Keys to the Species

of Oratosquilla

(Crustacea: Stomatopoda)

with Descriptions of

Two New Species

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1971 NUMBER 71
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 71

Raymond B. Manning Key S tO th e

of Oratosquilla

(Crustacea: Stomatopoda),

with Descriptions of

Two New Species

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Manning, Raymond B. Keys to the Species of Oratosquilla (Crustacea: Stomatopoda), with Descriptions of Two New Species. Smithsonian Contributions to

Zoology, 71:1-16. 1970. Eight species groups are recognized within the stomatopod crustacean genus Oratosquilla, which contains twenty-three species from

the Indo-West Pacific area. Keys to species groups and to species within each

group are presented. Oratosquilla tweediei, from Singapore, and O. ornata,

from Hong Kong, are newly described.

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 : 1971

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

Washington, D.C. 20402 - Price 80 cents (paper cover)
Raymond B. Manning Keys tO the

of Oratosquilla

(Crustacea: Stomatopoda),

with Descriptions of

Two New Species

Introduction

The genus Oratosquilla comprises many of the

large squillids from the Indo-West Pacific region

that formerly were placed in Squilla; Oratosquilla

was separated from Squilla by Manning (1968a).

The notes presented here originally were compiled

for a revision of the genus, but it has become apparent that much more material from localities

throughout the Indo-West Pacific region will have

to be studied before such a revision can be made.

It is hoped that this preliminary paper will stimulate interest in the group, aid identification of the

known species, and help to make more material

available so that a definitive revision can be prepared. Planned studies of the Albatross collections

from the Philippine Islands, as well as the collections made during the International Indian Ocean

Expeditions, should help to clarify the status of

several species of Oratosquilla not reported here.

Although the genus Oratosquilla as currently

composed appears to be homogeneous, eight distinct groups of species can be recognized, based

primarily on characters of the raptorial claw and

the carapace. Most of the species have 6 teeth on

the raptorial claw, but one, O. indica (Hansen,

1926), has 7 or 8, another, O. investigatoris (Lloyd,

1907), has 10 to 18, and three, O. gonypetes (Kemp,

1911) and allies, have but 5. The mandibular palp

Raymond B. Manning, Chairman, Department of Invertebrate

Zoology, National Museum of Natural History, Smithsonian

Institution, Washington, D.C. 20560

is usually present; it is suppressed in O. indica.

That species also differs from the other species in

the genus in having but two epipods; the other

species in the genus apparently have four epipods.

The number of epipods could not be verified in

the unique holotype of O. imperialis (Manning,

1965), in which two or three were visible; the

remainder may have been damaged.

The median carina of the carapace affords good

diagnostic features for distinguishing adults of most

of the remainder of the species. The median carina,

anterior to the cervical groove, usually terminates

anteriorly in a well-marked anterior bifurcation

(Figures la, 2a). In some species the branches of

the bifurcation are present, but the median carina

is interrupted at the base of the bifurcation (Figure 3a). In other species the branches of the bifurcation are completely suppressed (Figure 4a), although their position may be indicated by dark

pigment, by a median depression, or by low tubercles or short, interrupted portions of the carina.

The structure of the carina appears to be very

stable in almost all species, so far as is known,

between late juvenile stages and, perhaps, very old

stages. In young specimens of species of Oratosquilla, especially postlarvae, the median carina

and its bifurcation may not be distinct, as in the

holotype of O. calumnia. In some species in which

the anterior bifurcation is typically suppressed, as

in O. gonypetes, it may be present at some later

stage (Chopra, 1934; Manning, 1965, 1968b; Holthuis, 1967); O. gonypetes is the only species in

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

which the structure of the median carina is variable, and the possibility exists that the specimens

of that species reported from different areas actually represent distinct species.

By the recognition here of groups of species

within Oratosquilla, I do not intend to imply that

distinct subgenera should be recognized or that the

complex relationships of species within the genus

can be expressed adequately by the separation of

the species into a series of arbitrary groups. The

arrangement of species into these groups should

facilitate their recognition. Nine of the twentythree species of Oratosquilla have been described

in the last fifty years, and it seems likely that

several additional species will be recognized in the

future. Familiarity with the species groups as outlined below and use of the key to species groups,

as well as the keys to species within groups, should

aid recognition of undescribed species as well as

established ones.

Terms used in the keys and in the text have

been explained in detail in earlier papers (Manning, 1966; 1968b).

All of the specimens are in the Division of Crustacea, National Museum of Natural History, Smithsonian Institution, where they are listed under

United States National Museum (USNM) numbers.

The references given in brackets after each specific name in the keys are to papers in which additional information on each species can be found.

Acknowledgments

I thank L. S. Kornicker and J. Rosewater for their

comments on the manuscript. The illustrations

were prepared by my wife Lilly.

Key to Species Groups in Oratosquilla

1. Mandibular palp absent; submedian, intermediate, and lateral carinae of anterior 3

abdominal somites unarmed (2 epipods present; dactylus of claw with 7-8 teeth)

Indica Group

Mandibular palp present; some carinae of anterior 3 abdominal somites with posterior

spines (usually 4 epipods present; dactylus of claw with 5, 6, or 10-18 teeth) 2

2. Anterior bifurcation of median carina of carapace opening posterior to dorsal pit; cornea

small (corneal index 600 or more in adults), set transversely on stalk Nepa Group

Anterior bifurcation of median carina of carapace absent or opening anterior to dorsal

pit; cornea large (corneal index less than 600 in adults), set obliquely on stalk 3

3. Lateral process of fifth thoracic somite composed of 2 slender, sharp spines directed

laterally; lateral carinae of anterior 5 abdominal somites bicarinate Mikado Group

Lateral process of fifth thoracic somite not composed of 2 slender, sharp spines; lateral

carinae of anterior 5 abdominal somites, simple, not bicarinate 4

4. Cornea expanded, very large, width about one-third carapace length; dactylus of claw

with 10-18 teeth (rostral plate without median carina) Investigatoris Group

Cornea width less than one-third carapace length (if cornea equal to or greater than

one third carapace length, then rostral plate with median carina); dactylus of claw

with 5 or 6 teeth 5

5. Median carina of carapace entire, with well-formed anterior bifurcation (dactylus of claw

with 6 teeth) Oratorio Group

Median carina of carapace interrupted at base of bifurcation or branches of bifurcation

absent 6

6. Anterior branches of bifurcation of median carina of carapace present, well formed

(dactylus of claw with 6 teeth) Perpensa Group

Anterior branches of bifurcation of median carina of carapace completely absent, area of

bifurcation usually smooth, polished (occasionally arms of bifurcation represented by

broken lines of tubercles or by lines of pigment; entire branches rarely present) 7

7. Anterior width of carapace greater than one-half median length; dactylus of claw with

6 teeth Woodmasoni Group

Anterior width of carapace less than one-half median length; dactylus of claw with 5 teeth

Gonypetes Group
NUMBER 71

Indica Group

This group contains 0. indica (Hansen, 1926),

which differs from other species of Oratosquilla in

lacking both the mandibular palp as well as spines

on the carinae of the anterior three abdominal

somites and in having the dactylus of the raptorial

claw armed with 7-8 teeth, rather than 5 or 6. The

species was described in detail by Hansen (1926),

and Chopra (1934) provided additional information.

Nepa Group

This group contains two species, O. nepa (Latreille, 1828) and O. holoschista (Kemp, 1911).

The species can be recognized by: (1) the very

small eyes, with corneal indices usually 600 or more

in adults and the cornea set transversely on the

stalk; (2) the shape of the anterior bifurcation of

the carapace, which in both species opens posterior

to the dorsal pit on the median carina; and

(3) having 6 teeth on the claw. In all other

Key to Species of the Nepa Group

Portion of median carina of carapace between cervical groove and anterior bifurcation

simple, not bicarinate; submedian carinae of fourth abdominal somite armed posteriorly

O nepa (Latreille, 1828)

[Kemp, 1913]

Portion of median carina of carapace between cervical groove and anterior bifurcation

bicarinate; submedian carinae of fourth abdominal somite unarmed

O. holoschista (Kemp, 1911)

[Kemp, 1913]

species of Oratosquilla the anterior bifurcation,

when present, opens anteriorly to the dorsal pit.

Mikado Group

In the two species assigned here, O. stridulans

(Wood-Mason in Alcock, 1894) and O. mikado

(Kemp and Chopra, 1921), the two lobes of the

lateral process of the fifth thoracic somite are produced into two sharp spines and the lateral carinae

of the anterior five abdominal somites are bicarinate. Kemp and Chopra (1921) previously noted

that these features will distinguish these two species from the remainder of the species in the genus.

Both species have 6 teeth on the claw.

As I noted in 1965, Squilla zanzibarica Chopra,

1939, appears to be a synonym of the species now

known as O. mikado (Kemp and Chopra, 1921).

Key to Species of the Mikado Group

Rostral plate with median carina; undivided portion of median carina of carapace, anterior

to dorsal pit, about one-half as long as bifurcated portion

O. mikado (Kemp and Chopra, 1921)

Rostral plate without median carina; undivided portion of median carina of carapace, anterior to dorsal pit, about one-third as long as bifurcated portion

O. stridulans (Wood-Mason In Alcock, 1894)

[Kemp, 1913]

Investigatoris Group

This group also contains one species, O. investigatoris (Lloyd, 1907), which can be distinguished by

the large number of teeth on the dactylus of the

claw, 10-18. The eyes of O. investigatoris are comOratosquilla. Only in O. quadraticauda (Fukuda)

are the eyes of comparable size, about one-third

the carapace length; O. quadraticauda, however,

has but 5 teeth on the claw as well as a median

carina on the rostral plate. Kemp (1913) and

Chopra (1939) have provided additional data on

paratively larger than in most other species of O. investigatoris.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Oratorio Group

This group contains the four species with 6 teeth

on the claw, and in which the median carina of the

carapace is well formed for all of its length anterior

to the cervical groove; it is not interrupted at the

bifurcation, and both branches of the bifurcation

are distinct. The species included in this group are

O. oratoria (De Haan, 1844), O. mauritiana

(Kemp, 1913), O. kempi (Schmitt, 1929), and O.

calumnia (Townsley, 1953) . In all four species the

carpus of the raptorial claw is ornamented with

several tubercles. Oratosquilla oratoria and O.

calumnia are diagnosed herein.

Key to Species of the Oratoria Group

1. Submedian carinae of fourth, fifth, and sixth abdominal somites with posterior spines.. 2

Submedian carinae of fourth abdominal somite unarmed, those of fifth and sixth somites

only with posterior spines 3

2. Intermediate carinae of first abdominal somite armed posteriorly; anterior lobe of lateral

process of seventh thoracic somite sharp O. calumnia (Townsley, 1953)

Intermediate carinae of first abdominal somite unarmed; anterior lobe of lateral process

of seventh somite rounded

O. mauritiana (Kemp, 1913) [ = Squilla juxtaoratoria Ward, 1942]

[Manning, 1968b]

3. Lateral process of seventh thoracic somite not markedly bilobed; second and fifth abdominal somites each with dark dorsal patch in preservative (merus of claw lacking

inferodistal spine on outer face) O. kempi (Schmitt, 1929)

[Chopra, 1934]

Lateral process of seventh thoracic somite markedly bilobed; second and fifth abdominal

somites lacking dark dorsal patch (merus of claw with inferodistal spine on outer

face) O. oratoria (De Haan, 1844) [-Squilla affinis Berthold, 1845]

Oratosquilla calumnia (Townsley, 1953)

FIGURE 1

Squilla nepa.—Brooks, 1886, p. 25 [part, specimen from

Hawaii only; not S. nepa Latreille].

Squilla calumnia Townsley, 1953, p. 410, figs. 8-9.—Manning,

1968a, p. 135 [listed].

Squilla oratoria.—Kemp, 1913, p. 68 [part, specimens from

Hawaii only].—Edmondson, 1921, p. 287, fig. la.—Bigelow,

1931, p. 183—Townsley, 1953, p. 404, figs. 2-3 [not Squilla

oratoria De Haan].

MATERIAL.—1 tf, 26 mm; Hilo Dock, Hilo,

Hawaii; night light; Territory of Hawaii Division

of Fish and Game; 24 August 1948; holotype;

USNM 93097.—1 9, 89 mm; Hilo, Hawaii;

Albatross; 1901; USNM 64909.—1 damaged $,

CL 35.0 mm; Honolulu; Albatross; 1901; USNM

64908.—1 9, 175 mm; estuary, Oahu, Honolulu;

R. A. Kinzie III; USNM 125051.

DIAGNOSIS.—Rostral plate as long as or slightly

longer than broad, apex usually rounded; eye

large, cornea set obliquely on stalk, corneal indices

391-500; anterior width of carapace about half

median length; median carina of carapace entire,

bifurcation opening anterior to dorsal pit, distance

from pit to bifurcation less than distance from

bifurcation to anterior margin; raptorial claw with

6 teeth, dorsal ridge of carpus of claw bituberculate, inferodistal angle on outer face of merus with

blunt tooth; mandibular palp and 4 epipods present; lateral process of fifth thoracic somite produced into an anterior spine and a slender posterior spine, directed laterally; lateral process of

sixth thoracic somite bilobed, anterior lobe

rounded; lateral process of seventh thoracic somite

bilobed, anterior lobe angled, occasionally sharp;

lateral carinae of abdomen not bicarinate; abdominal carinae spined as follows: submedian (3) 4-6,

intermediate 1-6, lateral 1-6; telson lacking supplementary dorsal tubercles; telson denticles $-4,

6-8, 1; proximal segment of uropodal exopod less

than 1.5 times as long as distal segment; inner

spine of basal prolongation of uropod with

rounded lobe on outer margin, proximal margin of

lobe concave.

COLOR.—Posterior 3 thoracic and all abdominal

somites each with dark posterior line; second abdominal somite with rectangular patch of dark

chromatophores medially; distal half of proximal
NUMBER 71

FIGURE 1.—Oratosquilla calumnia (Townsley), female, TL 89 mm, Hawaii: a, anterior portion

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

thoracic somites; e, basal prolongation of uropod. (Setae omitted.)

segment and inner half of distal segment of uropodal exopod dark; distal half of endopod dark.

MEASUREMENTS.—Only male examined, TL 26

mm; females, TL 89-175 mm. Specimens ranging

from 120-200 mm in length have been recorded in

the literature. Other measurements of female, TL

175 mm: carapace length 36.5; anterior width of

carapace 18.4; cornea width 7.7; rostral plate

length 5.6, width 5.4; telson length 33.5, width

32.6.

REMARKS.—It is with some hesitation that I

identify the adult specimens from Hawaii reported

here with O. calumnia. The type of O. calumnia

is a juvenile male that shows almost none of the

diagnostic features of adults; the presence of six

teeth on the claw, the mandibular palp, four epipods, and the faintly bilobed lateral processes of

the exposed thoracic somites suggest that it can be

identified as an Oratosquilla. The carinae of the

carapace are indistinct and the anterolateral spines

of the carapace as well as most of the spines found

on the abdominal carinae are not developed.

The only other species of Oratosquilla recorded

from the Hawaiian Islands is O. quadraticauda

(Fukuda) (see Townsley, 1953, under Squilla

boops), which differs from the species reported

here in having but five teeth on the dactylus of the

claw. It seems likely that Townsley's juvenile holotype is conspecific with adults identified here as O.

calumnia. The identity of that juvenile probably

cannot be proved until similar specimens are related to an identifiable size.

Oratosquilla calumnia resembles O. oratoria in

having an entire median carina on the carapace,

but can be distinguished from O. oratoria by the

spined submedian carinae of the fourth abdominal

somite, which are never found in O. oratoria. Oratosquilla calumnia also differs from O. oratoria in
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

color pattern, for none of the specimens of the

latter species examined by me has the well-formed

rectangular patch of dark chromatophores on the

second abdominal somite characteristic of O.

calumnia.

Oratosquilla calumnia closely resembles O.

mauritiana in almost all respects. The two species

are so similar that they may prove to be terminal

populations of a widely distributed species in the

Indo-West Pacific region. Specimens have not been

recorded from intermediate areas, however, and,

until such are found, it seems better to recognize

both as distinct species. Oratosquilla mauritiana

appears to be a much smoother species, its dorsal

surface presenting, when dried, an appearance similar to O. woodmasoni rather than O. oratoria. A

good feature for distinguishing the two species is

the shape of the lateral processes of the exposed

thoracic somites. In O. calumnia the anterior lobe

of the lateral process of the sixth somite is broader

than that found in O. mauritiana; the anterior

lobe of the lateral process of the seventh somite is

sharp in O. calumnia, rounded in O. mauritiana.

The intermediate carinae of the first abdominal

somite are armed posteriorly in O. calumnia, unarmed in O. mauritiana.

DISTRIBUTION.—Pacific Ocean, where it has been

recorded from the Hawaiian Islands and Guam.

Oratosquilla oratoria (De Haan, 1844)

FIGURE 2

Squilla oratoria De Haan, 1844, atlas, pi. 51: fig. 2; 1849, p.

223.—Kemp, 1913, p. 66, pi. 5: figs. 54-56 [part; older

references].—Zaks, 1933, p. 73—Holthuis, 1941, p. 247

[older references].—Shiino, 1942, pp. 77 et seq., pis. 4-14

[embryology].—Gurney, 1946, p. 160 [listed; larvae].—Anonymous, 1949, p. 842, fig. 2418.—Liu, 1949, p. 36, figs, la-b —

PDawydoff, 1952, p. 145—PSerene, 1953, p. 507 [listed].—

Kobyakova, 1955, p. 199.—Utinomi, 1956, p. 91, pi. 46:

fig. 1 [color].—Kubo and Asada, 1957, pp. 254, 257, 258,

259, 264, 265, 266, 268, 270, 272, 287, 289, tables 16-18,

fig. 12 [ecology].—Kubo et al., 1959, pp. 1 et seq., figs.

1-16 [biology].—Utinomi, 1960, p. 114, pi. 57: fig. 8

[color].—Manning, 1965, p. 259—Lee and Wu, 1966, p.

50.—Manning, 1968a, p. 135 [listed].

Oratosquilla oratoria.—Manning, 1968a, figs, lc, 2d-f, 3b, 5e,

7c-d, 9e.

Not Squilla oratoria.—Edmondson, 1921, p. 287, fig. la.—

Bigelow, 1931, p. 183.—Townsley, 1953, p. 404, figs. 2-3

[= O. calumnia (Townsley, 1953) ].

MATERIAL.—53 tf, 88-180 mm; 43 $, 54-183

mm, in twenty-nine lots from scattered localities

between southern Japan and Hong Kong, including localities off Korea and China.

DIAGNOSIS.—Rostral plate broader than long,

lacking median carina, anterior margin truncate;

eye large, cornea set obliquely on stalk; anterior

width of carapace subequal to or slightly larger

than half of median length; median carina of

carapace entire, bifurcation opening anterior to

dorsal pit; raptorial claw with 6 teeth, dorsal ridge

of carpus tuberculate, inferodistal angle on outer

face of merus with blunt spine; mandibular palp

and 4 epipods present; lateral process of fifth thoracic somite produced into an anterior spine and

a posterior angled lobe; lateral processes of sixth

and seventh thoracic somites bilobed, anterior

lobes smaller than posterior; lateral carinae of

abdomen not bicarinate; abdominal carinae spined

as follows: submedian 5-6, intermediate 3-6, lateral

2-6; telson lacking supplementary dorsal tubercles;

telson denticles 3-5, 6-10, 1; proximal segment of

uropodal exopod subequal to distal segment in

length; inner spine of basal prolongation of uropod

with rounded lobe on outer margin, proximal margin of lobe concave.

COLOR.—Completely faded in the present material. Apparently the species lacks the dark dorsal

patches on the second or on the second and fifth

abdominal somites that may be found in several

other species in the genus. Color illustrations have

been given by Utinomi (1956, 1960).

MEASUREMENTS.—Males, TL 88-180 mm; females,

TL 54-183 mm. Other measurements of a male,

TL 153 mm: carapace length 31.3; anterior width

of carapace 17.1; cornea width 6.9; rostral plate

length 4.0, width 4.6; telson length 28.4, width 28.3.

REMARKS.—I have included a diagnosis and an

illustration for this well-known and relatively wellillustrated species, for there has been some confusion in the past over its identity; the brief diagnosis

given here should separate it from all known species of Oratosquilla and should allow its ready

separation from similar species that may occur with

it in the northwestern Pacific, including O. interrupta (Kemp), O. perpensa (Kemp), O. kempi

(Schmitt) , and O. imperialis (Manning).

Only three groups of species in Oratosquilla have

an entire median carina on the carapace, with the

anterior bifurcation not interrupted basally and

the anterior branches of the bifurcation wellformed. The sharply spined, laterally directed
NUMBER 71

FIGURE 2.—Oratosquilla oratorio (De Haan), female, TL 99 mm, Amoy, China: a, anterior

portion of body; b, eye; c, carpus of raptorial claw; d, lateral processes of fifth, sixth, and

seventh thoracic somites; e, basal prolongation of uropod. (Setae omitted.)

processes of the fifth thoracic somite will distinguish the two species of the Mikado Group, O.

mikado (Kemp and Chopra) and O. stridulans

(Wood-Mason), from the species in the Oratoria

Group. The two species of the Nepa Group, O.

nepa (Latreille) and O. holoschista (Kemp), differ from the species of the Oratoria Group in having much smaller eyes and in having the bifurcation of the median carina of the carapace opening

posteriorly to the dorsal pit on the carapace.

The four species of the Oratoria Group are very

similar in basic facies. Both O. calumnia (Townsley) , from Hawaii and Guam, and O. mauritiana

(Kemp), from the western Indian Ocean, differ

from O. oratoria in having the submedian carinae

of the fourth abdominal somite armed posteriorly.

The fourth species in the group, O. kempi

(Schmitt), which has approximately the same geographic range as O. oratoria, differs in having a

sharp median carina on the rostral plate and dark

dorsal patches on the second and fifth abdominal

somites. In addition, O. kempi lacks the spine on

the outer face of the merus of the claw and does

not have a strongly bilobed lateral process on the

seventh thoracic somite.

The corneal indices appear to be reliable characters in Oratosquilla; the following indices were

found in specimens of O. oratoria reported here.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Carapace

Length, in mm

10-15

16-20

21-25

26-30

31-35

36-40

41-45

Number of

Specimens

1

10

10

9

9

6

1

Corneal Index

Range

-

383-J56

394-486

435-482

453-530

486-521

_

Mean

349

413

426

456

490

502

510

I could detect no obvious differences in the

corneal indices of males and females or in specimens from different areas.

The biology of this species is reasonably well

known. Komai (1924) reported on embryology,

Komai and Tung (1929) gave an almost complete

account of larval development. Kubo and Asada

(1957) and Kubo et al. (1959) reported on the

ecology of the species in Tokyo Bay.

DISTRIBUTION.—Northwestern Pacific Ocean, from

Peter the Great Bay, Russia, and Japan southward

to Hong Kong, including Korea, Taiwan, and

China. Although it has been recorded from Viet

Nam by Gravier (1930, 1937), Dawydoff (1953),

and Serene (1937, as Squilla affinis; 1953), it was

not listed from there by Serene (1954). Its occurrence in the Philippine Islands, as recorded by

Roxas and Estampador (1930), requires verification. One specimen from the Philippines identified

by Miers (1880) as Squilla nepa was reidentified

by Kemp (1913) as S. oratoria; Kemp noted, however, that the specimen had an interrupted median

carina on the carapace and armed submedian

carinae on the fourth abdominal somite, so it may

be referable to O. fabricii (Holthuis).

Hansen (1926) recorded a small specimen, 18

mm in length, taken by the Siboga in the Molo

Strait, Indonesia; that record should be accepted

with caution, for specimens that small generally

do not exhibit many of the diagnostic features seen

in adults, and the record is well south of the documented range of the species.

Perpensa Group

This group comprises those species in which the

median carina of the carapace is interrupted at the

base of the anterior bifurcation; the carina, posterior to the bifurcation, and the branches of the

anterior bifurcation as well are well formed. Two

Key to Species of the Perpensa Group

1. Submedian carinae of fourth, fifth, and sixth abdominal somites armed posteriorly (dorsal

ridge of carpus of claw tuberculate) O. fabricii (Holthuis, 1941)

Submedian carinae of fourth abdominal somite unarmed 2

2. Lobe between spines of basal prolongation of uropod convex (dorsal ridge of carpus of

claw tuberculate) O. interrupta (Kemp, 1911)

[Kemp, 1913]

Lobe between spines of basal prolongation of uropod concave, distal margin rounded or

acute (dorsal ridge of carpus of claw smooth, entire) 3

3. Slender spine present between spines of basal prolongation of uropod (proximal segment

of uropodal exopod 1.5 times as long as distal) O. ornata, new species

At most a rounded lobe present between spines of basal prolongation of uropod (proximal

segment of uropodal exopod subequal in length to distal) 4

4. Second and fifth abdominal somites with dark, median, rectangular patch of chroma tophores (inferodistal angle on outer face of merus of claw rounded, unarmed)

O. imperials (Manning, 1965) [ = O. simulans (Holthuis, 1967)]

At most second abdominal somite with dark median patch of chromatophores (inferodistal angle on outer face of merus of claw with spine or blunt triangular prominence) 5

5. Rostral plate longer than broad; lateral carinae of second abdominal somite with posterior

spine O. inornate (Tate, 1883)

[Manning, 1966]

Rostral plate broader than long; lateral carinae of second abdominal somite unarmed

O. perpensa (Kemp, 1911) [ = O. anomala (Tweedie, 1935) ]

[Kemp, 1913; Manning, 1966]
NUMBER 71

subgroups can be recognized in the group, one

containing two species, characterized by a tuberculate ridge on the carpus on the raptorial claw,

and the other with four species, characterized by

an unbroken, smooth ridge on the carpus of the

claw. The first subgroup contains O. interrupta

(Kemp, 1911) and O. fabricii (Holthuis, 1941).

The second subgroup contains O. perpensa (Kemp,

1911), O. inornata (Tate, 1883), O. imperialis

(Manning, 1965), and O. ornata, new species, described below.

I can find no characters to separate O. imperialis

(Manning, 1965), described from Japan, from O.

simulans (Holthuis, 1967), based on material from

the Red Sea. Although I have not been able to

compare specimens of both species, comparison of

their descriptions leaves me no alternative to synonymizing them. The two species are almost identical

in morphology and color pattern. I reported that

the type of O. imperialis had 2 or 3 epipods; Holthuis reported 4 epipods in O. simulans. It is possible that the eipods of the unique type of O. imperialis were damaged. The distribution of O.

imperialis, known from localities in the Red Sea

and from Japan but not from intermediate localities, parallels that of O. mikado, which is known

from Japan and Zanzibar. It seems likely that these

distribution patterns, which are at best unusual in

Oratosquilla in which most species have restricted

ranges, reflect availability of collections or collecting effort rather than true discontinuous distribution patterns within the Indo-West Pacific region.

Oratosquilla ornata, new species

FIGURE 3

HOLOTYPE.—1 9, 43.5 mm; China Sea, off Hong

Kong; 21°53'N, 115°51'E; 62 fathoms; sand, green

mud; Albatross Station 5309; 4 November 1908;

USNM 77939.

DIAGNOSIS.—Rostral plate as long as broad, appearing elongate, lacking median carina, apex

rounded; eye large, cornea set obliquely on stalk;

corneal index 426; anterior width of carapace less

than half median length; median carina of carapace interrupted at base of bifurcation, bifurcation

opening anterior to dorsal pit; raptorial claw with

6 teeth, dorsal ridge of carpus entire, inferodistal

margin of outer face of merus at most angled; mandibular palp and 4 epipods present; lateral process

d

FIGURE 3.—Oratosquilla ornata, new species, female holotype, TL 43.5 mm, off Hong Kong:

a, anterior portion of body; b, eye; c, carpus of raptorial claw; d, lateral processes of fifth, sixth,

and seventh thoracic somites; e, basal prolongation of uropod. (Setae omitted.)
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of fifth thoracic somite produced into an anterior

spine and a posterior angled lobe; lateral processes

of sixth and seventh thoracic somites bilobed, anterior lobe of process of sixth somite truncate

laterally, almost as large as posterior lobe; lateral

carinae of abdomen not bicarinate; abdominal

carinae spined as follows: submedian 5-6, intermediate 3-6, lateral 2-6; telson without supplementary dorsal tubercles; telson denticles 4, 8, 1;

proximal segment of exopod longer than distal;

inner spine of basal prolongation of uropod with

sharp spine on outer margin, proximal margin of

spine concave.

DESCRIPTION.—Dorsal surface of body pitted,

rugose. Eye large, cornea bilobed, set very obliquely

on stalk; eyes not extending to end of first segment

of antennular peduncle; ocular scales truncate, inclined laterally, separate mesially; anterior margin

of ophthalmic somite flattened; corneal index 426.

Antennular peduncle slightly shorter than carapace; dorsal processes of antennular somite flattened, directed anterolaterally.

Antennal scale slender, curved, elongate, more

than half as long as carapace.

Rostral plate slender, length and width subequal, appearing elongate, upturned lateral margins converging on rounded apex; median carina

absent.

Carapace narrowed anteriorly, anterior width

less than half median length; anterolateral spines

strong but not extending to base of rostral plate;

median carina interrupted at base of anterior bifurcation, branches of bifurcation distinct; distance

from dorsal pit to anterior bifurcation less than

distance from bifurcation to anterior margin; portion of median carina posterior to cervical groove

with anterior bifurcation; intermediate carinae

not extending to anterior margin; posterolateral

margin of carapace evenly curved, not angled anteriorly; median posterior margin of carapace with

triangular projection.

Dactylus of raptorial claw with 6 teeth, outer

margin of dactylus evenly curved; dorsal ridge of

carpus undivided; inferodistal margin of outer face

of merus with unarmed angular lobe.

Mandibular palp and 4 epipods present.

Exposed thoracic somites with irregular submedian and intermediate carinae; lateral process

of fifth somite bilobed, anterior lobe a slender

spine, outer margin convex, directed anteriorly,

posterior lobe short, bluntly triangular, directed

laterally; lateral process of sixth somite bilobed,

anterior lobe truncate, almost as large as posterior,

apex acute but not spined; lateral process of seventh somite bilobed, anterior lobe triangular, posterior lobe larger, triangular; ventral keel of eighth

somite erect, triangular, apex rounded.

Anterior 5 abdominal somites each with 8 carinae, submedians slightly divergent on each somite;

sixth somite with 6 pairs of carinae; abdominal

carinae spined as follows: submedian 5-6, intermediate 3-6, lateral 2-6, marginal 1-5; sixth

somite with small ventrolateral spinule in front of

articulation of each uropod and with median tubercle on posterior margin of sternum.

Telson flattened, longer than broad, with 6

elongate marginal teeth; prelateral lobes present,

not markedly projecting laterally, shorter than

lateral carina; dorsal surface of telson ornamented

with median carina and carinae of marginal teeth,

supplementary tubercles or carinae absent; median

carina low, sharp, with obscure anterior notch and

long distal spine; outer submedian denticle

rounded, remainder sharp, 4, 8, 1; ventral surface

of telson with short but prominent postanal keel.

Uropods slender, elongate, proximal segment of

exopod longer than distal, with 9 graded spines on

outer margin, distalmost not extending to midlength of distal segment; endopod slender, inner

margin concave; lobe on inner spine of basal prolongation of uropod a slender erect spine, margin

concave.

COLOR.—Completely faded in holotype.

MEASUREMENTS.—Female holotype, TL 43.5 mm,

only known specimen; other measurements, in mm:

carapace length 9.8; anterior width of carapace 4.0;

cornea width 2.3; rostral plate length 1.4, width

1.4; telson length 8.2, width 6.9.

REMARKS.—Oratosquilla ornata can be distinguished from the other species of the Perpensa

Group, including O. perpensa, O. inornata, and O.

imperialis, by the presence of a slender spine

rather than a rounded lobe between the spines of

the basal prolongation of the uropod; as in the

case of most of the species assigned to this group,

the dorsal ridge on the carpus of the claw in O.

ornata is undivided. This new species also has a

much slenderer uropod than any of the members

of the group.

Tirmizi and Manning (1968) observed that a
NUMBER 71 11

slender spine replaced the rounded lobe between

the spines of the basal prolongation of the uropod

in O. hesperia from West Pakistan. In that species

the spines were present in juveniles with a total

length of approximately 50 mm or less, but in

specimens larger than 100 mm the rounded lobe

characteristic of adults was present. Alikunhi

(1967) also noted that the spine was present in

juveniles of O. woodmasoni from India. Thus the

holotype of O. ornata may prove to be a juvenile.

The basic facies of the unique holotype is adult,

however, for the body carination and spination,

usually poorly developed in specimens of this size,

resemble that of larger specimens of other species.

NAME.—The name is from the Latin, ornatus,

meaning decorate, alluding to the well-formed

carinae of the abdomen and the elongate uropod.

DISTRIBUTION.—Known only from the type locality, off Hong Kong in the China Sea, at a depth of

62 fathoms.

Woodmasoni Group

Members of this group can be characterized by

their broad, smooth carapace, with the anterior

width of the carapace greater than one-half its

median length and with the branches of the anterior bifurcation of the median carina of the

carapace poorly developed or completely absent.

The species assigned to this group include O.

woodmasoni (Kemp, 1911), O. massavensis (Kossmann, 1880), O. hesperia (Manning, 1968), and

O. tweediei, described below. Only two of the species, O. massavensis and O. hesperia, have the telson ornamented with dorsal tubercles on each side

of the median carina of the telson. As in the species

of the Oratoria Group, the carpus of the claw is

tuberculate in members of the Woodmasoni Group.

In all four of the species placed in this group

the body is relatively smooth, not so tuberculate

as in O. oratoria, and the carapace is noticeably

broader than in any of the other species in the

genus. The carapace of O. woodmasoni is comparatively the broadest; in that species, the anterior width, at the base of the anterolateral spines,

is usually greater than one-half the length of the

carapace and rostral plate combined. In the other

three species, the anterior length is greater than

one-half the median length of the carapace, excluding the rostral plate.

Further investigation may show that there are

several additional species in this complex; Chopra

(1934), Holthuis (1941), and Manning (1966) all

reported material from different localities that differs from Kemp's (1913) account of O. woodmasoni based primarily on material from India.

Oratosquilla massavensis is not known to occur

in the Indian Ocean adjacent to the Red Sea, but

it is apparently very abundant in the latter area

(Holthuis, 1967). It has been recorded from Viet

Nam by Serene (1953, 1954) ; but in view of its

apparent occurrence only in the Red Sea, Serene's

Key to Species of the Woodmasoni Group

1. Dorsal surface of telson with longitudinal row of tubercles, converging distally, on each

side of median carina 2

Dorsal surface of telson not ornamented with line of tubercles on each side of median

carina 3

2. Lateral margins of rostral plate straight; submedian carinae of abdomen divergent on

each somite, particularly on fourth and fifth somites; telson with 1 row of tubercles

flanking median carina on each side O. hesperia (Manning, 1968)

[Tirmizi and Manning, 1968]

Lateral margins of rostral plate sinuous; submedian carinae of abdomen subparallel;

telson with 2 rows of tubercles flanking median carina on each side

O. massavensis (Kossmann, 1880)

[Ingle, 1963]

3. Submedian carinae of fourth, fifth, and sixth abdominal somites each with posterior

spine; angular lobe present between apical spines of basal prolongation of uropod

O. tweediei, new species

Submedian carinae of fifth and sixth somites only armed with spines; low, inconspicuous rounded lobe present between spines of basal prolongation of uropod

O. woodmasoni (Kemp, 1911)

[Kemp, 1913]
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

records may be based on another undescribed species with a broad carapace, smooth body, and dorsal tubercles on the telson.

Oratosquilla tweediei, new species

FIGURE 4

HOLOTYPE.—1 9, 107 mm; Singapore; M. W. F.

Tweedie, collector; 1934; USNM 76026.

PARATYPE.—1 $, 96 mm; data as in holotype;

USNM.

DIAGNOSIS.—Rostral plate broader than long,

lacking median carina, apex truncate; eye large,

cornea set obliquely on stalk; corneal indices 332-

337; anterior width of carapace more than half

median length; anterior bifurcation of median

carina of carapace absent; raptorial claw with 6

teeth, dorsal ridge of carpus tuberculate, inferodistal angle on outer face of merus bluntly angled;

mandibular palp and 4 epipods present; lateral

process of fifth thoracic somite produced into an

anterior spine and a posterior angled lobe; lateral

processes of sixth and seventh thoracic somites bilobed, anterior lobes sharper and smaller than

posterior; lateral carinae of abdomen not bicarinate; abdominal carinae spined as follows: submedian 4-6, intermediate 3-6, lateral 2-6; telson

lacking supplementary dorsal tubercles; telson denticles 3, 9, 1; distal segment of uropodal exopod

longer than proximal; inner spine of basal prolongation of uropod with angled lobe on outer margin, proximal margin of lobe concave.

DESCRIPTION.—Dorsal surface of body smooth.

Eye large, cornea bilobed, set very obliquely on

stalk; eyes not extending beyond end of first segFIGURE 4.—Oratosquilla tweediei, new species, female holotype, TL 107 mm, Singapore: a, anterior portion of body; b, eye; c, carpus of raptorial claw; d, lateral processes of fifth, sixth, and

seventh thoracic somites; e, basal prolongation of uropod. (Setae omitted.)
NUMBER 71 13

ment of antennular peduncle; ocular scales truncate, inclined laterally, separate mesially; anterior

margin of ophthalmic somite rounded; corneal indices 332-337 in types.

Antennular peduncle subequal to carapace; dorsal processes of antennular somite blunt, truncate,

anterior margin almost transverse.

Rostral plate broader than long, trapezoidal, upturned lateral margins converging on truncate

apex; median carina absent.

Antennal scale slender, short, more than half as

long as carapace.

Carapace broad, anterior width greater than onehalf median length; anterolateral spines of moderate size, not extending to base of rostral plate;

median carina of carapace obscure anteriorly,

branches of anterior bifurcation only faintly indicated; portion of median carina posterior to cervical groove with anterior bifurcation; intermediate

carinae not extending to anterior margin; posterolateral margin of carapace evenly curved, not

angled anteriorly; posterior margin with median

projection.

Dactylus of raptorial claw with 6 teeth; outer

margin of dactylus sinuate; dorsal ridge of carpus

irregularly tuberculate, with 2-3 rounded distal

tubercles; inferodistal angle of outer face of merus

with blunt, triangular tooth.

Mandibular palp and 4 epipods present.

Posterior 3 thoracic somites with well marked,

smooth submedian and intermediate carinae; fifth

somite with faint indications of submedian carinae

and with irregular tubercles at level of intermediate carinae of sixth somite; lateral process of fifth

somite bilobed, anterior lobe a blunt spine directed

anterolaterally, posterior lobe slender, sharp, directed laterally; lateral process of sixth somite

bilobed, anterior lobe slender, sharp, recurved posteriorly, posterior lobe much larger, triangular,

apex acute; lateral process of seventh somite bilobed, anterior lobe small, spiniform, posterior lobe

much larger, apex acute; ventral keel of eighth

somite low, rounded.

Anterior 5 abdominal somites each with 8 carinae, submedians subparallel on anterior somites,

slightly divergent on fifth somite; sixth somite with

6 pairs of carinae; abdominal carinae spined as

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

2-6, marginal 1-5; sixth somite with small ventrolateral spinule in front of articulation of each

uropod and with low median carina on posterior

half of sternum.

Telson flattened, slightly broader than long, with

6 slender marginal spines; prelateral lobes present,

slightly projecting laterally, shorter than carina of

lateral tooth; dorsal surface of telson ornamented

with median carina and carinae of marginal teeth

only, supplementary dorsal tubercles or carinae

absent; median carina sharp, with anterior notch

and long posterior spine flanked distally with 1 or

2 irregular tubercles; denticles rounded, 3, 9, 1;

ventral surface with uninterrupted postanal keel.

Uropods stout, exopod with 7-9 short spines on

outer margin of proximal segment; distal segment

of exopod longer than proximal; endopod slender,

curved; basal prolongation with triangular projection on outer margin of inner spine, inner margin

of projecting lobe concave.

COLOR.—Almost completely faded in the types;

dark pigment present on distal half of proximal

segment and inner half of distal segment of uropodal exopod; distal two-thirds of uropodal endopod dark.

MEASUREMENTS.—Females only known, TL 96-

107 mm. Other measurements, in mm, of holotype,

TL 107 mm: carapace length 20.8; anterior width

of carapace 12.9; cornea width 6.2; rostral plate

length 3.1, width 4.8; telson length 18.8, width 19.2.

REMARKS.—Oratosquilla tweediei belongs to the

Woodmasoni Group of species in the genus, all of

which are characterized by the broad, smooth carapace on which the anterior bifurcation of the

median carina is absent or nearly so. Oratosquilla

tweediei agrees with O. woodmasoni (Kemp) and

differs from both O. massavensis (Kossman) and

O. hesperia (Manning) in lacking supplementary

dorsal tubercles on the telson; in both O. massavensis and O. hesperia the median carina of the

telson is flanked laterally by a line of tubercles that

converge distally under the apical spine.

Oratosquilla tweediei closely resembles O. woodmasoni but can be distinguished from it by the

following features: (1) the anterior width of the

carapace is greater than one-half the median

length, excluding the rostral plate; (2) the anterior

lobes of the lateral processes of both sixth and

seventh thoracic somites are sharper and more

prominent; (3) the lateral process of the eighth

somite is sharper (compare Figure Ad wtih Figure

63 in Kemp, 1913); (4) the submedian carinae of
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the fourth abdominal somite are armed posteriorly;

and (5) the lobe on the inner spine of the basal

prolongation of the uropod is much more prominent in O. tweediei than in O. woodmasoni (compare Figure 4e with Figure 65 in Kemp, 1913).

Oratosquilla woodmasoni and O. tweediei are

the eastern counterparts of O. hesperia and O.

massavensis in the Indo-West Pacific region; O.

hesperia is known from the northwestern Arabian

Sea, south to Madagascar, and O. massavensis apparently occurs only in the Red Sea (Tirmizi and

Manning, 1968). Oratosquilla woodmasoni is

known from localities between Hong Kong and

western India (Kemp, 1913) and O. tweediei is

known only from off Singapore.

This new species agrees with the specimen I

reported in 1966 from off Bowen, Queensland, and

differs from previous accounts of O. woodmasoni

in having a well-developed lobe on the outer margin of the inner spine of the basal prolongation of

the uropod. The Australian specimen, however,

differs in having the anterior branches of the bifurcation of the median carapace; these carinae are

completely lacking in both O. woodmasoni sensu

stricto and in O. tweediei. It seems likely that the

Australian specimen represents a distinct species.

The only other species in the genus with the submedian carinae of the fourth abdominal somite

armed are: (1) O. nepa (Latreille), which differs

in having very small eyes and in having the anterior

bifurcation of the median carina of the carapace

open posterior to the dorsal pit; (2, 3) O. mauritiana

(Kemp) and O. calumnia (Townsley), both of

which differ in having an entire median carina

on the carapace with a well-developed anterior

bifurcation; (4) O. fabricii (Holthuis), in which

the median carina is interrupted but the anterior

bifurcation is well formed; and (5, 6) O massavensis (Kossmann) and O. hesperia (Manning),

both of which have supplementary dorsal tubercles

on the telson. All of these species differ from O.

tweediei in other features as well, but these are

the most important differences.

NAME.—The species is named for M. W. F.

Tweedie, who collected the specimens and donated

them to the United States National Museum.

DISTRIBUTION.—Known only from the type locality, Singapore.

Gonypetes Group

This group contains three species, which have

but 5 teeth on the dactylus of the claw, O. quadraticauda (Fukuda, 1911), O. quinquedentata

(Brooks, 1886), and O. gonypetes (Kemp, 1911).

In other respects the species in the Gonypetes

Group resemble those in other groups with 6 teeth

on the claw, especially the species of the Woodmasoni Group, which lack a well-formed anterior

bifurcation on the median carina of the carapace.

As Holthuis (1967) commented, these species are

extremely difficult to identify when their raptorial

claws are missing.

In 1965, I pointed out that Squilla hoops Kemp

was a synonym of S. quadraticauda Fukuda.

On the basis of his studies of stomatopod larvae

aqd juvenile stages, Alikunhi (1967) suggested that

O. gonypetes and its allies [O. quadraticauda and

O. quinquedentata] are more closely related to the

"Chloridella" [now Clorida] and "raphidea" [now

Harpiosquilla] groups of species than are species

of the "nepa" [Nepa, Perpensa, and Woodmasoni]

groups.

Key to Species of the Gonypetes Group

1. Rostral plate with median carina; lateral process of seventh thoracic somite not bilobed

(eyes very large, cornea width about one-third carapace length)

O. quadraticauda (Fukuda, 1911) [ = Squilla boops Kemp, 1911]

[Kemp, 1913]

Rostral plate without median carina; lateral process of seventh thoracic somite bilobed

(eyes of moderate size, cornea width less than one-fourth carapace length) 2

2. Outer face of merus of claw with tooth at inferodistal angle; anterior lobe of lateral

process of seventh thoracic somite short, not exceeding half of length of posterior

lobe O. quinquedentata (Brooks, 1886)

Outer face of merus of claw unarmed inferodistally; anterior lobe of lateral process of

seventh thoracic somite more than half as long as posterior lobe. O. gonypetes (Kemp, 1911)

[Kemp, 1913]
NUMBER 71

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