Prog. Oceanog. Vol. 24, pp. 311-316, 1990.
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Biogeography of deep-sea stomatopod Crustacea, family
Bathysquillidae
RAYMONDB. MANNING1, ROY K. KROPP2 and JANEDOMINGUEZ2
1 Department oflnvertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, DC 20560, USA
ZBattelle Ocean Sciences, 1431 Spinnaker Drive, Ventura, CA 93001, USA
Abstract - One family of stomatopod crustaceans, the BathysquiUidae, occurs exclusively on
outer shelf and upper slope habitats, in depths between about 200 and 1500m. Stomatopoda
otherwise are largely restricted to shallow water habitats. The bathysquillids include four species in
three genera. Bathysquilla microps occurs in both Atlantic and Pacific oceans, B. crassispinosa is
found across the Indo-West Pacific from Japan to South Africa, Altosquilla soelae is known only
from the northwestern coast of Australia, and Indosquilla manihinei, originally reported from the
western Indian Ocean, is here reported from the Mariana Islands in the Pacific Ocean. The
bathysquillids, with a fossil species in the Eocene, represent the oldest extant stock of Stornatopoda.
The structure of the uropod in Indosquilla manihinei is intermediatebetween that of the fossil family
Sculdidae and the Recent stomatopods.
1. INTRODUCTION
Several zoogeographers, including BRINGS(1974) and EKMAN(1953) have suggested that the
continental slope now serves as a refugium for ancient relicts, including such organisms as the
fish Latimeria and the mollusk Neopilina. The stomatopod crustaceans, a small group of largely
tropical, shallow-water predators (MANmNG, 1969; ReAKA and MANNING, 1987) includes one
superfamily, the Bathysquilloidea, comprising one family, the Bathysquillidae, the four representatives of which are restricted to deep-sea habitats on the outer shelf and upper slope of the
Atlantic, Pacific, and Indian Oceans, in depths between about 200 and 1500m.
The four Recent superfamilies of Stomatopoda (MANNING, 1980) are characterized by a variety
of morphological characters, including the structure of the raptorial claw, the enlarged second
maxilliped, the form of the body, and the structure of the telson. More recently (MAN--G, SCI-I~F
and ABBOTT, 1984) the structure of the eyes has been shown to be strikingly different in members
of each of the four stomatopod superfamilies, with the eyes of the bathysquillids differing from
those of the other stomatopods in lacking facets and a distinctive middle band of specialized
facets, and in having the eye pigment poorly organized.
The bathysquillids, represented by one fossil species known from the Eocene of England
(QuAYLE,1987), are the only Recent stomatopods to show morphological ties to the fossil family
Sculdidae. Whereas almost all Recent stomatopods have the uropodal exopod divided into two
distinct segments, in the sculdids the uropodal exopod was unlsegmental (HoLTHUISand MArm~G,
311
312
R.B. MArm~Get al.
1969). The uropod of Indosquilla manihinei approaches this condition, for in that species the
uropodal exopod is divided into two segments by an indistinct suture, and the terminal segment
is very small compared with that of other Recent stomatopods.
Here we summarize all literature records for bathysquillids known to us, as well as available
information on distribution, depth, and size for each of the four known species. Additional
records are provided for B. crassispinosa, B. microps and I. manihinei.
Abbreviations used below include: leg. - collector; m - meters; mm - millimeters; and TL total length. Repositories for newly reported specimens are: BPBM - Bernice P. Bishop Museum,
Honolulu; MNHN, Mus6um National d'Histoire Naturelle, Paris; and USNM - National Museum
of Natural History, Smithsonian Institution, Washington.
1.1
Superfamily Bathysquilloidea Manning, 1967
Family Bathysquillidae Manning, 1967
Genus Altosquilla Bruce, 1985
Altosquilla soelae Bruce, 1985
Altosquilla soelae Bruce, 1985: 469, Figs. 1-3, 4A, 5
Previous Records: Indo-West Pacific: Northwest coast of Australia (BRUCE, 1985).
Size: Males (4 known), TL 88-120mm, female (1 known), TL 119mm.
Depth Range: All records are from outer shelf depths, 396-402m, 404m, 420-416m and 456458m.
Distribution: Known only from the northwestern Australian shelf (Fig.l) in depths between
about 400 and 450m.
1.2
Genus Bathysquilla Manning, 1963
Bathysquilla crassispinosa (Fukuda, 1909)
L ysiosquilla crassispinosa Fukuda, 1909:61
Previous Records: Indo-West Pacific: Japan (FuKUDA,1909, 1910, 1913; KOMAI,1927, 1938;
INGLEand MERRETT,1971; MANNINGand STRUHSAKER,1976; TAKEDAand YANAGISAWA,1985).
South China Sea (BRucE, 1985). Philippines (MoOSA, 1985). Madagascar (MANNINGand
STRUHSAKER,1976). Mozambique (MALEand MEmU~TT,1971). South Africa: North of Durban
(CALMAN,1923; KOMAI, 1927; GORDON, 1929; INGLEand MERgETW,1971); Off Durban (VON
BONDE, 1932; BARNARD,1950; INGLEand MERRET'r, 1971).
Additional Records: Madagascar, 22°21.9'S, 43°04.8'E, 350-420m, mud, leg. R. yon Cosel,
21 October 1986:2 males, TL 160 and 230mm (MNHN).
Size: Males (5 known), TL 127-285mm; females (8 known), TL 173-297mm.
Depth Range: Outer shelf and upper slope; this species has been taken in depths between about
200m and about 400m. Depth records include: 170-200m, 194-202m, 208-223m, 230-295m,
242m, 275m, 280-310m, and 350-420m.
Remarks: There appears to be no correlation between size and depth. MOOSA(1985) reported
a male TL 127mm from a depth of 170-200m, a female TL 260mm from a similar depth of 194202m, and off Madagascar two males TL 160mm and 230mm were taken together in 350-420m.
This is the largest species of bathysquillid.
Distribution: Known from widely scattered localities in the Indo-West Pacific, between Japan
and South Africa, including the South China Sea, the Philippines, Madagascar and Mozambique
(Fig. 1), in depths between about 200m and 400m.
8
o
i
~G. 1. Distribution of Recent Stomatopoda.
L~
314
1.3
R.B. MANN~Oetal.
Bathysquilla microps Manning (1961)
Lysiosquilla microps Manning, 1961:693, fig.5, pls. 10-11.
Previous Records: Western Atlantic: Santaren Channel, Bahamas (MANNING,1969; BULLIS
and THOMPSON, 1965). Off Cape Canaveral and SE of Tortugas, Florida (MANNING, 1961, 1969;
BULLISand THOMPSON,1965). Gulf of Mexico: South of Panama City, Florida; Bay of Campeche
(MANNINGand STRUHSAKER,1976). Off Suriname and French Guiana (/VIA_NNINGand STRUHSAKER,
1976; TAKEDA, 1983). Indo-West Pacific: Near Maui, Hawaiian Islands (MANNING and
STRUHSAKER,1976). Philippines (MoosA, 1985).
Additional records: Caribbean Sea: 10°26.4'N, 75°57.9'W to 10°22.9'N, 76°00.7'W, 12451519m, R/V Pillsbury sta. 388, 15 July 1966, 1 claw (USNM); 08°26'N, 54°17'W to 08°41'N,
54°19'W, 1235-1272m, R/V Pillsbury sta. 675, 12 July 1968, 1 male, TL 150mm (USNM);
07°56'N, 54°39'W to 08°08'N, 54°36'W, 1042-1070m, R/V Pillsbury sta. 673, 11 July 1968, 1
female, TL 166mm (USNM).
Hawaiian Islands: north shore of Kaui, 622m, shrimp trap, leg. P. Struhsaker, 31 August 1981,
1 male, TL 175mm (BPBM). French Frigate Shoals, ca. 640m, shrimp trap, leg. P. Struhsaker,
1 December 1981, 1 female, TL 84mm (BPBM).
Mariana Islands: leeward side of Guam between Agana and Agat, ca. 640m, May 1982, 1
female, TL 191 mm (USNM).
Size: Males (19 known), TL 53-255mm; females (10 known), TL 45-220mm.
Depth Range: Upper slope, in depths between 604 and 1519m, with 15 of the 23 records for
depth occurring between 600 and 800m.
Remarks: There appears to be no correlation between size and depth range; the largest and the
smallest specimens were taken in similar depths. This is the deepest living stomatopod.
Distribution: Western Atlantic, from the northern Gulf of Mexico and the Bahamas to
Suriname and French Guiana, and Pacific Ocean, from the Hawaiian Islands, the Mariana Islands,
and the Philippines (Fig. 1), in depths between 600 and about 1500m.
1.4
Genus lndosquilla Ingle and Merrett, 1971
Indosquilla manihinei Ingle and Merrett, 1971
lndosquilla manihinei Ingle and Merrett, 1971: 193, Figs. 1-9, pl. 1.
Previous records: Indo-West Pacific: South of Menai Island, Cosmoledo Atoll, western Indian
Ocean (INGLEand MERRETT,1971).
Additional records: Off Pagan, Mariana Islands, 18°05.2'N, 145°42.8'E, 458-586m, R/V
Townsend Cromwell cruise TC 83-05, sta. 20, 24 November 1983, 1 female, TL 176mm (USNM).
Size: Only known male, TL 160mm; only known female, TL 176mm.
Depth range: Upper slope, 420m and 458-586m.
Remarks: This is the first record for this species since its original description, and extends its
range into the Pacific Ocean.
Distribution: Indo-West Pacific, from the western Indian Ocean and the Mariana Islands in the
Pacific (Fig. 1), in depths between 400 and 600m.
REAKA and MANNING(1987) pointed out that Recent stomatopods exhibit typical Tethyan
distribution patterns, and the bathysquillids also follow this pattem. Like many stomatopods, the
bathysquiilids also show a relict pattern (Fig. 1), with what appear to be isolated populations in
each of the major world oceans.
The distribution pattern, the existence of an Eocene fossil representative of the family, the
Biogeography of deep-sea stomatopod Crustacea
315
degenerate eyes, and the similarity of the uropod structure in Indosquilla and the fossil sculdids
all support the hypothesis that the bathysquillids represent an ancient stock o f stomatopods that
are now restricted to the deep-sea.
INGLe and MERRET'r (1971: 197) suggested that I. manihinei was a neotenous stomatopod,
based on some o f its features that are shared with larvae. It could well be a m e m b e r o f very old
stock that exhibits neotenous features.
Although most bathysquillids have been taken in trawls, the specimens from the H a w a i i a n and
M a r i a n a Islands newly reported here were taken in deep trapping operations. This suggests that
the bathysquillids are active foragers and future trapping operations m a y well prove to be a
valuable source o f new material and new geographic records.
2. ACKNOWLEDGEMENTS
We thank R. Moffitt, National Marine Fisheries Service, for allowing us to examine the material from the
Marianas, R. Titgcn, Bernice P. Bishop Museum, for the loan of specimens from that collection;A. Crosnier, Mus6um
National d'Histoire Naturelle, Paris, for providing working space there; M.L. Realm, University of Maryland, for her
comments on an early draft of our manuscript; Lilly K. Manning for preparing the figure; and the Smithsonian
Institution, for support through its Research Opportunities Fund.
This is contribution number 256 from the University of Guam Marine Laboratory.
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