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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 521

Stomatopod Crustacea Collected by the Galathea Expedition, 1950-1952, with a List of Stomatopoda Known from Depths below 400 Meters

Raymond B. Manning

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1991

A B S T R A C T

Manning, Raymond B. Stomatopod Crustacea Collected by the Galathea Expedition, 1950–1952, with a List of Stomatopoda Known from Depths below 400 Meters. Smithsonian Contributions to Zoology, number 521, 18 pages, 12 figures, 1991.―Thirty-five species taken by the Galathea expedition are reported, most from shallow water stations. The seventeen species of Stomatopoda known with certainty to live in depths greater than 400 meters are listed. Acaenosquilla, new genus, is recognized for two species previously placed in Heterosquilloides, A. brazieri (Miers, 1880) and A. latifrons (De Haan, 1844). The collection includes a late larva of Bathysquilla crassispinosa (Fukuda, 1909), the first known larva of a bathysquillid. Five species are synonymized: Clorida javanica Moosa, 1974, with C. latreillei (Eydoux and Souleyet, 1842), Clorida merguiensis (Tiwari and Biswas, 1952) with C. verrucosa (Hansen, 1926), Gonodactylus childi Manning, 1971, with G. incipiens Lanchester, 1903, Heterosquilla (Heterosquilloides) zarenkovi Makarov, 1978, with Heterosquilloides insignis (Kemp, 1911), and both Squilloides latus spinosus Blumstein, 1970, and Squilloides espinosus Blumstein, 1974, with Lenisquilla lata (Brooks, 1886).

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication Data

Manning, Raymond B.

Stomatopod Crustacea collected by the Galathea Expedition, 1950–1952. with a list of Stomatopods known from depths below 400 meters / Raymond B. Manning.

p. cm. – (Smithsonain contributions to zoology; no. 521)

Includes bibliographical references.

1. Stomatopoda–Catalogs and colleclions–Denmark–Copenhagen. 2. Galathea Expedition (1950–1952) 3. Københavns Universitet. Zoologisk museum-Catalogs. I. Title. II. Series.

[QL444.M375] 591 s–d20 [595.3'82'0748913] 91-23081 CIP

art The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.

Stomatopod Crustacea Collected by the Galathea Expedition, 1950–1952, with a List of Stomatopoda Known from Depths below 400 Meters

Raymond B. Manning

____________

Raymond B. Manning, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Reviewers: Marjorie L. Reaka, University of Maryland; and David K. Camp, Florida Marine Research Institute.

Introduction

This report is based on the Stomatopoda collected during the voyage of the Danish vessel Galathea around the world between 1950 and 1952. Station data for the collections of the Galathea have been given by Bruun (1959), who listed the stations from below 400 meters, and by Wolff (1964), who provided data for the shallower stations. Five West African species taken by the Galathea, Eurysquilla galatheae Manning, 1977, Lysiosquilla hoevenii (Herklots, 1851), Nannosquilloides occultus (Giesbrecht, 1910), Protosquilla calypso Manning, 1974, and Rissoides africanus (Manning, 1974), were reported by Manning (1977), and the single Eastern Pacific species taken by the expedition, Squilla biformis Bigelow, 1891, was reported by Manning (1974b).

The majority of the stomatopods collected by the Galathea were taken in depths shallower than 400 meters. Only two species, Heterosquilloides insignis (Kemp, 1911) from 445 to 460 meters and Squilla biformis Bigelow, 1891, from 450 to 500 meters, were taken with certainty from below 400 meters. One specimen of the shallow-water Austrosquilla vercoi (Hale, 1924), was labelled as having been collected in 875 or 950 meters, but the station data accompanying the specimen are certainly erroneous.

The measurement given after the number of specimens taken at each station is total length (TL) in millimeters (mm), measured from the anterior edge of the rostral plate to a line drawn between the apices of the submedian spines of the telson. In the “Material” section, meters is abbreviated to m. Data given for each station are from the station data in Wolff (1964); if the depth on the specimen label is different, it, too, is given. All specimens are in the collections of the Zoological Museum, Copenhagen. Coordinates added by me are enclosed in parentheses and are from gazetteers of the United States Board on Geographic Names.

ACKNOWLEDGMENTS.—I thank Torben Wolff and Poul Chr. Jeppesen, Zoological Museum, Copenhagen, for much help during the course of this study.

A visit to the Zoological Museum in 1988 was supported by the Suzanne Liebers Erickson Exchange Program of the Smithsonian Institution. This research also was supported in part by the Smithsonian through the Scholarly Studies Program and former Secretary S. Dillon Ripley’s Fluid Research Fund; that support is gratefully acknowledged.

I thank Roy K. Kropp, Battelle Ocean Sciences, Rafael Lemaitre, Smithsonian Oceanographic Sorting Center, Marjorie L. Reaka, University of Maryland, and epecially David K. Camp, Florida Marine Research Institute, for reviewing the manuscript; Camp caught many inconsistencies and errors in a late draft of the manuscript, and his efforts materially improved this report.

The illustrations were prepared by my wife Lilly.

Superfamily BATHYSQUILLOIDEA Manning, 1967

Family BATHYSQUILLIDAE Manning, 1967

BATHYSQUILLIDAE Manning, 1967a:238.

Bathysquilla crassispinosa (Fukuda, 1909)

FIGURES 1, 2

Lysiosquilla crassispinosa Fukuda, 1909:61; 1910:146, pl. 10: fig. 4.—Kemp, 1913:117.

Bathysquilla crassispinosa.—Moosa, 1986:371, pl. 1: fig. a,b.—Bruce, 1988:87, figs. 1A-C, 2, 3A-H, 4A, 5, Frontispiece A [in color].—Manning, Kropp, and Dominguez, 1990:312.

FIGURE 1.—Bathysquilla crassispinosa (Fukuda, 1909); lateral view of larva, TL 40 mm, sta 200.

FIGURE 2.—Bathysquilla crassispinosa (Fukuda, 1909); larva, TL 40 mm, sta 200: a, carapace; b, raptorial claw; c, sixth abdominal somite, telson, and uropods; d, uropod; e, uropod, ventral view.

MATERIAL.—Sta 200, off Natal, South Africa, 29°39′S, 37°01′E, dip net at lantern light in about 5000 m, 18 Feb 1951: 1art larva, 40 mm.

REMARKS.—This is the first record of the larva of a bathysquillid. The presence of four pairs of marginal teeth on the telson and the pectinate propodus of the raptorial claw allow its identification as a bathysquillid. The specimen has the broad telson characteristic of Bathysquilla rather than the elongate telson found in Indosquilla. It is identified with B. crassispinosa as that is the only member of the genus known to occur in the area as an adult.

The specimen is a late larva, probably in the last pelagic larval stage preceding the molt to postlarva; the buds of the copulatory tubes are present. Among the features shown by this specimen are two dorsal spines on the carpus of the claw, the proximal very small, nine teeth on the dactylus of the claw, only one exposed thoracic somite, and an unsegmented uropodal exopod. Adults of B. crassispinosa have 9 to 11 teeth on the claw.

Perhaps the most striking feature of this specimen is the unsegmented uropodal exopod, a characteristic of the fossil family Sculdidae (see Manning, Kropp, and Dominguez, 1990), which possibly strengthens the hypothesis that the bathysquillids comprise a very old stock of stomatopods. However, in at least some stomatopod larvae the uropodal exopod is unsegmented in the last larval stage, as shown by the last larva of the lysiosquillid Heterosquilla tricarinata (Claus, 1871) (Greenwood and Williams, 1984, fig. 4P). In the last larval stage of squillids the uropodal exopod also may be unsegmented or the two segments are separated by an indistinct suture, as shown by Alikunhi (1952) for various species.

The original account of this species was published by Fukuda in Japanese in 1909, not in 1910 as often cited; the 1910 version is an English translation of the original account.

DISTRIBUTION.—Indo-West Pacific, from scattered localities between Japan and South Africa, including Madagascar (Manning, Kropp, and Dominguez, 1990) and in the Coral Sea, off Australia (Bruce, 1988); in slope depths between about 200 and about 400 (350–420) meters.

Superfamily GONODACTYLOIDEA Giesbrecht, 1910

Family EURYSQUILLIDAE Manning, 1977

EURYSQUILLIDAE Manning, 1977:33.

Eurysquilla galatheae Manning, 1977

Eurysquilla galatheae Manning, 1977:35, figs. 4, 5, 53.

MATERIAL.—Sta 116, Loanda-Lobito, Angola, 12°13′S, 13°27′E, 100 m, clayey sand, 20 Dec 1950: 1art, 31 mm (holotype).

DISTRIBUTION.—West Africa, from off Angola and off Gambia; sublittoral, in 18–100 meters.

Family GONODACTYLIDAE Giesbrecht, 1910

GONODACTYLINAE Giesbrecht, 1910:148.

Gonodactylus chiragra (Fabricius, 1781)

Squilla chiragra Fabricius, 1781:515.

Gonodactylus chiragra.—Kemp, 1913:155, fig. 2, pl. 9: fig. 107.

MATERIAL.—Sta 415, Tubajon Bay, Philippines, 10°20′N, 125°32′E, coral reef, tidal zone; 17–19 Jul 1951: 4art, 26–83 mm (1, 26 mm, in gastropod shell).

REMARKS.—This species was taken with G. mutatus and G. viridis at sta 415.

DISTRIBUTION.—Apparently this species is widely distributed in the Indo-West Pacific region, but most records require verification; shore.

Gonodactylus falcatus (Forskål, 1775)

Cancer falcatus Forskål, 1775:96.

Gonodactylus falcatus.—Manning, 1978a:4, figs. 1, 2a, 9.—Moosa, 1984:39.—Manning and Lewinsohn, 1986:7.—Moosa. 1989:224.

MATERIAL.—Sta 255, Mombasa, Kenya, 4°05′S, 39°41′E, coral reef, 0 m, 22 Mar 1951: 1art, 37 mm.

REMARKS.—This specimen has a median tubercle on the sixth abdominal somite, sharp central carinae on the telson, and a bilobed knob. It is clearly referable to G. falcatus.

This is the second record for this species in the Indian Ocean outside of the Red Sea proper and indicates that this species can be expected to occur elsewhere in the western Indian Ocean. Moosa (1984) reported it from Mauritius.

Moosa (1989) also reported this species from Japan, but as broad distibution patterns in this group of species appear to be the exception rather than the rule (Manning and Lewinsohn, 1986:9), his range extension for this species must be accepted with caution.

This species was taken with G. smithii at sta 255.

DISTRIBUTION.—Western Indian Ocean, from Mombasa, Kenya, Mauritius, the Red Sea, and, possibly, in the Pacific, from Japan; shore.

Gonodactylus incipiens Lanchester, 1903

FIGURE 3

Gonodactylus chiragra var. incipiens a Lanchester, 1903:451, pl. 23: fig. 10.

Gonodactylus incipiens.—Manning, 1967b:18, fig. 7.—Budiman and Moosa, 1983:399,400.

Gonodactylus childi Manning, 1971:75, fig. 1 [new synonymy].

MATERIAL.—Sta 682, Makaha’a at Nukualofa, Tonga Islands, 21°07′S, 175°10′W, 0.5–1 m, coral reef, 8 Mar 1952: 1art, 20 mm.

REMARKS.—Gonodactylus incipiens is one of the most common shore stomatopods in the central Pacific and, like G. affinis De Man, 1902, shows a wide range of variation in surface sculpture of the telson (see Moosa and Cleva, 1984a, fig. 2).

Gonodactylus childi Manning, 1971, originally described from Enewetak Atoll, is a synonym of G. incipiens. A second, similar species also described from Enewetak, Gonodactylus micronesica Mannning (1971:77, fig. 2) is a distinct species that lives in deeper water, 100 ft (about 30 m) as opposed to shore, and that has much longer accessory median carinae on the telson. In G. micronesica the accessory median carinae extend anteriorly beyond the posterior slope of the anterior submedian carinae; in G. incipiens they barely reach the posterior edge of the anterior submedian carinae.

DISTRIBUTION.—Known with certainty from several localities in the central Pacific, including Guam, Enewetak, the Ellice Islands, the Tuamotu Archipelago, and Australia, and in the Indian Ocean from Sri Lanka; shore.

FIGURE 3.—Gonodactylus incipiens Lanchester, 1903; male, TL 20 mm, sta 682: a, rostral plate and ocular scales; b, lateral processes of sixth and seventh thoracic somites, right side; c, sixth abdominal somite, telson, and uropod; d, submedian teeth of telson, ventral view; e, uropod, ventral view.

Gonodactylus lanchesteri Manning, 1967

Gonodactylus lanchesteri Manning, 1967b;11, fig. 4.—Manning and Lewinsohn, 1986:10.

MATERIAL.—Sta 224, Bay at Cape Diego (12°15′S, 49°20′E), Diego Suarez, Madagascar, 0.3 m, rocks with corals, 3 Mar 1951: 2art, 15 and 22 mm.

REMARKS.—There is also a male of this species, TL 27 mm, from the Red Sea at Abu Sardaf, Hurghada (as Ghardaqa, 27°14′N, 33°50′E), Egypt, in the collection of the Zoological Museum, Copenhagen.

DISTRIBUTION.—Western Indian Ocean, from the Red Sea to South Africa; shore.

Gonodactylus mutatus Lanchester, 1903

Gonodactylus chiragra var. mutatus Lanchester, 1903:450.

Gonodactylus mutatus.—Manning, 1978a:7, figs. 4, 5, 11.—Budiman and Moosa, 1983:399, 400.

MATERIAL.—Sta 415, Tubajon Bay, Philippines, 10°20′N, 125°32′E, coral reef, tidal zone, 17–19 Jul 1951: 1art, 31 mm.

REMARKS.—This species was taken with G. chiragra and G. viridis at sta 415.

DISTRIBUTION.—Western Indian Ocean to eastward to Vietnam, Thailand, and the Philippines; shore.

Gonodactylus smithii Pocock, 1893

Gonodactylus Smithii Pocock, 1893:475, pl. 20B: fig. 1.

Gonodactylus smithii.—Budiman and Moosa, 1983:399, 400.

MATERIAL.—Sta 255, Mombasa, Kenya, 4°05′S, 39°41′E, coral reef, 0 m, 22 Mar 1951: 1art, 32 mm.

REMARKS.—This specimen lacks the sharp, spiniform anterolateral angles of the rostral plate characteristic of G. acutirostris De Man, 1898 (see Manning and Lewinsohn, 1986, fig. 2a), and so is herein referred to G. smithii. Both species occur in the western Indian Ocean.

This species was taken with G. falcatus at sta 255.

DISTRIBUTION.—Western Indian Ocean to Vietnam, Indonesia, and Australia; shore.

Gonodactylus viridis Serène, 1954

Gonodactylus chiragra var. viridis Serène, 1954:7, 10, 74, 75, 76, fig. 13–3.

Gonodactylus viridis.—Manning, 1978c:4, fig. 2a–c.—Budiman and Moosa, 1983:399, 400.—Moosa, 1989:226.

MATERIAL.—Sta 415, Tubajon Bay, Philippines, 10°20′N, 125°32′E, coral reef, 17–19 Jul 1951: 3art, 21–33 mm, 2art, 26–50 mm.

REMARKS.—Gonodactylus viridis was taken with G. chiragra and G. mutatus at sta 415.

DISTRIBUTION.—Previously reported from Phuket Island, Thailand, in the Andaman Sea, and in the Pacific from Vietnam, Indonesia, the Philippines, and the Ryukyu Islands, Japan; shore.

Mesacturoides fimbriatus (Lenz, 1905)

Gonodactylus fimbriatus Lenz, 1905:388, pl. 47: fig. 11.—Kemp, 1913:175.—Manning. 1962:8.

Mesacturoides raymondi Tirmizi and Kazmi, 1980:61, figs. 1, 2.

Mesacturoides fimbriatus.—Manning, 1990:99.

MATERIAL.—Sta 256, Mombasa, Kenya, 4°05′S, 39°41′E, 1–2 m, corals, 22 Mar 1951: 1art, 17 mm, 1art, 12 mm.

REMARKS.—Manning (1990) showed that M. raymondi Tirmizi and Kazmi was a synonym of M. fimbriatus.

This species was taken with Chorisquilla spinosissima at sta 256.

DISTRIBUTION.—Western Indian Ocean, from Zanzibar, the Seychelles, Pakistan, and now from Mombasa, Kenya; shore.

Family ODONTODACTYLIDAE Manning, 1980

ODONTODACTYLIDAE Manning, 1980:366.

Odontodactylus brevirostris (Miers, 1884)

Gonodactylus brevirostris Miers, 1884:567, pl. 52: fig. C.

Odontodactylus brevirostris.—Kemp, 1913:143.—Manning, 1967c:22, figs. 6–8.

MATERIAL.—Sta 675, Raoul Island, Kermadec Islands, 29°13′S, 177°57′W, 60 m, stones, 3 Mar 1952: 1art, 29 mm.

DISTRIBUTION.—Western Atlantic and Indo-West Pacific, from the western Indian Ocean to Hawaii; sublittoral, in about 40 to 73–424 meters, usually in less than 100 meters. Juveniles have been taken at the surface.

Odontodactylus cultrifer (White, 1850)

Gonodactylus cultrifer White, 1850:96, pl. 16: figs. 1, 2.

Odontodactylus cultrifer.—Kemp, 1913:137.—Manning, 1967c:18, fig. 5.

MATERIAL.—Sta 428, Candos Bay, Mindanao, Philippines, 9°36′N 125°46′E, dip net at lantern light in 22 m, 30 Jul 1951: 1art, 74 mm.

DISTRIBUTION.—Western Pacific Ocean, from Australia and New Guinea northward to Japan; shallow sublittoral.

Family PROTOSQUILLIDAE Manning, 1980

PROTOSQUILLIDAE Manning, 1980:366.

Chorisquilla spinosissima (Pfeffer, 1888)

Gonodactylus spinosissimus Pfeffer, 1888:35.—Kemp, 1913:191, pl. 10: figs. 124, 125.

MATERIAL.—Sta 256, Mombasa, Kenya, 4°05′S, 39°41′E, 1–2 m, corals, 22 Mar 1951: 1art, 22 mm.

REMARKS.—There are also two males, one TL 20 mm, taken at Dana sta 3742 (anchorage at Biaro, Indonesia, 2°08.6′N, 125°21.7′E, 11 m, 6 Jul 1929), and a second, TL 21 mm, taken from reefs at the Murray Islands (9°55′S, 144°05′E) in 1888, in the Zoological Museum, Copenhagen.

This species was taken with Mesacturoides fimbriatus at sta 256.

DISTRIBUTION.—Indo-West Pacific, from the western Indian Ocean to Japan; shore to about 90 meters, usually in shallow water.

Protosquilla calypso Manning, 1974

Protosquilla calypso Manning, 1974a:8, fig. 1; 1977:50, figs. 10, 11, 53.

MATERIAL.—Sta 50, Islas das Rôlas, São Tomé, 00°00′N, 06°32′E, 5–8 m, coral, 29 Nov 1950: 3art, 17–20 mm, 8art, 11–23 mm.

DISTRIBUTION.—West Africa, from the offshore islands of the Gulf of Guinea; shore to about 69 (9–69) meters.

FIGURE 4.—Acaenosquilla latifrons (De Haan, 1844); female, TL 63 mm, Nagasaki, Japan (specimen in collection of the National Museum of Natural History, Smithsonian Institution): a, front; b, rostral plate, lateral view; c, eye; d, basal antennal segment, lateral view; e, raptorial claw; f, lateral processes of exposed thoracic somites; g, sixth abdominal somite, telson, and uropod; h, telson, ventral view; i, uropod, ventral view, inner stiff setae omitted.

Superfamily LYSIOSQUILLOIDEA Giesbrecht, 1910

Family LYSIOSQUILLIDAE Giesbrecht, 1910

LYSIOSQUILLINAE Giesbrecht, 1910:148.

Acaenosquilla, new genus

DEFINITION.—A lysiosquillid of moderate to large size, maximum length about 100 mm. Cornea bilobed. Rostral plate broader than long, with subapical ventral keel fitting between separate ocular scales. Antennal protopod with 1 ventral papilla. Claw with 6 or 7 (usually 6) teeth on dactylus and 4 movable teeth on propodus. Mandibular palp and 5 epipods present. Telson with 3-spined median elevation and 3 pairs of dorsal carinae; marginal armature consisting of row of small submedian denticles, 1 movable submedian tooth, 4 intermediate denticles, 1 intermediate tooth, 1 lateral denticle, and 1 lateral tooth. Ventral surface of telson with strong postanal tooth. Uropodal endopod lacking strong proximal fold on outer margin.

TYPE SPECIES.—Lysiosquilla latifrons De Haan, 1844 (Figure 4). The genus includes Acaenosquilla latifrons from Japan and A. brazieri (Miers, 1880) from Australia and New Zealand (see accounts of these species in Stephenson, 1962:38, 39).

ETYMOLOGY.—From the Greek, akaina (thorn) and the generic name Squilla, alluding to the postanal spine present in members of the genus. The gender is feminine.

REMARKS.—The combination of the ventrally-keeled rostral plate and the postanal spine will distinguish this genus from all other genera in the family.

Heterosquilloides insuetus Manning (1970b:81, fig. 2) from Australia and H. philippinensis Moosa (1986:386, fig. 5) from the Philippines also have a strong postanal spine but both species lack the ventral keel on the rostral plate and the antennal papillae; both species have a longer rostral plate and more dorsal spines on the telson. They should be referred to a new genus.

Acaenosquilla brazieri (Miers, 1880), new combination

Lysiosquilla Brazieri Miers, 1880:11, 125, pl. 1: figs. 3–6.

Heterosquilla (Heterosquilloides) brazieri.—Manning, 1966:125.

MATERIAL.—Sta 539, Coral Sea, 26°33′S, 153°31′E, 55 m (86 m on label), gravel, 5 Nov 1951: 1art, 23 mm.

REMARKS.—This small specimen is in very poor condition. The carinae that flank the median prominence of the telson in adults are not visible. There are 6 teeth on the claw, the penultimate smaller than the antepenultimate. The uropodal exopod has 6 outer spines and 7 inner stiff setae distally on the proximal segment.

DISTRIBUTION.—Previously recorded from localities in Australia and New Zealand; sublittoral, in depths of less than 100 meters.

Heterosquilloides insignis (Kemp, 1911)

FIGURE 5

Lysiosquilla insignis Kemp, 1911:94; 1913:126, pl. 9: figs. 99–102.

Heterosquilla (Heterosquilloides) zarenkovi Makarov, 1978:179, fig. 2 [new synonymy].

Heterosquilloides insignis.—Moosa, 1986:386, pl. 1: fig. c.

MATERIAL.—Sta 196, off Durban, South Africa, 29°55′S, 30°20′E, 445–460 m, sandy mud, 13–14 Feb 1951: 1art, 42 mm.

REMARKS.—I can see no differences between H. zarenkovi Makarov from Vietnam and H. insignis, which has been recorded from the Philippines by Moosa (1986), so I consider Makarov’s species to be a synonym of the latter. Even the sculpture of the telson in Makarov’s species appears to be identical with that shown here.

DISTRIBUTION.—Indian Ocean, from South Africa and the Andaman Islands, and Pacific Ocean from the Philippines and Vietnam; in depths between 275 and 415–510 meters.

FIGURE 5.—Heterosquilloides insignis (Kemp, 1911); male, TL 42 mm, sta 196: a, front; b, eye; c, sixth abdominal somite and telson; d, uropod, ventral view.

FIGURE 6.—Acanthosquilla derijardi Manning, 1970; female, TL 21 mm, sta 454: a, front; b, propodus and dactylus of raptorial claw; c, sixth abdominal somite, telson, and uropod; d, telson, ventral view; e, uropod, ventral view.

Lysiosquilla hoevenii (Herklots, 1851)

Squilla Hoevenii Herklots, 1851:17, 26, pl. 1: fig. 11.

Lysiosquilla hoevenii.—Manning, 1977:74, figs. 20, 21, 26a–c, 54.

MATERIAL.—Sta 48, São Tomé, 00°22′N, 06°46′E, 10 m, anchorage, 28 Nov 1950: 3art, 106–118 mm.

DISTRIBUTION.—West Africa, from the Cape Verde Islands to Angola; shallow water to 30 meters.

Lysiosquilla maculata (Fabricius, 1793)

Squilla maculata Fabricius, 1793:511.

Lysiosquilla maculata.—Kemp, 1913:111, pl. 8: figs. 86–91.—Manning, 1978b:3, figs. 1–3, 9.

MATERIAL.—Sta 269, Port Victoria (4°37′S, 55°27′E), Mahé, Seychelles, anchorage, dip net at lantern light in 28 m, 29 Mar 1951: 2art, 146–167 mm, 1art, 140 mm.

DISTRIBUTION.—Widely distributed in the Indo-West Pacific, from the western Indian Ocean to Japan; shallow water.

Family NANNOSQUILLIDAE Manning, 1980

NANNOSQUILLIDAE Manning, 1980:368.

Acanthosquilla derijardi Manning, 1970

FIGURE 6

Acanthosquilla derijardi Manning, 1970a:1430, 1434, fig. 2.—Moosa, 1973:14; 1975:8.—Holthuis, 1975:309.

(?)Acanthosquilla derijardi.—Moosa and Cleva, 1984a:427.

MATERIAL.—Sta 454, Java Sea, 5°23′S, 116°02′E, 60 m, coral-clay, 25 Aug 1951: 1art, 21 mm.

REMARKS.—This species was taken with Lenisquilla lata at Sta 454.

Moosa and Cleva (1984) questioned their record of this species from the Seychelles as their specimen was in poor condition.

DISTRIBUTION.—Known from scattered localities between the Red Sea and the Caroline Islands, including Madagascar and the Seychelles in the Indian Ocean, and from Borneo, Indonesia, the Philippines, and the Caroline Islands in the Pacific; shore to 60 meters.

Austrosquilla vercoi (Hale, 1924)

FIGURE 7

Lysiosquilla vercoi Hale. 1924:499, fig. 383, pl. 33: fig. 2.

Heterosquilla (Austrosquilla) vercoi.—Manning, 1966:130, fig. 10.

FIGURE 7.—Austrosquilla vercoi (Hale, 1924); female, TL 22 mm, sta 555: a, front; b, sixth abdominal somite, telson, and uropod.

MATERIAL.—Sta 555, Great Australian Bight, 37°31′S, 138°44′E, 875 m (950 m on label), sand, a little clay, 6 Dec 1951: 1art, 22 mm.

REMARKS.—This small specimen is clearly referable to A. vercoi but differs from adults in several features: the eye is broader, the antennal papillae are scarcely visible, the rostral spine is longer, the distal segment of the uropodal exopod is longer, and there are fewer stiff setae on the inner margin of the proximal segment of the uropodal exopod.

This species generally occurs in shallow water. The label data for both locality and depth certainly are erroneous.

DISTRIBUTION.—Australia and Tasmania; shore and shallow sublittoral.

Nannosquilloides occultus (Giesbrecht, 1910)

Lysiosquilla occulta Giesbrecht, 1910:3. 45, 52, 58–75, 125, 127, 131. pl. 6: figs. 1–3, 14, 17–20, 22–29, 33–35, 38, 39. pl. 7, pl. 8: figs. 1–39.

Nannosquilloides occulta.—Manning, 1977:91, figs. 27, 54.

MATERIAL.—Sta 89, off Congo River, 06°26′S, 11°56′E, 100 m, 9 Dec 1950: 1art, 15.5 mm.

Sta 115, Loanda-Lobito, Angola, 12°14′S, 13°25′E, 200 m, clayey sand, 20 Dec 1950: 1art, 15 mm.

DISTRIBUTION.—Mediterranean Sea and off West Africa, from Senegal to Angola; sublittoral.

Superfamily SQUILLOIDEA Latreille, 1803

Family HARPIOSQUILLIDAE Manning, 1980

HARPIOSQUILLIDAE Manning, 1980:367.

Harpiosquilla harpax (De Haan, 1844)

Squilla harpax De Haan, 1844: atlas, pl. 51: fig. 1; 1849:222.—Tiwari and

Harpiosquilla harpax.—Manning, 1969b:25, figs. 28–38.—Moosa, 1986:390.

MATERIAL.—Sta 305, Bay of Bengal, Ceylon-Calcutta, 20°51′N, 87°58′E, 43–52 m, mud, 26 Apr 1951: 5art, 70–105 mm, 3art, 77.5–98 mm.

REMARKS.—This species was taken with Lophosquilla tiwarii at sta 305.

DISTRIBUTION.—Widely distributed in the Indo-West Pacific region, from the western Indian Ocean to Japan; sublittoral, in depths of less than 100 m.

Family SQUILLIDAE Latreille, 1803

SQUILLARES Latreille, 1803:36.

Anchisquilla fasciata (De Haan, 1844)

Squilla fasciata De Haan, 1844: atlas, pl. 51: fig.4; 1849:224.—Kemp, 1913:34, pl. 1: figs. 21–23.

Anchisquilla fasciata.—Moosa, 1986:390.

MATERIAL.—Sta 451, Makassar Strait, 01°25′S, 117°05′E, 50–60 m, mud with shells, 23 Aug 1951: 2 damaged specimens.

Sta 531, 1 sea mile S of Port Moresby, New Guinea (9°30′S, 147°07′E), 30 m, mud, gravel, 25 Oct 1951: 1art, 24 mm.

DISTRIBUTION.—Known from scattered localities in the Indo-West Pacific, from the Red Sea to Japan; sublittoral, usually in depths shallower than 100 m.

Carinosquilla multicarinata (White, 1847)

Squilla multicarinata White, 1847:144, pl. 6: fig. 1.—Kemp, 1913:86, pl. 6: figs. 73–76.

MATERIAL.—Sta 360, 300 m W of Sultan Shoal Lighthouse (Sultan Shoal = 1°15′N, 103°39′E), Singapore, 25 m, muddy sand with fragments of corals and shells, 31 May 1951: 1art, 28 mm.

FIGURE 8.—Clorida gaillardi Moosa, 1986; female, TL 41 mm, sta 380: a, front; b, eye; c, raptorial claw; d, lateral processes of fifth to seventh thoracic somites; e, sixth abdominal somite, telson, and uropod; f, uropod, ventral view.

REMARKS.—Naiyanetr (1983:399) has provided keys to the species of Carinosquilla.

DISTRIBUTION.—Pacific Ocean, from Japan to Australia; sublittoral.

Clorida gaillardi Moosa, 1986

FIGURE 8

Clorida gaillardi Moosa, 1986:396, fig. 9.

MATERIAL.—Sta 380, off Kerteh, Trengganu, Malaysia, 04°43′N, 103°43′E, 48 m, 7 Jun 1951: 1art, 41 mm.

REMARKS.—This specimen agrees well with the original account of this species by Moosa. This species could easily be confused with Clorida latreillei (Eydoux and Souleyet, 1842), also taken by the Galathea, but differs in having a much larger lateral process on the fifth thoracic somite, more spines on abdominal carinae, less ornamentation on the telson, and a larger lobe between the spines of the basal prolongation of the uropod.

Clorida gaillardi was taken with C. verrucosa at sta 380.

DISTRIBUTION.—Previously known only from the Philippines and now from Malaysia; sublittoral, in 36–37 and 41 meters.

Clorida latreillei (Eydoux and Souleyet, 1842)

FIGURE 9

Clorida latreillei Eydoux and Souleyet, 1842:265, pl. 5: figs. 2–5.

Squilla latreillei.—Kemp. 1913:24, pl. 1: figs. 1–4.

Clorida javanica Moosa, 1974:76, figs. 2, 3 [new synonymy].

MATERIAL.—Sta 225, Bay at Cape Diego (12°15′N, 49°20′E), Diego Suarez, Madagascar, 6–8 m, sand with mud and stones, 3 Mar 1951: 1art, 57 mm.

REMARKS.—I believe that the features of Clorida javanica Moosa fall within the range of variation of characters of C. latreillei and that these taxa are synonymous. Moosa (1974:79) noted that C. javanica keyed out to C. latreillei in the key given by Manning (1968a:4–5). He also noted that C. javanica differed from C. latreillei in several features, as follows: the shape of the lateral process of the fifth thoracic somite; the lack of spines on the marginal carinae of the abdomen, except for that on the fifth somite; the absence of a postanal keel; and the color pattern.

The shape of the lateral process of the fifth thoracic somite and the numbers of spined marginal carinae on the abdomen are both variable features; the latter may differ from side to side in this species. Color pattern in preserved specimens may also be variable and is dependent at least in part on method of preservation.

This leaves the presence or absence of a postanal keel as the main character separating these two species. When present, the keel may be well developed and rather long or it may be low and short (Manning, 1969a:8). Numerous features of members of Clorida have proven to be variable, including presence or absence of a structure within a given species (see below, under C. verrucosa), and variability may be the rule rather than the exception in Clorida.

DISTRIBUTION.—Widely distributed in the Indo-West Pacific, from the western Indian Ocean to Japan; sublittoral.

FIGURE 9.—Clorida latreillei (Eydoux and Souleyet, 1842); female, TL 27 mm, sta 225: a, front; b, eye; c, raptorial claw; d, lateral processes of fifth to seventh thoracic somites; e, sixth abdominal somite, telson, and uropod; f, uropod, ventral view.

Clorida verrucosa (Hansen, 1926)

FIGURES 10, 11

Squilla microphthalma.—Kemp and Chopra, 1921:299 [part; not S. microphthalma H. Milne Edwards. 1837].

Squilla verrucosa Hansen, 1926:3, pl. 1: fig. 1.—Manning, 1968b:124.

Squilla merguiensis Tiwari and Biswas, 1952:350, fig. 1a.—Manning, 1968b: 124 [new synonymy].

Clorida verrucosa.—Manning, 1968a:5 [key].—Anonymous, 1972:27.—Blumstein, 1974:116, fig. 5.—Manning, 1976:10, fig. 5.—Makarov, 1979:42, fig. 2.

Clorida merguiensis.—Manning, 1968a:5 [key].—Blumstein, 1974:116, fig. 4.—Manning, 1976:8, fig. 4.—Makarov, 1979:44. fig. 3.—Moosa, 1986:400 [new synonymy].

MATERIAL.—Sta 294, off Tranquebar, India, 11°10′N, 79°59′E, 50 m, coarse sand and mud, 22 Apr 1951: 1art, 32 mm.

Sta 380, off Kerteh, Trengganu, Malaysia, 4°43′N 103°43′E, 48 m, 7 Jun 1951: 1art, 16 mm.

Sta 381 IX, Gulf of Thailand, 7°00′N, 103°18′E, 55 m, mud, a little sand and shells, 8 Jun 1951: 1art, 17 mm.

REMARKS.—These three specimens exhibit so much variability in characters that until now were considered to be diagnostic in members of this genus that I am convinced that Clorida verrucosa and Clorida merguiensis are synonyms. The two smaller specimens lack a mandibular palp, but it is clearly present in the larger female, as reported by Makarov (1979) in material from the Bay of Tonkin. The rostral carina is absent in two specimens, present in the male from the Gulf of Thailand. The postanal keel is well developed in the larger specimen, indistinct in the smaller ones. The larger specimen lacks submedian abdominal carinae, but they are present on the fifth and sixth somites in the adult male from sta 381 IX. The submedian carinae are spined only on the sixth somite in all specimens. I can find no characters that can be used to separate C. merguiensis from C. verrucosa.

Clorida verrucosa was taken with C. gaillardi at sta 380.

DISTRIBUTION.—Indo-West Pacific, from Malaya, Indonesia, the Philippines, Vietnam, and the Gulf of Siam in the Pacific, and from the Mergui Archipelago and southern India in the Indian Ocean; sublittoral, in depths between 13–31 and 70–140 meters.

Cloridopsis immaculata (Kemp, 1913)

Squilla Scorpio var. immaculata Kemp, 1913:45, pl. 2: fig. 31.

MATERIAL.—Sta 344, Singapore Island (l°22′N, 103°48′E), traps in prawn ponds in mangrove swamps, tidal zone, mud, 22 May 1951: 3, 82–86 mm.

DISTRIBUTION.—Indo-West Pacific, from Pakistan and India, and now from Singapore; shore.

Lenisquilla lata (Brooks, 1886)

FIGURE 12

Squilla lata Brooks. 1886:34, pl. 3: figs. 1–3.—Kemp, 1913:37, pl. 2: fig. 24.

Squilloides lata.—Manning, 1969a:13, fig. 4.

Squilloides latus spinosus Blumstein, 1970:223, figs. 4, 5 [new synonymy].

Squilloides espinosus Blumstein, 1974:121, fig. 7 [new synonymy].

Squilloides latus.—Blumstein, 1974:123.

Lophosquilla tiwarii.—Moosa and Cleva, 1984b:78 [error for Lenisquilla lata].

Lenisquilla spinosa.—Moosa, 1986:403.

FIGURE 10.—Clorida verrucosa (Hansen, 1926); female, TL 32 mm, sta 294: a, front; b, eye; c, raptorial claw; d, lateral processes of fifth to seventh thoracic somites; e, sixth abdominal somite, telson, and uropod; f, uropod, ventral view.

MATERIAL.—Sta 454, Java Sea, 05°23′S, 116°02′E, 60 m, coral-clay, 25 Aug 1951: 1 damaged, carapace length 6.4 mm.

REMARKS.—In his account of this species, Kemp (1913:38) reported on material from Burma, pointed out differences between that material and Brooks’ original description, and suggested that his Burmese specimens might belong to a distinct species. The differences discussed by Kemp included the apparent lack of a distal spine on the basal segment of the uropods and the presence of more spines on the abdominal carinae in Brooks’ material. Blumstein (1970) considered material from the Gulf of Tonkin to be conspecific with Kemp’s material from Burma and subspecifically distinct from Brooks’ species.

I examined the types of Squilla lata Brooks at The Natural History Museum, London (formerly British Museum (Natural History)), a male TL 74 mm (Figure 12) and a female TL 75 mm. The differences pointed out by Kemp are due to errors in Brooks’ original account. The types have a distinct distal spine on the basal segment of the uropod (not illustrated here), and have the following abdominal carinae armed with spines: submedian 6, intermediate 3–6, lateral 2–6, and marginal 1–5.

I consider Squilloides latus spinosus Blumstein, 1970 to be a synonym of Leniquilla lata (Brooks, 1886). Moosa (1986:404) already pointed out that Squilloides espinosus Blumstein, 1974, is based on a juvenile of Lenisquilla lata.

The synonymy given under the first reference to Lophosquilla tiwarii in Moosa and Cleva (1984b) makes it clear that they were dealing with Lenisquilla lata.

Lenisquilla lata was taken with Acanthosquilla derijardi at Sta 454.

DISTRIBUTION.—Indo-West Pacific, from the Arafura Sea, the Java Sea, Indonesia, Vietnam, Mozambique, and the Red Sea; sublittoral, in 60 to 150 meters.

Lophosquilla tiwarii Blumstein, 1974

Squilla costata.—Kemp, 1913:84, pl. 6: figs. 70–72 [part, specimen from Burma; not Squilla costata De Haan, 1844].

Lophosquilla tiwarii Blumstein, 1974:123, fig. 8.—Moosa, 1986:404.

MATERIAL.—Sta 305, Bay of Bengal, Ceylon-Calcutta, 20°51′N, 87°58′E, 43–52 m, mud, 26 Apr 1951: 1art, 50 mm, 1art, 51.5 mm.

REMARKS.—This species was taken with Harpiosquilla harpax at sta 305.

FIGURE 11.—Clorida verrucosa (Hansen, 1926); male, TL 17 mm, sta 381 IX: a, front; b, eye; c, propodus and dactylus of raptorial claw; d, lateral processes of fifth to seventh thoracic somites; e, sixth abdominal somite, telson, and uropod; f, basal prolongation of uropod, ventral view.

DISTRIBUTION.—Indo-West Pacific, from the Philippines, Indonesia, Vietnam, Burma, and now from the Bay of Bengal between Ceylon and Calcutta; sublittoral, in 36–37 to 182–187 meters.

Oratosquilla nepa (Latreille, 1828)

Squilla nepa Latreille, 1828:471.—Kemp, 1913:60, pl. 4: fig. 49.

Oratosquilla nepa.—Moosa, 1986:410.

MATERIAL.—Sta 341, Singapore (1°20′N, 103°50′E), bought in market, 21 May 1951: 1art, 101 mm.

Sta 373, off Kerteh, Trengganu, Malaysia, 4°30′N, 103°28′E, dip net at lantern light in 10 m, 6/7 Jun 1951: 1 broken specimen.

DISTRIBUTION.—Indo-West Pacific, from the western Indian Ocean to Hong Kong, Malaysia, and Vietnam; shallow sublittoral.

Oratosquilla turbata Manning, 1978

Oratosquilla turbata Manning, 1978d:35, fig. 20.

FIGURE 12.—Lenisquilla lata (Brooks, 1886); male syntype, TL 74 mm, Arafura Sea (specimen in The Natural History Museum, London): a, front; b, eye; c, lateral processes of fifth to seventh thoracic somites; d, telson; e, basal prolongation of uropod.

MATERIAL.—Sta 258, off Mombasa, Kenya, 4°06′S, 39°43′E, 53 m, coralline gravel with mud and shells, 22 Mar 1951: 1art, 29 mm.

Sta 259, off Mombasa, Kenya, 4°06′S, 39°43′E, 40 m, coralline sand and gravel with shells and a little mud, 22 Mar 1951: 2art (1 fragmented), 27 mm.

DISTRIBUTION.—Western Indian Ocean, from Madagascar and now Kenya; sublittoral, in 16 to 55 meters.

Oratosquilla woodmasoni (Kemp, 1911)

Squilla woodmasoni.—Kemp, 1911:99; 1913:74, pl. 5: figs. 63–65.

Oratosquilla woodmasoni.—Manning, 1978d:36, figs. 21, 22.—Moosa, 1986:410.

MATERIAL.—Sta 347, 1 kilometer E of Tanjong Rhu (1°18′N 103°52′E), off Singapore, 3 m, gravel and sand with mud, 23 May 1951: 1art, 80 mm.

DISTRIBUTION.—Widely distributed in the Indo-West Pacific, from the western Indian Ocean to the Philippines; shallow sublittoral.

Rissoides africanus (Manning, 1974)

Meiosquilla africana Manning, 1974a:16, fig. 5; 1977:117, figs. 38, 56.

MATERIAL.—Sta 122, NW of lighthouse, Lobito, Angola, 12°20′S, 13°40′E, 20 m, mud with sand and clay, 20 Dec 1950:1art, 27 mm.

DISTRIBUTION.—West Africa, from Mauritania to Angola; sublittoral, in 15–100 meters.

Squilla biformis Bigelow, 1891

Squilla biformis Bigelow, 1891:94; 1894:532, fig. 20, pl. 21.—Manning, 1974b:108.

MATERIAL.—Sta 741, Gulf of Panama, 07°28′N 79°36′W, 450 m (500 m on label), 15 May 1952: 3, 51–57 mm.

DISTRIBUTION.—Eastern Pacific, from the Gulf of California, Gulf of Panama, and off Peru.

Stomatopoda from Depths Greater than 400 Meters

Records of stomatopods reported from depths exceeding 400 meters that I have been able to find are summarized below. The depth of 400 meters is not considered to be any kind of natural boundary but was chosen to match the depth selected by the editors of the Galathea Reports as the upper limit of the deep-sea fauna (see Wolff, 1964).

Not included here are Chopra’s (1939) records of Gonodactylus chiragra (Fabricius, 1781) and Natosquilla investigatoris (Lloyd, 1907) from 1295 meters and the record given above of Austrosquilla vercoi from 875 meters, as both of these depths are well beyond the known depth ranges of these species. Gonodactylus chiragra and A. vercoi are both shore species and N. investigatoris is a pelagic species (Losse and Merrett, 1971).

The depth given in brackets following the species name is the overall depth range of the species.

BATHYSQUILLIDAE

Altosquilla soelae Bruce, 1985 [396–402 to 456–458 m]

396–402, 404, 416–420, and 456–458 m

Bruce, 1985:469

Bathysquilla crassispinosa (Fukuda, 1909) [170–200 to 350–420 m]

350–420 m

Manning, Kropp, and Dominguez, 1990:312

Bathysquilla microps (Manning, 1961) [604–769 to 1245–1519 m]

720–926 m

Takeda, 1983:171

732 and 915–952 m

Manning, 1961:693; 1969c:96; Bullis and Thompson, 1965:13

732–860 m

Bullis and Thompson, 1965:13; Manning, 1969c:96

604–769, 681–769, 723, 731–786, and 1281 m

Manning and Struhsaker, 1976:443

750–925 and 865–866 m

Moosa, 1986:371, 373

622, about 640, 1042–1070, 1245–1519, and 1235–1272 m

Manning, Kropp, and Dominguez, 1990:314.

Indosquilla manihinei Ingle and Merrett, 1971 [420 to 458–586 m]

420 m

Ingle and Merrett, 1971:193

458–586 m

Manning, Kropp, and Dominguez, 1990:314

EURYSQUILLIDAE

Eurysquilla chacei Manning, 1969 [419 m] 419 m

Manning, 1969c:257

Eurysquilla sewelli (Chopra, 1939) [695 m] 695 m

Chopra, 1939:166

LYSIOSQUILLIDAE

Heterosquilloides insignis (Kemp, 1911) [275 to 415–510 m]

430 m

Kemp, 1913:128; Ghosh and Manning, 1988:655

415–510 m

Moosa, 1986:386

445–460 m

herein

ODONTODACTYLIDAE

Odontodactylus brevirostris (Miers, 1884) [35 to 426–439 m]

426–439 m

Bigelow, 1931:150, as O. hansenii (Pocock, 1893); Manning, 1967c: 23, as 424–73 m, in error

PSEUDOSQUILLIDAE

Parasquilla ferussaci (Roux, 1828) [100 to 505–650 m]

252–550 m

Figueiredo, 1962:5

505–650 m

Arena and Li Greci, 1973:166

SQUILLIDAE

Fennerosquilla heptacantha (Chace, 1939) [105 to 458 m]

395–404 m

Manning, 1969c:171

421 m

Bullis and Thompson, 1965:13; Manning, 1969c: 171

458 m

Bullis and Thompson, 1965:13; Manning, 1969c:173

490–588, 740–785, and 753–804 m

Manning, 1981:298, as Kempina zanzibarica (Chopra, 1939)

Kempina stridulans (Wood–Mason, 1894 in Alcock, 1894) [122 to 439 m]

439 m

Kemp, 1913:80; Ghosh and Manning, 1988:659

Squilla biformis Bigelow, 1891 [156 to 474 m]

450 m

Manning, 1974b:108; herein

Squilla caribaea Manning, 1969 [190 to 366–439 m]

366–439 m

Manning, 1969c:235

Squilla intermedia Bigelow, 1893 [291 to 659–824 m]

366–412, 421, 439, 458, 531, and 659–824 m

Bullis and Thompson, 1965:13; Manning, 1969c:229

410,428–473,464–491, and 546 m

Manning, 1969c:229

458 m

Springer and Bullis, 1956:22; Bullis and Thompson, 1965:13; Manning, 1969c:229

619 m

Bigelow, 1894:530; Manning, 1969c:228

Squilloides leptosquilla (Brooks, 1886) [170–200 to 677–767 m]

344–403 and 677–767 m

Kemp, 1913:48

Squilloides tenuispinis (Wood-Mason, 1891 in Wood-Mason and Alcock, 1891) [165 to 439 m]

439 m

Kemp, 1913:49

DISCUSSION.—Only 17 of about 400 species of Stomatopoda (see Reaka and Manning, 1986) are known to occur in depths exceeding 400 meters, but 13 of the 17 have been taken in depths of less than 400 meters; most of the 17 species are sublittoral species that frequent the outer shelf and upper slope. All four bathysquillids have been taken in depths of more than 400 meters, but even one of these, B. crassispinosa, occurs in shallower water, too.

At the family level, representatives of only 6 of the 13 recognized families have been taken in more than 400 meters, the shore-dwelling gonodactylids, protosquillids, and nannosquillids being conspicuously absent from slope habitats. Other than the bathysquillids, 2 of 25 eurysquillids (8%), 1 of 26 lysiosquillids (4%), 1 of 5 odontodactylids (20%), 1 of 20 pseudosquillids (5%), and 8 of 149 squillids (5%) have been found in more than 400 meters. The low numbers of species living in such deep waters is reflected also at the generic level. Whereas all three bathysquillid genera (100%) are represented in these deep habitats, only 1 of 5 eurysquillid genera (20%), 1 of 5 lysiosquillid genera (20%), 1 of 4 pseudosquillid genera (25%), and 4 of 27 squillid genera (15%) even occur there. As one species of Odontodactylus has been taken below 400 meters, 100% of odontodactylid genera are found there.

There appears to be no clear geographic pattern to the occurrence of stomatopods in outer slope habitats. Only 1 of 30 eastern Atlantic (3%), 5 of 77 western Atlantic (6%), 1 of 50 eastern Pacific (2%), and 11 of about 250 Indo-West Pacific (4%) species occur in such depths, reflecting the shallow-water characteristics of the group.

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