Pacific Science (1989), vol. 43, no. 4 © 1989 by University of Press. All rights reserved

Phyllosoma Larvae and the Ocean Currents off the Hawaiian Islands!

B. F. PHILLIPs2 AND P. S. MCWILLIAM3

ABSTRACT: A total of 73 phyllosoma larvae of slipper or and one nisto stage were collected about 20 km offthe southwest coast of Oahu between August 1977 and October 1978. Larvae of regalis, aurora, squammosus, marginatus, and Panulirus penicillatus, all found as adults in the Hawaiian Islands, were present in the samples. In addition, larvae of Scyllarus demani and another species tentatively identified as Panulirus longipes femoristriga, not recorded as adults in the Ha­ waiian Islands, were present. The unidentified nisto, possibly of the Hawaiian , Scyllarus aurora, is described and illustrated.

CLARKE (1983) EXAMINED the bias due to avoid­ about 2.5 mm in diameter, and the tail section ance and mesh escapement of three sampling was of 505-jlm nitex plankton netting. TSK devices for the larvae, juveniles, and adults of flowmeters were mounted inside the frames of several species of midwater fishes. One of the the bongo nets and, for purposes ofcalculating three devices was a bongo net 1.25 m in diame­ volume filtered, were assumed to have mea­ ter. In addition to fishes, other were sured the actual flow through the nets. captured during the sampling program and in The nets were towed obliquely and without this paper the phyllosoma larvae of the Scyl­ opening-closing devices. Time-depth recorders laridae and Palinuridae, the slipper and spiny were attached. The bongo nets were weighted (rock) lobsters, are reported. Larvae ofthe im­ with about 200 kg of weight. They were portant commercial spiny and slipper lobsters launched and towed at ca. 1.75 m/sec for are known to be useful as indicators of the 90 min while cable was paid out at 8 m/min. origins of the water masses in which they are The ship was then slowed to ca. 1 m/sec and located (Murano 1957, Johnson and Brinton the nets retrieved at ca. 40 m/min. Descent 1963, McWilliam and Phillips 1983), and the times and wire out were carefully monitored; implications of the results are discussed in deviations from the appropriate schedule were relation to the oceanography ofthe Hawaiian rarely more than ± 10 m or ± I min. Total Islands. retrieval times varied by about 5 min between tows. All samples were taken about 20 km off the southwest coast of Oahu, Hawaii (21 °20• MATERIALS AND METHODS 30' N, 158°20-30' W), from the R. V. Kana The 1.25-m nets for the bongo nets were Keoki of the University ofHawaii. The water conical and about 7 m long. The first ca. 5 m depth at that location was 2000-4000 m. There was of knotless nylon mesh with openings were 14 cruises between August 1977 and October 1978 (Clarke 1983). Twenty-eight paired samples were collected from a maximum depth of 425 m (mean, 304

1 Manuscript accepted 30 September 1988. m; range, 175-425 m), and the volume filtered 3 2 Division of Fisheries, CSIRO Marine Laboratories, per haul was a mean of9434 m (range, 7350­ 3 P.O. Box 20, North Beach, W.A. 6020, Australia. 13,200 m ). 3 Division of Fisheries, CSIRO Marine Laboratories, Catches were initially preserved in 4% for­ G.P.O. Box 1538, Hobart, Tas. 7001, Australia. Present address: School of Biological Sciences, University of maldehyde and held in the same solution until Sydney, Sydney, N.S.W. 2006, Australia. sorting and measurement. The samples were 352 Phyllosoma Larvae and Ocean Currents-PHILLIPS AND MCWILLIAM 353 sorted at 10 x magnification under a dissecting Identification and Morphology ofthe microscope. Scyllaridae No difficulty was encountered in assigning the phyllosoma specimens to described species but the single nisto specimen could not be RESULTS identified to species. A total of73 phyllosoma larvae, comprising 11 scyllarids and 62 palinurids, and 1 nisto Description ofNisto of Scyllarus sp. was caught. The phyllosomata were identified using descriptions of the larvae given in the Figure I literature, including those of Johnson (1968, Total length 20.5 nun; carapace length (mea­ I97Ia,b), and by comparison with material in sured midline from base of rostral tooth) 6.5 a reference collection held by one of the au­ mm; carapace width 8.8 mm. thors (RF.P.). However, some palinurid speci­ A sharp rostral tooth is present. Anterior mens could not be assigned to species known and posterior branchial carinae are well de­ from Hawaii. veloped. The median carina is broad and dis­ continuous; the gastric section terminating Species ofthe Area anteriorly in a strong, sharp tooth; the cardiac section blunt anteriorly. The carapace bears a The following species of the Scyllaridae, distinctly arched posterior transverse groove, Synaxidae, and Palinuridae have been re­ and its posterior margin is incised medially corded as being found as adults in the Ha­ with a small, rounded notch. The lateral mar­ waiian area. gin of carapace is divided into three distinct regions, the anterior one terminating in a SCYLLARIDAE: strong, sharp anterolateral tooth followed by Scyllarides squamosus (H. Milne Edwards) five or six small, blunt teeth. The mediolateral (Holthuis 1947) and posterolateral regions of the margin bear (de Haan) (Morin and a sharp anterolateral tooth followed by four MacDonald 1984) to five and seven to eight small, blunt teeth, Arctides regalis Holthuis (Holthuis 1963) respectively. Scyllarus modestus Holthuis (Holthuis 1960) The distal squama (= sixth segment) ofthe rformerly identified as Scyllarus martensii antenna is incised with four broad and one Pfeffer by Rathbun (1906) (see Holthuis narrow lobes, each ending in an acutely angled 1960)] tip; a smaller sharp sixth lobe is present on the Scyllarus aurora (Holthuis) (Holthuis 1981) inner margin of this segment. The proximal [formerly Scyllarus timidus (see Holthuis squama (= fourth segment) of the antenna 1981)] bears a distinct spinose, dorsal ridge, also two antarcticus (Lund) (Holthuis prominent teeth on its anterior margin; its 1985) anterolateral corner is produced into a sharp SYNAXIDAE: tooth, and the lateral margin has two prom­ Palinurellus wieneckii (de Man) (Titgen and inent teeth followed by five to six small teeth Fielding 1986) posteriorly. The posterior margin ofabdominal terga of PALINURIDAE: somites I-III is incised with a small median Justitia mauritiana (Miers) (George and notch and rounded lateral notches. Abdominal Main 1967) terga of somites IV and V bear only rounded Panulirus marginatus (Quoy and Gaimard) lateral notches. None ofthe abdominal somites (George and Holthuis 1965) has a distinct median carina, but the median Panulirus penicillatus (Olivier) (Edmondson area of somites II and III is more elevated 1925) dorsally than those ofthe other segments. The 354 PACIFIC SCIENCE, Volume 43, October 1989

(a) (b)

5mm 5mm

pi

(c)

FIGURE 1. Nisto ofScyllarus sp.: (a) Ventral surface; (b) dorsal surface; (c) enlarged view ofsternal plates; (d) lateral view of abdomen. (bc = posterior section of branchial carina, ds = distal squama of antenna, In = lateral notch, mc = cardiac section of median carina, pI = pleuron, ps = proximal squama, rt = rostral tooth, sp = sternal spine, sV = fifth thoracic sternite, t = tubercle, tI = tergum ofabdominal somite I, tg = transverse groove.)

pleura of somites II and III end in a poste­ Indentification and Morphology ofthe riorly directed point; those of somites IV and Palinuridae V are more rounded distally. The posterolateral margins ofthe fifth tho­ Only Panulirus marginatus and P. penicil­ racic sternite are produced as a long spinous latus are recorded from the Hawaiian Islands, process with a distinct lateral tubercle on each and it is possible to separate P. marginatus anterolateral margin. No tubercles are ap­ and P. penicillatus phyllosoma specimens by parent on the other sternites. differences in the shape ofthe cephalic shield. In many features, this nisto closely resem­ The length/width measurements ofthe cephalic bles those of adult Scyllarus aurora described shield are normally greater in P. marginatus by Holthuis (1981), and ofthe few members of than in P. penicillatus. However, these ratios this recorded from Hawaiian waters, in some of the material originally assigned to this species seems to be the most likely candi­ these species did not fit the data of Johnson date for its identity. (1968). Other characteristics examined in- Phyllosoma Larvae and Ocean Currents-PHILLIPS AND MCWILLIAM 355 cluded the special arrangement ofthe mouth­ parts in relation to the maxillipeds and the number of pairs of swimming setae on the NNNt"'I""lOOOOO exopods of the third maxilliped and the first -;~--;..-;~~~~ four pairs of walking legs (pereiopods). ------A summary of the data for some of the specimens difficult to identify is given in Table 1, and one of these specimens is illustrated in 0,,",000000 Figure 2. NOOI:-'r'lvO\oo None ofthese characteristics is useful alone .....;rr)-.::tM~"'O"'OOO because relationships change as the larvae de­ velop. A particularly useful characteristic used 'Ot'N-.:;t\O("f')N*** * by Johnson (1968) to examine P. marginatus OMvNN-t"'I""lt"'f":l and P. penicillatus was the distance from the r-i""';""';""';"",;"",;"",;"",; midpoint between the coxal segments of the second maxillipeds to the mouthparts ,,",000000 \OO\v-OONO'r'l (labrum-first maxillae). This distance is ex­ OO""':""';""';NNN pressed as a proportion ofthe distance between the attachment ofthe coxae to the thorax. To 00,,",0,,",000 ("'f')r-O\O\v 0'\ lrlO'\ make measurements comparable and to allow 0000""';""';""';""'; for changes due to larval growth, the distance OooV)OOO\\OV) (a) between the coxae is always considered 1.0 ~t-:v:t-:~'"d:~~ and the proportional distance (b) extends ------from the midpoint between the coxae to the anterior margin of the labrum (see Figure 2). V)ooaaaao Data ofspecimens marked with an asterisk ("f')V)lrlMOO\O\N ""';NMMtrilritrir--: in Table 1do not fit the data for P. marginatus or P. penicillatus given by Johnson (1968). In addition, from stage VI onward the ratio of OV"lOtnoooo the cephalic shield width to thorax width is -vv)\ooooooo\ about 1: 1 in P. penicillatus, and in most cases N-.:ivitrio\oOo\~ the specimens treated in Table 1 have a ratio of 1: 1.13 or higher. Also, the size of all stage VI or older stages is less than the minimum size given by Johnson (1968) for equivalent stages of P. marginatus and P. penicillatus. After examination of the data and advice from Johnson (pers. comm.), it was concluded that at least one Panulirus species other than P. marginatus or P. penicillatus was present in the samples. Positive identification of single specimens is always difficult, and most speci­ mens were damaged. Hence, it is possible that some of the specimens could be the larvae of Panulirus longipes or ofa species for which the larvae are not yet described, but they are de­ finitely neither P. marginatus nor P. penicil­ latus. Itwas eventually decided that at least 11 of the specimens were the larvae of Panulirus longipes femoristriga (von Martens). The list of species and the total number in (,) ~' U ~ l lb )f

CO~a ...I....~m"p 2

'-- ....1' lmm

5mm

FIGURE 2. Panulirus longipes femoristriga stage V: (a) dorsal surface; (b) ventral surface; (c) mouthparts. The method of measuring the distance between the coxae (a) and from the midpoint between the coxal segments of the second max.illipeds to the mouthparts (labrum-first maxillae) (b) is indicated. (co = coxa, la = labrum, mxl = first \0 maxilla, mxp 2 = second maxilliped.) -00 \0 Phyllosoma Larvae and Ocean Currents-PHILLIPS AND MCWILLIAM 357

TABLE 2

PHYLLOSOMA LARVAE AND NISTOS COLLECTED IN ThAWLS OFF THE SOUTHWEST COAST OF OAHU BETWEEN AUGUST 1977 AND OCTOBER 1978

SPECIES NO. OF PHYLLOSOMATA NO. OF NISTOS

Scyllaridae Arctides regalis 4 Scyllarus aurora 3 Scyllarus demani 1 Scyllarus sp. o 3 Total 11 Palinuridae Panulirus marginatus 10 Panulirus marginatus? 26 Panulirus penicillatus 6 Panulirus penicillatus? 4 Panulirus penicillatus/marginatus? 5 Panulirus longipes femoristriga? 11 Total 62

the samples is given in Table 2. The data were Uchida, pers. comm.), and as far east as the too few for seasonal analysis. Tuvalu and the Tuamotu islands (Michel 1969). The late stages of development of both species ofthese larvae suggest that they could DISCUSSION have been transported over a considerable dis­ The specimen referred, after comparison tance from their point of hatching (Phillips with reference material, to Scyllarus demani and McWilliam 1986). Furthermore, their time Holthuis was a final (gilled) stage. S. demani of capture around Hawaii is consistent with has not previously been reported from Ha­ the timing of certain broad features of the waii. However it is reported from Australia oceanographic circulation of the region. (Ritz 1977) and Indonesia (Holthuis 1947). The oceanographic model of the Hawaiian The specimen strongly resembles the specimen region proposed by Seckel (1962: 407-408) described by Johnson (1971b) as Scyllarus sp. was an "envelope" of North Pacific Central C. from the South China Sea. Water moving up and down the chain of is­ Ofthe 11 specimens provisionally identified lands in association with the seasonal varia­ as Panulirus longipes femoristriga, 8 (stages tion in the intensity and position of surface IV-VII) were caught between August and boundary currents. This movement brings November, 1 (stage V) in January, 1 (stage water from the northern part ofthe Hawaiian VIII) in February, and 1(stage VII) in March. chain to the southern islands such as Oahu Johnson (1971 a) stressed the local origin of during the period of September or October the Hawaiian species described in that paper, through April or May. Studies by Robinson but P.longipesfemoristriga is not known from (1969) and Hasunuma and Yoshida (1978) Oahu or any other island in the chain. This have revealed that the surface flow from west species is widespread in the Indo-Pacific and to east across the North Pacific gyre is a coun­ is known from Amboina, Banda, Moluccas, tercurrent, now called the Subtropical Coun­ Papua New Guinea, Japan, Queensland, New tercurrent, which flows in the opposite di­ South Wales, New Caledonia, New Hebrides, rection to the major current system in the Fiji (George and Holthuis 1965, Pyne 1970), Hawaiian Archipelago, the North Pacific Guam and the Northern Mariana Islands (R. Equatorial Current. The Subtropical Counter- 358 PACIFIC SCIENCE, Volume 43, October 1989 current extends from the western boundary of occurrence offOahu. However, the possibility the Pacific to the Hawaiian Archipelago and that some oftheir late-stage larvae in the catch consists ofa train ofanticyclonic eddies about represent sporadic recruitment from a western 300-600 km across that travel east at vari­ source such as Johnston Island (some 687 km able speed but occasionally accelerate to over to the southwest) or the more westerly Wake 50 cm/s. Hence, phyllosoma larvae that were Island cannot be dismissed. carried outby the Subtropical Countercurrent Grigg (1981), in his discussion of possible from the western Pacific into the circulation sources of larval recruitment for Acropora, system of the Hawaiian region could even­ has estimated possible transport time by the tually become distributed around Oahu. Subtropical Countercurrent from Johnston Edmondson (1940), in reviewing the rela­ Island to French Frigate Shoals of 50 days tion of the marine fauna of Hawaii to that of and from Wake Island of 187 days. Both of the other sections ofthe Pacific area, reported these timespans are within the larval life span that the shoal-water fauna of Ha­ of most palinurid species (Phillips and Sastry waii is chiefly ofIndo-Pacific origin. Many of 1980), and both Johnston and Wake Islands the common shore species range all the way are possible sources of larval recruitment. from the Red Sea to Hawaii. A preponderance A timespan of 50 days from Johnston Is­ of Indo-West-Pacific elements in the Hawai­ land may be well within the normal larval life ian marine fauna is evident in more recent span of Scyllarus demani also. Laboratory­ data that indicate a relatively low percentage reared phyllosomata of this species averaged ofendemic species in most ofthe coastal, ben­ about 48 days from hatching to metamor­ thic invertebrate taxa ofcentral Pacific islands phosis (M. Ito, James Cook University of (Scheltema 1986) and especially ofthe Hawai­ North Queensland, pers. comm.). This single ian Archipelago (Kay and Palumbi 1987). This final phyllosoma of S. demani collected off also applies to the Palinuroidea (palinurid, Oahu may be an "expatriate" (Johnson and synaxid, and scyllarid lobsters) recorded to Brinton 1963) from a more westerly stock. Ifit date. Arctides regalis, Scyllarus modestus, S. did not hatch somewhere along the Hawaiian aurora, and Panulirus marginatus are the only chain, then its occurence here suggests that Hawaiian endemics of the 10 species listed settlement on the archipelago by additional here. The remaining species have West Pacific, West Pacific lobster species transported in the Indian Ocean, or Indo-West-Pacific distribu­ Subtropical Countercurrent is still possible. tions (Phillips et al. 1980), and Parribacus ant­ Johnson (1977) described a final-stage larva articus also has an Atlantic distribution (Hol­ of another Scyllarus species unknown from thuis 1985). Two of the species listed, Scyl­ the Hawaiian area that he believed was of larides haanii and Palinurellus wieneckii, local origin. are very recent first records (Morin and Amerson and Shelton (1976) did not report MacDonald 1984, Titgen and Fielding 1986). either Scyllarus demani or Panulirus longipes Therefore, it is possible that adults ofScyllarus jemoristriga as present at Johnston Atoll (Is­ demani and Panulirus longipes jemoristriga land) (although P. marginatus and P. penicil­ already inhabit the Hawaiian region. They latus were collected). However, no extensive may be found eventually in further field sur­ systematic sampling was carried out, and, as veys, or reported after taxonomic revisions of pointed out by the authors, further collecting lobster material held in earlier collections from would probably result in the location ofaddi­ the Hawaiian Archipelago. tional species. Because the larvae referred to P. longipes In general, the larvae ofthe Palinuridae (of jemoristriga included some mid- as well as late which P. longipes jemoristriga is a member) stages ofdevelopment, the presence ofa fairly have a more oceanic character and are the nu­ local (though undetected) parental stock, cou­ merically dominant phyllosomata in plankton pled with the seasonality of the Hawaiian cir­ samples taken in midocean (George 1974, culation pattern, seems the more likely expla­ Phillips and McWilliam 1986). Larvae of the nation for both their temporal and spatial smaller members of the Scyllaridae, particu- Phyllosoma Larvae and Ocean Currents-PHILLIPS AND MCWILLIAM 359

lady those ofthe genus Scyllarus, have a more dance estimates of small fishes by three coastal character and outnumber palinurid towed nets and preliminary results of the larvae in samples taken in shelfwaters (John­ use of small purse seines as sampling de­ son 1971b, Maigret 1978, McWilliam and vices. BioI. Oecanogr. 2: 311-340. Phillips 1983). Even the limited data of this EDMONDSON, C. H. 1925. Crustacea. Marine study conform to the general pattern. The Zoology ofTropical Central Pacific (Tana­ total catch numbered 11 scyllarid larvae (in­ ger Exped. Publ., 1). Bull. Bernice P. Bishop cluding only 1 Scyllarus phyllosoma) com­ Mus. 27: 3-62. pared with 62 palinurid larvae. All were col­ ---.1940. The relation ofthe marine fauna lected in oceanic waters. of Hawaii to that of other sections of the The oceanic larvae of Indo-West-Pacific Pacific area. Pages 593-598 in Proc. 6th species, therefore, seem to have a greater Pac. Sci. Congr., Berkeley. chance ofsettling on oceanic islands and atolls GEORGE, R. W. 1974. Coral reefs and rock of the central Pacific than larvae of more lobster ecology in the Indo-West Pacific re­ coastal character. This is evident in the species gion. Proc. Int. Coral Reef Symp. 1: 321­ composition of lobsters with an Indo-West­ 325. Pacific distribution recorded from the Hawai­ GEORGE, R. W., and L. B. HOLTHUIS. 1965. A ian region to date. The more coastal lobster revision of the Indo West Pacific Spiny larvae typically belong to the smaller scyllarid Lobsters of the Panulirus japonicus group. species, which have a relatively short pelagic Zool. Verh. Rijksmus. Nat. Hist. Leiden larval phase, with fewer larval stages. The 72: 1-36, 5 figs. larval life span of some Scyllarus species is GEORGE, R. W., and A. R. MAIN. 1967. The short (25 days to 3-4 months) compared with evolution ofspiny lobsters (Palinuridae): A that reported for Panulirus, Palinurellus (12 study of evolution in the marine environ­ months), and the few Scyllarides (8-9 months) ment. Evolution 21 :803-820. investigated (Phillips and Sastry 1980: table GRIGG, R. W. 1981. Acropora in Hawaii. Part IV). But oceanic larvae are not confined to the 2. Zoogeography. Pac. Sci. 35: 15-23. Palinuridae and Synaxidae. The larger scyl­ HASUNUMA, K., and K. YOSHIDA. 1978. Split­ larids, such as species of Scyllarides and ting of the subtropical gyre in the western Arctides as well as P. antarcticus, have as North Pacific. J. Oceanogr. Soc. Jpn. 34: many larval stages as the palinurids (Johnson 160-172. 1971a, Robertson 1969a,b) and, by inference, HOLTHUIS, L. B. 1947. Biological results ofthe a long pelagic larval phase. Snellius Expedition. XIV. The Macrura of the Snellius Expedition. 1. The Stenopidae, Nephropsidae, Scyllaridae and ACKNOWLEDGMENTS Palinuridae (with 11 plates and 2 text fig­ The material was kindly made available ures). Teminnckia VII: 1-178. by Thomas A. Clarke of the Department of ---. 1960. Preliminary descriptions ofone Oceanography and Hawaii Institute of Ma­ new genus, twelve new species and three new rine Biology. Richard Uchida ofthe National subspecies of scyllarid lobsters (Crustacea Marine Fisheries Service is acknowledged for Decapoda Macrura). Proc. BioI. Soc. Wash. his advice on the species ofPanulirus collected 73: 147-154. during surveys by the Fisheries Service. ---. 1963. Preliminary descriptions of some species ofPalinuridea (Crustacea De­ capoda, Macura Reptantia). Pmc. Ned. LITERATURE CITED K. Akad. Wet. Ser. C 66: 54-60. AMERSON, A. B., and P. C. SHELTON. 1976. ---. 1981. A new species of Scyllarus The natural history ofJohnston Atoll, Cen­ (Crustacea Decapoda Palinuridea) from tral Pacific Ocean. Atoll Res. Bull. 192: 1­ the Pacific Ocean. Bull. Mus. Natl. Hist. 479. Nat., Paris. 4th Ser., 3. Section A, no. 3: CLARKE, T. A. 1983. Comparison of abun- 847-853. 360 PACIFIC SCIENCE, Volume 43, October 1989

---. 1985. A revision of the family Scyl­ MURANO, R. 1957. Plankton as the indicator laridae (Crustacea: Decapoda: Macrura). 1. of water masses and ocean currents. Subfamily Ibacinae. Zool. Verh. Rijksmus. Oceanogr. Mag. 9: 55-63. Nat. Hist. Leiden 218: 1-130. PmLLIPS, B. F., J. S. COBB, and R. W. GEORGE. JOHNSON, M. W. 1968. Palinurid phyllosoma 1980. General biology. Pages 1-82 in J. S. larvae from the Hawaiian Archipelago Cobb and B. F. Phillips, eds. The biology (Palinuridea). Crustaceana Suppl. (Leiden) and management of lobsters. Vol 1. Physi­ 2:59-79. ology and behaviour. Academic Press, New ---. 1971a. The phyllosoma larvae ofslip­ York. per lobsters from the Hawaiian Islands and PHILLIPS, B. F., and P. S. MCWILLIAM. 1986. adjacent areas (Decapoda, Scyllaridae). The pelagic phase of spiny lobster develop­ Crustaceana (Leiden) 20:77-103. ment. Can. J. Fish. Aquat. Sci. 43:2153­ ---. 1971b. On palinurid and scyllarid 2163. lobster larvae and their distribution in the PHILLIPS, B. F., and A. N. SASTRY. 1980. South China Sea (Decapoda, Palinuridea). Larval ecology. Pages 11-57 in J. S. Cobb Crustaceana (Leiden) 21 :247-282. and B. F. Phillips, eds. The biology and ---. 1977. On a hitherto unknown phyl­ management of lobsters. Vol. 11. Ecology 10soma larval species of the slipper lobster and management. Academic Press, New Scyllarus (Decapoda, Scyllaridae) in the Ha­ York. . waiian Archipelago. Pac. Sci. 31: 187-190. PYNE, R. R. 1970. Tropical spiny lobsters, JOHNSON, M. W., and E. BRINTON. 1963. Bio­ Panulirus spp. of Papua and New Guinea. logical species, water-masses and currents. Search 1:248-253. Pages 381-414 in M. N. Hill, ed. The Sea: RATHBUN, M. J. 1906. The Brachyura and Ideas and observations on progress in the Macrura ofthe Hawaiian Islands. Bull. U.S. study of the seas. Vol 2. Interscience Pub­ Fish. Comm. 23:827-930, text fig. 1-79, lishers, New York. pI. 1-24. KAY, E. A., and S. R. PALUMBI. 1987. Endem­ RITZ, D. A. 1977. The larval stages of Scyl­ ism and evolution in Hawaiian marine in­ larus demani Holthuis with notes on the vertebrates. Trends Ecol. Evol. 2: 183-186. larvae of S. sordidus (Stimpson) and S. MCWILLIAM, P. S., and B. F. PmLLIPS. 1983. timidus Holthuis (Decapoda, Palinuridea). Phyllosoma larvae and other crustacean Crustaceana (Leiden) 32: 229-240. macrozooplankton associated with Eddy ROBERTSON, P. 1969a. Biological investiga­ 'J', a warm-core eddy offsoutheastern Aus­ tions of the deep sea. No. 48. Phyllosoma tralia. Aust. J. Mar. Freshwater Res. 34: larvae of a scyllarid lobster, Arctides 653-663. guineensis from the western Atlantic. Mar. MAIGRET, J. 1978 (1977). Contribution a l'e­ BioI. 4: 143-151. tude des langoustes de la cote occidentale ---. 1969b. The early larval development d'Afrique (crustaces, decapodes, Palinu­ ofthe scyllarid lobster Scyllarides aequinoc­ ridae). 4. Notes sur la croissance et l'aspect tialis (Lund) in the laboratory with a review des populations sur les cotes du Sahara. of the larval characters ofthe genus. Deep­ Bull. Inst. Fondam. Afr. Noire Ser. A 39(4): Sea Res. 16: 557-586. 841-894. ROBINSON, M. K. 1969. Theoretical predic­ MICHEL, A. 1969. Les larves phyllosomes du tions of subtropical countercurrent con­ genre Panulirus-Palinuridae-(Crustacea firmed by bathythermograph (BT) data. Decapoda) due Pacifique Tropical sud et Bull. Jpn. Soc. Fish Oceanogr. Spec. No. Equatorial. Cah. O.R.S.T.O.M. Ser. (Professor Uda's Commemorative Papers): Oceanogr. 7(4): 3-19. 115-121. MORIN, T. D., and C. D. MACDONALD. 1984. SCHELTEMA, R. S. 1986. Long-distance dis­ Occurrence ofthe slipper lobster Scyllarides persal by planktonic larvae of shoal-water haanii in the Hawaiian Archipelago. Proc. benthic invertebrates among Central Paci­ BioI. Soc. Wash. 97:404-407. fic Islands. Bull. Mar. Sci. 39: 241-256. Phyllosoma Larvae and Ocean Currents-PHILLIPS AND MCWILLIAM 361

SECKEL, G. R. 1962. Atlas of the oceano­ currence of Palinurellus wieneckii (de Man, graphic climate of the Hawaiian Islands 1881) in the Hawaiian Islands (Decapoda: region. U.S. Fish Wildl. Serv., Fish. Bull. Palinura: Synaxidae). J. Crustacean BioI. 61 :371-427. 6(2): 294-296. TITGEN, R. H., and A. FIELDING. 1986. Oc-