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Biogeography of deep-sea stomatopod Crustacea, family Bathysquillidae
Prog. Oceanog. Vol. 24, pp. 311-316, 1990.
0079 - 6611/90 $0.00 + .50 Pergamon Press pie
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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.
~G. 1. Distribution of Recent Stomatopoda. L~
i
o
8
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.
3. REFERENCES BAP~ARD,K.H. (1950) Descriptive list of South African stomatopod Cmstacea (mantis shrimps). Annals of the South African Museum, 38, 838-864. Bmc_~s, J.C. (1974) Marine zoogeography. McGraw-Hill, New York, 475pp. BRuce, A.J. (1985) Altosquilla soelae, new genus, new species, a bathysquiUid stomatopod from the Australian Northwest shelf. Journal of Crustacean Biology, 5, 468-475. BULLIS,H.R., JR. and J.R. THOMPSON(1965) Collections by the exploratory fishing vessels Oregon, Silver Bay, Combat and Pelican made during 1956 to 1960 in the southwestern North Atlantic. U.S. Fish and Wildlife Service, Special Scientific Report, Fisheries, 510, 130pp. CALMAN,W.T. (1923) Preliminary report on Cmstacea procured by the S.S. Pickle. VI. South African Fisheries and Marine Biological Survey, Research Report, 3, 1. EKMAN,S. (1953) Zoogeography of the Sea. Sidgwick & Jackson, London, 417pp. FUKUDA,T. (1909) The Stomatopoda of Japan. Dobutsugaku Zasshi, 21, 54-62, PI.4. (In Japanese) FtmOOA, T. (1910) Report on Japanese Stomatopoda, with descriptions of two new species. Annotationes Zoologicae Japonenses, 7, 139-152, P1.4. FUKUDA,T. (1913) On two species of Japanese stomatopods, with list of Stomatopoda found in Japanese seas. Dobutsugaku Zasshi, 25, 69-72. GORDON,I. (1929) Two rare stomatopods of the genus Lysiosquilla Dana. Annals and Magazine ofNaturalHistory, ser.10, 4, 460-462. HOLTHOlS,L. B. and R.B. ~ o (1969) Stomatopoda. In: Treatise on Invertebrate Paleontology, Part R, Arthropoda 4, R.C. MOOR~,editor, 535-552. INOLE,R.W. and N.R. MERReTr(1971) A stomatopod crustacean from the Indian Ocean, Indosquilla manihinei gen. et sp. nov. (Family Bathysquillidae) with remarks on Bathysquilla crassispinosa (FOKUDA,1910). Crustaceana, 20, 192-198, Pl.1. KOMALT. (1927) Stomatopoda of Japan and adjacent localities. Memoirs of the College of Science, Kyoto Imperial University, set.B, 3, 307-354, Pls.13-14. KOMAI, T. (1938) Stomatopoda occurring in the vicinity of the Kii Peninsula. Annotationes Zoologicae Japonenses, 17, 264-275. MAro¢~o,R.B. ( 1961) A new deep-water species of Lysiosquilla (Crustacea, Stomatopoda) from the Gulf of Mexico. Annals and Magazine of Natural History, ser.13, 3, 693-697, PIs. 10-11.
316
R.B. ~ o
et al.
R.B. (1963) Preliminary revision of the genera Pseudosquilla and LysosquiUa with descn'ptions of six new genera. Bulletin of Marine Science of the Gulf and Caribbean, 13, 308-328. ~ G , R.B. (1967) Preliminary account of a new genus and a new family of Stomatopoda. Crustaceana, 13, 238-239. MAtiNG, R.B. (1969) Stomatopod Crustacea of the western Atlantic. Studies in Tropical Oceanography, Miami, 8, 380pp. ~ o , R.B. (1980) The superfamilies, families, and genera of Recent stomatopod Crustacea, with diagnoses of six new families. Proceedings of the Biological Society of Washington, 93, 362-372. Ma_m~G, R.B., H. Scrn~ and B.C. ABBOa'r(1984) Eye structure and the classification of stomatopod Crustacea. Zoologica Scripta, 13, 41-44. MArnC~G,R.B. and P. S a ~ u H s ~ (1976) Occurrence of the Caribbean stomatopod, Bathysquilla microps, off Hawaii, with additional records for B. microps and B. crassispinosa. Proceedings of the Biological Society of Washington, 89, 439-449. MoosA, M.K. (1985) Stomatopod Crustacea. 11. R~sultats des Campagnes MUSORSTOM I and 11, Philippines, volume 2. Mdmoires du Musdum national d'Histoire naturelle, Paris, ser.A, Zoologie, 133, 367-414, PI.1. QUAYt~,W.J. (1987) English Eocene Crustacea (lobsters and stomatopod). Paleontology, 30, 581-612. ~ , M.L. and R.B. ~ o (1987) The significance of body size, dispersal potential, and habitat for rates of morphological evolution in stomatopod Crustacea. Smithsonian Contributions to Zoology, 448, 1-46. T~,.EOA,M. (1983) Crustaceans. In: Crustaceans and mollusks trawled of Suriname and French Guiana, Japan Marine Fishery Resource Research Centre, Tokyo, 19-185. T~W.OA,M. and F. YANAOtSAWA(1985) Rediscovery of two Carcinoplax crabs in the Kumanonada Sea, east of the Kii Peninsula, Central Japan, with notes on distribution of the related species. Bulletin of the Biogegraphical Society of Japan, 40, 57-62. VoN Boror~, C. (1932) Report No.9 for the year ending December 1931. Report of the Fisheries and Marine Biological Survey, Union of South Africa, 9, 1-128. ~ G ,
0079 - 6611/90 $0.00 + .50 Pergamon Press pie
Printed in Great Britain.
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.
~G. 1. Distribution of Recent Stomatopoda. L~
i
o
8
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.
3. REFERENCES BAP~ARD,K.H. (1950) Descriptive list of South African stomatopod Cmstacea (mantis shrimps). Annals of the South African Museum, 38, 838-864. Bmc_~s, J.C. (1974) Marine zoogeography. McGraw-Hill, New York, 475pp. BRuce, A.J. (1985) Altosquilla soelae, new genus, new species, a bathysquiUid stomatopod from the Australian Northwest shelf. Journal of Crustacean Biology, 5, 468-475. BULLIS,H.R., JR. and J.R. THOMPSON(1965) Collections by the exploratory fishing vessels Oregon, Silver Bay, Combat and Pelican made during 1956 to 1960 in the southwestern North Atlantic. U.S. Fish and Wildlife Service, Special Scientific Report, Fisheries, 510, 130pp. CALMAN,W.T. (1923) Preliminary report on Cmstacea procured by the S.S. Pickle. VI. South African Fisheries and Marine Biological Survey, Research Report, 3, 1. EKMAN,S. (1953) Zoogeography of the Sea. Sidgwick & Jackson, London, 417pp. FUKUDA,T. (1909) The Stomatopoda of Japan. Dobutsugaku Zasshi, 21, 54-62, PI.4. (In Japanese) FtmOOA, T. (1910) Report on Japanese Stomatopoda, with descriptions of two new species. Annotationes Zoologicae Japonenses, 7, 139-152, P1.4. FUKUDA,T. (1913) On two species of Japanese stomatopods, with list of Stomatopoda found in Japanese seas. Dobutsugaku Zasshi, 25, 69-72. GORDON,I. (1929) Two rare stomatopods of the genus Lysiosquilla Dana. Annals and Magazine ofNaturalHistory, ser.10, 4, 460-462. HOLTHOlS,L. B. and R.B. ~ o (1969) Stomatopoda. In: Treatise on Invertebrate Paleontology, Part R, Arthropoda 4, R.C. MOOR~,editor, 535-552. INOLE,R.W. and N.R. MERReTr(1971) A stomatopod crustacean from the Indian Ocean, Indosquilla manihinei gen. et sp. nov. (Family Bathysquillidae) with remarks on Bathysquilla crassispinosa (FOKUDA,1910). Crustaceana, 20, 192-198, Pl.1. KOMALT. (1927) Stomatopoda of Japan and adjacent localities. Memoirs of the College of Science, Kyoto Imperial University, set.B, 3, 307-354, Pls.13-14. KOMAI, T. (1938) Stomatopoda occurring in the vicinity of the Kii Peninsula. Annotationes Zoologicae Japonenses, 17, 264-275. MAro¢~o,R.B. ( 1961) A new deep-water species of Lysiosquilla (Crustacea, Stomatopoda) from the Gulf of Mexico. Annals and Magazine of Natural History, ser.13, 3, 693-697, PIs. 10-11.
316
R.B. ~ o
et al.
R.B. (1963) Preliminary revision of the genera Pseudosquilla and LysosquiUa with descn'ptions of six new genera. Bulletin of Marine Science of the Gulf and Caribbean, 13, 308-328. ~ G , R.B. (1967) Preliminary account of a new genus and a new family of Stomatopoda. Crustaceana, 13, 238-239. MAtiNG, R.B. (1969) Stomatopod Crustacea of the western Atlantic. Studies in Tropical Oceanography, Miami, 8, 380pp. ~ o , R.B. (1980) The superfamilies, families, and genera of Recent stomatopod Crustacea, with diagnoses of six new families. Proceedings of the Biological Society of Washington, 93, 362-372. Ma_m~G, R.B., H. Scrn~ and B.C. ABBOa'r(1984) Eye structure and the classification of stomatopod Crustacea. Zoologica Scripta, 13, 41-44. MArnC~G,R.B. and P. S a ~ u H s ~ (1976) Occurrence of the Caribbean stomatopod, Bathysquilla microps, off Hawaii, with additional records for B. microps and B. crassispinosa. Proceedings of the Biological Society of Washington, 89, 439-449. MoosA, M.K. (1985) Stomatopod Crustacea. 11. R~sultats des Campagnes MUSORSTOM I and 11, Philippines, volume 2. Mdmoires du Musdum national d'Histoire naturelle, Paris, ser.A, Zoologie, 133, 367-414, PI.1. QUAYt~,W.J. (1987) English Eocene Crustacea (lobsters and stomatopod). Paleontology, 30, 581-612. ~ , M.L. and R.B. ~ o (1987) The significance of body size, dispersal potential, and habitat for rates of morphological evolution in stomatopod Crustacea. Smithsonian Contributions to Zoology, 448, 1-46. T~,.EOA,M. (1983) Crustaceans. In: Crustaceans and mollusks trawled of Suriname and French Guiana, Japan Marine Fishery Resource Research Centre, Tokyo, 19-185. T~W.OA,M. and F. YANAOtSAWA(1985) Rediscovery of two Carcinoplax crabs in the Kumanonada Sea, east of the Kii Peninsula, Central Japan, with notes on distribution of the related species. Bulletin of the Biogegraphical Society of Japan, 40, 57-62. VoN Boror~, C. (1932) Report No.9 for the year ending December 1931. Report of the Fisheries and Marine Biological Survey, Union of South Africa, 9, 1-128. ~ G ,