On the Distribution and Fishery Potential of the Japanese Red Crab Chaceon granulatus in the Palauan Archipelago, Western Caroline Islands

L. C. HASTIE and W. B. SAUNDERS

Introduction terest has grown in the potential for (Sakai,1978; Cayre et al., 1979; King, commercial exploitation of deep-water 1984; Saunders, 1984; Hayasaka, 1985; As the world demand for crustacean geryonid crabs. These animals are Manning and Holthuis, 1989). Exten­ meat of all kinds continues to rise, in- widely distributed in the world's sive exploratory deep-water trapping oceans, commonly occurring on conti­ surveys have been undertaken in this nental slopes at depths of 100-2,000 m region and are well documented L. C. Hastie is with the Department of Zoology, (Manning, 1990). Several members of (Struhsaker and Aasted, 1974; King, University of Aberdeen, Aberdeen AB9 2TN, l the genus Chaceon (formerly Geryon ), 1981, 1986; Gooding, 1984; Moffit and Scotland, and W. B. Saunders is with the De­ partment of Geology, Bryn Mawr College, Bryn typically medium to large crabs with Polovina, 1987). However, these have Mawr, PA 19010. high quality flesh, are considered tended to concentrate on the two marketable. caridean shrimp species, Heterocarpus Presently, there are two major ensifer and H. laevigatus. geryonid fisheries in the Atlantic. Off A recent study of this nature by the northeastern United States is an es­ Saunders et aI., (1989) revealed a popu­ ABSTRACT-A deep-water trapping sur­ tablished fishery for the red crab, lation of the Japanese red crab, vey in the Palauan archipelago, Western Caroline Islands, has revealed an abun­ Chaceon quinquedens, which has op­ Chaceon granulatus, on the deep fore­ dance of the Japanese red crab. Chaceon erated since the early 1970's (Arm­ reef slopes around Palau, Western granulatus. The recorded depth range (250­ strong, 1990). A similar species from Caroline Islands. Preliminary evidence 900 m) is similar to that ofother geryonids, west African waters, C. maritae, has suggests that it may be of sufficient but the large numbers of females caught below 700 m is atypical. Mean yields in also been significantly exploited density to support a local fishery. It is excess of 5 kg crabs plus 1 kg shrimp, (Melville-Smith, 1988). On information possible that similar populations of Heterocarpus laevigatus, by-catch per trap­ from FAO (1986) and Melville-Smith geryonids exist on the slopes of sev­ night were attainable at optimum depths. (1988), it can be estimated that the to­ eral Indo-Pacific archipelagoes. This Chaceon granulatus is apparently a very tal world geryonid catch exceeded paper is largely based on observations large geryonid, with maximum weights of 2.02 kg and 1.51 kg recorded for male and 12,000 t in 1981. Interest in these fish­ of C. granulatus made during red crab female specimens, respectively. A range of eries has resulted in exploratory sur­ explorations in Palau. A brief review body colors was observed: Orange-red veys for several Atlantic species, in­ of the present status, distribution, and shades appear to dominate the deeper wa­ cluding C. afjinis (Shelton, 1979), C. fishery potential of Pacific geryonids ters (below 500 m) while yellow-tan colors !enneri are more abundant in the upper reaches. (Wenner et al., 1987; Kendall, in general is included. Preliminary evidence suggests that 1990; Lindberg et aI., 1990; Wenner, Chaceon granulatus is highly marketable, 1990), C. notialis (Barea and Defeo, Procedures and the infrastructure in Palau is such that 1985; Defeo et aI., 1991), and, in other A total of 103 traps was set between crabs could either be marketed fresh lo­ areas, C. quinquedens (McElman and cally or airfreighted to Japan as a quick­ 170 m and 900 m depth at five stations Elner, 1982) and C. maritae (Cayre et frozen product. The high post-trapping sur­ around Palau (Fig. I), between 1987 and aI., 1979). vival rates observed indicate that 1988. Several trap designs were initially maintaining crabs in live-holding tanks may There is an apparently similar, wide tested (Saunders et al., 1989). Quantita­ be afeasible option. The large catches and distribution of geryonid crabs in the tive samples were obtained by using a quality of deep-water crabs taken suggests tropical and subtropical Pacific that the Palauan population of Chaceon large box trap (2x Ix I m) with 25 cm granulatus may be able to support a small­ diameter funnels at each end. About I scale fishery. It is not yet known whether 'The family geryonidae was revised by Manning kg of bait, usually frozen skipjack tuna, this population is unusually large or and Holthuis (1989) to include three genera: whether these findings typify the deep fore­ Geryon (2 species), Chaceon (21 species), and Katsuwonus pelamis, was wrapped in reeffauna of the region. Zariquieyon (I species). chicken wire and suspended in the cen-

26 Marine Fisheries Review equipped with a gasoline pot hauler. An Echotec CV950 VDU fathometer was available for depth determina­ tion. Catches, stored in a large ice cooler onboard, were later sorted in the laboratory. The crabs were sexed, ..'--. weighed, and measured, and the shrimp by-catch data (species, num­ ber, weight) were also collected. All "­ data were logged into a Macintosh Plus microcomputer using Cricket ~IA \~ Statworks software. ~ \\...\~'1t Results :( {~I Distribution and Catch Rates I; A total of 98 retrieved traps (5 were ./ ~ '''''( lost) yielded 238 red crabs (C. ) r\j \~ .~- granulatus, 268 kg) and 1,550 shrimps .,/,':'--- .ill (15 species, 55 kg). Chaceon was found /~/ ~.•• ·~U over a considerable depth range (250­ West Passage :") ... ~,): !iY 900 m, with corresponding water tem­ peratures of 5-10°C (Fig. 2; Saunders et aI., 1989)); the lower limit of the depth range was not established. Fig­ ure 3 shows the relative numbers of N crabs caught at 50 m intervals. There .is a marked increase below 650 m. The ; ~ ~a ,~ sex ratio recorded overall was 1.329: 1 ~ \:~l~ in favor of males, which were caught ~Iong Figure I.-Map more frequently over most of their "i ;;;t'-.JC::i:J( showing deep­ \,\0J) \;;M~rr':/.\ \'.-~gUlpeIU water trapping range (350-850 m). However, female \-v '"'f'1 (/ \ stations around crabs were more abundant in the deeper ... \, ;.f 0 toIutremdiu Palau, Western sets and tended to dominate catches ~ ~ • .,11 N d' Caroline Islands. from below 700 m. A few females were \~i '7, ' gareme IU @!'k1 Traps were set also caught in the shallowest sets (ca. 103 times at 170-900 m; at 250 m), resulting in a greater depth 1) Mutremdiu range than that recorded for males. All i~~' Bay (38 sets), the berried females were caught 2) Augulpelu between 650 m and 770 m depth Reef (7 sets), 3) Ngaremediu (Table I). Reef (19 sets), In terms of catch per unit effort o 5 10 ml ...... ~,-+-1-~i...... 4) Aulong Bay (CPUE), high yields ( >5 kg per trap­ o 5 10 Icm (7 sets), and night) were attained below 500 m 5) Toagel Mlun­ gui (West Pas­ (Fig. 2). Mean shrimp by-catches of sage) (32 sets). around I kg per trap-night were also obtained between 600 and 700 m. These were dominated by the caridean Heterocarpus laevigatus, which com­ ter of the trap; mesh baffles were in­ usually soaked overnight for 15-25 hours posed 92% (51 kg) of the total shrimp cluded to impede egress (this design is and pulled the following day. yield by weight. A smaller species, referred to as the standard trap hence­ The survey vessel was the MRD H. ensifer, dominated the shallower 2 forth). Polypropylene rope (4-8 mm di­ Mesekiu, a diesel 10m Yanmar , catches of shrimp (170-400 m), but ameter) was used in conjunction with catch rates were generally poor surface buoys and marker flags. Scrap­ (Fig. 2). Large numbers of the deep­ 2Mention of trade names or commercial firms water eel, Synaphobranchus affinis, iron weights and anchors were attached does not imply endorsement by the National Ma­ to reduce trap dragging. Set traps were rine Fisheries Service, OAA. were also taken, below 400 m.

54(1). /992 27 6 --- 30 Observed body colors ranged from \ , light yellow to deep red. Differences between sexes were not detected (Table 25 5 \ 2). However, orange-red shades appear -~ I to dominate the deep waters (below 500 -Cl , m), while yellow-tan colored animals c: , 4 20 () a. are more abundant in the upper reaches (1j , .2­ ~ (Table 3). Most individuals were uni­ -... 3 \ 15 a. formly colored, although a few dark -Cl .., ~ , E red specimens had black patches on the , Q) - '. I- carapace, claws, and legs. Darker crabs 2 '~- 10 "0 ...... -..... also tended to have harder, worn shells Q) ...._­ .... and frequently exhibited heavy barnacle ------.... >­ 5 settlement. Marketability 0 Yields of quality white meat were 0 200 400 600 800 1000 easily extracted from the claws, legs, and bodies of a number of fresh crabs Depth (m) which had been steam-cooked. This Figure 2.-CPUE values obtained for Chaceon granulatus (black squares) be­ was found to be palatable and similar tween 170 and 900 m depth. Shrimp by-catch values for Heterocarpus laevigatus in texture and flavor to that of lobster, (diamonds) and H. ensifer (white squares) are included. Based on standard trap Homarus sp. Some brown meat, data. The broken line represents a water temperature profile for Palau (May­ also found in the bodies, was of poor October 1987), based on average readings from four trap sites (Saunders et al., 1989). quality. A high percentage of crabs appeared to survive trapping operations. A Physical Characteristics (P< 0.0 1) in size and body weight were sample of 43 hauled crabs were exam­ Figure 4 shows the relative wet detectable (Student's t test). The rela­ ined after 30 minutes on ice, and re­ weights of male and nonovigerous fe­ tionships between carapace width and corded as dead or alive, depending on male crabs examined. On average, body weight were also tested, showing any signs of movement in the claws, males are larger and heavier than fe­ significant correlations (P

1 Table 1.-5ummarized size and weight data recorded for Chaceon granulalus specimens.

Carapace width (mm) Weight (kg) o ...j.:.:(2;.:.),,"",.t,...:...:~ Sex No. Mean S.D. Range Mean S.D. Range 200 250 300 350400450 500 550 600650700750 800 850 900 Male 105 1532 12.401 124-179 1.219 0.329 0.58-2.02 Depth (m) Female! 68 146.6 11.472 114-174 0.963 0.224 0.50-151 Figure 3.-Numbers of Chaceon granulatus caught at 50 m Female' 11 147.6 10.782 134-170 depth intervals, based on standard trap data. Male and female 1.061 0.208 073-1.46 crabs are represented by black and hatched columns respec­ lNonovigerous. tively. Effort (trap-nights) is shown in parentheses. 20vigerous.

28 Marine Fisheries Review 30 Table 2.-0bserved body colors (carapace, claws, legs) of 51 Chaceon granu!afus speci­ a mens caught at stations 4 and 5.

Color No. %of Males %01 Females

20 Orange-red 31 60% (24) 64%( 7} Dark red 9 18%(7) 18%(2) C :> Yellow-tan 10 20%(8) 18%(2) 0 Brown 1 2%( 1) 0%(0) '-'

10 similar to those reported for C. quinquedens (Gray, 1970; Meade and Gray, 1973; Haefner and Musick, 1974; O~.--!""'"- McElman and Elner, 1982) and C. 02 0 ~ , 2 1.4 13 2.0 2.2 24 26 Weight (kg) maritae (Cayre et ai, 1979; Melville­ Smith, 1986), as well as other species under economic consideration in cer­ 30.,...------, tain areas (Wenner et aI., 1987; Defeo et aI., 1991). Although the lower depth limit was not established, the observed depth range 20+----­ of Chaceon granulatus (240-900 m) ap­

C pears to be geryonid-typical. Sexual zo­ :> o nation has been frequently reported for '-' geryonids (Haefner and Musick, 1974; 10+--­ Wigley et a!., 1975; Cayre et aI., 1979; Beyers and Wilke, 1980; McElman and Elner, 1982; Barea and Defeo, 1985;

07.-_""1 Melville-Smith, 1988), with most work­ 0.2 0.4 0.6 0.8 1.0 1.4 1.6 18 20 2.2 2.4 213 ers concluding that females are typically Weight (kg) more abundant at shallower depths. Figure 4.-Histograms of wet weights recorded However, our findings suggest that, for a) male and b) nonovigerous female crabs within the population of Chaceon captured during the survey. granulatus around Palau, the reverse may be the case. A few C. fenneri fe­ males trapped by Wenner et aI. (1987) in the South Atlantic Bight were deeper 8 than most of the males, suggesting that 1) Y = - 8.4981 + 3.0956x RA 2 = 0.810 sexual zonation may depend on several 2) Y = - 6.7935 + 2.7360x RA 2 = 0.826 factors, such as species, water tempera­ tures, seasonal effects, and spawning migrations. Overall sex ratios in favor of males have been reported in trapping surveys for C. quinquedens (Gray, 1970; 7 Haefner and Musick, 1974; McElman 01 o and Elner, 1982), C. maritae (Melville­ ...J Smith, 1988), C. fenneri (Wenner et aI., 1987), and C. notialis (Defeo • et aI., 1991). To an extent, trapping may

Table 3.-Percent of observed body colors (pooled sexes) of Chaceon granu!afus recorded at different 6+----~---,....------,r------.-----. depth intervals. 4.7 4.8 4.9 5.0 5.1 5.2 <500 m ------500-700 m >700 m Log carp. width (mm) Color No. % No. % No. % No. Orange-red 31 25% (2) 60% (18) 85% (11) Figure 5.-Bivariate size (carapace width)-weight scatterplots Dark red 9 25% ( 2) 17% (5) 15% (2) for 1) male and 2) nonovigerous female crabs. Corresponding Yellow-tan 10 50% (4) 20% ( 6) 0% (0) Brown regression lines and correlation coefficients are displayed. 1 0% ( 0) 3% ( 1) 0% (0)

54( 1),1992 29 be sex-selective, in that female crabs present infrastructure in Palau is such tation. Collectively, however, this area carrying ripe eggs may not be attracted that crabs could either be marketed must be considerable. to bait; the possible exclusion of larger fresh locally or as a quick-frozen prod­ Acknowledgments males due to funnel diameter restric­ uct airfreighted to Japan (Y. Akiba, tions should also be considered. Nippa Fishing Corporation, Tokyo, per­ This study was supported by a Haefner and Musick (1974) observed sonal commun.). The caridean shrimp Saltonstall-Kennedy Grant to the Pa­ a predominance of female C. quin­ by-catches are also valuable and could cific Fisheries Development Founda­ quedens in trawls at shallower depths, be marketed similarly. These options tion, Honolulu, Hawaii, and the assis­ while Melville-Smith (1988) has re­ could be assessed in an economic fea­ tance of Carol Odo of that organization ported that C. maritae females are sibility study, based on extensive CPUE is sincerely acknowledged. Toshiro sometimes more abundant at certain data. Given the high post-trapping sur­ Paulis, Chief, Marine Resources Divi­ depths and localities. The larger num­ vival rates observed, a further option sion, Palau, was the project coordina­ ber of male crabs caught during this might be to maintain crabs in live-hold­ tor, and Pablo Siangldeb, T. Remoket, survey may have been influenced by ing tanks. M. Kotaro, T. Boisek, and E. Ngotel the trapping effort, which was concen­ The catches and quality of deep-wa­ assisted extensively. G. A. Heslinga, trated between 500 m and 750 m depth ter crabs taken indicate that the Palauan Micronesian Mariculture Demonstra­ (Fig. 3). population of C. granulatus may be tion Center, provided extensive support. According to records (Shroeder, able to support a small-scale fishery. It The National Marine Fisheries 1959; McRae, 1961; Kjennerud, 1967; is not known, presently, whether this Service's Honolulu Laboratory staff, Mason and Davidson, 1969; Haefner population is unusually large or including Robert Moffit, Steve Ralston, and Musick, 1974; Sakai, 1978; Cayre whether these findings typify the deep Daryl Tagami, and Jeffrey Polovina, et aI., 1979; McElman and Elner, 1982; fore-reef fauna of the region. Certainly, were particularly helpful. Raymond B. Barea and Defeo, 1985; Wenner et aI., this species and its close congener, C. Manning, U.S. National Museum of 1987; Manning and Holthuis, 1989), hicolor have been recorded in several Natural History, Washington, D.C., C. granulatus appears to be similar in Pacific localities (Fig. 6; Sakai, 1978; kindly assisted with crustacean identi­ size and weight to C. affinis, C. bi­ Cayre et aI., 1979; King, 1984; fications, and specimens have been de­ color, and c.fenneri, and considerably Hayasaka, 1985; Manning and posited at the U.S. National Museum larger than C. quinquedens, C. maritae, Holthuis, 1989). and at the Bishop Museum, Honolulu. and C. notialis. Larger attainable sizes Geryonid crabs are typically slow Additional details are provided by of male geryonids have previously been growing (Lux et aI., 1982; Van Saunders et aI., 1989, Final Report, reported for all the aforementioned spe­ Heukelem et aI., 1983; Melville-Smith, Project 63A (available from the Pacific cies (Schroeder, 1959; McRae, 1961; 1989), and major recruitment events are Fisheries Development Foundation, Sakai, 1978; Cayre et aI., 1979; thought to occur very rarely (Hines, P.O. Box 2359, Honolulu, HI 96804). Shelton, 1979; McElman and Elner, 1990). Deep-water communities in gen­ 1982; Barea and Defeo, 1985; Wenner eral are particularly vulnerable to over­ Literature Cited et aI., 1987), and so C. granulatus ap­ fishing (Ralston, 1986), and the pro­ Annstrong, D. A. 1990. Commentary on crab pears typical in this respect. tection of these resources by strict management and the east coast United States Presently, there is debate as to management regimes will be crucial, geryonid fisheries. In W. J. Lindberg and E. whether body coloration is useful in particularly in Palau and other similar L. Wenner (Editors), Geryonid crabs and as­ sociated continental slope fauna: A research distinguishing geryonid species (Man­ Indo-Pacific archipelagoes. Although workshop report, p. 23-29. Fla. Sea Grant ColI. ning and Holthuis, 1989; Manning, these deep-water crab stocks are likely Tech. Rep. 58. 1990), or whether it is too variable to be small and restricted in compari­ Barea, L., and O. Defeo. 1985. Primeros ensayos de captura del crustaceo batial Geryon within certain species to be used as a son to those on the continental slopes, quinquedens Smith, en al area comun de pesca taxonomic character (Hines, 1990). Our they should be easier to define and Argentino-Uruguaya. Contrib. Dep. Oceanogr. (F.H.c.) Montevideo 2: 189-203. findings support the latter view, at least hence manage effectively. Biomass es­ Beyers, C. J. de B., and C. G. Wilke. 1980. Quan­ for C. granulatus. Manning (1990) has timates of several geryonid stocks have titative stock survey and some biological and noted that red geryonid species are of­ been calculated by various techniques, morphometric characteristics of the deep-sea red crab Geryon quinquedens off South West ten found in deeper water than tan ones; including trapping (Cayre et aI., 1979; Africa. Fish. Bull. S. Afr. 13:9-19. this tendency has now been recorded Defeo et aI., 1991), trawling (Beyers Cayre, P., P. Le Loeuff, and A. Intes. 1979. intraspecifically for C. granulatus. and Wilke, 1980), effective area fished Geryon quinquedens Ie crab rouge profond. Biologie, peche, conditionnement, potentialites Preliminary evidence suggests that (EFA) (McElman and Elner, 1982; d'explotation. Peche Marit. 1210,8 p. C. granulatus is highly marketable. Melville-Smith, 1986; Defeo et aI., Defeo, 0., V. Little, and L. Barea. 1991. Stock assessment of the deep-sea red crab Chaceon Handling and processing techniques for 1991), and photography (Wigley et aI., notialis in the Argentinian-Uruguayan Com­ geryonids are well documented 1975; Melville-Smith, 1983, 1985). In­ mon Fishing Zone. Fish. Res. 11:25-39. (McRae, 1961; Haefner and Musick, dividual archipelagoes in the Western FAO. 1986. Yearbook of fishery statistics. Catches and landings, 1984. Food Agric. Or­ 1974; Meade and Gray, 1973; Cayre et Pacific possess relatively small areas gan., U.N., Yearb. Fish. Stat. 58,451 p. aI., 1979; Melville-Smith, 1988). The at suitable depths for geryonid exploi­ Gooding, R. M. 1984. Trapping surveys for the

30 Marine Fisheries Review deep-water caridean shrimps Heterocarpus Cent. S. Pac. Occass. Pap. 4, 96 p. King, M. G. 1981. Deep-water shrimp resources laevigatus and H. ensifer, in the northwestern Hines, A. H. 1990. Commentary on life history in Vanuatu; a preliminary survey off Port Villa. Hawaiian islands. Mar. Fish. Rev. 46(2): 18-26. and ecology of deep-sea crabs of the family Mar. Fish. Rev. 43(12):7-10. Gray, G. w., Jr. 1970. Investigation of the basic Geryonidae. In W. J. Lindberg and E. L. 1984. The species and depth distri­ life history of the red crab (Ceryon Wenner (Editors), Geryonid crabs and associ­ bution of deep-water caridean shrimps quinquedens). R.I. Dep. Nat. Resour. Cornplet. ated continental slope fauna: A research work­ (Decapoda, Caridea) near some southwest Pa­ Rep. NOAA-740627 14, 32 p. shop report, p. 30-38. Fla. Sea Grant Coil. cific islands. Crustaceana 47(2): 174-191. Haefner, P. A., Jr., and 1. A. Musick. 1974. Tech. Rep. 58. ---- .1986. The fishery resources of Pa­ Observations on distribution and abundance of Kendall, D. 1990. An assessment of the Georgia cific Island countries, Part I: Deep-water red crabs in Norfolk Canyon and adjacent con­ golden crab fishery. In W. J. Lindberg and E. shrimps. Food Agric. Organ., U.N., Fish. Tech. tinental slope. Mar. Fish. Rev. 36( I):31-34. L. Wenner (Editors), Geryonid crabs and as­ Pap. 27,45 p. Hayasaka, S. 1985. Marine ecological studies on sociated continental slope fauna: A research Kjennerud, J. 1967. A find of Ceryon affinis the habitat of Nautilus pompilius in the envi­ workshop report, p. 18-19. Fla. Sea Grant Coil. Milne-Edwards and Bouvier, 1984 (Crustacea rons of Viti Levu, Fiji. Kagoshima Univ. Res. Tech. Rep. 58. Decapoda) off the coast of Norway. Sarsia 29: 193-198. Lindberg, W. J., F. D. Lockhart, N. J. Blake, R. B. Erdman, H. M. Perry, and R. S. Waller. 1990. Patterns of population structure and abundance for golden and red crabs in the east­ ern Gulf of Mexico. In W. J. Lindberg and E. L. Wenner (Editors), Geryonid crabs and associated continental slope fauna: A research workshop report, p. 8-9. Fla. Sea Grant CoIl. Tech. Rep. 58. a Lux, F. E., A. R. Ganz, and W. F. Rathjen, 1982. Marking studies of the red crab Ceryon a quinquedens Smith off southern New England. a J. Shellfish Res. 2:71-80. Manning, R. B. 1990. Studies on systematics of geryonid crabs. In W. J. Lindberg and E. L. Wenner (Editors), Geryon id crabs and associ­ ated continental slope fauna: A research work­ shop report, p. 1-2. Fla. Sea Grant Coil. Tech. Rep. 58. ---- and L. B. Holthuis. 1989. Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae). Proc. BioI. Soc. Wash. 102(1):50-77. Mason, J., and C. Davidson. 1969. Ceryon affinis A. Milne-Edwards and Bouvier, 1894, in Eu­ ropean waters (Decapoda, Brachyura). Crustaceana 16(2):208-210. McElrnan, J. F., and R. W. Elner. 1982. Red crab (Ceryon quinquedens) trap survey along the edge of the Scotian Shelf, September 1980. Can. Tech. Rep. Fish. Aquat. Sci. 1084, 12 p. McRae, E. D., Jr. 1961. Red crab explorations off the northeastern coast of the United States.

~ Commer. Fish. Rev. 23(5):5-10. .'-~ Meade, T./L., and G. W. Gray, Jr. 1973. The red ., .:".. •. ... crab. Univ. R.I. Mar. Tech. Rep. 11,21 p. Melville-Smith, R. 1983. Abundance of deep sea crab Ceryon maritae in South West African 'I,. .. waters from photography. S. Afr. J. Mar. Sci. 1:123-131. . a " ----. 1985. Density distribution by depth of Ceryon maritae on the northern crab rl.,;.' ... grounds of South West AfricalNamibia deter­ mined by photography in 1983, with notes on the portunid crab Bathynectes piperitus. S. Afr. J. Mar. Sci. 3:55-62. aa ----. 1986. Red crab Ceryon maritae den­ sity in 1985 by technique of effective area fished per trap on the northern fishing grounds off South West Africa. S. Afr. J. Mar. Sci. 4:257-263. ----. 1988. The commercial fishery for and population dynamics of red crab Ceryon '~ maritae off South West Africa. 1976-1986. S. Afr. J. Mar. Sci. 6:79-95. ----. 1989. A growth model for the deep­ sea red crab (Ceryon maritae) of South West Africa/Namibia (Decapoda, Brachyura). Crustaceana 56:279-292. Figure 6.-Distribution of Chaceon granulatus (white squares) and C. hicolor Moffitt, R. B., and J. J. Polovina. 1987. Distri­ (black squares) in the Western Pacific Ocean, based on the records of Sakai bution and yield of the deep-water shrimp (l978), Cayre et al. (l979), Hayasaka (1985), Manning and Holthuis (l989), Heterocarpus resource in the Marianas. Fish. and Saunders et al. (l989). Bull. 85:339-349.

54( IJ, 1992 31 Ralston, S. 1986. An intensive fishing experi­ Schroeder, W. C. 1959. The lobster, Homarus Wenner, E. L. 1990. Distribution and abundance ment for the caridean shrimp, Heterocarpus americanus, and the red crab, Geryon of golden crab, Chaceon jenneri, in the South laevigatus, at Alamagan Island in the Mariana quinquedens, in the offshore waters of the west­ Atlantic Bight. In W J. Lindberg and E. L. Archipelago. Fish. Bull. 84:927-934. ern North Atlantic. Deep-Sea Res. 5:266-282. Wenner (Editors), Geryonid crabs and associ­ Sakai, T. 1978. Decapod crustacea from the Em­ Shelton, R. G. W. 1979. Deep water crabs at ated continental slope fauna: A research work­ peror Seamount chain. Res. Crust. 8(Suppl.): 1­ Rockall. Scott. Fish. Bull. 45, 45 p. shop report, p. 6-7. Fla. Sea Grant ColI. Tech. 39. Struhsaker, P., and D. C. Aasted. 1974. Deep­ Rep. 58. Saunders, W. B. 1984. The role and status of water shrimp trapping in the Hawaiian Islands. ----, G. F. Ulrich, and J. B. Wise. 1987. Nautilus in its natural habitat: Evidence from Mar. Fish. Rev. 36( I0):24-30. Exploration for golden crab, Geryon jenneri deep-water camera photosequences. Paleo­ Tomiyama, T. 1983. Fisheries in Japan: Crab. Jpn. in the South Atlantic Bight: Distribution, popu­ biology 10:469-486. Mar. Prod. Photo Mater. Assoc., Tokyo, 178 p. lation structure, and gear assessment. Fish. ---- , L. C. Hastie, and T. Paulis. 1989. Van Heukelem, W, M. C. Christman, C. E. Bull. 85:547-560. Deep-water shrimp survey and feasibility Epifanio, and S. D. Sulkin. 1983. Growth of Wigley, R. L., R.B. Theroux, and H. E. Murray. study, Republic of Palau, Western Caroline Is­ Geryon quinquedens (Brachyura: Geryonidae) 1975. Deep-sea red crab (Geryon quinquedens) lands. Pac. Fish. Develop. Found., Honolulu. juveniles in the laboratory. Fish. Bull. 81 :903­ survey off the northeastern United States. Mar. Final Rep., Project 63A, 120 p. 905. Fish. Rev. 37(8): 1-21.

32 Marine Fisheries Review

-(: U.S. GOVERNMENT PRINTING OFFICE: 1992-790-090/60002 REGION NO.8