crustacean research, no. 39: 3736 - 5352, 2010 37 Larval stages of the crinoid-associated squat lobster, Allogalathea elegans (Adams & White, 1848) (: : ) described from laboratory- reared material Yoshihisa Fujita

Abstract.– T h e c o m p l e t e l a r v a l of Lauriea gardineri (Laurie, 1926) and development of Allogalathea elegans Phylladiorhynchus integrirostris (Dana, 1853) (Adams & White, 1848) is described (see Fujita & shokita, 2005; Fujita, 2007), all and illustrated from laboratory-reared from coral reefs of the ryukyu Islands. material. This species has four zoeal t his paper describes the larval stages and one megalop. Diagnostic zoeal development of Allogalathea elegans for characters of the genus Allogalathea are the first time. Diagnostic zoeal characters of compared with those of the other galatheid the genus Allogalathea are compared with genera for which the larval morphology those of the other galatheid genera for which is known. Zoeas of A. elegans are readily the larval morphology is known. Megalopal distinguished from those of other morphology of A. elegans is also compared galatheids by the combination of setation with those of the two sympatric galatheids, of the maxillular and maxillar endopods. Galathea amboinensis and G. inflata. Megalopal morphology of A. elegans is remarkably different from those of the two Materials and Methods sympatric galatheids, Galathea amboinensis and G. inflata, in dentition on the lateral spent female margins of the rostrum and carapace. an ovigerous female of Allogalathea elegans was found solitarily on the host crinoid, Oxycomanthus bennetti (Müller, Introduction 1841), on the onna coast (udui) of okinawa Allogalathea elegans (adams & White, Island, ryukyu Islands, on 27 september 1848) is one of the well-known crinoid- 2000. however, this identification is associated squat lobsters, inhabiting shallow provisional, because cabezas et al. (in press) waters in the Indo-West Pacific from the are revising the genus, in which three new eastern coast of africa to Fiji Islands and species are described, along with A. elegans. from Japan to southern australia (Baba et al., three of the four species are being reported 2008, 2009). this species is also common in from Japan, so diagnostic characters of the the ryukyu Islands, Japan where four other spent female used in this study are given crinoid-symbiotic squat lobsters, all belonging in order to allow correct identification of to the genus Galathea, are sympatric, often the present material after the revisional sharing the same crinoid hosts (Baba & work has been published: (1) body orange Fujita, 2008). During my current studies on in base colour, with broad, yellow median the biology of the ryukyuan galatheids, I longitudinal stripe flanked by one dark brown have reported crinoid-galatheid associations and one white stripe on each side; (2) rostrum and larval development of G. amboinensis 2.1 times longer than broad; (3) pereopod De Man, 1888 and G. inflata Potts, 1915 1 palm 1.6 times longer than dactylus; (4) (see Fujita & Baba, 1999; Fujita et al., 2001, pereopod 2 merus lacking distinct spines on 2003). Meanwhile, I have also reported the dorsal margin, length 2.7 times breadth; and larval development of Sadayoshia edwardsii (5) epipods present on pereopods 1-2 . (Miers, 1884), and the zoeal morphology 38 LarVaL staGes oF ALLOGALATHEA ELEGANS 37

rearing methods and descriptions of larvae between 13 and 16 days. hatching of larvae occurred on 6 october 2000, and 45 zoeas were recorded. almost Description of larvae all of the hatched zoeas were mass-cultured Allogalathea elegans (adams & White, 1848) in a circular plastic tank (30 cm in diameter) Figs. 1–10, tables 1–2 containing 8-liter of filtered, uV treated First Zoea seawater. In addition, five zoeas were kept size. cL 1.18-1.25 mm, tL 2.64-2.68 mm. individually in 50 ml glass beakers in order to carapace (Figs. 1a, 2a, 2f). typical determine the normal number of their larval galatheid larval form, with posteriorly stages and duration. the salinity and water produced acute spine on each side; temperatures of the seawater were 34.0–35.0 posterodorsal margin with 17-18 small teeth, ‰ and 25.5–27.5 ºc, respectively. Food given posteroventral margin with 18-25 small teeth; throughout all the larval stages was freshly rostrum spiniform, covered with denticle- hatched Artemia sp. nauplii. approximately like small spines, extending to level of tip of one-third of the water in the tank was changed antennal scaphocerite; eyes sessile. daily. antennule (Fig. 3a). elongated, slightly the zoeal and megalopal specimens were swollen medially and narrowing distally, fixed and preserved in 50 % ethylene glycol terminally bearing 3 aesthetascs and 3 setae for morphological observations. Dissections (comprising 2 simple and 1 plumose setae); and measurements were carried out under one long plumose seta on future endopod bud. a nikon sMZ-10 binocular microscope antenna (Fig. 3f). Protopod with spinulose by using fine entomological needles. distal process; endopod fused to protopod, Measurements of carapace length (cL), as long as distal process of protopod, distally total length (tL), and postorbital carapace bearing one terminal seta; scaphocerite length (PcL, measured only in megalops) posteriorly produced into an acute spine, follow Fujita et al. (2001). Drawings were covered with small spines, and mesial margin made using a nikon optiphot-2 microscope. with 10 plumose setae, ventral surface heavily Descriptions were based on two specimens dentate. of each zoeal stage and one specimen of Mandibles (Fig. 4a). asymmetrically the megalop. Body somites are described dentate; without palp. from anterior to posterior, appendages from Maxillule (Fig. 5a). coxal endite with 7 endopod to exopod, and segments and plumodenticulate setae; basial endite with setae from proximal to distal. terminology 2 cuspidate and 3 plumodenticulate setae; generally follows Gore (1979) and Ingle endopod unsegmented, with 1 short seta (1991). proximally and 1+4 setae distally (setal the spent female and complete larval formula, 1+1+4). series (undissected samples) are deposited Maxilla (Fig. 6a). coxal and basial endites in the ryukyu university Museum, Fujukan, bilobed, with 8+4 and 5+4 plumodenticulate okinawa, Japan (ruMF) under the following setae, respectively; endopod unsegmented, registration numbers: ruMF-Zc-01174 for with 3 subdistomesial, 2 distomesial, and the spent female, ruMF-Zc-1175-1178 for 4 distolateral setae (setal formula, 3+2+4); first to fourth zoeas, RUMF-ZC-1179 for the scaphognathite with 4 marginal plumose megalop. setae, posteriorly ending in elongate plumose process. First Maxilliped (Fig. 7a). coxa with 2 results terminal setae; basis with 12 ventral setae Allogalathea elegans passed through four arranged 3+3+3+3; endopod 5-segmented, zoeal stages before attaining the megalopal with setation of 3, 2, 1, 2, 4 +I (I=dorsal (decapodid) stage. however, all the megalops plumose seta); exopod medially constricted, could not metamorphose to the first juvenile with 4 natatory plumose setae terminally. stage. the duration of the zoeal stages from second Maxilliped (Fig. 8a). coxa naked; hatching to the end of the fourth stage ranged basis with 1+2 setae on distoventral margin; 38 Y. FuJIta 39

Fig. 1. Allogalathea elegans (Adams & White, 1848): a, first zoea, lateral; b, second zoea, lateral; c, third zoea, lateral; d, fourth zoea, lateral. scale bars = 0.5 mm.

endopod 4-segmented, with 2, 2, 2, 4+I setae; short plumose seta (=anomuran hair), third to exopod as in first maxilliped. seventh articulated, long and stout plumose third Maxilliped (Fig. 9a). small setae with small spines. biramous bud. Pereopods. not visible. second Zoea abdomen (Figs. 1a, 10a). Five somites; size. cL 1.48-1.51 mm, tL 2.97-3.04 mm. each somites covered with numerous small carapace (Figs. 1b, 2b, 2g). Posterodorsal spines; posterodorsal margins without distinct and posteroventral margin of carapace with teeth; pair of posterolateral spines on somites 3-14 and 16-19 teeth, respectively; dorsal 4-5; pair of short setae on posterodorsal surface of carapace with 3 pairs of short margin of somites 2-5; pleopods absent. setae, one pair at midpoint of carapace, two telson (Fig. 10a, 10b). trigonal form, pairs between eyes; rostrum depressed, lateral deeply concave on median part of posterior margin with 23-27 teeth; eyes now stalked. margin, dorsal surface covered with numerous antennule (Fig. 3b). Protopod with small spines, posterior margin with 7 + 7 3 plumose setae on distal one-fourth, 4 processes (telsonal formula; I +ii + 3-7), first aesthetascs plus 4 setae on distal end; (lateralmost) spines fused to telson, second endopodal bud slightly developed, with a 40 LarVaL staGes oF ALLOGALATHEA ELEGANS 39

Fig. 2. Allogalathea elegans (Adams & White, 1848): a-d, carapace, first to fourth zoeas, dorsal; e, megalop, dorsal; f, rostrum, first zoea, dorsal; g, same, second zoea, dorsal; h, same, megalop, dorsal. Scale bars: a-e = 0.5 mm, f-g = 0.1 mm.

terminal plumose seta. absent; scaphocerite similar to first zoea. antenna (Fig. 3g). Protopod with Mandibles (Fig. 4b). as in first zoea; additional robust spine at ventrodistal end; without palp. endopod developed, slightly longer than in Maxillule (Fig. 5b). coxal endite the previous stage, distal plumose seta now unchanged; basial endite with 4 cuspidate 40 Y. FuJIta 41

Fig. 3. Allogalathea elegans (Adams & White, 1848), antennule (a–e) and antenna (f–j): a, f, first zoea; b, g, second zoea; c, h, third zoea; d, i, fourth zoea; e, j, megalop, middle segments of antenna omitted. scale bars = 0.1 mm.

and 3 plumodenticulate setae; endopod First Maxilliped (Fig. 7b). coxa, basis and unchanged. endopod unchanged; exopod with 7 plumose Maxilla (Fig. 6b). c o x a l e n d i t e natatory setae. unchanged; proximal lobe of basial endite second Maxilliped (Fig. 8b). coxa and unchanged, distal lobe with 5-6 setae; basis unchanged; second segment of endopod endopod unchanged; scaphognathite now with 1 dorsal plumose seta; exopod with 7 with 6 marginal plumose setae. natatory setae. 42 LarVaL staGes oF ALLOGALATHEA ELEGANS 41

Fig. 4. Allogalathea elegans (Adams & White, 1848), mandible (R, right side; L, left side) : a, first zoea; b, second zoea; c, third zoea; d, fourth zoea; e, left mandible of megalop. scale bars = 0.1 mm.

third Maxilliped (Fig. 9b). endopod carapace (Figs. 1c, 2c). almost as in developed, naked or with 1-2 terminal setae; previous stage; rostrum with 23-27 lateral exopod with 6 natatory setae. teeth; posterodorsal and posteroventral Pereopods (Fig. 9f). unsegmented buds. margins of carapace with 3-5 and 12 teeth, Abdomen (Fig. 1b). Similar to first zoea; respectively. no pleopod bud. antennule (Fig. 3c). Basal part of protopod Telson (Fig. 10c). Similar to first zoea, but slightly swollen, with 1 short seta, midpoint posterior plumose proccesses arranged as 8+8 of protopod with1 short seta, terminally with (telsonal formula; I +ii + 3-8); pair of short 1 long plumose seta directly proximal to setae on dorsal surface. base of endopod and 4 short plumose setae at junction of exopod; endopod similar to that third Zoea of first zoea; exopod now articulated, with 3 size. cL 1.78-1.85 mm; tL 3.55-3.70mm. rows of lateral aesthetascs numbering 2, 3, 42 Y. FuJIta 43

Fig. 5. Allogalathea elegans (Adams & White, 1848), maxillule: a, first zoea; b, second zoea; c, third zoea; d, fourth zoea; e, megalop. scale bars = 0.1 mm.

2, terminally with 3 aesthetascs and 4 setae Maxillule (Fig. 5c). coxal endite with 8 (including 1 plumose). plumodenticulate setae; basial endite with antenna (Fig. 3h). endopod much 5 cuspidate and 3 plumodenticulate setae; longer than in previous stage, with 1 short endopod unchanged. plumose seta subdistally; mesial margin of Maxilla (Fig. 6c). coxal and basial scaphocerite with 11 plumose setae; no other endites with 8+4 and 7+7 setae, respectively; changes. endopod unchanged; scaphognathite with 10 Mandibles (Fig. 4c). small palp bud newly marginal plumose setae, posterior plumose appeared. process shorter than in previous stage. 44 LarVaL staGes oF ALLOGALATHEA ELEGANS 43

Fig. 6. Allogalathea elegans (Adams & White, 1848), maxilla: a, first zoea; b, second zoea; c, third zoea; d, fourth zoea; e, megalop. scale bars = 0.1 mm.

First Maxilliped (Fig. 7c). almost as in Pereopods (Fig. 9g). More developed than previous stage. in the previous stage, first pereopod bifid second Maxilliped (Fig. 8c). third distally, showing chelate appearance. segment of endopod with 1 dorsal plumose abdomen (Figs. 1c, 10d). six somites; seta, setal fomula now 2, 2+I, 2+I, 4+I; no biramous uropods newly appeared, exopod other changes. well developed, with 10-11 marginal plumose third Maxilliped (Fig. 9c). similar to setae, endopod naked bud; no pleopods. second zoea; endopod articulated. telson (Fig. 10d). Posterior processes still 44 Y. FuJIta 45

Fig. 7. Allogalathea elegans (Adams & White, 1848), first maxilliped: a, first zoea; b, second zoea; c, third zoea; d, fourth zoea; e, megalop. scale bars = 0.1 mm.

8+8, but fourth process on each side robust, small teeth, respectively; rostrum with 23-26 fixed spine covered with numerous minute lateral teeth. spines (telsonal formula; I+ii+3+IV+5-8) ; 2 antennule (Fig. 3d). Proximal half of pairs of short setae on posterodorsal surface. protopod more swollen than in previous stage, with 2 short distal setae, distal half of Fourth Zoea protopod with 2 short setae at midpoint, 1 size. cL 1.96-2.10 mm; tL 3.94-4.15 long plumose seta directly proximal to base mm. of endopod and 4 short plumose setae at carapace (Figs. 1d, 2d). almost as junction of exopod; endopod more elongate in previous stages; posterodorsal and than in previous stage, terminal plumose seta posteroventral margins with 0-3 and 11-12 now absent; exopod with 4 rows of lateral 46 LarVaL staGes oF ALLOGALATHEA ELEGANS 45

Fig. 8. Allogalathea elegans (Adams & White, 1848), second maxilliped: a, first zoea; b, second zoea; c, third zoea; d, fourth zoea; e, megalop. scale bars = 0.1 mm.

aesthetascs numbering 4, 3, 3, 2, terminally Mandibles (Fig. 4d). Palp well developed, with 3 aesthetascs, 3 simple setae, and 1 but unsegmented. plumose seta. Maxillule (Fig. 5d). coxal endite with 9 antenna (Fig. 3i). endopod well plumodenticulate setae; basial endite with developed, 3.5 times as long as ventodistal 7 cuspidate and 3 plumodenticulate setae; spine of protopod; mesial margin of endopod unchanged. scaphocerite with 12 plumose setae; no other Maxilla (Fig. 6d). coxal endite with 13+5 changes. setae; basial endite with 8+7 setae; endopod 46 Y. FuJIta 47

Fig. 9. Allogalathea elegans (Adams & White, 1848), third maxilliped (a–e) and pereopods (f–j): a, first zoea; b, second zoea; c, third zoea; d, fourth zoea; e, megalop; f, second zoea; g, third zoea; h, fourth zoea; i, megalop, third pereopod; j, megalop, fifth pereopod. Scale bars = 0.1 mm.

unchanged; scaphognathite ending posteriorly in previous stage, fifth pereopod now distally in plumose process, with 19 plumose setae bifid. on lateral margin, 2 plumose setae on mesial abdomen (Figs. 1d, 10e, 10f). Biramous margin. pleopods now appeared on somites 2-5; First Maxilliped (Fig.7d). unchanged, as exopod of uropod with 12-13 marginal in previous stage. plumose setae, endopod with 7 marginal second Maxilliped (Fig. 8d). unchanged. plumose setae. third Maxilliped (Fig. 9d). endopod well telson (Fig. 10e). Longer than broad; developed and indistinctly segmented, with 3 pairs of small setae on dorsal surface; no 2-3 setae; no other changes. other distinct changes. Pereopods (Fig. 9h). More developed than 48 LarVaL staGes oF ALLOGALATHEA ELEGANS 47

Fig. 10. Allogalathea elegans (Adams & White, 1848): a, abdomen and telson, first zoea, dorsal; b, distal part of telsonal process, first zoea, dorsal; c, telson, second zoea, dorsal; d, uropod and telson, third zoea, dorsal; e, same, fourth zoea, dorsal; f, pleopods on abdominal somites 2-5, fourth zoea, ventral; g, uropod and telson, megalop, dorsal; h, pleopods on abdominal somites 2-5, megalop, ventral. scale bars = 0.1 mm. 48 Y. FuJIta 49

Megalop (Decapodid) second segment with 1 short seta, third size. cL 1.03-1.13 mm; tL 3.34-3.45 segment with 2 terminal setae; endopod mm. 2-segmented, proximal segment with 2 setae, carapace (Figs. 2e, 2h). similar to adult distal segment with 3 terminal setae and with form in general, lateral margin with 8 spines; 3 subterminal setae; exopod 5-segmented, dorsal surface with several transverse striae, first segment naked, second, third and fourth with scattered setae as illustrated; rostrum segments with 7+5, 3+3, 3 lateral aesthetascs, triangular with 5 small lateral spines. respectively, fifth segment with 2 (1 long and antennule (Fig. 3e). Biramous; peduncle 1 short simple) setae. 3-segmented, proximal segment robust, antenna (Fig. 3j). Peduncle 4-segmented, with 3 distal spines: large distodorsal and first segment without distinct spine, second distolateral, and small distomesial, dorsal segment with distomesial and distolateral surface with 2 rows of setae in crescent spines, distomesial larger, third segment with arrangement suggesting opening of statocyst, distomesial spine; flagellum composed of 14

table 1. summary of dimensional and morphological characters in each zoeal stage of Allogalathea elegans (adams & White) reared under laboratory conditions. abbreviations: a, aesthetascs; ps, plumose seta; s, seta; sp= spine. telsonal formula follows that of Gore (1979).

Zoea I Zoea II Zoea III Zoea IV

cL (mm) 1.18-1.25 1.48-1.51 1.78-1.85 1.96-2.10 tL (mm) 2.64-2.68 2.97-3.04 3.55-3.70 3.94-4.15 rostral teeth absent 23-27 23-27 23-26 carapace with posterodorsal teeth 17-18 3-14 3-5 0-3 posteroventral teeth 18-25 16-19 12 11-12 eyes sessile stalked stalked stalked antennule: exopod tip 3a+3s(1ps) 4a+4s(1ps) 3a+4s(1ps) 3a+4s(1ps) lateral row absent absent (2+3+2)a (4+3+3+2)a antenna: scaphocerite setae 10ps 10ps 11ps 12ps Mandible: palp absent absent present present Maxillule: coxal endtite 7s 7s 8s 9s basal endite 2sp+3s 4sp+3s 5sp+3s 7sp+3s endopod (1+1+4)s (1+1+4)s (1+1+4)s (1+1+4)s Maxilla: coxal endtite (8+4)s (8+4)s (8+4)s (13+5)s basal endite (5+4)s (5+5-6)s (7+7)s (8+7)s endopod (3+2+4)s (3+2+4)s (3+2+4)s (3+2+4)s scaphognathite 4ps+I 6ps+I 11ps+I 22ps+I First maxilliped: coxa 2s 2s 2s 2s basis (3+3+3+3)s (3+3+3+3)s (3+3+3+3)s (3+3+3+3)s endopod 3s, 2s, 1s, 2s, 4s+I 3s, 2s, 1s, 2s, 4s+I 3s, 2s, 1s, 2s, 4s+I 3s, 2s, 1s, 2s, 4s+I exopod 4ps 7ps 7ps 7ps second maxilliped: basis (1+2)s (1+2)s (1+2)s (1+2)s endopod 2s, 2s, 2s, 4s+I 2s, 2s+I, 2s, 4s+I 2s, 2s+I, 2s+I, 4s+I 2s, 2s+I, 2s+I, 4s+I exopod 4ps 7ps 7ps 7ps third maxilliped: exopod 0 (bud) 6ps 6ps 6ps Pereopods: absent small bud 1st bifid 1st & 5th bifid abdomen: numbers of somites 5 5 6 6 somites with posterolateral spines 4-5 4-5 4-5 4-5 somites with posterodorsal teeth absent absent absent absent Pleopods: absent absent absent present uropods: endopod absent absent naked 7ps exopod absent absent 11ps (12-13)ps telson: telsonal formula I+ii+3-7 I+ii+3-8 I+ii+3+IV+5-8 I+ii+3+IV+5-8 dorsal setae absent 1 pair 2 pairs 3 pairs 50 LarVaL staGes oF ALLOGALATHEA ELEGANS 49

articles, each naked or with 1-8 setae distally. gonopores. Mandibles (Fig. 4e). symmetrically scoop- abdomen (Figs. 2e, 10g, 10h). six like process; palp 3-segmented, basal segment somites, scatteredly setose on dorsal surface with 2 setae, distal part of third segment with as illustrated. Biramous pleopods present 9 stout, serrate setae. on somites 2-5; exopod with natatory setae, Maxillule (Fig. 5e). coxal endite numbering 9, 9, 10, 9, respectively; endopods with 24 setae, 3 long setae on proximal successively longer posteriorly, with 2 or 3 lobe; basial endite with 17 cuspidate and small hooks. 6 plumodenticulate setae distally, and 1 uropod and telson (Fig. 10g). telson plumose seta on proximal lateral margin; subquadrangular, posterior margin emarginate endopod with 1 proximal seta. with 6 pairs of long, plumose setae, Maxilla (Fig. 6e). coxal endite bilobed, dorsal surface with sparse short setae as with 32+14 setae; basial endite bilobed, with illustrated; uropods biramous, exopod with 12+18 setae; endopod unsegmented, with 3 19-20 plumose setae along entire margin, proximal and 1 distal seta; scaphognathite and 2 simple setae and 4 or 5 spines on with 35 plumose setae marginally. posterodorsal surface, endopod shorter than First Maxilliped (Fig. 7e). coxal and exopod, with 12 plumose setae marginally, 8 basial endites with 10 and 29 setae on ventral setae and 3 spines on dorsal surface. margins as illustrated, respectively; epipod distinct, laminated lateral to the coxal endite; Discussion endopod unsegmented, terminally with 0-2 setae; exopod indistinctly 2-segmented, with the zoeas of Allogalathea elegans 2 (1 midlateral and 1 terminal) plumose setae. share important larval characters with other second Maxilliped (Fig. 8e). Bilobed; coxa galatheids (other than Munidopsis) as listed with 1 distomesial seta; basis with 4 setae; by Konishi & saito (2000), in particular, the the epipods absent; endopod 5-segmented, carapace with a pair of posterolateral spines proximal first segment (ischium) indistinctly and posterodorsal and posteroventral teeth, separated from basis, with 3 setae, second and the maxillar scaphognathite bearing a segment (merus) longer than first segment, long plumose process on the posterior margin. with 6 setae, third segment (carpus) with 2 Zoeal morphology described on the basis of setae at dorsodistal end; fourth and terminal the laboratory-reared materials is known for segments (propodus and dactylus) with 18 species of 11 galatheid genera: Agononida 6 and 9 setae inclusive of stout pectinate Baba & de saint Laurent, 1996, Babamunida setae, respectively; exopod incompletely cabezas, Macpherson, & Machordom, 2008, 2-segmented, proximal segments with 2 long Cervimunida Benedict, 1902, Galathea plumose and 4 short setae as illustrated, distal Fabricius, 1793, Lauriea Baba, 1971, Munida segment with 8 terminal plumose setae. Leach, 1820, Munidopsis Whiteaves, 1874, third Maxilliped (Fig. 9e). Bilobed; Neonida Baba & de saint Laurent, 1996, coxa with 8 setae; basis with 3 setae; epipod Phylladiorhynchus Baba, 1969, Pleuroncodes possibly present (difficult to confirm in this stimpson, 1960, and Sadayoshia Baba, 1969 study); gill buds not discernible; endopod (see Gore, 1979, Konishi & saito, 2000; 5-segmented, basal segment (ischium) with Fujita, 2007; Fujita & shokita, 2005; Fujita crista dentata bearing 8 denticles, with 4 setae et al., 2001, 2003; Guerao et al., 2006). Zoeal ventrally; second segment (merus) as long characters of A. elegans are summarized as as ischium (when measured along lateral follows: (1) the anterolateral spine of the midline), with distal spine at extensor margin, carapace is absent through the zoeal stages; (2) 3 (2 long and 1 small) setae; third to fifth the rostrum is spiniform in the first zoea, and segments (carpus, propodus and dactylus) in the subsequent stages it is depressed and with 6, 9, 9 setae, respectively; exopod with 8 flattened, with lateral teeth; (3) the maxillular terminal plumose setae. endopod is unsegmented, with the setation Pereopods (Figs. 2e, 9i, 9j). similar of 1+1+4 through the zoeal stages; (4) the to adult form in general; no male/female maxillar endopod bears 3+2+4 setae through 50 Y. FuJIta 51 2s *11 short absent absent (3+4)s 7+7[z1] 7+7[z1] 9+9[z4] (1+1+4)s 1 segment 1 8+8[z2-z3] (3,2,1,2,4)s Sadayoshia (3+3+3+3)s absent[z1-z2 6th somite [z3-z4] somite 6th amuelsen (1972); *8 *8 (1972); amuelsen s *10 absent absent (1+4)s (3+4)s 7+7[z1] 8+8[z2] 9+9[z3] 2s[z3-5] 1s[z1-z2] 1s[z1-z2] 1 segment 1 12+12[z5] (2,2,1,2,4)s long[z1-z2] long[z1-z2] (2+3+3+2)s short[z3-z5] absent[z1-z2] absent[z1-z2] Pleuroncodes 10-12+10-12[z4] 6th somite [z3-z4] somite 6th ars (1889), (1889), ars s 1s *9 7+7 short absent absent (1+4)s (3+2+4)s 1 segment 1 (2,2,1,2,4)s (2+3+3+2)s 4th-5th somites 4th-5th Phylladiorhynchus aito (2000); *7, *7, (2000); aito 2s *8 s 7+7 long absent absent absent (3+4)s Neonida 0, (1+4)s 0, 2 segments 2 (3,2,1,2,4)s (2+3+3+3)s are known only the firstzoeal stage (see Fagetti, 1960; *7 hokita (2005). (2005). hokita short naked 1s[z1] 1s[z1] 2s[z2] 1s[z1] 3s[z2] absent absent absent s present no data no (0,1,2,3)s (0,1,2,3)s 1 segment 1 Munidopsis 13-15+13-15[z1] 14-16+14-16[z2] Phylladiorhynchus oberts (1973), Konishi & Konishi (1973), oberts r 2s *6 absent absent (1+4)s (3+4)s Munida , , and 7+7[z1] 8+8[z2] 1 segment 1 10+10[z3] (3,2,1,2,4)s long[z1-z2] long[z1-z2] or (3+2+4)s or short[z3-z5] absent[z1-z2] (2-3+3+3+3)s 11-12+11-12[z4] 12-13+12-13[z5] 6th somite [z3-z4] somite 6th Neonida uus (1934), (1934), uus , , h 2s *5 7+7 short absent absent present Lauriea aito (2000), Guerao et Guerao aito *2, al. (2000), et *4, Guerao (2006); Gore (1960); *3, al. (1979), Fagetti (2006); (3+2+4)s 1s, (1+4)s 1s, 2 segments 2 (3,2,1,2,4)s s (3+3+3+3)s Lauriea ampodonico (1971); *11, Fujita & & Fujita *11, (1971); ampodonico , , c 2s *4 short absent absent present 7+7[z1] or absent or Galathea 8+8[z2-z5] 2 segments 2 (3,2,1,2,4)s (3+2-3+3-4)s (2-3+3+3+3)s (0-1)s, (1+3-4)s (0-1)s, Cervimunida , , *3 7+7 7+7 long absent absent absent (3+4)s no data no (1+1+4)s 1 segment 1 (2,2,1,2,4)s (2+3+3+3)s Cervimunida eferences: *1, Konishi & *1, Konishi eferences: r 2s Babamunida *2 7+7 long , , absent absent absent (3+4)s (1+1+4)s 1 segment 1 (3,2,1,2,4)s (3+3+3+3)s Babamunida 2s *1 7+7 long absent absent absent (3+4)s Agononida 0, (1+4)s 0, Agononida 2 segments 2 (3,2,1,2,4)s (2+3+3+3)s 2s short absent absent 7+7[z1] nger (1990), Fujita et al. (2001, 2003); *5, Fujita (2007); *6, *6, (2007); Fujita *5, 2003); (2001, al. et Fujita (1990), nger (1+1+4)s (3+2+4)s 1 segment 1 absent[z1] 8+8[z2-z4] (3,2,1,2,4)s (3+3+3+3)s a Allogalathea present study present present[z2-z4] aito, 2000; Guerao et Guerao 2000; aito, al., 2006). s hristiansen & hristiansen he he larval morphologies of Konishi & Konishi c t & Fagetti (1960), Boyd *10, (2007); Fujita *9: (2006); al. et Guerao bdomen: able 2. Morphological differences in zoeal stages among twelve galatheid genera, which larval morpology has been described. Abbreviations: s, seta; z1-z5, first to fifth zoeas. zoeas. fifth to first z1-z5, seta; s, Abbreviations: described. been has morpology larval which genera, galatheid twelve among stages zoeal in differences Morphological 2. able eferences teeth lateral ostrum: arapace: elson: t Genus r r c spine anterolateral endopod: Maxillular segments of number setation endopod: Maxillar maxilliped: First coxa basis endopod margins) (ventral a dorsal with somites spine posteromedian t processes posterior spine lateral 52 LarVaL staGes oF ALLOGALATHEA ELEGANS 51

the zoeal stages; (5) the first maxilliped has two setae on the coxa and 12 (3+3+3+3) setae Literature cited on the basis, through the zoeal stages; (6) the endopod of the first maxilliped has a setal Baba, K., & Fujita, Y., 2008. squat lobsters of formula 3,2,1,2,4 through the zoeal stages; (7) the genus Galathea (Decapoda: anomura: the abdominal segments are unarmed through Galatheidae) associated with comatulid the zoeal stages; (8) the telsonal formulae crinoids from the ryukyu Islands, Japan. are I+ii+3–7, I+ii+3–8, I+ii+3+IV+5–8, research, 37: 43–62. I+ii+3+IV+5–9, in the first, second, third and ———, Macpherson, e., Poore, G. c. B., fourth zoeas, respectively; and (9) the lateral ahyong, s. t., Bermudez, a., cabezas, spines of the telson are much shorter than P., Lin, c. W., nizinski, M., rodrigues, the posterior plumose processes through the c., & schnabel, K. e., 2008. catalogue zoeal stages (see tables 1 and 2). the setation of squat lobsters of the world (crustacea: of the maxillular endopod as displayed by D e c a p o d a : a n o m u r a - f a m i l i e s A. elegans is known only for the first zoeas chirostylidae, Galatheidae and Kiwaidae). of Cervimunida johni Porter, 1903 and Zootaxa, 1905: 1-220. Babamunida javieri (Macpherson, 1994), and ———, ———, Lin, c. W., & chan, t. Y., in all zoeal stages of Sadayoshia edwardsii 2009. crustacean Fauna of taiwan. squat (Miers, 1884) (see Fagetti, 1960; Fujita & lobsters (chirostylidae and Galatheidae). shokita, 2005; Guerao et al., 2006). however, national taiwan ocean university, A. elegans zoeas differ considerably from Keelung, 311pp. these three species by the setation of the Boyd, c. M., 1960. the larval stages maxillar endopod (see table 2). of Pleuroncodes planipes stimpson In the ryukyu Islands, five sympatric (crustacea, Decapoda, Galatheidae). crinoid-associated galatheids, Galathea Biological Bulletin, 118: 17–30. amboinensis, G. continua, G. inflata, G. cabezas, P., Macpherson, e., & Machordom, leptocheir and A. elegans, are found , often a., 2008. a new genus of squat lobster sharing the same crinoid hosts (Fujita & (Decapoda: anomura: Galatheidae) Baba, 1999; Baba & Fujita, 2008; Fujita, from the south West Pacific and Indian unpublished data). the Allogalathea elegans ocean inferred from morphological and megalop is readily distinguished from molecular evidence. Journal of crustacean megalop of G. amboinensis and G. inflata by Biology, 28: 68–75. the following characters: (1) the rostrum is ——— , ——— , & ——— , i n p r e s s . about as long as the remaining carapace and Allogalathea (Decapoda: Galatheidae): has five lateral teeth (much shorter rostrum a monospecific genus of squat lobsters? with 4 lateral teeth in Galathea); and (2) the Zoological Journal of Linnean society. carapace lateral margin possesses eight spines christiansen, M. e., & anger, K., 1990. (six in G. amboinensis or seven spines in G. complete larval development of Galathea inflata). intermedia Lilljeborg reared in laboratory culture (anomura: Galatheidae). Journal of crustacean Biology, 10: 87–111. acknowledgements Fagetti, e., & campodonico, I., 1971. Larval I am indebted to Dr. M. osawa of the development of the red crab Pleuroncodes university of the ryukyus for reading an m o n o d o n (Decapoda a n o m u r a : early draft of the manuscript and making Galatheidae) under laboratory conditions. valuable comments, and akane Ito for kind Marine Biology, 8: 70–81. help in the collection. thanks are Fujita, Y., 2007. First zoeas of two shallow- extended to Prof. K. Baba and anonymous water galatheids, Lauriea gardineri reviewers for their valuable comments and (Laurie, 1926) and Phylladiorhynchus suggestions on the manuscript. this study was integrirostris (Dana, 1853) (crustacea: supported in part by the research Institute of Decapoda: anomura: Galatheidae). Marine Invertebrates. Proceedings of the Biological society of 52 Y. FuJIta 53

Washington, 120: 74–85. the deep-sea squat lobsters, Agononida ———, & Baba, K., 1999. two galatheid incerta (henderson, 1888) and Munida associates of crinoids from the striola Macpherson and Baba, 1993 with ryukyu Islands (Decapoda: anomura: notes on larval morphology of the family Galatheidae), with their ecological notes. (crustacea: anomura: Galatheidae). crustacean research, 28: 112–124. Zoological science, 17: 1021–1029. ———, & shokita, s., 2005. the complete Miers, J. e., 1884. crustacea. In: report on larval development of Sadayoshia the zoological collections made in the edwardsii (Miers, 1884) (Decapoda: Indo-Pacific ocean during the voyage of anomura: Galatheidae) described from h.M.s. "alert", 1881-82. London, pp. laboratory reared material. Journal of 178–322, 513–575. natural history, 39: 865–886. roberts, P. e., 1973. Larvae of Munida ———, Baba, K., & shokita, s., 2001. Larval s u b r u g o s a ( W h i t e , 1 8 4 7 ) f r o m development of Galathea inflata Potts, Perserverance harbour, campbell Island. 1915 (Decapoda: anomura: Galatheidae) Journal of the royal society of new described from laboratory-reared material. Zealand, 3: 393–408. crustacean research, 30: 111–132. samuelsen, t. J., 1972. Larvae of Munidopsis ———, ———, & ———, 2003. Larval tridentata (esmark) (Decapoda, anomura) development of Galathea amboinensis reared in the laboratory. sarsia, 48: 91–98. (Decapoda: anomura: Galatheidae) sars, G. o., 1889. Bidrag til Kundskaben om under laboratory conditions. crustacean Decapodernes Forvandlinger. II. Lithodes- research, 32: 79–97. Eupagurus-Spiropagurus-Galathodes- Gore, r. h., 1979. Larval development G a l a t h e a - M u n i d a - P o r c e l l a n a - of Galathea rostrata under laboratory (Nephrops). archiv for Mathematik og conditions, with a discussion of larval naturvidenskab, 13: 133–201. development in the Galatheidae (crustacea Wilkens, h., Parzefall, J., & ribowski, a., anomura). Fishery Bulletin, united states, 1990. Population biology and larvae of the 76: 781–806. anchialine crab Munidopsis polymorpha Guerao, G., Macpherson, e., samadi, s., (Galatheidae) from Lanzarote (canary richer de Forges, B., & Boisselier, M.- Islands). Journal of crustacean Biology, c., 2006. First stage zoeal descriptions of 10: 667–675. five species from Western pacific (Crustacea: Decapoda: Anomura). Zootaxa, 1227: 1–29. addresses: university education center, huus, J., 1934, Zur morphologisch- university of the ryukyus, 1 senbaru, systematischen und biologischen Kenntnis nishihara-cho, okinawa 903-0213, Japan. der nordischen Munida-larven (crustacea Marine Learning center, 2-95-101 Miyagi, Decapoda). Bergens Museums Årbok, 8: chatan-cho, okinawa 904-0113,Japan. 1–32. e-mail: [email protected] Ingle, r., 1991. Larval stages of northeastern atlantic crabs: an Illustrated Key, Received: 5 april 2010. London: chapman and hall, 363 pp. Accepted: 2 november 2010. Konishi, K., & saito, t., 2000. Larvae of 54