dorsal

patch, that on second in middle of somite; that

on fifth on posterior margin, often subdivided into

2 triangular patches. Posterior margin of each abdominal somite and submedian and intermediate

carinae lined with dark pigment, area between

submedian and intermediate carinae with dark

longitudinal line. Sixth abdominal somite with triangular black spot below lateral carinae. Median

carina of telson with 2 dark spots, anterior rectangular, posterior triangular, outer edges connected

by faint dark line. Surface pits and carinae of marginal teeth of telson dark. Distal half of proximal

segment and inner y3 of distal segment of uropod

dark (most of distal segment in some specimens).

Basal prolongation of uropod with dark line at

about midlength of inner ventral surface.

MEASUREMENTS.—Males, TL 30-52 mm; females,

TL 43^7 mm. Other measurements of male paratype, TL 39 mm: CL 8.8, anterior width 4.2; cornea

width 2.1; antennular peduncle length 9.9; rostral

plate length 1.5, width 1.4; telson length 7.2, width

7.0.

REMARKS.—This species closely resembles O.

gonypetes, and part of the material reported herein

had been identified with that species by Kemp

(1911, 1913), Kemp and Chopra (1921), and Hansen (1926). This species differs from O. gonypetes

in numerous features: the lateral carinae of the abdomen anterior to the third somite and the marginal carinae of the first somite are unarmed, the

proximal segment of the uropodal exopod is relatively longer, and the lobe on the basal prolongation of the uropod is comparatively much larger.

The long, slender proximal segment of the uropodal exopod is a good recognition character.

The specimens from Kabosa Island, Burma, apparently were taken together with O. gonypetes.

The small male from Siboga sta. 2 was identified

with O. perpensa by Hansen (1926).

ETYMOLOGY.—The name is from the Latin subtilis ("thin"), alluding to the shape of the uropodal

exopod.

DISTRIBUTION.—Known from four localities in

the Philippine Islands, Indonesia, off Burma, and

off Madras, India, in depths between 37 and 91.5 m.
NUMBER 272

35

14. Oratosquilla turbata, new species

FIGURE 20

Squilla gonypetes.—Manning, 1968b:23, fig. 7 [not Squilla

gonypetes Kemp, 1911].

Oratosquilla gonypetes.—Manning, 1970:1430 [not Squilla

gonypetes Kemp, 1911].

MATERIAL.—Bane de Pracel [17°00'S, 43°30'E], W coast of

Madagascar; 55 m; muddy sand; A. Crosnier, leg.; Jun 1959;

holotype: 1 $, TL 53 mm (USNM 124672). Data same; paratype: 1 $, TL 45 mm (USNM 155310). Data same; paratypes: 1 $, TL 51 mm; 1 $, TL 39 mm (MNHNP). Baie

Moramba [14°55'X, 47°18'E], NW coast of Madagascar; 30 m;

muddy sand; A. Crosnier, leg.; dredge; 1 Mar 1958: 1 $ postlarva, TL 15 mm (USNM 124671). Madagascar; 23°21'30"S,

43°37'57"E; 16 m; muddy sand; R. Derijard, leg., sta D-57;

28 Sep 1963; paraiype: 1 $, TL 40 mm (MNHNP).

DESCRIPTION.—Size small to moderate, total length

of adults to 53 mm. Body appearing smooth, surface punctation scarcely visible under magnification.

Eye (Figure 206) moderate to small, cornea bilobed, set obliquely on stalk. Eyes not extending

to end of first segment of antennular peduncle.

Corneal indices 396-457.

Antennular peduncle longer than carapace. Dorsal processes of antennular somite produced into

blunt lobes directed anterolaterally.

Rostral plate (Figure 20a) slightly broader than

long, appearing elongate, lateral margins converging on rounded apex. Median carina absent.

Anterior width of carapace slightly less than

half median length. Anterolateral spines (Figure

20a) strong, extending to, but usually not overreaching, base of rostral plate. Median carina interrupted at anterior bifurcation, branches of bifurcation low but distinct. Intermediate carinae not

turning toward laterals, not extending to anterior

margin.

Dactylus of claw with 5 teeth (6 on 1 side in 1

specimen), outer margin evenly curved, with obtuse projection basally. Dorsal ridge of carpus (Figure 20c) undivided. Inferodistal angle on outer face

of merus unarmed.

Exposed thoracic somites (Figure 20d) each with

unarmed submedian and intermediate carinae, intermediates of fifth somite irregular. Lateral process

of fifth somite bilobed, anterior lobe a sharp, anterolaterally directed spine, posterior lobe smaller,

broad, quadrangular or trapezoidal, directed laterally. Lateral process of sixth somite strongly bilobed, anterior lobe large, subrectangular, posterior

lobe subtriangular, apex blunt, rounded. Lateral

process of seventh somite distinctly bilobed, anterior lobe smaller than that of sixth somite, triangular, apex rounded.

Submedian carinae slightly divergent on fifth

abdominal somite. Abdominal carinae spined as

follows: submedian 5-6, intermediate 4-6, lateral

3-6, marginal 1-5.

Telson flattened, slender, appearing elongate,

length and width subequal. Prelateral lobe subequal in length to margin of lateral tooth. Denticles (Figure 20e) 4, 7, 1, outer submedian and

outer intermediate enlarged, rounded, inner submedian and some intermediates spine tipped. Ventral surface with short postanal keel.

Uropod broad, proximal segment as long as or

slightly shorter than distal, with 8-10 movable

spines on outer margin, distalmost short, not extending to midlength of distal segment. Lobe on

outer margin of inner spine of basal prolongation

(Figure 20/) small, not so broad as adjacent spine,

margin concave, apex rounded.

COLOR.—Thoracic and abdominal somites with

dark, irregular posterior line. Second abdominal

somite with dark transverse patch. Fifth abdominal

somite with pair of dark, triangular submedian

patches. Sixth abdominal somite black posterolaterally. Telson with rectangular anterior and triangular posterior dark patches along median carina.

Uropodal exopod with distal part of proximal segment, inner half of distal segment, and distal half

of endopod dark (from Manning, 1968b).

MEASUREMENTS.—Males, TL 40-53 mm; females,

TL 39-45 mm; postlarva, TL 15 mm. Other measurements of male holotype, TL 53 mm: CL 12.4,

anterior width 4.6; cornea width 2.8; antennular

peduncle length 12.8; rostral plate length 3.6, width

3.8; telson length 9.7, width 9.7.

REMARKS.—This species, the third representative

of the gonypetes complex, gonypetes group, most

closely resembles O. gonypetes, differing from it

in several features, some of which I listed in 1968:

the branches of the anterior bifurcation of the

median carina of the carapace are more distinct;

the posterior lobe of the lateral process of the fifth

thoracic somite in much broader, and is quadrangular rather than triangular in shape; intermediate

carinae anterior to the fourth abdominal somite

and lateral carinae anterior to the third are un-
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 20.—Oratosquilla turbata, new species, male holotype, TL 53 mm, Madagascar: a, anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

armed; and the dark squares on the fifth abdominal

somite are represented by dark triangles. The lobe

between the spines of the basal prolongation of

the uropod is slender, as in O. gonypetes.

Oratosquilla turbata also resembles O. subtilis

in having fewer abdominal carinae armed than

O. gonypetes, but in O. turbata the posterior lobe

of the lateral process of the fifth thoracic somite

is much broader, the uropod is broader, with a

shorter proximal segment on the exopod, and the

lobe between the spines of the basal prolongation

of the uropod is much smaller.

The male from the Derijard collection is unusual

in that one claw is armed with 5 teeth, the other

with 6. In all other respects it agrees with the remainder of the material. Whether such variation

can be expected to occur throughout the gonypetes

group remains to be determined. If so, it could

drastically alter species concepts there.

ETYMOLOGY.—The name is from the Latin turbatus ("confused").

DISTRIBUTION.—Known only from localities off

Madagascar, in depths between 16 and 55 m, on

muddy sand.

15. Oratosquilla woodmasoni (Kemp, 1911)

FIGURES 21-22

Squilla wood-masoni Kemp, 1911:99; 1913:74, pi. 5: figs.

63-65.

Squilla woodmasoni.—Manning, 1966:100, fig. 5.

Oratosquilla tweediei Manning, 1971:12, fig. 4.

Oratosquilla jakartensis Moosa, 1975:13, fig. 1.

MATERIAL.—Hong Kong [Hong Kong Island, 22°15'N,

114°H'E]; syntype of Squilla woodmasoni: 1 $, TL 78.5 mm

(ZMBS). Singapore [01°17'N, 103°51'E], Malaysia; M.W.F.

Tweedie, leg.; 1934; holotype of Oratosquilla tweediei: 1 9,

TL 107 mm (USNM 76026). Data same; paratype of Oratosquilla tweediei: 19, TL 96 mm <USNM 127449). Puri

[19°48'N, 85°51'E], Orissa coast, India; F. H. Gravely, leg.;

syntypes of Squilla woodmasoni: 1 $, TL 80 mm (USNM

125746). Data same; syntypes of Squilla woodmasoni: 1$,

TL 78 mm; 1 $, TL 73 mm (USNM 143581). Orissa State

^l'WN , 84°00'E], India; F. H. Gravely; 1886; syntypes of

Squilla woodmasoni: 1$, T L 81 mm; 1$ , T L 81 mm

(MCSN). Madras [13°05'N, 80°17'E], India; syntypes of

Squilla woodmasoni: 1 $, TL 88 mm; 1 $, TL 96 mm

(BMNH 1910.11.14.11-12). Madras, India; syntype of Squilla

woodmasoni: I $, TL 93 mm (ZSI 3098/5). Data same; syntype of Squilla woodmasoni: I $, TL 94 mm (USNM 143580).

DESCRIPTION.—Size moderate, total length of
NUMBER 272

37

a

FIGURE 21.—Oratosquilla woodmasoni (Kemp, 1911), male syntype, TL 93 mm, Madras: a,

anterior part of body; b, eye; c, carpus of claw; d, lateral processes of fifth, sixth, and seventh

thoracic somites; e, submedian teeth and denticles of telson, ventral view; /, basal prolongation

of uropod, ventral view.

adults to about 110 mm. Surface of body smooth,

highly polished.

Eye (Figure 21 b) large to moderate, cornea bilobed, set very obliquely on stalk. Eyes extending

almost to end of first segment of antennular peduncle. Anterior margin of ophthalmic somite

rounded, usually with median spinule. Corneal

indices 294-355.

Antennular peduncle as long as, or slightly

longer than, carapace. Dorsal processes of antennular somite produced into triangular lobes, apices

sharp but not spiniform, directed anterolaterally.

Rostral plate (Figure 21a) short, broader than

long, trapezoidal or subquadrate, apex truncate

or slightly rounded. Median carina absent.

Anterior width of carapace more than half median length. Anterolateral spines (Figure 21a) small,

not extending to base of rostral plate. Median

carina with at most low prominences indicating

position of bifurcation, branches of bifurcation

often indicated by pigment only, median depression usually present between branches. Intermediate carinae parallel to laterals, falling short of

anterior margin.

Dactylus of claw with 6 teeth, outer margin

strongly sinuous. Dorsal ridge of carpus (Figure

21/) irregularly tuberculate, usually with 2 large

tubercles. Inferodistal angle on outer face of merus

produced into blunt spine.

Exposed thoracic somites (Figure 2Id) with low,

unarmed submedian and intermediate carinae,

those of fifth somite scarcely if at all discernible.

Lateral process of fifth somite bilobed, anterior

lobe a short, blunt spine, directed anterolaterally;

posterior lobe smaller, triangular, apex acute but

rounded. Lateral process of sixth somite bilobed,

anterior lobe small, slender, subtriangular, apex

acute, posterior lobe much larger, triangular, apex

acute, occasionally sharp. Lateral process of seventh

somite bilobed, anterior lobe much smaller, slenderer than that of sixth somite, usually ovate or

subtrapezoidal, posterior lobe much larger, apex

acute, occasionally sharp but not spiniform.

Submedian carinae slightly divergent on fifth

abdominal somite. Abdominal carinae spined as

follows: submedian (4) 5-6, intermediate 3-6, lateral

2-6, marginal 1-5.

Telson flattened, broader than long. Prelateral

lobe subequal in length to margin of lateral tooth.

Denticles (Figure 21e) subquadrate or rounded,
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2-A, 9-10, 1. Ventral surface with short postanal

keel.

Uropod broad, proximal segment of exopod

shorter than distal, with 8-9 movable spines on

outer margin, proximal directed dorsally, distal not

extending to midlength of distal segment. Lobe

on outer margin of inner spine of basal prolongation (Figures 21/, 22) variable in size and shape,

not so broad as adjacent spine, margin concave,

apex rounded, angled, or spined.

COLOR.—Body appearing dusky, completely covered with minute dark chromatophores, concentration denser middorsally on abdomen. Posterior

margins of thoracic and abdominal somites and

bases of teeth of telson dark. Articulation of segments of uropodal exopod with prominent dark

spot. Distal half of uropodal endopod dark. Uropod bright blue in life.

MEASUREMENTS.—Males, TL 78-93 mm; females

TL 73-107 mm. Kemp (1911, 1913) mentioned

specimens as large as 109 mm. Other measurements

of male syntype, TL 78 mm: CL 15.9, anterior

width 9.4; cornea width 5.1; antennular peduncle

length 17.2; rostral plate length 2.4, width 3.1;

telson length 14.6, width 15.0.

REMARKS.—Examination of the syntypes of O.

woodmasoni has convinced me that O. tweediei

Manning, 1971, must be considered as its synonym.

The latter was based on specimens from Singapore

in which the submedian carinae of the fourth abdominal somite were armed and in which the lobe

between the spines of the basal prolongation of

the uropod was produced into sharp spines. Apparently these features vary geographically in O. woodmasoni. In Indian specimens the submedian carinae

of the fourth abdominal somite are unarmed; they

are usually armed in Indo-Malayan specimens, but

some specimens from that region may have only

one of the submedian carinae of the fourth somite

terminating in a spine. The lobe between the

spines of the basal prolongation is very low in

Indian specimens (Figures 21a, 22a) (not so low

as shown in Kemp, 1913, pi. 5: fig. 65), is produced

into a spine in most Indo-Malayan specimens (Figure 22b, d), and is produced into an angled lobe

in Australian specimens (Figure 22c). The anterior

margin of the ophthalmic somite is always broadly

rounded and usually has a median apical spinule,

but that spinule is not always developed. Finally,

dark antennules and blue uropods are characteristic

of O. woodmasoni (H. Dingle, pers. comm.); the

uropods of fresh specimens morphologically identifiable with O. tweediei clearly show blue pigment.

I believe that O. jakartensis Moosa, 1975, based

on two juveniles, 40 to 54 mm long, also is identifiable with O. woodmasoni. In O. jakartensis the

anterior margin of the ophthalmic somite is described as being notched, but it is shown as

rounded, with an apical spinule in Moosa's fig. 1

(a lapsus similar to that which I made in the origFIGURE 22.—Oratosquilla woodmasoni (Kemp, 1911), outline of basal prolongation of uropod:

a, male syntype of 5. woodmasoni, TL 78 mm, Puri; b, female holotype of O. tweediei, TL 107

mm, Singapore; c, male, TL 113 mm, Moreton Bay; d, female holotype of O. jakartensis, TL 54

mm, Indonesia (from Moosa, 1975, fig. Id).
NUMBER 272 39

inal account of O. tweediei). Further, the submedian

carinae of the fourth abdominal somite are unarmed and the lobe between the spines of the

basal prolongation of the uropod is produced into

a sharp spine (Figure 22d). Another feature by

which Moosa distinguished O. jakartensis from O.

woodmasoni, a more prominent anterior lobe on

the lateral process of the sixth thoracic somite, is

variable in the material reported here.

Apparently O. woodmasoni can exhibit considerable geographical variation, and this must be taken

into account in identifying material of this species.

DISTRIBUTION.—Indo-West-Pacinc region, from

localities in the western Pacific through the Indian

Ocean to East Africa. Little is known of its depth

range and habitat.

Two New Genera for Species Previously

Referred to Oratosquilla

Kempina, new genus

DEFINITION.—Size moderate to large. Body very

rugose, surface eroded and pitted, carinae strongly

developed. Anterior margin of ophthalmic somite

unarmed (Figure 23a, d). Eye small to very small,

cornea bilobed, set almost transversely on stalk.

Ocular scales subquadrate, separate. Carapace with

d f

FIGURE 23.—Two species of Kempina, new genus: a-c, K. mikado (Kemp and Chopra, 1921,

fig. 2); d-f, K. stridulans (Wood-Mason, 1894) (from Kemp and Chopra, 1921: fig. 1).
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

normal complement of carinae (median, intermediates, laterals, reflected marginals), median with well

developed anterior bifurcation. Mandibular palp

present. 4 epipods present. Dactylus of claw with

6 teeth. Lateral process of fifth thoracic somite

(Figures 23c, /, 24) a single slender spine, directed

laterally; slender ventral spine, directed anterolaterally, also present, visible in dorsal view. Lateral

processes of sixth and seventh thoracic somites

(Figures 23c, /, 24a) strongly bilobed. Abdomen

with 8 carinae on anterior 5 somites. Telson flattened, with median carina and carinae of marginal

teeth, lacking supplementary dorsal carinae; 3 pairs

of marginal teeth present, submedians with fixed

apices; prelateral lobes present. Basal prolongation

of uropod produced into 2 strong spines, inner

margin tuberculate or crenulate, unarmed.

FIGURE 24.—Kempina mikado (Kemp and Chopra,

1921), female, TL 115.5 mm, Japan, lateral process

of fifth thoracic somite: a, dorsal view; b, lateral

view.

TYPE-SPECIES.—Squilla mikado Kemp and

Chopra, 1921.

OTHER SPECIES.—Squilla stridulans Wood-Mason

in Alcock, 1894, and Squilla zanzibarica Chopra,

1939; the latter probably is a synonym of S. mikado

but perhaps should be considered distinct until

material of the two can be compared directly.

ETYMOLOGY.—The genus is named for Stanley

W. Kemp, whose basic studies on Indo-West-Pacific

stomatopods laid the foundation for subsequent

work on the group. The gender is feminine.

AFFINITIES.—Kempina shows strong affinities with

the Atlantic-East Pacific genus Squilla Fabricius,

1787, with which it shares numerous features: eye

size and shape, normal complement of carinae on

the body, single lateral process on the fifth thoracic

somite, mandibular palp, and six teeth on the

dactylus of the claw. It differs from Squilla in having the surface of the body much more rugose, the

carinae of the body better developed, and in the

strongly bilobed lateral processes of the six and

seventh thoracic somites. Only in the western Atlantic Squilla heptacantha (Chace, 1939) is the anterior lobe of the lateral processes of the sixth thoracic somite as comparably well developed as in

the species of Kempina, and S. heptacantha, like

Kempina, has only four epipods. Of the other

genera of squillids, only the dissimilar Pterygosquilla Hilgendorf, 1890, has such a slender, laterally directed lateral process on the fifth thoracic

somite.

Natosquilla, new genus

DEFINITION.—Size moderate. Body very smooth,

not strongly carinate. Anterior margin of ophthalmic somite (Figure 25a), usually with 3 points. Eye

large (Figure 25a), cornea indistinctly bilobed, set

obliquely on stalk. Ocular scales small, angled or

rounded, separate. Carapace with normal complement of carinae (median, intermediates, laterals,

reflected marginals), median suppressed anterior to

dorsal pit, lacking distinct anterior bifurcation.

Mandibular palp present. 4 epipods present. Dactylus of claw with 10-18 teeth (Figure 25b). Lateral

process of fifth thoracic somite (Figure 25c, d) appearing bilobed (in dorsal view), lobes separate,

anterior lobe directed anterolaterally, posterior lobe

(Figure 25d) in higher plane, directed laterally.

Lateral process of sixth somite distinctly bilobed,

that of seventh obscurely so. Abdomen with 8 carinae on anterior 5 somites. Telson flattened, with

median carina and carinae of marginal teeth, lacking supplementary dorsal carinae; 3 pairs of marginal teeth present, submedians with fixed apices;

prelateral lobes present. Basal prolongation of uropod produced into 2 strong spines, inner margin

tuberculate or crenulate, unarmed.

TYPE-SPECIES.—Squilla investigatoris Lloyd, 1907.

OTHER SPECIES.—None.

ETYMOLOGY.—The name is derived from the
NUMBER 272

41

FIGURE 25.—Natosquilla investigator is (Lloyd, 1907), male, TL 89 mm, Zanzibar: a, anterior

part of body; b, claw; c, lateral processes of fifth, sixth, and seventh thoracic somites; d, lateral

process of fifth thoracic somite, lateral view.

Latin natus ("swim"), in combination with the

generic name Squilla, alluding to the swarming

behavior of the type-species.

AFFINITIES.—Natosquilla shares several features

with Busquilla Manning, 1978, especially the enlarged eyes, the suppression of the anterior bifurcation of the median carina of the carapace, and

the shape of the lateral process of the fifth thoracic

somite. However, in Natosquilla the anterior margin of the ophthalmic somite has three points or

denticles rather than a single median spinule, the

ocular scales are much smaller, there are 10-18

rather than five teeth on the claw, and the anterior

lobes of the lateral processes of the sixth and seventh thoracic somites are greatly reduced; the lateral process of the seventh somite is obscurely

bilobed in N. invesligatoris, distinctly bilobed in

species of Busquilla.

The morphological differences in representatives

of the two genera suggest that they are not closely

related and that they independently acquired the

enlarged eyes. In Natosquilla the eye size may be

an adaptation for their observed schooling and

swarming behavior at the surface in very deep

water (Losse & Merrett, 1971). So far as is known,

however, the two species of Busquilla are bottom

dwellers, like most stomatopods; swarming or schooling behavior has not been reported for either

species.

The enlarged eyes and the large number, 10-18,

of teeth on the claw will serve to distinguish Natosquilla from all other stomatopod genera.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Range of variation of corneal indices by size in 0. solicitans and O. hindustanica

(numbers in parentheses = the mean and the number of specimens)

0. solicitans O. hindustanica

Carapace

length (mm)

11

12

13

14

15

16

17

18

19

20

21

Round rostral

plate, large

uropod lobe

(Figure 15a)

;

400

380

Round rostral

plate, small

uropod lobe

(Anping)

290

313

304

-

-

Flat rostral

plate, large

uropod lobe

(Figure 156)

340-355

(346;4)

339

369

370

-

Round rostral

plate, small

uropod lobe

(Figure 15c)

349

350

368

-

Flat rostral

plate, small

uropod lobe

(Figure \5d)

368-384

(376;2)

366

388

383

Round rostral

plate, small

uropod lobe

(Figure 15^)

309-326

(320 ;6)

335-365

(337 ;4)

332-358

(344;4)

338-353

(349; 6)

354-373

(36O;7)

375

412
Literature Cited

Ahmed, M. M.

1971. A New Species of Oratosquilla (Crustacea: Stomatopoda) from Arabian Gulf. Pakistan Journal of Scientific Industrial Research, 14(3):251—253, figure 1.

Alcock, A.

1894. On the Results of the Deep-Sea Dredging during

the Season 1890-91 (Concluded). In Natural History

Notes from H. M. Indian Marine Survey Steamer

"Investigator," Commander R. F. Hoskyn, R. N.,

Late Commanding, Series II, No. 1. Annals and

Magazine of Natural History, (6)13:400-411, 1 figure.

Bigelow, R. P.

1893. Preliminary Notes on the Stomatopoda of the Albatross Collections and on Other Specimens in the

National Museum. Johns Hopkins University Circulars, 12(106): 100-102.

Blumstein, R.

1974. Stomatopod Crustaceans From the Gulf of Tonkin

with the Description of New Species. Crustaceana,

26(2): 113-126, figures 1-10.

Brooks, W. K.

1886. Report on the Stomatopoda Collected by H.M.S.

Challenger during the Years 1873-76. The Voyage of

H.M.S. Challenger, Zoology, 16:1-116, plates 1-16.

Chace, Fenner A., Jr.

1939. Preliminary Descriptions of One New Genus and

Seventeen New Species of Decapod and Stomatopod

Crustacea. In Reports on the Scientific Results of

the First Atlantis Expedition to the West Indies,

under the Joint Auspices of the University of Havana and Harvard University. Memorias de la Sociedad Cubana de Historia Natural, 13(1):31—54.

Chhapgar, B. F., and S. R. Sane

1968. The Stomatopoda of Bombay. Journal of Biological

Sciences (Bombay), 9 (1-2)[1966]:43-46.

Chopra, B.

1934. On the Stomatopod Crustacea Collected by the

Bengal Pilot Service off the Mouth of the River

Hughli, Together with Notes on Some Other Forms.

Records of the Indian Museum, 36:17-43, figures

1-5.

1939. Stomatopoda. Scientific Reports, The John Murray

Expedition, 1933-34, 6(3): 137-181, figures 1-13.

Dawydoff, C.

1952. Contribution a l'ttude des Invertebr^s de la Faune

Marine Benthique de l'lndochine. Bulletin Biologiqtw de France et de la Belgir/ue, supplement,

37:1-158.

Fabricius, J. C.

1787. Mantissa Insectorum sistens eorum Species nuper

detectas adiectis Characteribus Genericis, Differentiis Specificis, Emendationibus, Observationibus.

Volume 1, xx+ 348 pages. Hafniae: Christ. Gottl.

Proft.

Fukuda, T.

1911. Nippon-san Kokyakurui tsuiho. Dobutsugaku Zasshi,

22:173-175, 1 plate. [Text in Japanese.]

Gravier, Ch.

1937. Stomatopodes des Cotes d'Indo-chine. Annales de

•I'Institut Oceanographique, Monaco, new series,

17(3): 175-211, figures 1-23.

de Haan, W.

1833-1850. Crustacea. In Von Siebold, Fauna Japonica

sive Descriptio Animalium, quae in Itinere per

Japoniam, jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent, Suscepto,

Annis 1823-1830 Collegit, Notis, Observationibus, et

Adumbrationibus Illustravit. i-xvii + i-xxxi + ix-xvi

+243 pages, plates A-J, L-Q, 1-55, circ. tab. 2.

Lugduni-Batavorum: A. Arnz.

Hilgendorf, F.

1890. Eine neue Stomatopoden-Gattung Pterygosquilla.

Sitzungsberichte der Gesellschaft naturforschender

Freunde zu Berlin, 1890:172-177, 1 figure.

Hansen, H. J.

1926. The Stomatopoda of the Siboga Expedition. SibogaExpeditie, monographe 35:1-48, plates 1-2.

Holthuis, L. B.

1941. The Stomatopoda of the Snellius Expedition. In

Biological Results of the Snellius Expedition, XII.

Temminckia, 6:241-294, figures 1-9.

1967. The Stomatopod Crustacea Collected by the 1962

and 1965 Israel South Red Sea Expeditions. The

Second Israel South Red Sea Expedition, Report

No. 1. Israel Journal of Zoology, 16:1-45, figures 1-7.

Kemp, S.

1911. Preliminary Descriptions of New Species and Varieties of Crustacea Stomatopoda in the Indian Museum. Records of the Indian Museum, 6(2):93-100.

1913. An Account of the Crustacea Stomatopoda of the

Indo-Pacific Region, Based on the Collection in the

Indian Museum. Memoirs of the Indian Museum,

4:1-217, 10 figures, plates 1-10.

Kemp, S., and B. Chopra

1921. Notes on Stomatopoda. Records of the Indian Museum, 22(4):297-311, figures 1-4.

Lee, Sin-Che, and Shi-Kuei Wu

1966. The Stomatopod Crustacea of Taiwan. Bulletin of

the Institute of Zoology, Academia Sinica, 5:41-58,

figs. 1-8.

Lloyd, R. E.

1907. Contributions to the Fauna of the Arabian Sea,

with Descriptions of New Fishes and Crustacea.

Records of the Indian Museum, 1:1-12-

43
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Losse, G. F., and N. R. Merrett

1971. The Occurrence of Oratosquilla investigatoris (Crustacea: Stomatopoda) in the Pelagic Zone of the Gulf

of Aden and the Equatorial Western Indian Ocean.

Marine Biology, 10(3):244-253, figures 1-5.

Makarov, R.

1971. Fauna and Distribution of Red Sea Stomatopods.

In Kovalevsky, editor, Benthos of the Shelf of the

Red Sea, pages 141-155. Kiev, U.S.S.R.: Akademia

nauk SSSR. [In Russian.]

Manning, Raymond B.

1965. Stomatopoda from the Collection of His Majesty

The Emperor of Japan. Crustaceana, 9(3): 249-262,

figures 1-2, plates 11-12-

1966. Notes on Some Australian and New Zealand Stomatopod Crustacea, with an Account of the Species

Collected by the Fisheries Investigation Ship Endeavour. Records of the Australian Museum, 27(4):

79-137, figures 1-10.

1968a. A Revision of the Family Squillidae (Crustacea,

Stomatopoda), with the Description of Eight New

Genera. Bulletin of Marine Science, 18(1): 105-142,

figures 1-10.

1968b. Stomatopod Crustacea from Madagascar. Proceedings of the United States National Museum,

124<3641):1-61, figures 1-16.

1969. Notes on Some Stomatopod Crustacea from Southern Africa. Smithsonian Contributions to Zoology,

1:1-17, figures 1-4.

1970 [1969]. Some Stomatopod Crustaceans from Tulear,

Madagascar. Bulletin du Museum National d'Histoire Naturelle, series 2, 41(6):1429-1441, figures 1-3.

1971. Keys to the Species of Oratosquilla (Crustacea:

Stomatopoda), with Descriptions of Two New Species. Smithsonian Contributions to Zoology, 71:1-16,

figures 1-4.

1976. Redescriptions of Oratosquilla indica (Hansen) and

Clorida verrucosa (Hansen), with Accounts of a

New Genus and Two New Species (Crustacea, Stomatopoda). Beaufortia, 25(318):1-13, figures 1-5.

1978. New and Rare Stomatopod Crustacea from the IndoWest-Pacific Region. Smithsonian Contributions to

Zoology, 264:36 pages, 16 figures.

Moosa, M. Kasim

1973. Type Specimens of Crustacea Formerly in the National Museum, Singapore. Journal of the Singapore

National Academy of Science, 3(2): 142-149.

1975. Notes on Stomatopod Crustacea from Seribu Islands

and Adjacent Waters with a Description of a New

Species. Penelitian Laut di Indonesia (Marine Research in Indonesia), 15:1-20, figure 1.

Nobili, G.

1903. Crostacei di Singapore. Bollettino dei Musei di

Zoologia ed Anatomia Comparata delta R. Universita de Torino, 18(455): 1-39, 1 plate.

Odhner, Teodor

1923. Indopazifische Stomatopoden. Meddelanden fr&n

Goteborgs Musei Zoologiska Avdelning, 30:1-16, figures 1-10.

Parisi, B.

1922. Elenco degli Stomatopodi del Museo di Milano.

Atti della Societa Italiana di Scienze Naturali, e del

Museo Civile di Storia Naturale, Milano, 61:91-114,

figures 1-7.

Schmitt, Waldo L.

1931. Chinese Stomatopods Collected by S. F. Light. Lingnan Science Journal, 8(1929): 127-155, plates 16-19.

Stephenson, W.

1952. Faunislic Records from Queensland, Part I: General Introduction; Part II: Adult Stomatopoda

(Crustacea). Papers, Department of Zoology, University of Queensland, 1(1):1-15.

1953. Notes on the Australian Stomatopoda (Crustacea) in

the Collections of the Queensland Museum. Memoirs of the Queensland Museum, 13(l):40-49.

Stephenson, W., and F. McNeill

1955. The Australian Stomatopoda (Crustacea) in the Collections of the Australian Museum, with a Check

List and Key to the Known Australian Species.

Records of the Australian Museum, 23(5):239-265,

figure 1.

Stephenson, W., W. T. Williams, and S. D. Cook

1974. The Benthic Fauna of Soft Bottoms, Southern Moreton Bay. Memoirs of the Queensland Museum,

17(1):73-128, figures 1-6.

Sunier, A.L.J.

1918. The Stomatopoda of the Collection of the "VissirijStation" at Batavia. Contributions a la Faune des

Indes Neerlandaisesj l(4):62-75, figures 1-4.

Tate, R.

1883. Descriptions of Some New Species of Squilla from

South Australia. Transactions and Proceedings of

the Royal Society of South Australia, 6:48-53,

plate 2.

Tweedie, M.W.F.

1934. Notes on Stomatopoda in the Raffles Museum. Bulletin of the Raffles Museum, 9:33^1.

1935. Two New Species of Squilla from Malayan Waters.

Bulletin of the Raffles Museum, 10:45-^52, plate 1.
REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION

Manuscripts intended for series publication receive substantive review within their

originating Smithsonian museums or offices and are submitted to the Smithsonian Institution

Press with approval of the appropriate museum authority on Form SI-36. Requests for

special treatment—use of color, foldouts, casebound covers, etc.—require, on the same

form, the added approval of designated committees or museum directors.

Review of manuscripts and art by the Press for requirements of series format and style,

completeness and clarity of copy, and arrangement of all material, as outlined below, will

govern, within the judgment of the Press, acceptance or rejection of the manuscripts and art.

Copy must be typewritten, double-spaced, on one side of standard white bond paper,

with 1% " margins, submitted as ribbon copy (not carbon or xerox), in loose sheets (not

stapled or bound), and accompanied by original art. Minimum acceptable length is 30 pages.

Front matter (preceding the text) should include: title page with only title and author

and no other information, abstract page with author/title/series/etc., following the established format, table of contents with indents reflecting the heads and structure of the paper.

First page of text should carry the title and author at the top of the page and an unnumbered footnote at the bottom consisting of author's name and professional mailing address.

Center heads of whatever level should be typed with initial caps of major words, with

extra space above and below the head, but with no other preparation (such as all caps or

underline). Run-in paragraph heads should use period/dashes or colons as necessary.

Tabulations within text (lists of data, often in parallel columns) can be typed on the text

page where they occur, but they should not contain rules or formal, numbered table heads.

Formal tables (numbered, with table heads, boxheads, stubs, rules) should be submitted as camera copy, but the author must contact the series section of the Press for editorial attention and preparation assistance before final typing of this matter.

Taxonomic keys in natural history papers should use the alined-couplet form in the

zoology and paleobiology series and the multi-level indent form in the botany series. If

cross-referencing is required between key and text, do not include page references within the

key, but number the keyed-out taxa with their corresponding heads in the text.

Synonymy in the zoology and paleobiology series must use the short form (taxon,

author, yearpage), with a full reference at the end of the paper under "Literature Cited."

For the botany series, the long form (taxon, author, abbreviated journal or book title, volume,

page, year, with no reference in the "Literature Cited") is optional.

Footnotes, when few in number, whether annotative or bibliographic, should be typed

at the bottom of the text page on which the reference occurs. Extensive notes must appear at

the end of the text in a notes section. If bibliographic footnotes are required, use the short

form (author/brief title/page) with the full reference in the bibliography.

Text-reference system (author/year/page within the text, with the full reference in a

"Literature Cited" at the end of the text) must be used in place of bibliographic footnotes in

all scientific series and is strongly recommended in the history and technology series:

"(Jones, 1910:122)" or ".. . . Jones (1910:122)."

Bibliography, depending upon use, is termed "References," "Selected References," or

"Literature Cited." Spell out book, journal, and article titles, using initial caps in all major

words. For capitalization of titles in foreign languages, follow the national practice of each

language. Underline (for italics) book and journal titles. Use the colon parentheses system

for volume/number/page citations. '10(2):5-9." For alinement and arrangement of

elements, follow the format of the series for which the manuscript is intended.

Legends for illustrations must not be attached to the art nor included within the text but

must be submitted at the end of the manuscript—with as many legends typed, doublespaced, to a page as convenient.

Illustrations must not be included within the manuscript but must be submitted sepa

rately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.)

can be intermixed throughout the printed text. They should be termed Figures and should

be numbered consecutively. If several "figures" are treated as components of a single laiger

figure, they should be designated by lowercase italic letters (underlined in copy) on the illustration, in the legend, and in text references: "Figure 9Jb." If illustrations are intended to be

printed separately on coated stock following the text, they should be termed Plates and any

components should be lettered as in figures: "Plate 9b_." Keys to any symbols within an

illustration should appear on the art and not in the legend.

A few points of style: (1) Do not use periods after such abbreviations as "mm, ft,

yds, USNM, NNE, AM, BC." (2) Use hyphens in spelled-out fractions: "two-thirds." (3)

Spell out numbers "one" through "nine" in expository text, but use numerals in all other

cases if possible. (4) Use the metric system of measurement, where possible, instead of

the English system. (5) Use the decimal system, where possible, in place of fractions.

(6) Use day/month/year sequence for dates: "9 April 1976." (7) For months in tabular listings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc.

Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front

matter and ALL legends, etc., at the back of the text—in the following order: (1) title page,

(2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes,

(7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.
i