Distribution and Ecology of Deep-Water Caridean Shrimps (Crustacea; Natantia) Near Tropical Pacific Islands
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BULLETIN OF MARINE SCIENCE, 41(2): 192-203, 1987 DISTRIBUTION AND ECOLOGY OF DEEP- WATER CARIDEAN SHRIMPS (CRUSTACEA; NATANTIA) NEAR TROPICAL PACIFIC ISLANDS Michael King ABSTRACT Deep-water caridean shrimps have been caught in baited traps set as deep as 800 m off many tropical Pacific Islands. Studies on the biology and distribution of several species (most belonging to the genus Heterocarpus) have been of both commercial and academic interest. The widespread distribution and abundance of these shrimps has encouraged some com- mercial attempts at exploitation, but the economics of the fishing operations have been marginal. Some of the studies on the ecology of deep-water shrimps have produced findings which run counter to competition-based ecological theory. Adult predation may decrease with increasing depth, allowing deeper-water species to have an extended lifespan, an in- creased degree of iteroparity, and a corresponding increase in lifetime reproductive effort. Furthermore, the probabilities of larval survival, which appear to decrease with increasing depth, may be offset by the production of larger eggs by deeper-water species. Small numbers of deep-water caridean shrimps were found during trawling surveys in Hawaiian waters during 1967 and 1968 (Struhsakerand Yoshida, 1975). These species were later found to be susceptible to trapping (Clarke, 1972; Struh- saker and Aasted, 1974). Deep-water shrimps were subsequently found in the northern Pacific islands of Guam (Wilder, 1977) and the Northern Marianas (Moffitt, 1983). The most extensive surveys have been carried out in Hawaii and the Marianas where traps have been set in many different locations (Oishi, 1983; Ralston, in prep. I; Dailey and Ralston, in prep.2; Moffitt and Polovina, in prep.3). South of the equator, deep-water shrimps have been found in Fiji (Brown and King, 1979; King, 1984) and New Caledonia (Intes, 1978). The Fijian surveys are notable in that they were carried out over an extended time period in one location, beyond the barrier coral reef near Suva. The research vessel TAINUI obtained small numbers of deep-water shrimps in Tahiti in 1978 (P. Hatt, pers. comm. 1980). Catches of shrimps were made during a preliminary deep-water trapping survey in Vanuatu in 1980 (King, 1981 a); further surveys were conducted during 1981 and 1982 (De Reviers et al., 1982). Specimens of caridean shrimps were taken in Western Samoan waters by the French research vessel CORIOLIS(Anon., 1977) and during a fisheries resource survey in 1980 (King, 1980; 1984). In 1981, a preliminary survey was completed in Tonga (King, 1981b; 1984). During a demonstration in the use of traps in a consultancy in Papua New Guinea, small numbers of caridean shrimps were caught (King, 1982). At least 17 species of deep-water caridean shrimps have been found in Pacific islands. Some larger species, particularly those of the pandalid genus Heterocarpus, have elicited some commercial interest (King, 1981c; 1986). Some aspects of deep-water shrimp biology have also been of theoretical interest in the study of I S. Ralston. An intensive fishing experiment for the caridean shrimp Helerocarpus fael'jgalus at Alamagan Island in the Mariana Archipelago. Pacific Scicncc. 2 M. D. Dailey and S. Ralston. Aspects of the reproductive biology, spatial distribution, growth and monality of the deepwater caridean shrimp Helerocarpus lae\'igatus in HawaiI. 'R. B. Moffitt and J. J. Polovina. Distribution and yield of the deepwater shrimp resource in the Marianas. 192 KING: DEEP. WATER SHRIMPS FROM PACIFIC ISLANDS 193 the evolution of life-history patterns at different depths (King and Butler, 1985). This paper reviews what is known of the biology and ecology of deep-water caridean shrimps in Pacific islands. METHODS Most surveys carried out from Pacific islands were similar in that baited traps were set, either singly or in a string, along transects running from depths of about 250 m to about 800 m and left in the sea overnight. A typical trap arrangement is shown in Figure I. Traps of several different types have been tested and used in Pacific islands; a typical trap type is shown in Figure I. Most traps were made from steel rod frames and covered with galvanized wire or plastic mesh and operate on the principle that shrimps, attracted to the bait hanging in the traps, enter cone-shaped entrances worked into the trap sides or top. Full descriptions of gear and methods used are given in King (1986). RESULTS Species Found.-At least 17 species of caridean shrimps have been found during trapping surveys in Pacific islands. A list of all species found in Fiji, Vanuatu, Western Samoa, Tonga, the Marianas and Hawaii is given in Table 1; surveys in Vanuatu, Western Samoa and Tonga were preliminary. Descriptions and a key to the species found in southwestern Pacific islands are given in King (1984). Most published work refers to the distribution and biology of the Heterocarpus species which, because of their large size, are generally regarded as having the best commercial potential. Of the Heterocarpus species, H. ensifer has been found in only small quantities in Fiji, Vanuatu, Tonga (King, 1984) and New Caledonia (Intes, 1978) but in larger quantities in northern Pacific islands such as Hawaii (Struhsaker and Aasted, 1974), Guam (Wilder, 1977), and the Marianas. Individuals of this species caught in the southern Pacific Ocean appear smaller than those caught in the northern Pacific. H. sibogae, on the other hand, is more commonly found in the southern Pacific. H. gibbosus, which forms a large part of the catch in Fiji, has only occasionally been found in other southern Pacific islands and rarely in the northern Pacific. H. laevigatus is widely distributed in Pacific islands, and because of its large size and abundance is of greatest interest in fisheries development. Several other organisms have been commonly caught in traps set for deep-water shrimps. The more notable of these include snapper (Lutjanidae), particularly Ete/is and Pristipomoides species (in depths less than 300 m) and several species of eels at all depths. In Fiji, Nautilus pompilius has been taken (up to 12individuals per trap) in depths of less than 380 m (King, 1984). Another nautilus, N. mac- romphalus which is unique to New Caledonia, has been caught in deep-water traps (Intes, 1978). The deep-sea red crab, Geryon has been caught in depths greater than 600 m in both Fiji and New Caledonia. Amphipods and isopods, including a giant Bathynomus species, sometimes enter traps in large numbers and devour any bait not protected by plastic or small mesh containers. Distribution by Depth. - The distribution of deep-water shrimps is highly related to depth, with each particular species occupying different but overlapping depth ranges. As traps are set at increasing depths and distances from the shore or reef, catches of each species reach a maximum before decreasing and being replaced by another, and often larger, species in deeper water. In general, shallow water catches (from less than 400 m) consist of small shrimps such as Parapandalus serratifrons and Plesionika longirostris. Medium-sized Het- 194 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.2, 1987 8 mm ROPE Figure 1. The general arrangement of a trap fishing rig with a typical four-entrance box trap shown in the inset. The length of line between the buoys and the anchor is equivalent to the depth of water plus an excess of at least 25%. erocarpus species predominate in catches deeper than 400 m. One of the largest species found, Heterocarpus laevigatus, is common in depths of more than 500 m. The general depth distribution of the eight more common species of deep- water shrimps is shown in Figure 2. It should be noted, however, that the depth of maximum abundance of each species may vary between islands; the reported depth distributions of one of the most common species, Heterocarpus laevigatus, are compared in Figure 3. It is difficult to compare the relative catch rates of deep-water shrimps in different islands due to the effects of using different traps, baits and soak times in the various surveys. Mean catch rates at optimum depth ranges, in areas where surveys have been conducted using small traps (volume between 0.2 and 0.3 m3), vary from about 1 to 3 kg per trap per night. Larger traps, more recently preferred by commercial fishermen in Hawaii, are reported to catch at least five times more shrimps than small traps (Methot, 1984). The Fijian surveys have provided some information on the possible seasonality KING: DEEP· WATER SHRIMPS FROM PACIFIC ISLANDS 195 Table I. Caridean shrimps found in deep-watertrappingsurveys in Fiji (Fij), Vanuatu (Van), Western Samoa (Sam), Tonga (Ton), Marianas (Mar) and Hawaii (Haw). The trap abundance of each species is indicated as C (common), 0 (occasional), R (rare) or no letter (not recorded). From King (1984), Anon. (1984), and R. Moffitt (pers. comm. 1986) Species Fij Van Sam Ton Mar Haw Parapandalus (=Plesionika*) serratlfrons C C C C Plesionika longirostris (=edwardsii*) C C C C C 0 Plesionika rostricrescentis R R Plesionika ensis 0 0 0 0 R R Plesionika ocellus R Plesionika sindoi (=P. ocellus*) R Plesionika martia 0 R 0 Periclimenes sp. R Eugonatus crassus R Heterocarpus ensifer 0 0 0 C C Heterocarpus sibogae C C C C R Heterocarpus gibbosus C 0 R R 0 Heterocarpus laevigatus C C C C C C Heterocarpus dorsalis 0 C 0 Heterocarpus lepidus 0 Heterocarpus longirostris C Heterocarpus tricarinatus R *' Revised names (Chace, 1985). in catches of deep-water shrimps. Figure 4 shows the mean catch rates of three species, Heterocarpus sibogae, H. gibbosus and H. laevigatus, by depth during trapping surveys down the outer reef slope near Suva in Fiji (King, 1984).