RAYMOND B. MANNING A Review of the

Genus Harpiosquilla

r (Crustacea, Stomatopoda),

r

 with Descriptions of

Three Mew Species

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 36
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 36

Raymond B. Manning A Revie w of th e

Genus Harpiosquilla

(Crustacea, Stomatopoda),

with Descriptions of

Three New Species

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1969
ABSTRACT

Manning, Raymond B. A Review of the Genus Harpiosquilla (Crustacea, Stomatopoda), with Descriptions of Three New Species. Smithsonian Contributions to

Zoology, 36:1-41. 1969.—Available specimens of the large squillid Harpiosquilla

from the collections of the Australian Museum, Sydney, and the Division of Crustacea,

National Museum of Natural History, Smithsonian Institution, show that the genus

comprises seven species; three of these species are newly described. The species of

Harpiosquilla occur throughout the Indo-West Pacific region, from Japan and

Australia westward to the Red Sea and South Africa; some species, particularly

H. annandalei, H. harpax, and H. raphidea are widely distributed in the region

whereas others, including H. indica, new species (India), H. japonica, new species

(Japan), and H. stephensoni, new species (Australia), exhibit more limited distribution

patterns. All available literature is summarized, and the descriptive accounts are

accompanied by notes on biology, development, and distribution.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1969

For sale by the Superintendent of Documents, U.S. Government Printing Office

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Contents

Page

Introduction 1

Acknowledgments 1

Systematic Account 1

Genus Harpiosquilla Holthuis, 1964 1

Key to Species of Harpiosquilla 5

1. Harpiosquilla annandalei (Kemp, 1911) 6

2. Harpiosquilla raphidea (Fabricius, 1798) 9

3. Harpiosquilla japonica, new species 15

4. Harpiosquilla stephensoni, new species 17

5. Harpiosquilla melanoura Manning, 1968 21

6. Harpiosquilla harpax (de Haan, 1844) 25

7. Harpiosquilla indica, new species 33

General Remarks 36

Bibliography 37

Raymond B. Manning A Revie w of the

Genus Harpiosquilla

(Crustacea, Stomatopoda),

with Descriptions of

Three New Species

Introduction

Examination of two species of Harpiosquilla for a report on the stomatopods of Madagascar (Manning,

1968b) revealed that a single specimen of Harpiosquilla from Australia identified by me in 1966 as H.

harpax was actually an undescribed species. Through

the kindness of John C. Yaldwyn, formerly of the

Australian Museum, I was able to examine all material of Harpiosquilla in the collection of that institution. The series included H. harpax, H. melanoura,

which was described from Madagascar in 1968, and

an undescribed species, named H. stephensoni below.

Although Australian specimens of both H. harpax and

H. raphidea had been recorded in the collections of the

Australian Museum, the latter species apparently does

not occur in Australian waters. The surprising absence

of H. raphidea, supposedly the most widely distributed

species of the genus, in Australian waters, led me to

examine the series of Harpiosquilla in the Division of

Crustacea, National Museum of Natural History,

Smithsonian Institution (SI Crust.) and the collections made by the International Indian Ocean Expedition (HOE). Two other undescribed species were

found in these collections. The present review of the

species of Harpiosquilla results from the study of these

three collections.

Raymond B. Manning, Chairman, Department of Invertebrate Zoology, National Museum of Natural History,

Smithsonian Institution, Washington, D.C. 20560.

Terms and measurements used herein have been explained in detail in earlier papers (Manning, 1966,

1968b).

Acknowledgments

I am indebted to John Yaldwyn for the loan of the

extensive series of Harpiosquilla from the Australian

Museum (AM) and for promoting the exchange of

several specimens from that series with the Division

of Crustacea, Smithsonian Institution. L. B. Holthuis,

Rijksmuseum van Natuurlijke Historie, Leiden

(RMNH), made available the lectotype of Harpiosquilla harpax, without which the identity of several

of the species recognized below would have remained

in doubt. The line drawings were made by my wife

Lilly with the support of the Office of Oceanography

and Limnology, Smithsonian Institution. I also thank

Fenner A. Chace, Jr. and David L. Pawson for their

comments on the manuscript.

Systematic Account

Genus Harpiosquilla Holthuis, 1964

Alimerichthus Claus, 1871 p. 147 [p. 39 on separate; a junior

homonym of Alimerichthus Guerin-M6neville, 1855].—

Holthuis and Manning, 1964, pp. 138, 143.—Evans and

China, 1966, p. 205 [luted] [name no. 1822 on Official

Index of Rejected and Invalid Generic Names in

Zoology].

Harpiosquilla Holthuis, 1964, p. 140.—Manning, 1968a, p.

121.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DIAGNOSIS.—Eye large, T-shaped, cornea bilobed

and much broader than stalk; ocular scales separate;

carapace usually with median, intermediate, lateral

and reflected marginal carinae; median carina not bifurcate at either end; posterolateral margins of

carapace with deep excavations, anterior margin of

each excavation angled; mandibular palp present; 5

epipods present; dactylus of raptorial claw with teeth,

outer margin of dactylus usually with prominent

angled or rounded lobe in adult males; opposable margin of propodus of raptorial claw with spaced erect

spines rather than closely set pectinations; lateral process of fifth thoracic somite usually an inconspicuous

lobe, fifth somite usually also with ventral pair of

spines; lateral processes of sixth and seventh thoracic

somites not strongly bilobed, usually acute and sharp

posterolaterally; abdomen broad, with 8 pairs of

carinae on first 5 somites, 6 on the sixth, submedians

usually low, absent in some species; telson without supplementary dorsal ornamentation other than median

carina and carinae of marginal teeth; 3 pairs of marginal teeth present on telson, submedians with fixed

apices, prelateral lobes present; inner spine of basal

prolongation of uropod the longer, inner margin

crenulate or serrate.

TYPE-SPECIES.—Squilla harpax de Haan, 1844, by

original designation. The gender is feminine.

NUMBER OF SPECIES.—Seven, of which three are

described as new herein.

RANGE.—Indo-West Pacific region, from Japan and

Australia in the Pacific to the Red Sea and South

Africa. Also see General Remarks (p. 36).

REMARKS.—Until 1952, only two of the species currently assigned to Harpiosquilla, H. annandalei and H.

raphidea, were recognized by students of the group.

In that year K. K. Tiwari and S. Biswas, studying the

collections of the Zoological Survey of India, pointed

out that raphidea comprised two species, raphidea

sensu stricto and harpax, a species described from

Japan by W. de Haan in 1844. They gave the diagnostic features of the two species and included a list of

localities based on materials in the Zoological Survey.

Another species, H. melanoura, was described by me

in 1968(b), and three additional new species are described below. The genus now comprises seven species.

Most accounts in the literature are too brief to allow

us to determine which of the seven species might have

been reported by earlier workers as Squilla raphidea.

Where possible, however, I have tried to identify records in the literature and assign them to one of the

species recorded here. Sometimes the accounts give

information, either in the text or in the figures, which

makes this possible; in other cases I have been able to

examine specimens previously recorded, as in the collections of the Australian Museum which proved to

include three species, all incorrectly identified as H.

raphidea; and in some cases, notably records of H.

raphidea from China and Japan, corrections can be

made on the basis of geography. Harpiosquilla raphidea does not occur in the Red Sea and is not known to

occur north of the Philippine Islands in the Pacific

Ocean, so records from those areas must be referable

to other species. Under the accounts of both H. harpax

and H. raphidea I have included a section on "Synonymy" to explain my identification of the old records.

Early records which have not been verified by reexamination of the material must be accepted with

caution.

Harpiosquilla includes the largest known stomatopods, the largest species being H. raphidea (Fabricius),

which attains a total length of 335 mm. Their large

size and the deep posterolateral excavations on the

margins of the carapace make adult members of the

genus very easy to recognize. Another characteristic

of species of Harpiosquilla is the row of spaced, erect

spines on the opposable margin of the propodus of the

claw. These spines apparently replace the closely set

pectinations found on the propodus of all other members of the family Squillidae. Similar spines are also

found on the propodus of the two species of Bathysquilla, the only genus in the Bathysquillidae. Bathysquilla and Harpiosquilla, however, were derived from

separate stocks and apparently have developed the remarkable armature of the claw independently. The

function of the spines on the propodus is unknown;

they may aid in holding prey. Their number and arrangement may have some value as a specific character

for H. raphidea, for in that species, as pointed out by

Tirmizi and Manning (1968), there is usually no more

than one smaller spinule or denticle between the major

erect spines. In the other species there are usually

several smaller spinules and denticles between the

major spines.

Adults of Harpiosquilla show no apparent close relationship to any other genus in the family Squillidae;

the genus occupies an isolated position in the family.

On the basis of his studies on the larval development

of stomatopods, K. H. Alikunhi (1952, p. 313) made
NUMBER 3 6

the following observations: "The larvae of S.[quilla]

raphidea occupy a peculiar position. Though described

as belonging to the Alimerichthus group, they show

some conspicuous differences from the typical Alimerichthus of the 'Chloridella' group, viz.; larger size;

broad shield-like carapace and absence of free teeth

other than the terminal on the dactylus. In the number of marginal spinules on the carapace (4 + 3) these

larvae perhaps occupy an intermediate position between Alimerichthii of the 'Chloridella' and Scorpio

groups on the one hand and the typical Alima larvae

on the other." Since that was written, each of these

groups referred to above have been recognized as distinct genera. Thus Harpiosquilla, on the basis of

larvae, may occupy a position intermediate between

(a) Clorida and Cloridopsis, and other small-eyed

genera, and (b) Oratosquilla and its allies. Its precise

relationship to any of these genera cannot be determined with available information.

There is no fossil record of the genus.

That H. harpax is generically distinct from Squilla

was recognized perhaps as long as 100 years ago by

the zoologist William Stimpson; he used the generic

name Prolepta on the label of the specimen collected

by the North Pacific Exploring Expedition. As far as

I can determine, that name has not been published.

Inasmuch as we know relatively little about the

biology of stomatopods in general, we are fortunate in

having some basic information available on the early

ontogeny and postlarval development of Harpiosquilla, primarily as a result of the work of K. H.

Alikunhi and K. B. Nair in India. Possibly this is due

to the easily recognizable facies of Harpiosquilla

adults. It is ironic that we cannot determine with certainty which of the species of Harpiosquilla they

studied. Nair (1941) reported on extrusion of eggs and

embryological development in a Harpiosquilla from

Madras, and Alikunhi, in a long series of studies on

larvae culminating in his extensive work on postlarval

development of stomatopods published in 1967, examined material from Madras and other localities. Both

H. harpax and H. raphidea could occur there, so I

have here referred their papers to whichever name was

used in their works.

The characteristic larvae of Harpiosquilla, first reported by Claus (1871) as Alimerichthus, are discussed in some detail by Alikunhi (1952, 1959). They

are easily recognized by the broad carapace with one

or two strong, laterally directed lateral spines. Alikunhi (1967) has also given a very detailed account

of growth changes in the postlarvae. He was able to

hold one specimen in the laboratory for 282 days; the

specimen underwent 14 molts.

All of the descriptions given here are based on subadults or adults. I have seen no very young specimens

of any species and can only predict some possible differences in major morphological features between juveniles and adults; Alikunhi (1967) has noted some

characteristics of juveniles. The eyes of juveniles are

undoubtedly larger than those of adults; within the

size range of the specimens examined the corneal indices of smaller specimens are always smaller than those

of larger specimens (Table 1); this is the normal pattern found also in other squillids. The body carinae of

young specimens are not so well developed as in adults;

this could cause a problem in recognizing small specimens of H. harpax, for example, in which the submedian carinae of the abdomen of adults are at best low

and poorly developed. In H. stephensoni, new species,

the lateral process of the fifth thoracic somite of the

smallest specimens examined is not as markedly angled

nor does it project as far laterally as in the largest specimens; thus the lateral spine on that somite, characteristic of both H. annandalei and H. raphidea, may be

but a blunt lobe in juveniles.

Alikunhi (1967, p. 903) made the following observations on postlarvae and early juveniles of

Harpiosquilla:

. . . the early post-larva is characterised by the absence of

anterolateral spines on the carapace and the presence of

terminally articulated submedian spines on the telson. Though

rudimentary spines appear on the anterolateral corners of

the carapace during the second or third post-larval stage, the

terminal articulation of the submedian spines of the telson

continues to be present even after the ninth post-larval moult.

In the species of the nepa group the terminal articulation

of the submedian spines disappears with the first post-lai.al

moult; i.e., it is present only during the first post-larval stage.

In this feature also H. raphidea differs from S. nepa or S.

woodmasoni but resembles S. latreiUei (Chloridella group).

Adults are easily recognized by the presence of an

inflated median carina on the telson; often the submedian carinae of the sixth abdominal somite are also

inflated in adults of both sexes. Adult males of most

species may be distinguished by the presence of a prominent angled or rounded projecting lobe on the outer

margin of the dactylus of the raptorial claw; the bases

of some of the teeth of the claw may also be enlarged.

The chelae and telsons of both sexes have been figured

where possible.
The color pattern of preserved specimens is a good

specific character, particularly the coloration of the

uropod. In H. melanoma the distal segment of the uropodal exopod is entirely black. In H. annandalei the

same segment is black with a white midrib and this

species also has distinctive paired submedian spots on

the telson. In H. harpax, H. indica, new species and H.

stephensoni, new species, only the inner half of the

distal segment of the uropodal exopod is dark, but

there are other less distinctive differences in color pattern. I have not been able to determine with certainty

the color pattern of the uropod in either H. japonica,

new species, or H. raphidea.

The armature of the abdominal carinae appears to

be a very good specific character. In H. annandalei, for

example, the submedian carinae of both the fifth and

sixth abdominal somites are armed posteriorly; in all

other species only the submedians of the sixth somite

are provided with spines. Similarly, in H. raphidea and

H. annandalei the intermediate carinae of all six abdominal somites are spined posteriorly, but in H. indica

and H. japonica, the intermediate carinae of the first

and second somites are unarmed and different patterns

of abdominal armature are present in the other three

species. In general, the number of armed carinae increases with age.

The relative distinctness of the submedian carinae

of the last three thoracic and first five abdominal

somites also may be of help in the recognition of species.

In H. raphidea these carinae are strongly developed,

more so than in any of the other species. The submedian carinae of the body in both H. japonica and

H. stephensoni are not so well developed, being lower

and less well defined, but they are still readily distinguishable. In H. harpax the submedian carinae are

very low, poorly defined, and often can be detected

only when the dorsal surface is dried. The submedian

carinae are completely absent in both H. indica and

H. melanoura; the latter species is also the only one

which lacks a median carina on the carapace.

The marginal denticles of the telson are sharper and

better defined in H. annandalei, H. harpax, H. indica,

new species, and H. melanoura than in the remainder

of the species.

The eyes of each species are of different sizes and

these differences are reflected in the corneal indices,

summarized for all species in Table 1. Because of the

change in the index with size I have separated the

indices in the table by arbitrary subgroups based on

rarapace length. The overall range given at the bottom

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Corneal Indices of species of Harpiosquilla

Carapace

length

in (mm)

11-15

16-20

21-25

26-30

31-35

36-40

41-45

46-50

51-55

^6-60

Overall

range

H. annandalei

Range

280-291

260-287

-

-

-

-

-

-

-

240-291

No.

1

3

6

-

-

-

-

-

-

-

—

Mean

262

285

278

-

-

-

-

-

-

-

—

H. raphidea

Range No.

-

-

-

_

-

1

1

1

475-504 3

380-530 -

Mean

-

-

-

-

-

410

486

469

489

—

Carapace

length

in (mm)

11-15

16-20

21-25

26-30

31-35

36-40

41-45

46-50

51-55

56-60

Overall

range

H. japonica

Range No. Mean

_ _ _

_ _ _

1 328

325-337 2 331

_

_ _

_

_

_

325-337 -

H. stephensoni

Range

_

-

-

-

349-392

-

396-420

-

410-425

336-434

No.

_

-

1

3

1

2

1

2

—

Mean

_

-

-

336

364

366

408

434

418

—

Carapace

length

in (mm)

11-15

16-20

21-25

26-30

31-35

36-40

41-45

46-50

51-55

56-60

Overall

range

H. melanoura

Range No.

1

237-246 3

1

1

-

-

-

_ _

- _

229-271 -

Mean

229

240

252

271

-

-

-

_

_

—

H. harpax

Range

251-292

263-301

269-311

269-320

305-358

294-344

-

-

_

_

251-388

No.

9

4

10

18

16

5

1

1

_

_

—

Mean

275

283

290

306

327

323

329

356

_

-

—

H.

indica

_

-

-

-

290

-

-

-

-

-

290
NUMBER 3 6

of each column includes records of eye size given in

the literature.

One feature of telson morphology is helpful in

species recognition, particularly when more than one

species is present in a sample. The carinae of the marginal teeth are dorsally tuberculate or nodulose in all

species, and the known species show a wide range of

size and prominence of this feature. In H. raphidea

the tubercles are large, inflated, and few in number;

the opposite extreme can be seen in H. harpax and H.

melanoura in which the tubercles are small, often

sharp, and very numerous. Harpiosquilla annandalei,

H. indica, new species, H. japonica, new species, and

H. stephensoni, new species, fall between these two extremes; in H. annandalei the tubercles are large, low,

and sparse. Specimens of H. stephensoni, new species,

and H. harpax (or any of those with very fine tuberculation) can always be separated on the basis of the

tuberculation of the marginal carinae. This feature is

shown in the plates in which the telsons of each species

have been illustrated. The species, if placed in a series

based on the number of tubercles, would be arranged

as follows: H. raphidea (few tubercles), H. annandalei, H. japonica, H. stephensoni, H. indica, H. harpax, and H. melanoura (many tubercles). The marginal carinae of the telson are sharpest in H. annandalei, H. harpax, and H. melanoura.

Another feature which has not been used previously

as a specific character is the relative length of the

lateral and marginal carinae of the telson. In adults of

H. raphidea both carinae are inflated and may fuse

completely. In the remainder of the species they are

sharper and distinct throughout their length. The marginal carina of H. melanoura is almost three times as

long as the lateral carina; in H. harpax it is more than

twice but less than three times as long as the lateral

carina, and in the remainder of the species it is less

than twice as long. It seems to be shortest in H. indica,

for in the only known specimen of that species the

marginal carina is no more than one and one-half

times as long as the lateral carina. The species with the

longest marginal carinae, H. harpax and H. melanoura, can be distinguished from all other species on

the basis of this character. k

The ventral keel of the eighth thoracic somite seems

to have limited value as a specific character; it is sharp

in annandalei, rounded in the remainder of the species.

Two of the species, H. annandalei and H. raphidea,

differ from the other five in having the intermediate

carinae of the thoracic somites armed posteriorly; the

remaining species in the genus have unarmed intermediate carinae on the thoracic somites.

The species seem to fall into two natural groups

based on the shape of the rostral plate. In H. harpax,

H. indica, new species, H. melanoura, and H.

raphidea the rostral plate is elongate, anteriorly tapering to a slender median projection. In H. annandalei,

H. japonica, new species, and H. stephensoni new

species, the plate is short, with the length and width

subequal or the width greater, and the apex is rounded

or obtusely angled, lacking a noticeable median

projection.

The key to species is based on adults.

Key to Species of Harpiosquilla

1. Submedian carinae distinct on posterior 3 thoracic and first 5 abdominal somites. (Carapace

with median carina.) 2

1'. Submedian carinae indistinct or lacking on posterior 3 thoracic and first 5 abdominal somites.

(Rostral plate longer than broad, with apical projection; raptorial dactylus of male with

prominent angular projection on outer margin; fifth thoracic somite laterally acute; ventral keel of eighth thoracic somite rounded; intermediate carinae of thoracic somites

unarmed posteriorly; submedian carinae of fifth abdominal somite unarmed posteriorly. ).5

2(1). Intermediate carinae of thoracic somites and first abdominal somite spinous posteriorly.

(Raptorial dactylus with 8 teeth; fifth thoracic somite laterally acute; intermediate

carinae of second abdominal somite spinous posteriorly.) 3

2'. Intermediate carinae of thoracic somites and first abdominal somite unarmed posteriorly.

(Rostral plate as broad as long, without apical projection; ventral keel of eighth thoracic

somite rounded; submedian carinae of fifth abdominal somite unarmed posteriorly; marginal carina of telson less than twice as long as lateral carina.) 4

3(2). Rostral plate as broad as long, without apical projection; raptorial dactylus of adult male

without angular projection on outer margin; ventral keel of eighth thoracic somite sharp;

submedian carinae of fifth abdominal somite spinous posteriorly; size moderate, total

length less than 150 mm. (Marginal carina of telson less than twice as long as lateral

carina; distal segment of uropodal exopod black with white midrib.). .. . 1. H. annandalei

356-607 O - 69 - 2
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3'. Rostral plate longer than broad, with long apical projection; raptorial dactylus of adult

male with prominent angular projection on outer margin; ventral keel of eighth thoracic

somite rounded; submedian carinae of fifth abdominal somite unarmed posteriorly;

size large, total length up to 335 mm 2. H. raphidea

4(2')- Raptorial dactylus with 8 teeth; intermediate carinae of second abdominal somite unarmed posteriorly; size moderate, total length less than 175 mm. (Fifth thoracic somite

rounded laterally.) 3. H. japonica, new species

4'. Raptorial dactylus with 7 teeth; intermediate carinae of second abdominal somite spinous

posteriorly; size large, total length up to 315 mm. (Raptorial dactylus of adult male

with prominent angular projection on outer margin; distal segment of uropodal exopod

dark colored on inner half.) 4. H. stephensoni, new species

5(1'). Carapace without median carina; distal segment of uropodal exopod entirely black. (Raptorial dactylus with 8 teeth; no submedian carinae on thoracic and first 5 abdominal

somites; intermediate carinae of first abdominal somite unarmed posteriorly; marginal

carina of telson more than twice as long as lateral carina; size moderate, total length

less than 170 mm.) 5. H. melanoma

5'. Carapace with median carina; distal segment of uropodal exopod dark colored on inner

half only 6

6(5'). Raptorial dactylus with 8 teeth; indistinct submedian carinae on thoracic and first 5

abdominal somites; intermediate carinae on second abdominal somite spinous posteriorly;

marginal carina of telson more than twice as long as lateral carina; size moderate to

large, total length up to 248 mm 6. H. harpax

6'. Raptorial dactylus with 9 teeth; no submedian carinae on thoracic and first 5 abdominal

somites; intermediate carinae on second abdominal somite unarmed posteriorly; marginal carina of telson less than twice as long as lateral carina; size moderate, total length

of mature individuals 150 mm or more 7. H. indica, new species

1. Harpiosquilla annandalei (Kemp, 1911)

FIGURES 1-3

Squilla annandalei Kemp, 1911, p. 99; 1913, p. 92, pi. 7 (figs.

78-80).—Sunier, 1918, p. 71.—Kemp and Chopra, 1921,

pp. 298 [listed], 307.—Chopra, 1934, p. 27, fig. 2; 1939,

p. 159, fig. 7.—Foxon, 1939, p. 259 [discussion of larva].—

Serene, 1954, pp. 6, 8 [listed].—Stephenson and McNeill,

1955, p. 255 [key].—Holthuis, 1964, p. 140 [listed].—

Manning, 1968a, p. 121 [listed].

Squilla annendalei.—Alikunhi, 1952, pp. 267, 269 [discussion

of larvae; erroneous spelling].

Harpiosquilla annandalei.—Manning, 1965, p. 250, pi. lla.—

Lee and Wu, 1966, p. 51, figs. 6A-B.—Manning, 1968b,

p. 14 [key].—Tirmizi and Manning, 1968, p. 31 [discussion].

MATERIAL.—1 $ , 71 mm; Tung Kang, Taiwan;

74-93 m; S. Lee, S. K. Wu; 5 March 1965; SI Crust.

113649.—1 4, 107 mm; 2$ , 109-117 mm; Andaman

Sea, Mergui Archipelago; 09°13'N, 97°51'E; 58-60

m; ANTON BRUUN Cruise 1, Sta. 20; IIOE; 23 March

1963; SI Crust. 125350.—3 9, 72-93 mm; Bay of

Bengal, off Burma; 19°32'N, 92°52'E; 53 m; ANTON

BRUUN Cruise 1, Sta. 49; IIOE; 6 April 1963; SI

Crust. 125348.—1 $ , 99 mm; Gulf of Oman, off Iran;

25°45-50'N, 57°O7'E; 92-95 m; clay, mud, sand,

minute gastropod shells; ANTON BRUUN Cruise 4B,

Sta. 255A; IIOE; 30 November 1963; SI Crust.

125349.-1$, 97 mm; Gulf of Oman, off Iran;

26°10-13'N, 57°02'E; 55-64 m; green mud; ANTON

BRUUN Cruise 4B, Sta. 256A; IIOE; 30 November

1963; AM.—2^, 129-135 mm; Gulf of Oman, off

Arabia; 25°12'N, 56°47-51'E; 206 m; grey soft mud;

ANTON BRUUN Cruise 4B, Sta. 263A; IIOE; 2 December 1963; SI Crust. 125716.

DIAGNOSIS.—Size moderate, adults with TL less

than 150 mm; antennular peduncle longer than carapace and rostral plate combined; corneal indices ranging between 240 and 291 (Table 1); rostral plate (Figure 2) with length and width subequal or width

greater, apex rounded or obtusely angled, lacking

slender anterior projection; carapace with median

carina; opposable margin of propodus of claw with

smaller spines and denticles between largest spines;

dactylus of claw with 8 teeth, outer margin of dactylus

lacking prominent angular projection in adult males:

fifth thoracic somite (Figure 3) acutely angled or

spined laterally; posterior 3 thoracic somites with

submedian and intermediate carinae, intermediates

with posterior spines; ventral keel of eighth thoracic
NUMBER 3 6

somite sharp; all six abdominal somites with submedian carinae, abdominal carinae spined as follows:

submedian 5-6, intermediate 1-6, lateral 1-6, marginal 1-5; denticles 5-7, 10-11, 1, inner submedians

largest, rounded, remainder spiniform; marginal

carina less than twice as long as carina of lateral tooth;

postanal carina short, not extending halfway between

anus and posterior margin; distal segment of uropodal

exopod black with white midrib.

COLOR.—Second and third segments of antennular

peduncle with proximal dorsal black spot and distal

black ring, proximal spot fainter on third segment;

dark transverse bar present dorsally between ophthalmic and antennular somites; medan portion of rostral

plate outlined in dark pigment; carapace with anterolateral margins, posterior median margin, grooves, and

carinae outlined with dark pigment; merus of claw

with inner subdistal black spot and with proximal and

distal dark spot on dorsal depression; posterior 3 thoracic and all 6 abdominal somites with dark posterior

line; submedian and intermediate carinae of carapace

darker than adjacent surface; second abdominal

somite with transverse dark bar middorsally; first and

third to fifth abdominal somites each with traces of

broken transverse bar; telson with pair of proximal,

submedian dark circles, margin darker than center;

pits, arranged in concentric rows on dorsum of telson,

dark; distal segment of uropodal exopod black with

white midrib; distal half of uropodal endopod with

diffuse dark pigment. The overall color pattern is

shown on Figure 1.

SIZE.—Males, TL 71-135 mm; females, TL 72-117

mm. Males of TL 106-123 mm and females of TL

43-115 mm have been recorded in the literature.

Other measurements of a male, TL 107 mm: carapace length 22.1; cornea width 8.5; antennular peduncle length 26.5; rostral plate length 2.7, width

3.3; raptorial propodus length 27.9; fifth abdominal

somite width 20.0; telson length 17.8, width 16.8.

DISCUSSION.—Harpiosquilla annandalei can be

distinguished readily from all other species in the genus

by a variety of characters, including the length of the

antennular peduncle (longer than the carapace and

rostral plate combined), the sharp ventral keel on the

eighth thoracic somite, the armed submedian carinae

of the fifth abdominal somite, and the color pattern

of the telson and uropod (Figure 1). None of these

features is found in any other species of Harpiosquilla

now known.

FIGURE 1.—Harpiosquilla annandalei (Kemp). Male, TL

135 mm, Gulf of Oman: dorsal view.
8

Harpiosquilla stephensoni, new species, and H. japonica, new species, also have a short rostral plate but

in neither of those species are the intermediate carinae

of the thorax or the submedian carinae of the fifth

abdominal somite armed posteriorly. Harpiosquilla stephensoni, new species, and H. raphidea also have an

acute or even spiniform lateral process on the fifth

thoracic somite; H. raphidea is the only other species

of the genus in which the intermediate carinae of the

last three thoracic somites are armed posteriorly and

in which all intermediate carinae of the abdomen also

are spined.

FIGURE 2.—Harpiosquilla annandaUi (Kemp). Male, TL

135 mm, Gulf of Oman: anterior portion of carapace.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Corneal indices of H. annandalei are summarized

in Table 1. The eyes of the specimens reported herein

are similar to those of H. harpax of comparable size,

but the overall range of corneal indices, 240-291, in

contrast to that observed in H. harpax, 251-358, suggests that the eyes of H. annandalei are larger than

those of H. harpax.

Chopra (1934, 1939) has pointed out that the anterior lobes of the lateral processes of the sixth and

seventh thoracic somites are variable in shape and in

some specimens may be subdivided secondarily. Similar variation in the shape of the anterior lobes of these

somites is evident in the specimens reported herein.

The lateral processes in H. annandalei (Figure 3) are

more strongly bilobed than in any other species of the

genus.

Harpiosquilla annandalei also has a higher and

sharper median carina on the telson than any other

species of Harpiosquilla; Lee and Wu (1966) commented on the shape of the carina in their specimens

from Taiwan, and they noted that the carina, in lateral

view, is convex anteriorly, concave posteriorly. The

relative thinness of the median carina does not seem to

be affected by age, and the median carina is not particularly inflated in either large males or large females.

One specimen, from ANTON BRUUN Sta. 49, a female

TL 72 mm, had nine teeth on the dactylus of one

claw, the normal complement of eight teeth on the

other dactylus.

In all specimens examined the intermediate carinae

of the last three thoracic somites and of all six abdominal somites were armed posteriorly.

SEXUAL DIMORPHISM.—This species does not exhibit

secondary sexual dimorphism to the extent found in

other species such as H. harpax and H. raphidea. The

two males reported herein from ANTON BRUUN Sta.

263A are the largest recorded for the species and they

show no marked sexual dimorphism. The outer edge

of the dactylus of the claw in the largest male, TL 135

mm, is faintly sinuous but not markedly different in

shape from the claw of the females. The teeth on the

raptorial claw of these two males are slightly more

inflated basally than in smaller males or in any females,

but the difference in shape is relatively minor.

BIOLOGY.—Virtually nothing is known about the

biology of this species. In general, it occurs in deeper

water, 15-206 meters, than either H. harpax or H.

raphidea. Specimens collected from the ANTON BRUUN

were taken on mud or grey mud and sand. Serene
NUMBER 3 6 9

(1954) recorded the species from sandy mud, in 15-25

meters, in the Baie de Cauda, Viet-Nam. The specimen

reported by Chopra (1939) was taken on green mud

in 201 meters in the Gulf of Oman. Harpiosquilla

annandalei was taken together with H. melanoura al

ANTON BRUUN Sta. 20, in 5&-60 meters off the Mergui

Archipelago and with H. harpax at ANTON BRUUN

Sta. 256A, in 55-64 meters in the Gulf of Oman.

DEVELOPMENT.—Larval stages of H. annandalei

FIGURE 3.—Harpiosquilla annandalei (Kemp). Male, TL

135 nun, Gulf of Oman: lateral processes, of fifth, sixth,

and seventh thoracic somites.

have not been identified with certainty. Foxon (1939)

identified larvae from the Red Sea, Arabian Sea, and

Gulf of Oman with Alima multispinus (Claus) She

noted that the larvae could be separated into two

groups, referred to by her as typical and atypical, and

suggested that they might be the larvae of H. raphidea

and H. annandalei (note that neither of these species

is known to occur in the Red Sea proper and H. raphidea is not known with certainty to occur in the Gulf

of Oman.)

Alikunhi (1952, p. 269) pointed out that "while

the atypical form is perhaps identical with S. {Alima)

raphidea, the typical larva might well belong to the

very closely related S. annendalei [sic]." Ingle and

Delia Croce (1967) corroborate Alikunhi's identification of Foxon's atypical larvae with H. raphidea. This

seems highly unlikely if the atypical larvae came from

the Red Sea rather than the Arabian Sea; however, the

larvae identified with H. raphidea by Alikunhi could

be the larvae of H. harpax, and this might explain the

presence of this larval form in the Red Sea,

TYPE.—The four syntypes are in the Zoological

Survey of India [Indian Museum], Calcutta.

TYPE-LOCALITY.—Gulf of Martaban, Burma, in 53-

67 fathoms.

DISTRIBUTION.—Indo-West Pacific region, from

scattered localities between Japan and the Gulf of

Oman in depths between 15 and 206 meters. Records

for adults in the literature are as follows: JAPAN:

Hayama, Sagami Bay (Manning, 1965). TAIWAN:

Tungkang market, 40-50 fms (Lee and Wu, 1966).

VIET-NAM: Baie de Cauda, 15-25 m (Serene, 1954).

JAVA SEA: 30-35 fms (Sunier, 1918). BAY OF BENGAL:

off Kabusa Island, Mergui Archipelago, 33 fms (Kemp

and Chopra, 1921); Gulf of Martaban, Burma, 53-67

fms (Kemp, 1911,1913); Sandheads, mouth of Hughli

River [Hooghly], India, less than 20 fms (Chopra,

1934). ARABIAN SEA; Gulf of Oman, 201 m (Chopra,

1939). Some of the larvae reported from the Red Sea,

Arabian Sea, and Gulf of Oman by Foxon (1939) may

be referable to this species.

2. Harpiosquilla raphidea (Fabricius, 1798)

FIGURES 4-9

Squilla raphidea Fabricius, 1798, p. 416.—Latreille, 1802, p.

279; 1828, p. 471 .—Milne-Edwards, 1837, p. 524.—Gibbes,

1850, p. 199 (discussion; p. 35 on separate].—Miers,

1880, p. 27 [part].—Preudhomme de Borre, 1882, p. cxi.—

de Man, 1888, p. 296.—Thallwitz, 1892, p. 55.—Hender-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

son, 1893, p. 453.—Sharp, 1893, p. 108.—Thurston, 1895,

p. 120.—de Man, 1898, p. 694.—Lanchester, 1900, p. 264;

1901, p. 553.—Nobili, 1903, p. 38.—Lanchester, 1906, p.

133.—Kemp, 1913, p. 88 [part]; 1915, p. 172; 1918, p.

297.—Sunier, 1918, p. 70, fig. 3.—Hansen, 1921, p. 79 —

Kemp and Chopra, 1921, p. 298 [listed].—Smedley, 1927,

p. 231.—Roxas and Estampador, 1930, p. 101.—Chopra,

1934, p. 27 [part].—Tweedie, 1934, p. 40.—Foxon, 1939,

p. 259 [discussion of larvae].—Gravely, 1941, p. 74.—

Holthuis, 1941, p. 256 [part].—Nair, 1941, p. 544, figs. 3 -

10, 21, 23-30, pi. 29 (figs. 34-35), pi. 30 (figs. 37-38, 40)

[embryology].—Chacko, 1942, p. 404.— Alikunhi and

Aiyar, 1942, p. 56, figs. 3-4 [larvae]; 1943, p. 81 [growth].

—Alikunhi, 1950, p. 103 [discussion; larvae]; 1952, p. 265,

fig. 9 [larvae].—Tiwari and Biswas, 1952, p. 356, figs. 3a,

c, e.—Ingle, 1963, p. 14 [key].—Zimsen, 1964, p. 653

[listed].—Holthuis, 1964, p. 140 [listed].—Alikunhi, 1965,

p. 35 [postlarva; listed].—Manning 1968a, p. 121 [listed].

Squilla raphidia.—Bosc, 1801-1802, p. 122 [erroneous

spelling].

Squilla mantis (B) var. major Lamarck, 1818, p. 187 [in

synonymy].—Deshayes and Milne-Edwards, 1838, p. 322;

1839, p. 373 [in synonymy].

Squilla Mantis.—Latreille, 1818, p. 6, pi. 324 [not Squilla

mantis (Linnaeus)].

Squilla Raphidia.—Bosc, 1830, p. 95 [erroneous spelling].

Squilla Raphidea.—Herklots, 1861, p. 152 [p. 39 on separate;

listed].

Alimerichthus Claus, 1871, p. 147, pi. 8 (fig. 30) [larva; p. 39

on separate].—?Brooks, 1886, p. 96 [discussion].

Alimerichthus pyramidalis Lanchester, 1903, p. 457 [larva].—

Gurney, 1946, p. 161 [references].

Squilla raphidea var. africana Balss, 1910, p. 8, fig. 2b.

Alima pyramidalis.—Gurney, 1946, p. 159 [references].

Squilla (Alima) raphidea.—Alikunhi, 1952, p.269 [discussion

of larvae].

?Harpiosquilla raphidea.—Alikunhi, 1967, p. 894, figs. 114-

125, pi. 2 [larvae and young; part?].

Harpiosquilla raphidea.—Chhapgar and Sane, 1968, p. 45

[key.]—Manning, 1968b, p. 14 [key].—Tirmiri and Manning, 1968, p. 31, fig. 12.

not Squilla raphidea.—Berthold, 1845, p. 47; 1847, p. 29 —

White, 1847, p. 84.—Bigelow, 1894, pp. 511 [key], 535 —

Tattersall, 1906, p. 166.—Balss, 1910, p. 8, fig. 2a.—Parisi,

1922, p. 103.—Komai, 1927, p. 323.—Komai, Akatsuka,

and Ikari, 1927, p. 295.—Komai and Ikari, 1929, p. 121.—

Gravier, 1930, p. 525.—Serene, 1937, p. 68 [listed].—

Komai, 1938, p. 268.—Suvatti, 1938, p. 52.—Chopra, 1939,

p. 158—Liu, 1949, p. 43, pi. 6 (figs. 15-17).—Anonymous, 1949, p. 843, fig. 2421.—Barnard, 1950, p. 851, figs,

lc, g.—Suvatti, 1950, p. 132.—Dawydoff, 1952, p. 145.—

Serene, 1953, p. 507; 1954, pp. 6, 8, 62, pi. 4 (figs. 1-6).—

Millard and Harrison, 1954, p. 176 [listed].—Day and Morgans, 1956, p. 306 [listed].—Utinomi, 1956, p. 91, pi. 46

(fig. 2) [colored figure].—Yamaji, 1959, fig. on p. 68.—

Utinomi, 1960, p. 114, pi. 57 (fig. 7) [colored figure].—

Chuang, 1961, pp. 181, 206, pi. 81 (fig. 4), lowest fig. on

pi. 82.—Stephenson, 1962, p. 34.—Crosnier, 1965, p. 61

[listed]. [All = H. harpax (de Haan).]

not Squilla raphidea.—?Tate, 1883, p. 48.—Stephenson,

1952, p. 4; 1953, p. 43. [All probably = H. stephensoni,

new species.]

not Chloridella raphidea.—Rathbun, 1902, p. 55. [=//.

japonica, new species.]

not Squilla raphidea.—Fukuda, 1913, pp. 70, 72, fig. on p. 71.

[=H. japonica, new species.]

not Squilla raphidia.—Torralbas, 1917, p. 621 [p. 81 on separate] [=Squilla empusa Say, 1818].

not Squilla raphidoea.—Serene, 1939, p. 349 [erroneous spelling; ?=//. harpax].

not Squilla raphidea.—Stephenson and McNeill, 1955, p. 239

[a mixture of H. harpax (de Haan), H. melanoura Manning, and H. stephensoni, new species].

MATERIAL.—1 9 , 216 mm; Sandakan, North Borneo, Indonesia; Herre, col.; 30 June 1929; SI Crust.

125360.—29, 250-277 mm; Thailand; Hugh M.

FIGURE 4.—Harpiosquilla raphidea (Fabricius). Female,

TL 250 mm, Thailand: anterior portion of body.
NUMBER 36 11

FIGURE 5.—Harpiosquilla raphidea (Fabricius). Male, TL

310 mm, off Bombay: raptorial claw (pro pod us length

72.5 mm).

FIGURE 6.—Harpiosquilla raphidea (Fabricius). Female,

TL 250 mm, Thailand: raptorial claw (propodus length

69.7 mm).

Smith; SI Crust. 93736.—1 9, 270 mm; Pak Poon

[Pakpoon, western side of Gulf of Siam, Nakon Sritamarat Province], Thailand; Hugh M. Smith; 19 October 1923; SI Crust. 69530.—1 S , CL 56.0 mm; same;

SI Crust. 106388.—1 $, 310 mm; taken with a 60

mile radius of Bombay, India; C. L. Kaufmann; SI

Crust. 104363.

DIAGNOSIS.—Size large, adults with TL to 335 mm;

antennular peduncle shorter than carapace; corneal

indices ranging between 380 and 530 (Table 1);

rostral plate (Figure 4) longer than broad, with

slender anterior projection; carapace with median

carina; upper margin of propodus of claw usually with

1 smaller spine or denticle between largest spines;

dactylus of claw with 8 teeth, outer margin with prominent angular projection in adult males (Figure 5);

fifth thoracic somite (Figure 7) with lateral spine;

posterior 3 thoracic somites with submedian and intermediate carinae, intermediates spined posteriorly;

ventral keel of eighth thoracic somite rounded; all 6

abdominal somites with submedian carinae, abdominal

carinae spined as follows: submedian 6, intermediate

1-6, lateral 1-6, marginal 1-5; denticles rounded,

4-5, 8, 1; marginal carina about twice as long as carina

of lateral tooth, usually fused with it in large specimens; postanal carina extending about halfway between anus and posterior margin; proximal portion

of inner half of distal segment of uropodal exopod with

dark pigment.

COLOR.—The color pattern is not well marked in

any of the specimens examined. Posterior 3 thoracic

and first 5 abdominal somites each with dark posterior

line; fifth thoracic somite with dark anterior line;

telson with proximal pair of dark spots; uropodal

exopod with dark spot, primarily on inner surface,

on distal third of proximal segment; some dark pigment on inner proximal third of distal segment of

uropodal exopod, but inner half of distal segment not

completely dark.

Tirmizi and Manning (1968, p. 33) noted that a

fresh specimen from Karachi, West Pakistan was colored as follows:

. . . ocular peduncles light pink; posterior border of carapace with a black band; antennal scale yellowish, outlined

with dark pigment; claw with merus pink, also marked with a

greenish yellow patch; distal end of propodus with bright

yellow streak; thorax and abdomen appearing speckled; last

3 thoracic and first 3 abdominal somites pink, last 3 abdominal somites more cream-colored; tips of spines of last 4

abdominal somites yellow; posterior margin of first 4 abdominal somites black; carinae of telson bluish, apices of teeth

yellow; telson with pair of submedian yellow-brown spots;
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

uropod lightly marked with yellow and black, inner half of

distal segment of exopod grayish, outer half yellow.

SIZE.—Only intact male examined, TL 310 mm;

females, TL 216-277 mm. Males ranging between

197-275 mm TL and females ranging between 160-

288 mm TL have been recorded in the literature.

Kemp (1913) recorded a specimen of TL 335 mm but

did not specify its sex.

Other measurements of a female, TL 277 mm:

carapace length 57.0; cornea width 12.0; antennular

peduncle length 42.0; rostral plate length 13.6, width

8.2; raptorial propodus length 81.4; fifth abdominal

somite width 59.8; telson length 52.4, width 54.3.

DISCUSSION.—Several of the features of H. raphidea allow its ready separation from the other known

species of the genus. It shares with H. annandalei the

sharp lateral process on the fifth thoracic somite

(Figure 7), and the armed intermediate carinae of

the posterior three thoracic and all six abdominal

somites; these two features will distinguish it from

FIGURE 7.—Harpiosquilla raphidea (Fabricius). Female

TL 250 mm, Thailand: lateral processes of fifth, sixth, and

seventh thoracic somites.

the five other species. It differs from H. annandalei in

several characters over and above color pattern and

size of adults, for the submedian carinae of the fifth

abdominal somite are unarmed, the rostral plate is

elongate, with a slender median projection (Figure 4),

and adult males of H. raphidea have the large angled

prominence on the outer margin of the dactylus of the

claw (Figure 5). As noted under the Discussion of

H. annandalei (p. 7), the color pattern of that species

is quite distinctive.

Harpiosquilla raphidea is the largest known stomatopod; Kemp (1913) recorded a specimen from Bombay with a total length of 335 mm. In very large

specimens, as in the male, TL 310 mm, recorded here

from Bombay, the marginal teeth of the telson are

markedly compressed and inflated. The lobe between

the spines of the basal prolongation of the uropod

becomes comparatively smaller with increasing size.

Inflation of the carinae of the marginal teeth of the

telson in adults smooths out the normal dorsal rugosity

of these carinae.

Two authors have recorded specimens with nine

rather than eight teeth on the dactylus of the raptorial

claw. Lanchester (1906) noted that a male from

Patani Bay, Halmahera, had nine spines on the left

dactylus and Roxas and Estampador (1930) reported

that their specimens had nine, rarely eight teeth, on

the claw. The latter authors may have been dealing

with another species, for their specimens were under

160 mm TL.

The structure of the dorsal organ of this species

was discussed by Hansen (1921).

SYNONYMY.—Inasmuch as H. raphidea was the

only species in the genus which was recognized between 1798 and 1911, when Kemp described annandalei, and which was considered to include all specimens distinct from annandalei between 1911 and 1952,

it is not surprising to find that the literature is quite

confused; many of the records of raphidea in the literature could refer to one or more of the six other

species recognized herein. Some of the records here

referred to raphidea may include that species and other

species as well.

Many of Kemp's (1913) records were verified by

Tiwari and Biswas (1952) in their account in which

H. harpax (de Haan) was shown to be distinct from

H. raphidea; the latter authors also verified identifications of some material reported on by Chopra

(1934). These verified records have been used to com-
NUMBER 3 6 13

pile the distribution of the species as recorded in the

literature.

Balss (1910) distinguished his variety africana from

raphidea sensu stricto by the shape of the rostral plate

(which he illustrated), the presence of well-developed

keels on the abdomen and telson, and the well-developed teeth of the telson. His raphidea sensu stricto is

apparently H. harpax whereas his africana is raphidea

sensu stricto.

Sunier (1918) has clearly illustrated a specimen of

raphidea; the distinctive lateral process of the fifth

thoracic somite and telson are easily discernible.

In his account of postlarval development and growth

of raphidea, Alikunhi (1967) notes that according to

the features discussed by Tiwari and Biswas (1952) his

juveniles should be referred to harpax. He mentions,

however, that at one stage the juveniles have all of the

intermediate carinae of the abdomen provided with

spines. This is a feature always found in H. raphidea

and rarely found in H. harpax, for the intermediate

carinae of the first abdominal somite are usually unarmed in H. harpax. It seems likely that Alikunhi was

dealing with specimens of both species. Adults of both

species occur on the eastern coast of India.

As noted under Distribution (p. 15), most records

in the literature require verification.

SEXUAL DIMORPHISM.—As in most other species of

Harpiosquilla, large males exhibit marked changes in

the morphology of the raptorial claw and the telson; to

a lesser extent the carinae of the sixth abdominal

somite are also affected. The outer margin of the dactylus of the claw in the male (Figure 5) has a large,

angled protuberance and the bases of the teeth on the

inner margin of the dactylus, particularly the second

through the fifth tooth, have basal inflations. The

carinae of the sixth abdominal somite, especially the

submedians, are inflated in large adults of both sexes,

but they are more inflated in the males; in some specimens the posterior spines of the submedian teeth are

obsolete. In adult males (Figure 8*), the median carina

of the telson is so inflated that it may obliterate the

distal dorsal spine. The carinae of the marginal teeth

are also inflated; usually the marginal and lateral

carinae of the telson are completely fused. Claws (Figures 5, 6) and telsons (Figures 8, 9) of both males and

females have been illustrated. Although the carinae of

the telson are also inflated in adult females, as in the

specimen from Thailand, TL 250 mm, illustrated here,

the extent of inflation is never so great as in males of

similar size.

There is no information available on the size at

which males begin to exhibit dimorphism, but Alikunhi

(1967, p. 903) noted that: "Sexual maturity is not

FIGURE 8.—Harpiosquilla raphidea (Fabricius). Male, TL

310 mm, off Bombay: posterior portion of body (telson width

55.8 mm).

FIGURE 9.—Harpiosquilla raphidea (Fabricius). Female,

TL 250 mm, Thailand: posterior portion of body (telson

width 48.1 mm).

356-607 O - 69 - 3
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

attained even after the 15th moult after metamorphosis. Other specimens up to 137.0 mm in length

were also found immature in March."

Kemp (1918) noted that one male from Tale Sap,

Thailand, TL 200 mm, lacked an angular projection

on the claw.

BIOLOGY.—As in the other species of Harpiosquilla,

very little is known about the biology of H. raphidea.

It occurs in relatively shallow water; the few depth

records in the literature indicate that it has been taken

in depths between 1 and 28 fathoms. Kemp (1918)

noted that it was taken in brackish water, specific

gravity 1.0085, at Tale Sap, Thailand.

Several authors have reported specimens from local

fish markets, so it may have economic importance in

some areas and it may be collected in connection with

commercial fishing operations. It has been recorded

from markets in Karachi, West Pakistan (Tirmizi and

Manning, 1968), Singapore (Tweedie, 1934), Sarawak (Tiwari and Biswas, 1952), Divisoria Market,

Manila (Kemp, 1915), and markets in Manila (Roxas

and Estampador, 1930) ; the specimens from Thailand

reported by Kemp (1918) were taken in fishermen's

nets.

Smedley (1927) commented that raphidea was the

commonest of local mantis shrimps at Morib, on the

west coast of the Malay peninsula.

Sunier (1918) found fragments of the shrimp

Penaeus indicus Milne-Edwards in the stomachs of

two specimens and Brachyurum [Brachyura?] in the

stomach of a third.

There is no information available on other stomatopods or invertebrates associated with this species.

Postlarval development and changes with age have

been discussed in some detail by Alikunhi (1967).

DEVELOPMENT.—Nair (1941) discussed egg extrusion and embrological development in a species from

Madras, India, identified by him as Squilla raphidea;

he also made observations on the embryology of Oratosquilla holoschista (Kemp), O. nepa (Latreille),

and O. woodmasoni (Kemp).

Alikunhi (1952) has given a detailed account of a

larval form identified by him as raphidea; Alikunhi

and Aiyar (1942, 1943) and Alikunhi (1950) had

previously reported the metamorphosis of a larva into

postlarval and juvenile raphidea. In an abstract of a

paper presented at the Symposium on Crustacea sponsored by The Marine Biological Association of India,

Alikunhi (1965) noted that raphidea was among eighteen species from the Madras coast, the postlarval development of which was well known.

In the paper presented at the Symposium and published in Part II of the Proceedings of the Symposium

on Crustacea, Alikunhi (1967) has presented results of

the most detailed study of growth in stomatopods ever

made. Aspects of postlarval growth are reported in

varying detail for eighteen species, including raphidea.

Of particular interest is the fact that the movable submedian teeth of the postlarva are retained beyond that

stage in Harpiosquilla. Although Alikunhi identifies his

material with raphidea and mentions that by the

twelfth molt the intermediate carinae of the first and

second abdominal somites are armed posteriorly, a

characteristic of adult raphidea, he notes on p. 904

that according to the diagnostic characters of harpax

and raphidea the reared material actually belongs to

harpax. It seems highly likely that both species were

represented in his material.

Alikunhi (1952) identified his larva with Alimerichthus sp. Claus (1871) from the Indian Ocean and with

Alimerichthus pyramidalis Lanchester, 1903 from

North Male Atoll, Maldive Islands; Lanchester had

identified his larva with Claus' species and with an

Alimerichthus reported by Brooks (1886, pp. 95-98).

Brook's larvae, however, were from the Atlantic and

are not likely to be the larvae of an Indo-West Pacific

species.

Alikunhi noted that his series of larvae from Madras

included two distinct forms. He identified his typical

larva with H. raphidea and stated (1952, p. 267) :

The post-larva obtained by moult from one such larva more

or less agrees with S. raphidea. However, since the early postlarvae in most cases do not show the full adult characters and

since S. raphidea and S. annendalei [sic] are very closely

related, it is a possibility that the atypical larva probably

belongs to the latter species. This is, however, just a surmise

which is not based on any positive evidence.

Alikunhi (1952, p. 269) also noted that Erichthus

multispinus Claus, identified with H. raphidea by

Foxon (1939), "appears to be a very different species,

probably not related to S. raphidea, even though it also

possesses laterally directed spinules on the carapace.

Borradaile's [1907] specimens are perhaps identical

with the specimen figured by Claus."

TYPE.—Probably not extant. Zimsen (1964) did not

locate the type in her comprehensive survey of

Fabrician types.

TYPE-LOCALITY.—Oceano Indico D. Daldorff. I. K.

Daldorff collected material for Fabricius at Fredericks-
NUMBER 3 6 15

nagore, on the eastern coast of India. Fredericksnagore,

located on the Hooghly River, north of Calcutta, is

now known as Serampore (Webster's Geographical

Dictionary, Rev. Ed., 1966).

DISTRIBUTION.—Indo-West Pacific region, from

Indonesia and the Gulf of Siam westward to East

Africa. Most of the records in the literature probably

refer to H. harpax, H. indica, new species, or H.

melanoura as well as H. raphidea; from most accounts

it is not possible to determine which species was actually being recorded. The following records are

probably referable to H. raphidea: SOUTH CHINA

SEA: NE of Tegal, 06°53'N, 110°21'E, 6.5-9 fms

(Sunier, 1918). INDONESIA: Indischen Archipel (de

Man, 1898); Java Sea, between Singapore and Surabaya [Surabaja, Java] (? part; de Man, 1898); Borneo

Bank, Makassar Strait, 15 fms; Madura Strait, ca. 16

fms; Java Sea, N. of western Madura, 7.5 fms; vicinity

of Madura; near Tandjong Priok, Batavia [Djakarta],

less than 1 fm; same, 6-7 fms; beyond western entrance

of Banka Strait, Sumatra, 5-8 fms; outside of estuary

of Rokan River, Sumatra, 18-20 fms; Bagan Si Api

Api, eastern Sumatra; east coast of Sumatra (all

Sunier, 1918). MALAYSIA : Singapore (Kemp, 1913;

Tiwari and Biswas, 1952); Penang, 11 fms (Sunier,

1918); Kuching Fish Market, Sarawak (Tiwari and

Biswas, 1952). THAILAND: (?) Tale Sap (Kemp,

1918; Tiwari and Biswas, 1952). INDIAN OCEAN:

(Fabricius, 1798). BAY OF BENGAL: north and south

of Eastern Channel (Tiwari and Biswas, 1952). EAST

PAKISTAN: COX'S Bazaar (Tirmizi and Manning,

1968). INDIA: Sunderbans [Sundarbans] (Tiwari

and Biswas, 1952); Sandheads, mouth of River Hughli

[Hooghly; including Saugor Island] (Kemp, 1913;

Chopra, 1934; Tiwari and Biswas, 1952); Calcutta

Bazaar (Tiwari and Biswas, 1952); Chandipur, Balasore (Tiwari and Biswas, 1952); offPuri (Kemp, 1913;

Tiwari and Biswas, 1952); Bombay (Kemp, 1913;

Chhapgar and Sane, 1968). WEST PAKISTAN:

Karachi (Tirmizi and Manning, 1968). EAST

AFRICA: (Balss, 1910).

The following records need to be verified; all fall

within the known range of H. raphidea but it is not possible to determine with certainty the identity of the species reported by each author. PHILIPPINE ISLANDS:

Philippines (Holthuis, 1941); Manila Bay (Roxas and

Estampador, 1930) ; Divisoria Market, Manila (Kemp

1915) ; Cebu (Roxas and Estampador, 1930) ; Davao

Gulf, Mindanao (Roxas and Estampador, 1930). INDONESIA: Dutch East Indies; Skroe, Dutch New

Guinea; Amboina; Haroekoe (all Holthuis, 1941);

Patani Bay, Halmahera (Lanchester, 1906); Moluccas

(Holthuis, 1941); Kisar, near Timor (Holthuis,

1941); Borneo (Miers, 1880); Sandakan, northeast

Borneo; Balikpapan, east Borneo; Java; Java Sea

(all Holthuis, 1941); Madura (Thallwitz, 1892);

Batavia [DJakarta] (Preudhomme de Borre, 1882) ;

Bay of Batavia [Djakarta]; Tandjong Priok, near Batavia [Djakarta]; Cheribon, Java; Deli, east Sumatra;

mouth of Djambi River, east coast of Sumatra; Padang,

west coast of Sumatra; coast of Atyeh, Sumatra; Telok

Dalam, Nias (all Holthuis, 1941). MALAYASIA:

Singapore (Nobili, 1903); Siglap, Singapore

(Tweedie, 1934); Morib (Smedley, 1927); Kota

Bharu (Lanchester, 1901); [?] Moratabas (Lanchester,

1900); Malacca (Sharp, 1893). THAILAND: Singora

(Lanchester, 1901). INDIAN OCEAN: (BOSC, 1801—

1802, 1830; Latreille, 1802; Lamarck, 1818; MilneEdwards, 1837; Deshayes and Milne-Edwards, 1838,

1839). ANDAMAN ISLANDS: Port Blair (Kemp,

1913). BURMA : Mergui Archipelago (de Man, 1888;

Kemp, 1913); off Irrawaddy Delta, 15°20'N, 94°55'E,

20 fms (Kemp, 1913); Rangoon (Kemp, 1913);

Akyab, Tenasserim (Kemp, 1913). INDIA: no spe-=

cific locality (Sharp, 1893) ; Sunderbunds [Sundarbans] (Henderson, 1893); off Gopalpur, Ganjan coast,

25-28 fms (Kemp, 1913); Indes Orientales, Pondichery (Latreille, 1828); Madras (Henderson, 1893;

Nair, 1941; Gravely, 1941); Pamban, Gulf of Manaar

[Mannar] (Thurston, 1895); Krusadai Island, Gulf of

Mannar (Chacko, 1942); Bombay Harbour, Bombay

[part?] (Kemp, 1913). CEYLON: (Kemp, 1913).

WEST PAKISTAN: Karachi (Kemp, 1913). PERSIAN

GULF: 28°59'N, 50°05'E, 25 fms (Kemp, 1913).

Larvae attributed to H. raphidea have been recorded

from the following localities: INDIAN OCEAN: (Claus,

1871). INDIA: Madras (Alikunhi and Aiyar, 1942,

1943; Alikunhi, 1950, 1952, 1965, 1967. MALDIVE

ISLANDS: North Male Atoll, 27-35 fms (Lanchester,

1903). EAST AFRICA: Mozambique Channel (Ingle

and Delia Croce, 1967).

3. Harpiosquilla japonica, new species

FIGURES 10-11

Chloridella raphidea.—Rathbun, 1902, p. 55 [not Squilla

raphidea Fabricius].

Squilla raphidea.—Fukuda, 1913, pp. 70, 72, figure on p. 71

[not S. raphidea Fabricius] [in Japanese].
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HOLOTYPE.—1 9 , 172 mm; Wakanoura, Kii, Japan;

Jordan and Snyder, col.; 1900; SI Crust. 26340.

PARATYPES.—1 3 , 165 mm; 1 $ , 135 mm; data as

in holotype; SI Crust. 125718.

DIAGNOSIS.—Size moderate, adults with TL less

than 175 mm; antennular peduncle not so long as

carapace; corneal indices ranging between 325 and

337 (Table 1) ; rostral plate (Figure 10) with length

and width subequal, apex rounded or obtusely angled,

lacking slender anterior projection; carapace with

median carina; upper margin of propodus of claw with

smaller spines and denticles between largest spines;

dactylus of claw with 8 teeth; fifth thoracic somite

(Figure 11) rounded laterally; posterior 3 thoracic

somites with submedian and intermediate carinae, none

armed posteriorly; ventral keel of eighth thoracic

somite rounded; all 6 abdominal somites with subFIGURE 11.—Harpiosquilla japonica, new species.

Female holotype, TL 172

mm, Wakanoura: lateral

processes of fifth, sixth,

and seventh thoracic

somites.

MOURE 1U.—tiarpiosquilla japonica, new species. Female

holotype, TL 172 mm, Wakanoura: anterior portion of

body.

median carinae, abdominal carinae spined as follows:

submedian 6, intermediate 3-6, lateral 1-6, marginal

1-5; denticles 4-6, 10-11, 1; marginal carina of telson

less than twice as long as carina of lateral tooth; postanal keel extending about halfway between anus and

posterior margin; color of uropodal exopod unknown.

COLOR.—Completely faded in the type-specimens.

SIZE.—Only male examined, TL 165 mm; females

TL 135-172 mm.

Other measurements (in mm) of male: carapace

length 34.1; cornea width 10.5; antennular peduncle

length 32.4; rostral plate length 5.4, width 5.4; fifth

abdominal somite width 35.3; telson length 31.3, width

30.3.

DISCUSSION.—Harpiosquilla japonica, new species,

resembles both H. annandalei and H. stephensoni, new

species, and differs from the remainder of the species

in the genus in having a short rostral plate, as broad

as long, without an anterior projection. Harpiosquilla

annandalei differs in numerous features, as noted under the discussion of that species, and it can be distinguished immediately from H. japonica by the

presence of armed submedian carinae on the fifth abdominal somite. Harpiosquilla stephensoni and H.

japonica are very similar in general facies, but they

can be distinguished readily by the characters used in

the key, namely (1) the presence of eight teeth on the

claw of H. japonica, seven on H. stephensoni; (2) the

intermediate carinae of the second abdominal somite

are unarmed in H. japonica, spined posteriorly in H.

stephensoni, and (3) the lateral process of the fifth
NUMBER 3 6 17

abdominal somite is rounded in H. japonica, angled

or spined in H. stephensoni. The last feature may be

of limited value in distinguishing specimens of H.

japonica from smaller specimens of H. stephensoni,

for, as noted under the Discussion (p. 19) of the latter

species, small adults may not have the projecting lateral process well developed.

Although the corneal indices of H. japonica, new

species, overlap those of the smallest specimen of H.

stephensoni, new species, examined for this study, the

eyes of the two species differ in shape when two specimens of similar size are compared. The lobes of the

cornea are more inflated in H. stephensoni and the

eye appears smaller in that species.

I have previously assumed that records of H. raphidea from Japanese waters are referable to H. harpax,

for H. raphidea is not known to occur that far northward. However, some of the specimens reported in

the literature may well be H. japonica, new species.

Fukuda (1913) clearly shows a short rostral plate,

short marginal carinae on the telson, and relatively

strong submedian carinae on the abdomen, all of

which are characteristic of H. japonica rather than

H. harpax.

Although Rathbun (1902) reported five specimens

in the lot collected by Jordan and Snyder, only three

are now in the collections of the Division of Crustacea,

National Museum of Natural History, Smithsonian

Institution. The other two specimens probably were

returned to Stanford University and may now be

housed in the collections of the California Academy of

Sciences.

SEXUAL DEMORPHISM.—The male paratype lacks

raptorial claws, so no comparison of that appendage

can be made with the claw of the female (only one

female, the holotype, has claws). The median carina

of the telson in the male is slightly more inflated than

that of each female.

BIOLOGY.—Unknown.

DEVELOPMENT.—Unknown.

TYPES.—The holotype and two paratypes are in

the Division of Crustacea, National Museum of Natural History, Smithsonian Institution.

TYPE-LOCALITY.—Wakanoura, Kii, Japan.

NAME.—The name refers to the occurrence of the

species off Japan.

DISTRIBUTION.—Known only from Japan. Fukuda's

article is in Japanese so I cannot determine whether

he mentioned any specific localities.

4. Harpiosquilla stephensoni, new species

FIGURES 12-17

?Squilla sp. [Alima pyramidalis].—Foxon, 1932, p. 381 [larva;

? several species; identification questioned by Ingle and

Delia Croce, 1967].

Squilla raphidea.—Tate, 1883, p. 48 [?].—Stephenson, 1952,

p. 4; 1953, p. 43 [part?].—Stephenson and McNeill, 1955,

p. 239 [part] [not S. raphidea Fabricius].

Harpiosquilla harpax.—Manning, 1966, p. 87, fig. 1 [not S.

harpax de Haan].

Harpiosquilla sp.—Manning, 1968a, fig. 4.

HOLOTYPE.—1 9,315 mm; Port Curtis, Queensland,

Australia; seine net over flat; no other data; AM P9664.

PARATYPES.—1 $ , 220 mm; near Emery Point, Darwin Harbour, Northern Territory; taken in seine net

FIGURE 12.—Harpiosquilla stephensoni, new species.

Male paratype, TL 272 mm, Port Curtis: anterior portion

of body.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Harpiosquilla stephensoni, new species. Male

paratype, TL 272 mm, Port Curtis: raptorial claw (propodus

length 63.0 mm).

FIGURE 14.—Harpiosquilla stephensoni, new species. Female holotype TL 315 mm, Port Curtis: raptorial claw (propodus length 79.2 mm).

off beach; Capt. E. F. Wells, col.; early 1954; AM

P-12382.—1 5 , 170 mm; same data; SI Crust. 111354.

—1 9 , 168 mm; Darwin Harbour, Northern Territory;

no other data; AM P-12770.—1 9 , 240 mm; same; in

60 ft; C. W. Holman, col.; 2 July 1962; AM P-14924.

—4 $ , 136-202 mm; 1 9 , 135 mm; southern Gulf of

Carpentaria, Queensland; 14 fms or less; CSIRO

Prawn Survey; December 1963; AM P-15503 to P15507.—1 $, 208 mm; southeastern Gulf of Carpentaria, Queensland; 14 fms or less; CSIRO Prawn Survey; December 1963; AM P-15509.—1 broken 9,

CL 51.6 mm; Cairns Inlet, northern Queensland; AM

P-9685.—1 5, 245 mm; Port Curtis area, Queensland; M. Ward and W. Boardman, col.; July 1929;

AM P-14925.—1 9 , 277 mm; same; AM P-14926.—

1 $ , 272 mm; Port Curtis, Queensland; SI Crust.

125720.

DIAGNOSIS.—Size large, adults with TL to 315 mm;

antennular peduncle shorter than carapace; corneal

indices ranging between 336 and 434 (Table 1) ; rostral

plate (Figure 12) as long as or slightly longer than

broad, apex obtusely angled or rounded, not projecting

anteriorly; carapace with median carina; upper margin of propodus of claw with smaller spines and denticles between largest spines; dactylus of claw with 7

teeth, outer margin of dactylus (Figure 13) with

prominent obtusely rounded projection in adult males;

fifth thoracic somite (Figure 15) angled laterally or

with lateral acute projection; last 3 thoracic somites

with submedian and intermediate carinae, intermediates unarmed; ventral keel of eighth thoracic somite

rounded; submedian carinae present on all 6 abdominal somites, abdominal carinae armed as follows: submedian 6, intermediate 2-6, lateral 1-6, marginal 1-5;

denticles 4-6, 6-11, 1, usually 5-6, 7-8, 1; marginal

carina less than twice as long as carina of lateral tooth;

postanal keel extending slightly more than halfway

between anus and posterior margin; at most inner half

of distal segment of uropodal exopod dark.

COLOR.—Black transverse line present on dorsum

between antennular and ophthalmic somites; antennal

scale lacking any marked concentration of dark pigment; carinae, grooves, and both anterior and posterior

margins of carapace dark; display patch on dorsal surface of merus yellow, with proximal dark oval bar;

last 3 thoracic and first 5 abdominal somites with dark

posterior line and irregular dark area on dorsum; telson

with pair of oval or elongate dark spots on anterior

dorsal surface; proximal portion of anterior margin of

uropod dark; proximal segment of uropodal exopod

with inner dark patch, distal segment with dark, diffuse

pigment on inner half or third; distal half of endopod

darker than proximal, but apex light

J. C. Yaldwyn made available a color slide taken by

D. F. McMichael of one specimen, AM P-15507.

Most of the pattern mentioned above is visible. The

background color of the body is ivory. The major difference from the pattern on the same specimen in preservative is that there is but one anterior, elongate

patch of dark pigment on the telson; it is not broken

into two submedian patches.
NUMBER 36 19

SEE.—Males TL 136-272 mm; females TL 135-315

mm. Only H. raphidea, among the currently known

species of Harpiosquilla, grows to such a large size; it

is known to reach 335 mm in total length.

Other measurements of female holotype, TL 315

mm: carapace length 59.1; cornea width 14.4; antennular peduncle length 46.7; rostral plate length 10.9,

width 9.9; raptorial propodus length 79.2; fifth abdominal somite width 61.4; telson length 57.2, width

55.6.

DISCUSSION.—The short rostral plate of this species (Figure 12), a feature which it shares with H.

annandalei and H. japonica, new species, will distinguish it from H. harpax, H. indica, new species, H.

melanoura, and raphidea. H. stephensoni, new species,

differs from H. annandalei in many features, including size of adults and color pattern, but the absence of

spines on the submedian carinae of the fifth abdominal

•".•••?. - • •. . • .

FIGURE 15.—Harpiosquilla stephensoni, new species. Male

paratype, TL 272 mm, Port Curtis: lateral processes of fifth,

sixth, and seventh thoracic somites.

somite in H. stephensoni is the best way to separate the

two species.

Harpiosquilla stephensoni, is very similar to H.

japonica, described above. However, H. japonica differs in having eight teeth on the claw, unarmed intermediate carinae on the abdomen anterior to the third

somite, and the lateral process of the fifth thoracic

somite is rounded, not angled posterolaterally. The

submedian carinae of the abdomen are more prominent in H. stephensoni than in H. japonica.

Although the corneal indices of H. stephensoni and

H. japonica overlap, the eyes of H. japonica appear to

be larger when specimens of similar size are compared.

Harpiosquilla japonica is probably a smaller species,

for the single male examined, TL 165 mm, shows signs

of inflation of the median carina of the telson, a secondary sexual feature characteristic of adult males.

See also the remarks under the account of H. japonica,

new species (p. 15).

Adult specimens (TL 200-225 mm or more) differ

from younger ones in having the lateral process of the

fifth thoracic somite shaped differently. In adults (Figure 15) the edge of the lateral process slopes posterolaterally to an acute angle, the apex of which may

form a blunt tubercle or spine. The spine, if present, is

never so well developed as in H. annandalei or H.

raphidea. The lateral process of subadult specimens

is not so markedly angled posterolaterally. However,

juveniles of H. stephensoni can be distinguished by the

other features cited above.

Corneal indices for H. stephensoni, new species, are

summarized in Table 1. They overlap those of H.

harpax to some extent, but the eyes of the largest specimens of H. harpax are about the same size as those of

the smallest specimens of H. stephensoni, and the lobes

of the cornea are not as inflated as in the latter species.

The eyes of the smaller specimens of H. raphidea reported herein are similar in size to those of the largest

specimen of H. stephensoni, but the eyes of large specimens of H. raphidea are much smaller.

Specimens recorded as S. raphidea by Stephenson

and McNeill (1955) from Darwin Harbour (2 9) ,

to belong to H. stephensoni. Most of the remainder of

the specimens reported by Stephenson and McNeill

are referred herein to H. harpax; one specimen proved

to be H. melanoura. It is likely that some of the

specimens from Queensland recorded as S. raphidea by

Stephenson (1952, 1953) belong to H. harpax, but I
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 16.—HarpiosquiUa stephensoni, new species. Male

paratype, TL 272 mm, Port Curtis: posterior portion of body

(telson width 45.3 mm). FIGURE 17.—Harpiosquilla stephensoni, new species. Female

holotype, TL 315 mm, Port Curtis: posterior portion of body

(telson width 55.6 mm).

include his records here because of the apparent

abundance of H. stephensoni in Queensland waters.

In 1966 I reported on a specimen of H. stephensoni

as H. harpax and noted that it differed from other

specimens available for study in several features. Two

of these, the convex lobe on the basal prolongation of

the uropod and the elongate dark spots on the telson,

have proved to be of little value. The two specimens

from off Emery Point, Darwin Harbour, show both

of these features but in other respects they appear to

be typical subadult specimens of H. stephensoni. No

other specimens have elongate spots on the telson but

several show variation in shape of the lobe between

the spines of the basal prolongation. I can only conclude that these features vary in this species.

The carinae of the marginal teeth of the telson in

H. stephensoni are lined dorsally with large tubercles;

the carinae appear nodulose in lateral view (Figures

16, 17). These tubercles are well defined in large specimens are are not obliterated in them as in H. raphidea.

The carinae are inflated and the apices of the teeth

are upturned. In these features H. stephensoni resembles H. raphidea, but the tubercles are much larger in

the latter species.

The marginal denticles of the telson are less numerous, in general, than in other species of Harpiosquilla.

They are also sharper and more distinct in smaller

specimens than in the largest ones examined.

SEXUAL DIMORPHISM.—Males of TL 180 mm or

more have secondary sexual characteristics similar to

those found in some other species. There is a pronounced lobe, often irregular in outline, on the outer

margin of the dactylus of the claw (Figure 13); it is

not so sharply angled as in males of H. raphidea. The

bases of the second to fifth dactylar teeth are inflated.

The submedian carinae of the sixth abdominal somite

and the median carina of the telson are also inflated in

adult males (Figure 16), but the median carina of the

telson in large females may also be very swollen. Claws

(Figures 13, 14) and telsons (Figures 16,17) of a large

male and female have been illustrated.

BIOLOGY.—Relatively little information is available.

Specimens have been taken in shallow water, 25 meters

or less; three specimens were taken in seine nets. The

range of the species in Australia overlaps that of H.

harpax but whether both species occur together is not

known.

One of the specimens from the southern Gulf of

Carpentaria, AM P-15507, has numerous spherical

egg cases, approximately 1.5 mm in diameter, attached

to both the dorsal and ventral surfaces of the body;

they are most numerous on the posterior portion. Many

of the capsules had ruptured, but one was found to

contain three small gastropods, each about 1.1 mm

long. Commensals have not been recorded from members of this genus, and egg cases of other organisms are
NUMBER 3 6 21

rarely found on stomatopods. At the time of this writing, the egg cases had been identified tentatively by Dr.

Joseph Rosewater, Division of Mollusks, Smithsonian

Institution, as those of marginellid gastropods. Both

marginellids and Harpiosquilla are burrowers, and

possibly the adult gastropod used the Harpiosquilla in

its burrow (?) as a substrate for attachment of its egg

capsules.

DEVELOPMENT.—Larvae of this species are not

known with certainty. Foxon (1932) reported on

larvae from the Great Barrier Reef which she identified with Alimerichthus pyramidalis Lanchester; she

also synonymized A. unidens Lanchester, 1903, and

Alimerichthus a Tattersall, 1906, with A. pyramidalis.

A. pyramidalis was subsequently identified with H.

raphidea by Alikunhi (1952) who also pointed out that

A. unidens and Tattersall's larva as well could be attributed to other species. Alikunhi (1952, p. 268) also

noted: "That Foxon was dealing with a collection of

species under the name A. pyramidalis is, therefore,

obvious. Since larvae measuring 18 mm in length are

reported in this collection, it is likely that S. raphidea

is also represented in it."

However, Ingle and Delia Croce (1967), who reexamined Foxon's material, stated (p. 67):

Alikunhi (1952, p. 265) attributes Foxon's (1932, p. 381)

material of Alima pyramidalis (Lanchester) to the adult

species Squilla raphidea Fabricius. These particular specimens in the B.M. (N.H.) Collections do not quite correspond

to Alikunhi's description and figures as most of them have

spinule no. 5 .. . poorly developed and one or more teeth on

the inner margin of the raptorial dactylus.

The possibility remains that Foxon was dealing with

some larvae of either H. harpax or H. stephensoni, new

species, both of which are common on the coast of

Australia.

TYPE.—The holotype and most of the paratypes are

in the Australian Museum, Sydney. One male and one

female paratype are in the collections of the Division

of Crustacea, National Museum of Natural History,

Smithsonian Institution.

TYPE-LOCALITY.—Port Curtis, Queensland, Australia.

NAME.—The species is named for Professor W.

Stephenson of the University of Queensland who has

been instrumental in the development of our knowledge of Australian stomatopods.

DISTRIBUTION.—Known only from the coastal

waters of Australia, where it has been taken from

scattered localities between Darwin, Northern Territory, and Port Curtis, Queensland; shallow water to

14 fathoms. The following records in the literature

may be based on material of this species. AUSTRALIA:

Darwin, Northern Territory (Manning, 1966); probably Port Darwin, Northern Territory (Tate, 1883) ;

Queensland (Stephenson, 1952); Cooktown, Queensland (Stephenson, 1953); Tambian Beach, Keppel

Bay, Queensland (Stephenson and McNeill, 1955) ;

Peel Island, Moreton Bay, Queensland (Stephenson,

1953); Woody Point Pier, Moreton Bay, Queensland,

10-11 m (Stephenson, 1952); 3 miles south of Woody

Point Pier, Moreton Bay, Queensland, 6 m (Stephenson, 1952); 4 miles east of Scarborough Pier, Moreton

Bay, Queensland, 10 m (Stephenson, 1952); and

Southport, Queensland (Stephenson, 1953).

Stephenson (1952, 1953) may also have had H.

harpax represented in his collections from Queensland.

Larvae of this species from the Great Barrier Reef

may have been reported by Foxon (1932) as Alima

pryramidalis.

5. Harpiosquilla melanoura Manning, 1968

FIGURES 18-27

Squilla raphidea.—Stephenson and McNeill, 1955, p. 239

[part] [not S. raphidea Fabricius].

Harpiosquilla melanoura Manning, 1968b, pp. 14 [key] 18,

fig. 5.

MATERIAL.—1 $ , 161 mm; Rose Bay, Port Jackson,

New South Wales, Australia; in prawn net; 10 March

1925; AM P-7929.—1 9, 151 mm; Andaman Sea,

Mergui Archipelago; 09°13'N, 97°51'E; 58-60 m;

ANTON BRUUN Cruise 1, Sta. 20; IIOE; 23 March

1963; SI Crust. 125719.—19, 128 mm; Andaman

Sea, off Burma; 14°07'N, 97°05'E; 69-73 m; ANTON

BRUUN Cruise 1, Sta. 38; IIOE; 30 March 1963; SI

Crust. 125358.

In addition to these specimens I have examined the

type-series, 2 S , TL 102-129 mm, 2 9, TL 126-136

mm, from the Bane de Pracel, western coast of

Madagascar.

DIAGNOSIS.—Size moderate, adults with TL less

than 170 mm; antennular peduncle not so long as carapace and rostral plate combined; corneal indices ranging between 229 and 271 (Table 1); rostral plate

(Figure 19) longer than broad, lateral margins tapering to slender median projection; carapace without

median carina; upper margin of propodus of claw with

smaller spines and denticles between largest spines;
22

FIGURE 18.—Harpiosquilla melanoura Manning. Female,

TL 151 mm, Mergui Archipelago: dorsal view.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dactylus of claw with 8 teeth, outer margin of dactylus with prominent angular projection in adult males;

fifth thoracic somite (Figure 21) rounded laterally;

posterior 3 thoracic somites lacking submedian carinae;

intermediate carinae of thoracic somites unarmed;

ventral keel of eighth thoracic somite rounded; first to

fifth abdominal somites lacking submedian carinae,

abdominal carinae spined as follows: submedian 6,

intermediate (2) 3-6, lateral 1-6, marginal 1-5; denticles 5-7, 9-12, 1; marginal carina of telson almost 3

times as long as carina of lateral tooth; postanal keel

not extending halfway between anus and posterior

margin; distal segment of uropodal jet black.

COLOR.—Inner and outer surfaces of antennular

peduncles with distal black spot on each segment; anterolateral angle of antennal scale dark, anterointernal

angle dark in some specimens; basal portion of rostral

plate with oval area outlined with dark pigment; anterolateral margins, gastric grooves, and posterior

median margin of carapace dark, median area, anterior

to cervical groove, with dark oval patch; merus of claw

with 3 dark spots, 1 at inner distal margin, 1 on proximal edge of dorsal depression, and 1 subdistally on

outer surface; propodus of claw with inner and outer

dark spot; posterior 3 thoracic and anterior 5 abdominal somites (sixth in some specimens) with posterior black lines; second abdominal somite with thin,

transverse dark bar, traces of dark pigment also present

medially on third to fifth somites; dorsal surface of

telson with proximal submedian pair of oval dark spots,

surface pits also dark; ventral surface of telson dark

along midline, with dark patch at base of each tooth;

anterior edge of basal segment of uropod dark; distal

portion of proximal segment of uropodal exopod dark,

distal segment of exopod jet black; distal half or twothirds of endopod dark. The overall color pattern is

shown in Figure 18.

SIZE.—Only male examined, TL 161 mm; females

TL 128-151 mm. Including the type-series from Madagascar, males are known to range between TL 102-161

mm, females between TL 126-151 mm.

Other measurements of a female, TL 151 mm:

carapace length 31.2; cornea width 11.5; antennular

peduncle length 31.7; rostral plate length 6.4, width

5.0; raptorial propodus length 43.4; fifth abdominal

somite width 32.2; telson length 28.5, width 27.6.

DISCUSSION.—Harpiosquilla melanoura is the only

species in the genus lacking both the median carina of

the carapace and the submedian carinae of the last
NUMBER 3 6 23

FIGURES 19-23.—Harpiosquilla melanoura Manning. Male holotype, TL 129 mm, Bane de

Pracel, Madagascar: 19, anterior portion of body; 20, eye; 21, lateral processes of exposed

thoracic somites; 22, last two abdominal somites, telson, and uropod; 23, uropod, ventral view

(setae omitted; from Manning, 1968b).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

three thoracic and first five abdominal somites. It further differs from all other species of Harpiosquilla now

known in having the entire surface of the distal segment of the uropodal exopod jet black in color.

Harpiosquilla melanoura most closely resembles H.

indica, new species, described herein, but the latter

species has a well-developed median carina on the

carapace. Other differences are discussed under H.

indica (p. 33).

Corneal indices for H. melanoura are summarized

in Table 1. The eyes of this species are similar in size

to those of H. harpax but are slightly larger and more

inflated.

The marginal carinae of the telson are comparatively longer in H. melanoura than in other species of

Harpiosquilla for each is almost three times as long

as the carina of the lateral tooth. H. harpax is the

only other species in which the marginal carina is

more than twice as long as that of the lateral tooth.

Of the seven specimens of this species now known,

only one has the intermediate carinae of the second

abdominal somite armed posteriorly. In the other

specimens the intermediate carinae of the third to sixth

somites only are provided with posterior spines.

SEXUAL DIMORPHISM.—Large males (TL 129 mm

or more) (Figure 24) have a well-developed obtuse

projection on the outer margin of the dactylus of the

claw and the bases of individual teeth on the claw

are noticeably inflated. Claws of a male and a female

are shown in Figures 24 and 25. Males also differ from

females in having the submedian carinae of the sixth

abdominal somite and the median carina of the telson more inflated (Figures 26,27).

BIOLOGY.—Essentially unknown. Most specimens

have been taken in depths between 55 and 73 meters;

they have not been taken in the littoral habitat as have

both H. harpax and H. raphidea. Specimens from

Madagascar were taken on sand or muddy sand.

Harpiosquilla melanoura was taken along with H.

annandalei at ANTON BRUUN Sta. 20 off the Mergui

Archipelago.

DEVELOPMENT.—Unknown.

TYPES.—Division of Crustacea, National Museum

of Natural History, Smithsonian Institution. Paratypes

are in the Smithsonian Institution and the Museum

National d'Histoire Naturelle, Paris.

TYPE-LOCALITY.—Bane de Pracel, western coast of

Madagascar.

DISTRIBUTION.—Indo-West Pacific region, from

Rose Bay, Port Jackson, New South Wales, Australia

(Stephenson and McNeill, 1955, as S. raphidea);

Mergui Archipelago; off Burma; and the Bane de

Pracel, off the western coast of Madagascar (Manning,

1968b). It has not been recorded previously east of

Madagascar.

FIGURE 24.—Harpiosquilla melanoura Manning. Male, TL

161 mm, Port Jackson, New South Wales: raptorial claw

(propodus length 39.0 mm).

FIGURE 25.—Harpiosquilla melanoura Manning. Female,

TL 151 mm, Mergui Archipelago: raptorial claw (propodus

length 43.4 mm).
NUMBER 3 6 25

FIGURE 26.—Harpiosquilla melanoura Manning. Male, TL

161 mm, Port Jackson, New South Wales: posterior portion

of body (telson width 30.0 mm).

6. Harpiosquilla harpax (de Haan, 1844)

FIGURES 28-38

Squilla harpax de Haan, 1844, atlas, pi. 51 (fig. 1); 1849,

text, p. 222.—Tiwari and Biswas, 1952, p. 358, figs. 3b,

d, f.—Barnard, 1955, p. 49—Alikunhi, 1959, p. 132, fig13 [larva].—Ingle, 1963, pp. 14, 18, figs. 9, 59.—Holthuis,

1964, p. 140 [listed].—Manning, 1968a, p. 121 [listed],

fig. 3a.

Squilla raphidea.—Berthold, 1845, p. 47; 1847, p. 29.—

White, 1847, p. 84.—Miers, 1880, p. 27 [part].—Bigelow,

1894, pp. 511 [key], 535.—de Man, 1898, p. 694 [part ?].—

Tattersall, 1906, p. 166.—Balss, 1910, p. 8, fig. 2a.—Kemp,

1913, p. 88, pi. 7 (fig. 77) [part].—Parisi, 1922, p. 103.—

Komai, 1927, p. 323.—Komai, Akatsuka, and Ikari, 1927,

p. 295.—Komai and Ikari, 1929, p. 121.—Gravier, 1930,

p. 525.—Chopra, 1934, p. 27 [part].—Serene, 1937, p. 68

[listed].—Komai, 1938, p. 268.—Suvatti, 1938, p. 52.—

Chopra, 1939, p. 158.—Holthuis, 1941, p. 256 [part].—

Liu, 1949, p. 43, pi. 6 (figs. 15-17).—Anonymous, 1949,

p. 843, fig. 2421.—Barnard, 1950, p. 851, figs, lc, g.—

Suvatti, 1950, p. 132.—Dawydoff, 1952, p. 145.—Serene,

1953, p. 507; 1954, pp. 6, 8, 62, pi. 4 (figs. 1-6).—

Millard and Harrison, 1954, p. 176 [listed].—Stephenson

and McNeill, 1955, pp. 239 [part], 255 [key].—Day and

FIGURE 27.—Harpiosquilla melanoura Manning. Female,

TL 151 mm, Mergui Archipelago: posterior portion of body

(telson width 27.6 mm).

Morgans, 1956, p. 306 [listed].—Utinomi, 1956, p. 91,

pi. 46 (fig. 2) [color figure].—Yamaji, 1959, fig. on p.

68.—Utinomi, 1960, p. 114, pi. 57 (fig. 7) [color figure].—

Chuang, 1961, pp. 181, 206, pi. 81 (fig. 4), lowest fig. on

pi. 82.—Stephenson, 1962, p. 34.—Crosnier, 1965, p. 61

[listed]. [All not Squilla rap hide a Fabricius.]

Squilla obsoleta White, 1847, p. 84 [nomen nudum; in

synonymy].

Squilla Harpax.—Herklots, 1861, p. 152 [p. 39 on separate].

Squilla raphidea van—Gravier, 1937, p. 186, figs. 8-10.

Squilla gilesi.—Boone, 1938, pp. 201, 203, pi. 71 [not S.

gilesi Kemp].

Squilla raphidoea.—Serene, 1939, p. 349 [erroneous spelling].

?Squilla sp. [Alima multispinus].—Foxon, 1939, p. 258, fig.

2B [larva] [not Erichthus multispinus Claus, 1871].

?Squilla sp.—Gohar and Al-Kholy, 1957, p. 106 [larvae; part,

several species mixed ?].

Squilla hanpax.—Alikunhi, 1959, p. 132 [discussion; erroneous spelling].

Harpiosquilla harpax.—Lee and Wu, 1966, p. 51, figs.

7A-F.—Manning, 1967, p. 103.—Holthuis, 1967, p. 15.—

Chhapgar and Sane, 1968, p. 45 [key].—Manning, 1968b,

pp. 14 [key], 15, fig. 4.—Tirmizi and Manning, 1968, p.

33, fig. 13.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

?Alima stages of Harpiosquilla raphidea-harpax group.—

Ingle and Delia Croce, 1967, p. 6, figs. 19, 24 [larvae].

not Harpiosquilla harpax.—Manning, 1966, p. 87, fig. 1

[=H. stephensoni, new species].

MATERIAL.—1 3, 152.5 mm; Japan; lectotype of S.

harpax; RMNH 28H.—1 3,175 mm; China or Japan;

u.s.s. PALOS; SI Crust. 5146.—1 3, CL 20.2 mm; between Kobe and Yokahama, Japan; 34°06'15"N,

134°57'50"E; 40-37 fms; fine gray shell mud; ALBATROSS Sta. 4963; 27 August 1906; SI Crust. 76058.—

1 3, 85 mm; Takao, Taiwan; F. Baker, col.; December

1914; SI Crust. 47935.—1 3 ,109 mm; same; SI Crust.

47928.—13, 146 mm; Tungkang market, Taiwan;

40-50 fms; S. Lee and S. K. Wu; 5 March 1965; SI

Crust. 113645.-1$, 215 mm; Hong Kong; North

Pacific Exploring Expedition; SI Crust. 2108.—13,

137 mm; Makung, Penghu Island, Pescadores Islands,

China Sea; 19 April 1961; SI Crust. 124762.—1 3 ,

132 mm; Nhatrang Bay, Viet-Nam; DE LANESSAN,

col.; 1930; SI Crust. 124721.—2 3 (1 broken), 132

mm; 2 9 (1 broken), 109 mm; Sandakan Fish Market,

North Borneo; Chin Phui Kong, col.; 5 November

1956; AM P-14923.—1 broken 9, CL 39.5 mm; off

North East Island, near Groote Eylandt, western Gulf

of Carpentaria, Northern Territory, Australia; 25 fms;

CSIRO Prawn Survey; 4 September 1963; AM P -

15508.—19, 248 mm; Tin Can Bay, near Gympie,

Queensland, Australia; taken in net by C. Lea; 7 July

1952; AM P-12256.—1 broken 3, CL 36.2 mm; Cape

Flattery, Queensland, Australia; AM G-1792.—19,

225 mm; The Basin, Newcastle Harbour, New South

Wales, Australia; from prawn trawl; A. A. Racek, col.;

27 May 1954; AM P-12512.—13, CL 29.6 mm;

Hawkesbury River, N.S.W., Australia; hooked by line

fisherman; March 1952; AM P-12230—1 broken S,

CL 27.5 mm; Bobbin Head, near Jerusalem Bay,

Hawkesbury River, N.S.W., Australia; taken from

stomach of fish, flathead; May 1952; AM P-12229.—

IS, 188 mm; Jerusalem Bay, Hawkesbury River,

N.S.W., Australia; hooked by line fisherman; 18 May

1952; SI Crust. 124765.—19, 143 mm; Jerusalem

Bay, Hawkesbury River, near Sydney, N.S.W., Australia; R. Woodger, col.; AM P-15399.—19, 180 mm;

Jerusalem Bay, Hawkesbury River, near Sydney,

N.S.W., Australia; handline from 30 ft of water on

hook baited with small fish; A. B. Iverach, col.; 24

January 1965; AM P-15402.—19, 175 mm; Cowan

Creek, Hawkesbury River, N.S.W., Australia; 1 February 1953; SI Crust 124766.—13, 165 mm; same,

from stomach of 71 lb flathead fish; March 1960; AM

P-13324.—1 9, 190 mm, same; handline and baited

hook; L. Kelly, col.; 23 April 1960; AM P-15400.—

1 9, 185 mm; same; handline and baited hook;

A Leslie, col.; 18 March 1961; AM P-15401.—

19, 195 mm; same; handline from 15 ft water on

small hook baited with prawn; 26 April 1965; AM

P-15403.—IS, 110 mm; Lem Sing, Thailand; 10

January 1924; SI Crust. 124761.—1 9 , 168 mm;

Tachalom, Gulf of Siam; trap; Hugh M. Smith; 21

July 1923; SI Crust. 69533.—19, 157 mm; South

Kang Kradan, Thailand; Hugh M. Smith; 1923; SI

Crust. 69534.—19, 153 mm; beach north of Singora,

Thailand; Hugh M. Smith; 5 October 1923; SI Crust.

69535.—IS, 87 mm; Bay of Bengal; 15°04'N,

95°51'E; 29—33 m; ANTON BRUUN Cruise 1, Sta. 41A;

IIOE; 31 March 1963; SI Crust. 125356.—1 9, 153

mm; Sandheads, mouth of River Hooghly [Hughli],

India; LADY FRASER; SI Crust. 119312.—IS, 120 mm;

2 9 , 129-173 mm; India; 17°54-57'N, 72°23-27'E;

46-55 m; green mud, shells; ANTON BRUUN Cruise 4B,

Sta. 201A; IIOE; 13 November 1963; SI Crust.

125352.—23, 71-146 mm; 29, 122-164 mm; off

Bombay, India; 18°47.3-49/N, 72°36-38'E; 12.5-14

fms; M.V.F. JANJIRA; Field No. 9682; IIOE; 12 May

1964; SI Crust. 125354.—73, 72-120 mm; 5 9,

64-140 mm; India; 20°30/N, 70°50-54/E; 33 m;

brown sticky mud; ANTON BRUUN Cruise 4B, Sta 204A;

IIOE; 15 November 1963; SI Crust. 125355.—1 9,

165 mm; northwestern India; 22°52-54'N, 68°34-

36'E; 15 m; soft mud; ANTON BRUUN Cruise 4B, Sta.

223A; IIOE; 19 November 1963; SI Crust. 125353.—

19, 148 mm; Gulf of Oman; 26°10-13'N, 57°02'E;

55-64 m; green mud; ANTON BRUUN Cruise 4B, Sta.

256A; IIOE; 30 November 1963; SI Crust. 125351.—

5 3 , 139-166 mm; 89 , 134-185 mm (in 2 lots);

Archico Bay, south of Massawa, Ethiopia, Red Sea;

Israel South Red Sea Exped., Sta. E62/4118; 8 April

1962; SI Crust. 124763,124764.

DIAGNOSIS.—Size moderate to large, TL to 248 mm;

antennular peduncle usually shorter than carapace;

corneal indices ranging between 251 and 388 (Table

1); rostral plate longer than broad (Figure 31), with

slender anterior projection; carapace with median

carina; opposable margin of propodus of claw with

smaller spines and denticles between largest spines;

dactylus of claw with 8 teeth, outer margin of dactylus

with prominent angular projection in adult males;
NUMBER 3 6 27

fifth thoracic somite (Figure 32) rounded laterally;

posterior 3 thoracic somites with submedian and intermediate carinae, none armed posteriorly; ventral keel

of eighth thoracic somite rounded; abdomen with submedian carinae on all somites, abdominal carinae

spined as follows: submedian 6, intermediate (1)

2-6, lateral 1-6, marginal 1-5; denticles 4-6, 10-12

(14), 1; marginal carina of telson more than twice as

long as carina of lateral tooth; postanal keel extending

about halfway between anus and posterior margin;

distal segment of uropodal exopod with inner half

black.

COLOR.—In preservative, black transverse line on

dorsum between antennular and ophthalmic somites;

antennal 6cale with diffuse dark pigment, concentrated

along anterior margin; carinae, grooves, and anterior

and posterior margin of carapace dark, pair of dark

spots also present on lateral plates of carapace at level

of cervical groove; display patch on dorsal surface of

merus yellow, with proximal oval dark patch and

distal dark spot; last 3 thoracic and first 5 abdominal

somites with dark posterior line; telson with pair of

dark circles or square spots on anterior dorsal surface;

distal segment of uropodal exopod with diffuse dark

chromatophores on inner half, distal third of proximal

segment also dark; distal half of endopod dark, with

light midrib in some specimens; apices of teeth of

uropods and telson yellow in some specimens. The

overall color pattern is shown in Figure 28.

The notes on color in life given below from Komai

(1927, p. 323) and Utinomi (1956, 1960) may refer

to this species but may also refer to H. japonica, new

species.

When fresh, the entire surface is pale and slightly ochraceous, besprinkled with minute dark spots. A pinkish color is

found on the carapace, raptorial limbs and on the thoracic

and abdominal somites. A conspicuous round patch of chestnut-brown color occurs on either side of the base of the middorsal carina of the telson.

According to Utinomi (1956) this species is pinkish,

setae of scale and uropod yellow, telson with pair of

proximal red spots. He (1960, p. 114) also reports:

Antennal scale blue, setae yellow; propodus yellow distally,

apices of teeth of chela yellow; submedian carinae of first

to fifth abdominal somites and sixth to eighth thoracic somites

yellow, lateral carinae on body with more red; submedian

carinae of sixth abdominal somite green; median carina of

telson and pits green; carinae of marginal teeth green, apices

yellow; most of uropod yellow.

Tirmizi and Manning (1968, p. 33) reported the

following color pattern in fresh specimens from Karachi, West Pakistan:

FIOURE 28.—Harpiosquilla harpax (de Haan). Male, TL

188 mm, Jerusalem Bay, New South Wales: dorsal view.
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 29-34.—Harpiosquilla harpax (de Haan). Female, TL 118 mm, Ambaro Bay, Madagascar: 29, anterior portion of body; 30, outline of eye; 31, outline of rostral plate; 32, outline

of lateral processes of exposed thoracic somites; 33, last two abdominal somites, telson, and

uropod; 34, uropod, ventral view. (Setae omitted; from Manning, 1968b).
NUMBER 36 29

. . . eyestalks bright yellow; antennal peduncle and scale

with pink patches; carinae of carapace lined with dark spots;

merus and carpus of raptorial claw with pink patches; posterior margin of each of thoracic and abdominal somites

lined with black spots; lateral portions of abdominal somites

pink; sixth abdominal somite pink between submedian carinae; spines on abdomen, telson, and uropod yellow; telson

with grey pits and carinae except for red median carina;

proximal submedian patches of telson maroon (black in

preservative); base of uropod pink.

SIZE.—Males, TL 71-188 mm; females, TL 64-248

mm. Males ranging in TL between 80 and 195 mm and

females TL 112 to 215 mm have been recorded in the

literature (size data from corroborated distribution

records only).

Other measurements of male lectotype, TL 152.5

mm: carapace length 29.1; cornea width 9.2; antennular peduncle length 28.1; rostral plate length 5.0,

width 4.8; raptorial propodus length 38.4; fifth abdominal somite width 30.6; telson length 29.4, width

26.2.

DISCUSSION.—Harpiosquilla harpax, the commonest and most widely distributed species of the genus,

can be distinguished with ease from the other species.

The apical median projection of the rostral plate at

once separates it from H. annandalei, H. japonica,

new species, and H. stephensoni, new species; in all

of those species the rostral plate is blunt apically, either

broadly rounded or obtusely rounded. Harpiosquilla

harpax is a smaller species than H. raphidea, and the

carinae on the body are not nearly so well developed

as in that species; in H. harpax the intermediate carinae of the last three thoracic somites are unarmed

whereas they are provided with spines in H. raphidea,

and the lateral process of the fifth thoracic somite is

rounded in H. harpax, spined laterally in H. raphidea.

The faint submedian carinae on the abdomen will distinguish H. harpax from both H. indica, new species,

and H. melanoura; the latter species lacks the median

carina on the carapace as well. In H. indica, the intermediate carinae of the first and second abdominal

somites are unarmed; the carinae of the second abdominal somite are always armed in adults of H.

harpax.

The color pattern of H. harpax, in preserved specimens, is very similar to that of both H. indica and

H. stephensoni, but very distinct from those of H.

annandalei and H. melanoura. In H. harpax, the inner

half of the distal segment of the uropodal exopod is

covered with diffuse dark pigment. In H. annandalei

that segment is black with a white midrib and in H.

melanoura it is entirely black. The color pattern of the

uropod is not nearly so well marked in H. raphidea as

in any of these species. The color pattern of H. japonica, is not known.

The shape and length of the marginal carinae of

the telson are also distinctive in H. harpax which has

a slender marginal carina over twice as long as the

carina of the intermediate tooth. In H. melanoura the

marginal carina is three times as long as the intermediate, and in the remainder of the species it is less than

twice as long.

Harpiosquilla harpax is not n small species but it

apparently does not get nearly so large as do either

H. raphidea or H. stephensoni. The maximum observed size in H. harpax is 248 mm in contrast with

335 mm for H. raphidea and 315 mm for H.

stephensoni.

The intermediate carinae of the first abdominal

somite were armed in only three of these specimens,

a female from Luzon, a female from India, and a

male from Viet-Nam. In addition, the small specimen

(CL 20.2 mm) from Japan lacked spines on the intermediate carinae of the first two abdominal somites. All

but one of the specimens from the collections of the

Australian Museum had both intermediate carinae of

the first abdominal somite unarmed posteriorly. In all

other respects the specimens from Australia agree well

with the remainder of the specimens from other localities available for study.

Holthuis (1967) noted that the anterior projection

of the rostral plate was a variable feature in his specimens from the Red Sea. There is also some variation

in relative length of the anterior projection in specimens from Australia. The plate in H. harpax, however,

never terminates so bluntly as it does in H. annandalei,

H. stephensoni, and H. japonica.

All available specimens of H. harpax were examined

to determine the extent of variation of the corneal

indices, if any. Although the samples from any area

were small, no obvious variation by sex, size, or location could be detected. The eye is remarkably uniform in size and shape over a wide geographic area.

Corneal indices of all species are summarized in

Table 1.

All specimens examined by me had eight teeth on

the dactyl us of the claw. Parisi (1922) recorded one

specimen from Suruga Gulf, Japan, with but seven

teeth on one claw.
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 35.—Harpiosquilla harpax (de Haan). Male, TL

188 mm. Jerusalem Bay, New South Wales: raptorial claw

(propodus length 42.9 mm).

FIGURE 36.—Harpiosquilla harpax (de Haan). Female, TL

175 mm, Cowan Creek, New South Wales: raptorial claw

(propodus length 45.9 mm).

It has been pointed out already that Alikunhi's

(1967) excellent account of growth is probably based

on both H. harpax and H. raphidea. Certainly some

of the observations made by Alikunhi on changes with

age apply to this species as well as to H. raphidea.

SYNONYMY.—It has been very difficult to determine

which of the early records of S. raphidea actually belonged to H. harpax. Several criteria have been used

but only a few records could be checked so most of the

references referred herein to H. harpax require

verification.

Geography has been the basis for assigning some

records of H. harpax rather than to H. raphidea. The

latter species is not known to occur with certainty as

far north as China or Japan, as far south as Australia,

or as far west as South Africa. Most locality records

from Japan for H. raphidea probably are referable to

H. harpax, but some may be referable to H. japonica,

new species. Most records from Japan require

verification.

In some cases I have examined material previously

recorded in the literature, although only in the case

of the collections from the Australian Museum and

the Smithsonian Institution have I been able to examine the bulk of the collection. Some of the material

recorded by the following authors has been examined:

de Haan, 1844, 1849; Holthuis, 1941, 1967; Lee and

Wu, 1966; Boone, 1938; Gravier, 1930; Millard and

Harrison, 1954; Stephenson and McNeill, 1955. I

have accepted the records of Tiwari and Biswas (1952)

and those of Kemp (1913) and Chopra (1934) which

they corroborated. The records of Ingle (1963) and

the earlier records corrected by him have also been

assigned to H. harpax.

The numerous records from Viet-Nam by Gravier

(1930), Serene (1937, 1939, 1953, 1954) and

Dawydoff (1952) are assigned here for two reasons.

Their accounts concur with my observations on H.

harpax rather than H. raphidea, and we have no authenticated records of H. raphidea from east of Borneo

and north of Thailand. Serene's (1954) observations

on sexual dimorphism agree well with what I have

observed in H. harpax. All of these authors, however,

could have been dealing with both species.

Gravier (1937) recorded three specimens from the

Baie d'Along [Tonkin], Viet-Nam, as S. raphidea var.
NUMBER 3 6 31

His illustrations clearly show that his specimens are

referable to H. harpax, for the thorax and telson of

the illustrated specimen are typical of that species.

De Man (1898) noted that a small male, TL 11 cm,

from the Java Sea had poorly developed submedian

carinae on the abdomen and lacked the spine on the

fifth thoracic somite; he attributed these features to

individual aberration. He was almost certainly dealing with H. harpax in this case; he may also have

examined some H. raphidea.

Gravier (1937) also commented that another specimen collected by Dawydoff differed from his material

in several features. I cannot determine from his account the distinguishing features of DawydofFs

material.

Tattersall (1906) reported that a male from Ceylon

had an obtuse lateral process on the fifth thoracic

somite; he, too, was probably dealing with H. harpax.

Judging from the size of specimens reported from

Queensland, Australia, TL to 222 mm, some of the

specimens identified by Stephenson (1952, 1953) as

S. raphidea may actually be referable to H. stephensoni, new species, and H. harpax.

Suvatti's records of S. raphidea from Thailand

(1938, 1950) are apparently based on collections made

there by Hugh M. Smith. Material from the localities

given by Suvatti is in the Smithsonian Instituion and

is clearly referable to H. harpax. Although Smith also

collected H. raphidea in Thailand, those specimens

were not recorded by Suvatti.

The specimens from Singapore reported by Boone

(1938) as S. gilesi are almost certainly H. harpax;

although a lateral spine is shown on the fifth thoracic

somite by Boone (op. cit., pi. 72), the specimens actually lack lateral spines. It seems likely that the ventral spines on that somite are exaggerated in the figure.

Boone's records have been corrected by both Holthuis

(1941) and Manning (1967).

The specimen figured by Chuang (1961) on his

plate 82 shows no trace of submedian carinae on the

abdomen or of a lateral spine on the fifth thoracic

somite. For these reasons I have assigned his records to

H. harpax, which has very low submedian carinae on

the abdomen. Chuang (1961, p. 181) commented on

the presence of middorsal carinae on the last two thoracic somites as a distinguishing feature of S. raphidea;

none of the species of Harpiosquilla are ornamented

with middorsal carinae on either the thorax or

abdomen.

FIGURE 37.—Harpiosquilla harpax (de Haan). Male, TL

188 mm, Jerusalem Bay, New South Wales: posterior portion

of body (telson width 33.9 mm).

FIGURE 38.—Harpiosquilla harpax (de Haan). Female, TL

175 mm, Cowan Creek, New South Wales: posterior portion

of body (telson width 32.2 mm).
32

Liu (1949), on the other hand, shows strong submedian carinae on the thorax and abdomen and short

marginal carinae on the telson, but mentions that the

fifth thoracic somite has no lateral process; an apical

projection is shown on the rostral plate. It seems likely

that the submedian carinae of the body and the carinae

of the telson were exaggerated in the illustration and

that he was dealing with H. harpax.

SEXUAL DIMORPHISM.—Males of H. harpax exhibit

the secondary sexual characters also found in most

other species of Harpiosquilla. The obtuse prominence

on the outer margin of the claw in adult males can be

observed at a size of about 120 mm TL (CL ca. 22

mm) (Figure 35). It becomes larger and more angular with increasing age. It is not so well developed in

H. harpax at TL 150 mm as it is in H. indica, new

species. The bases of the second through the sixth dactylar teeth are inflated, as illustrated by Serene (1954,

pi. 4, fig. 4). The submedian carinae of the sixth abdominal somite and the median carina of the telson

may be inflated in adults of both sexes; often the median carina of the telson is more inflated in females

than in males (Figures 37, 38). The carinae of the

marginal teeth of the telson are not markedly inflated

in adults of either sex.

BIOLOGY.—As is the case in the other species of

Harpiosquilla, we can glean relatively little information on the biology of H. harpax from the literature and

from specimen-associated data. It is a widely distributed species in the Indo-West Pacific region, and it

occurs in shallow water, between 2 and 93 meters.

Chuang (1961) recorded it from the sublittoral zone

in Singapore. It apparently has a preference for mud

bottom, for recorded bottom types include green mud,

shell; brown sand; grey sand; brown sticky mud; soft

mud; green mud; muddy sand; and sandy shore. It

was taken together with H. annandalei in 55-64 meters

in the Gulf of Oman by the ANTON BRUUN ; in general,

it occurs in shallower water than does H. annandalei.

Harpiosquilla harpax has been recorded from fish

markets from three areas, the Tungkang Market, Taiwan (Lee and Wu, 1966), the Sandakan Fish Market,

Borneo, and Fish Harbour, Karachi, West Pakistan

(Tirmizi and Manning, 1968). In Pakistan, at least,

it is not sought for food but is brought to shore along

with the trash from commercial shrimping operations.

The specimens from Australia were taken in prawn

nets, on hook and line by fishermen, and from the

stomach of a fish known locally as flathead. It may

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

enter water of lowered salinity, for it seems to be abundant in different sections of the Hawkesbury River,

New South Wales.

DEVELOPMENT.—The larvae of H. harpax are not

known with certainty. Foxon (1939) reported larvae

from the Red Sea, the northern Arabian Sea, and the

Gulf of Oman; she noted that the larvae were of two

types, with one (typical form) or two (atypical form)

laterally directed spines on the lateral margin of the

carapace. Both forms occurred in all three areas. She

suggested that these might be the larvae of raphidea

and annandalei, and pointed out the similarity of the

larva reported by her to Erichthus multispinus Claus.

In his discussion of the development of raphidea,

Alikunhi (1952, p. 269) noted that "The atypical form

of Alima multispinous described by Foxon (1939) also

shows close similarity to the final pelagic larva of

S. raphidea," differing in having one rather than two

laterally directed spinules on the carapace. Alikunhi

further pointed out that the typical form reported by

Foxon differed from known Harpiosquilla larvae in

having four free teeth on the dactylus of the claw.

Alikunhi then suggested that the atypical form

might be identified with raphidea whereas the typical

form might be the larva of annandalei. Ingle and Delia

Croce (1967, p. 67) reexamined Foxon's specimens

and stated, "The specimens identified by Foxon (1939,

p. 258) as atypical forms of Alima multispinosa

(Claus) agree with Alikunhi's figures and description

of late S. raphidea larvae."

Inasmuch as H. raphidea is not known to occur in

the Red Sea and H. harpax is the only Harpiosquilla

which occurs in the Red Sea, northern Arabian Sea,

and the Gulf of Oman, it seems likely that Foxon's

atypical larva may be the larva of H. harpax. Whether

or not the other larva reported by Foxon is even identifiable with Harpiosquilla remains to be seen.

Some of the larval forms from the Red Sea identified as Squilla sp. by Gohar and Al-Kholy (1957) may

be identifiable with H. harpax (see their figs. 5-7, pi.

xu). Judging from the illustrations, their Squilla sp.

includes the larvae of more than one species.

Alikunhi (1959) recorded young larvae of harpax

from off the Mahanadi Estuary, India.

The larval stages of Harpiosquilla reported from

the Mozambique Channel by Ingle and Delia Croce

(1967) may be referable to H. harpax or H. melanoura,

both of which occur as adults off Madagascar, or to H.

raphidea, which occurs on the east coast of Africa.
NUMBER 3 6 33

The alima larva from Japan figured by Yamaji

(1959) adjacent to his figure of raphidea does not

resemble the larva of Harpiosquilla.

Some of the observations made by Alikunhi (1967)

on Harpiosquilla postlarvae may apply to H. harpax

as well as to H. raphidea.

TYPE.—The lectotype and paralectotypes are in the

Rijksmuseum van Natuurlijke Historie, Leiden, The

Netherlands.

TYPE-LOCALITY.—Japan.

DISTRIBUTION.—Indo-West Pacific region, from

Japan and Australia in the Pacific to the Red Sea and

South Africa in the Indian Ocean. Shallow water to

93 m. The following records in the literature are probably correct: JAPAN: NO specific locality (de Haan,

1844, 1849; Herklots, 1861; Holthuis, 1941; Manning,

1968b).—TAIWAN: Tungkang Fish Market, 40-50

fms (Lee and Wu, 1966).—CHINA: Hongkong (Bigelow, 1894; Kemp, 1913; Tiwari and Biswas, 1952).—

VIET-NAM: Baie d'Along [Tonkin] (Gravier, 1937);

Baie Nhatrang, 50 m (Gravier, 1930).—PHILIPPPINE

ISLANDS: NO specific locality (Miers, 1880; Ingle,

1963).—MALAYSIA: Singapore (Boone, 1938; Tiwari

and Biswas, 1952; Manning, 1967); Siglap, Singapore

(Chuang, 1961); Morib (Chuang, 1961).—INDONESIA: Java Sea, between Singapore and Surabaya

[Surabaja] (de Man, 1898).—AUSTRALIA (all Stephenson and McNeill, 1955) : Cape Flattery and Tin Can

Bay, Queensland; Cowan Creek, Jerusalem Bay, and

Hawk River, New South Wales.—THAILAND: Tachalom and Gulf of Siam, north of Singora (Suvatti,

1938,1950).—ANDAMAN ISLANDS: Port Blair (Tiwari

and Biswas, 1952).—INDIAN OCEAN: (White, 1847;

Miers, 1880; Ingle, 1963).—INDIA: mouth of River

Hughli [Hooghly, including Sandheads] (Kemp, 1913;

Tiwari and Biswas, 1952); off Madras coast (Kemp,

1913); Madras Presidency (Tiwari and Biswas, 1952);

Bombay (Chhapgar and Sane, 1968) ; off Bombay,

20-25 fms (Tiwari and Biswas, 1952).—CEYLON:

southwest part of Palk Bay, off Adam's Bridge and

Rameswaram Island, 7-9 fms (Tattersall, 1906).—

WEST PAKISTAN: Karachi (Tirmizi and Manning,

1968).—GULF OF OMAN: (Chopra, 1939; Ingle,

1963).—RED SEA: southern Red Sea (Ingle, 1963);

Archico Bay, south of Massawa, Ethiopia, 8—17 fms

(Holthuis, 1967).—ZANZIBAR: (Miers, 1880; Ingle,

1963).—MADAGASCAR: Ambaro Bay, Nosy Be, 2-5 m

(Manning, 1968b) ; Baie de Narendry, 6 m (Manning,

1968b).—SOUTH AFRICA: Richards Bay (Millard and

Harrison, 1954); Durban (Barnard, 1950, 1955; Day

and Morgans, 1956).

The following records, listed in the literature as

raphidea, may be referable to H. harpax; records for

Japan might be dealing with H. japonica, new species,

and those from Australia may be referable to H. stephensoni, new species. JAPAN: NO specific locality

(Anonymous, 1949; Utinomi, 1956, 1960; Yamaji,

1959); Suruga Gulf (Kemp, 1913; Parisi, 1922); Kii

Peninsula (Komai, Akatsuka, and Ikari, 1927); Seto,

Kii Province (Komai, 1927, 1938); Tanabe Bay, Kii

Peninsula (Komai and Ikari, 1929); Tanabe (Komai,

1938); Wakanoura (Komai, 1938); Kasiwazima, Tosa

Province (Komai, 1927); Misaki (Komai, 1927);

Nagasaki (Balss, 1910; Parisi, 1922; Komai, 1927).—

TAIWAN: northern Formosa (Komai, 1927); Takao

(Balss, 1910).—CHINA: no specific locality (Berthold,

1845, 1847); Hongkong (Liu, 1949) .—VIET-NAM:

Indo-China (Gravier, 1930; Serene, 1937, 1953); Baie

de Cauda, 15-25 m (Serene, 1954); Baie de Nhatrang,

8-12 m (Serene, 1939); Baie d'Along [Tonkin] (Dawydoff, 1952); Baie de Tourane, 25-30 m; Embouchure

du Bassac; Cap Saint Jacques; Baie de Hone Coke,

entree de Port Dayot; Poulo Condore Islands (all

Gravier, 1930).—SUBLICHES OSTASIENS: (Balss,

1910).—AUSTRALIA: Wyndham, Western Australia

(Stephenson, 1962).—INDIA: Sandheads (Chopra,

1934) .—MADAGASCAR : (Crosnier, 1965).

Larvae which may prove to be the larvae of H. harpax have been reported from the Red Sea by Foxon

(1939) and Gohar and Al-Kholy (1957), the northern

Arabian Sea and Gulf of Oman by Foxon (1939), the

Mozambique Channel by Ingle and Delia Croce

(1967), and perhaps, from the east coast of India by

Alikunhi (1952, 1959). See also the section on larvae

for both H. harpax (p. 25) and H. raphidea (p. 9).

7. Harpiosquilla indica, new species

FIGURES 39-43

HOLOTYPE.—1 <J , 150 mm; 10 miles due south of

Mandapam Camp, South India; 12 fms; otter trawl;

L. P. Woods on Norwegian trawler M-2; Sta. LW-20

(GA 64-7); HOE; 14 February 1964; SI Crust.

125717.

DIAGNOSIS.—Size moderate, male with adult characters at TL 150 mm; antennular peduncle shorter

than carapace; corneal index 290; rostral plate (Figure

40) longer than broad, lateral margins tapering to
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

slender median projection; carapace with median carina; upper margin of propodus of claw with smaller

spines and denticles between largest spines; dactylus of

claw with 9 teeth, outer margin with prominent angular projection in adult male; fifth thoracic somite

rounded laterally (Figure 42); posterior 3 thoracic

somites lacking submedian carinae; intermediate

carinae of thoracic somites unarmed; ventral keel of

eighth thoracic somite rounded; first to fifth abdominal

somites lacking submedian carinae, abdominal carinae

spined as follows: submedian 6, intermediate 3-6, lateral 1-6, marginal 1-5; denticles 5, 11, 1; marginal

carina of telson not more than 1.5 times as long as

carina of lateral tooth; postanal keel short, not extending halfway between anus and posterior margin; at

most inner half of distal segment of uropodal exopod

dark.

COLOR.—Inner and outer surfaces of antennular

peduncles lined with dark pigment; antennal scale outlined with dark chromatophores; dark transverse bar

present dorsally between antennular and ophthalmic

somites; rostral plate outlined with dark pigment, basal

portion of plate with oval ring of dark chromatophores; margins, carinae, and grooves of carapace outlined with dark pigment; meral depression of claw with

proximal and distal dark spots; posterior 3 thoracic

FIGURE 39.—Harpiosquilla indica, new species,

holotype, TL 150 mm: dorsal view.

Male FIGURE 40.—Harpiosquilla indica, new species. Male

holotype, TL 150 mm: anterior portion of carapace.
NUMBER 36 35

and all abdominal somites with dark posterior line and

intermediate carinae lined with dark pigment; submedian area of thoracic and abdominal somites with

diffuse dark pigment; telson with marginal carinae and

dorsal pits darker than remaining surface; ventral surface of telson with bases of teeth and submedian distal

area darker than remainder of surface; anterior edge

of basal segment and proximal segment of uropodal

exopod with diffuse dark spot; distal fourth of proximal

segment of uropodal exopod with diffuse dark pigment,

inner half of distal segment black; distal half of endopod dark with apex lighter along midline. The overall

color pattern is shown in Figure 39.

SIZE.—Only specimen known, the male holotype,

TL 150 mm. Other measurements (in mm) : carapace

length 30.5; cornea width 10.5; antennular peduncle

length 27.0; rostral plate length 6.8, width 5.4; raptorial propodus length 39.4; fifth abdominal somite

width 33.0; telson length 29.6, width 28.0.

DISCUSSION.—Harpiosquilla indica, new species,

most closely resembles H. melanoura, differing from

the latter species as follows: (1) there are nine teeth

on the dactylus of the claw rather than eight; (2) there

is a well-developed median carina on the carapace;

(3) only the inner half of the distal segment of the

uropodal exopod is black; and (4) the marginal carina

of the telson is comparatively shorter, being not more

than one and one-half times rather than nearly three

times as long as the carina of the lateral tooth.

The lack of submedian carinae on the posterior three

thoracic and anterior five abdominal somites, the unarmed intermediate carinae anterior to the third abdominal somite, and the shorter marginal carina of

the telson will distinguish H. indica from H. harpax,

which it otherwise closely resembles. In H. indica the

projection on the outer margin of the dactylus of the

male is much more angular than that found in H.

harpax, and the median carina of the telson in the

holotype of H. indica is more inflated than in males of

either H. harpax or H. melanoura of similar size.

Harpiosquilla japonica differs from H. indica in

lacking the median projection on the rostral plate and

in having submedian carinae on the posterior three

thoracic and all six abdominal somites.

The eye of the holotype of H. indica is similar in

size to that of H. melanoura and noticeably larger than

FIGURE 41.—Harpiosquilla indica, new species. Male

holotype, TL 150 mm: raptorial claw (propodus length

39.4 mm).

FIGURE 42.—Harpiosquilla indica, new species. Male

holotype, TL 150 mm: lateral processes of fifth, sixth, and

seventh thoracic somites.
36

FIGURE 43.—Harpiosquilla indica, new species. Male

holotype, TL 150 mm: posterior portion of body (telson

width 28.0 mm).

the eyes of either H. stephensoni or H. harpax of similar size.

The color pattern of H. indica is very similar to that

of H. harpax. The holotype of H. indica lacks the

paired submedian spots on the telson but that may be

an artifact of preservation, and it also lacks the concentration of dark chromatophores on the anterolateral

margin of the antennal scale which is characteristic

of H. harpax.

SEXUAL DIMORPHISM.—The holotype has a pronounced projection on the dactylus of the claw (Figure

41) which is much more angular than that found in

H. harpax. The basal inflation of the teeth of the claw,

swollen submedian carinae of the sixth abdominal somite, and inflated median carina of the telson (Figure

43) are well developed in the single male specimen

available for study.

BIOLOGY.—Unknown.

DEVELOPMENT.—Unknown.

TYPE.—The holotype is in the Division of Crustacea,

National Museum of Natural History, Smithsonian

Institution.

TYPE-LOCALITY.—Ten miles due south of Mandapam Camp, South India, in 12 fathoms.

NAME.—The name refers to the occurrence of the

species off India.

DISTRIBUTION.—Known only from the type-locality.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

General Remarks

The species of Harpiosquilla are large squillids which

are found in relatively shallow water throughout most

of the Indo-West Pacific region. All species occur on

the shelf; the maximum depth recorded is 206 meters

for H. annandalei. That species, which has been taken

in depths between 15 and 206 meters, also shows the

widest depth range. H. melanoura has been taken in

depths between 55 and 73 meters. The remaining

species, H. harpax, H. indica, new species, H. japonica,

new species, H. stephensoni, new species, and H. raphidea, generally occur in depths of 51 meters or less,

although H. harpax has been collected as deep as 91

meters. At least three species, H. harpax, H. stephensoni, and H. raphidea, may occur in the sublittoral

zone.

Judging from the limited information available both

in the literature and from data associated with specimens reported here, the species of Harpiosquilla all

prefer a soft, level bottom; they do not appear to be

associated with coral reefs, or with rough, rocky bottom. At least one species, H. raphidea, is tolerant of

lowered salinities.

The genus is not known to occur in Oceania. This

may be a result of the paucity of collections from that

area but more likely reflects the lack of suitable habitat

there.

In a study of certain penaeid shrimps from the IndoMalayan area, Hall (1962) commented on the existence of an apparent distributional barrier in the Malacca Straits. On the basis of this he cast suspicion

on the validity of all littoral penaeid species reported

to range throughout the Indo-West Pacific region. This

does not seem to be true in Harpiosquilla, for four of

the seven known species occur on both sides of the

Indo-Malayan area, and two of these four species have

broad distributional patterns in the Indo-West Pacific

region. H. harpax, for example, extends from Japan to

Australia in the western Pacific, and from there westward to the Red Sea and South Africa. H. annandalei

has a similar distribution pattern, based on many fewer

samples, but it does not occur in Australia and has

not been recorded with certainty west or south of the

Gulf of Oman. H. raphidea is known to occur from

Borneo and Thailand to India, West Pakistan, and

East Africa; it is not known with certainty to occur

north of Borneo and Thailand in the Pacific and there

are no apparent reasons for its relatively limited distribution in the western Pacific or its absence in Aus-
NUMBER 36 37

tralian waters. H. melanoura occurs in New South

Wales, Australia, the Andaman Sea, and Madagascar.

The remaining species of Harpiosquilla may be indigenous to the areas from which they are now known.

H. indica, new species, is known only from the type

locality off southern India, H. japonica, new species,

only from Japan, and H. stephensoni, new species, from

the Northern Territory and Queensland, Australia.

The latter two species are very similar and possibly

represent the terminal portions of a more widely distributed ancestral stock. Among the stomatopods, a

similar Japan-Australia link is shown by two species of

Heterosquilla, H. latifrons from Japan and H. brazieri

from Australia (Manning, 1966).

Within Australia the two most abundant species,

H. harpax and H. stephensoni, have an interesting distributional patern. H. stephensoni is known with certainty from Darwin Harbour, Northern Territory to

Port Curtis, Queensland; i.e., it is known only from

the tropical waters of the eastern and northeastern

coast on the Solanderian Province (Knox, 1963). If,

as I have assumed, Stephenson's (1952, 1953) records

for Squilla raphidea prove to be based on H. stephensoni rather than H. harpax, its distribution on the eastern Australian coast will be extended to Southport,

Queensland, in the northern edge of the Peronian

Province. H. harpax, however, extends from the Gulf

of Carpentaria, on the Queensland side, south to the

vicinity of Sydney in the cooler Peronian Province,

is much more abundant in the southern portion of its

range. It is nevertheless surprising to find H. harpax,

the most abundant species of the genus, all of which

otherwise are tropical, in the cooler, warm-temperate

environs of Sydney. Perhaps its ability to adapt to a

wider range of conditions in part helps to explain its

overall wide distribution.

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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