crustacean research, no. 39: 5355 - 64,66 2010 55 The larval development of Chirostylus stellaris Osawa, 2007 (Crustacea: Anomura: Chirostylidae) described from laboratory reared material

Yoshihisa Fujita & Paul F. clark

Abstract.– T h e c o m p l e t e l a r v a l a complete larval description of all stages, development of the anomuran chirostylid, discuss the significance of larval appendage Chirostylus stellaris Osawa, 2007, is morphology described and compare described and illustrated from laboratory- Chirostylus larval characters with other reared material. This species has two chirostylid genera including Gastroptychus zoeal stages and one megalop. The zoeal caullery, 1896, Uroptychus henderson, 1888 phase was completed within 48 hours and and Eumunida smith, 1883. the megalopl phase lasted approximately three weeks. The setae on the coxal, Materials and Methods basial and endopodal segments of the maxillule, maxilla, first maxilliped and an ovigerous Chirostylus stellaris was second maxilliped were absent in zoeal collected from cape Maeda-misaki, okinawa stages suggesting that they did not feed Island, ryukyu Islands on 21 June 2003. the during this phase. In fact C. stellaris zoeas female was maintained in a 1.4-liter plastic showed lecithotrophic and abbreviated aquarium filled with seawater for about 20 development. The zoeas of Chirostylus days until releasing zoeas on 11 July 2003. are distinguishable from those of other nine zoeas were kept individually in 50 ml chirostylid genera such as Gastroptychus, glass beakers to determine the number of Uroptychus, and Eumunida. However, no larval stages and their duration. the salinity significant differences were found among and water temperature of the seawater were zoeas of three Chirostylus species including 34.5–35.0 Psu and 26.0–27.0ºc, respectively. C. ortmanni, C. sandyi and C. stellaris. the zoeas did not feed. Megalops were fed freshly hatched Artemia sp. nauplii and commercial tropical fish food (“tetraMin”, Introduction spectrum Brands, Inc.). Chirostylus ortmann, 1892 now contains the larvae were fixed and preserved in seven species, all from the Indo-Pacific 50 % ethylene glycol for morphological (Baba, 2009). however, larval information observations. Dissections and measurements of Chirostylus is known only from the first were carried out under a nikon sMZ- zoeas of two species: Chirostylus dolichopus 1000 binocular microscope by using fine ortmann, 1892, reported by ogawa & entomological needles. Drawings were made Matsuzaki (1992) (= currently C. ortmanni using a nIKon oPtIPhot-2 microscope. Miyake & Baba, 1968; see osawa & Zoeal appendages were described based on nishikiori, 1998) and C. ortmanni reported the malacostracan somite plan, from anterior by clark & ng (2008) (=currently C. sandyi to posterior as described by clark et al. (1998). Baba, 2009). recently an ovigerous female of carapace length (cL) was measured from C. stellaris osawa, 2007 was collected from the basal part of the rostrum to the posterior the ryukyu Islands, Japan and transported to midpoint of the carapace for zoeas, and laboratory where the larvae were reared to from the tip of the rostrum to the posterior first crab stage. midpoint of the carapace for megalop. the the purpose of this study is to provide measurements of the antennal endopod and 5654 Y. FuJIta & P. F. cLarK

Fig. 1. Chirostylus stellaris osawa, 2007. survival and duration of larval stages reared under laboratory conditions at 26.0–27.0°c in seawater.

protopodal process each were taken from its after the 22nd day. base to the tip. setal terminology generally follows Ingle (1991). Description of larvae the spent female (the paratype of Chirostylus stellaris osawa, 2007 Chirostylus stellaris) was deposited in the (Figs. 2–6) coastal Branch of natural history Museum First zoea and Institute, chiba (cMnh-Zc 2134) (see Material examined: 2 specimens (including osawa, 2007). the complete larval series one exuvia). (dissected and undissected specimens) was size: cL 1.22 mm. deposited in the ryukyu university Museum, carapace (Fig. 2a, B, c): multispinulate Fujukan, okinawa, Japan (ruMF) under (approximately 62 spines) globose carapace; the following registration numbers: ruMF- prominent, extremely long dorsal and lateral ZC-01229–01230 for first and second zoeas, spines absent; rostral spine present and ruMF-Zc-01231 for megalops. spinulate, subequal to antennal protopod in length; anterodorsal and posterodorsal setae absent; ventral margin without setae; eyes results sessile. the larval development of Chirostylus antennule (Fig. 3a): biramous; peduncle stellaris comprised two zoeal, indicating unsegmented; endopod unarticulated, abbreviated development, and one megalopal unsegmented, without setae; exopod stage. The first zoea molted within 12 hours to unsegmented, with 3 proximal, 3 (1+2) second stage zoea, then after 48 hours all the median and 5 terminal aesthetascs. second stage zoea metamorphosed to megalop antenna (Fig. 3B): biramous; protopod (Fig. 1). the maximum duration of the zoeal with distally multispinulate process; endopod phase was 48 hours after hatching. the first with 2 terminal denticles, length about half megalop appeared approximately 36 hours (52%) that of protopodal process. after the female released the first stage zoeas. Mandible: (Fig. 3c): almost symmetrical; the megalopal phase lasted approximately 15 incisor margins smooth; unsegmented palp days until the first juvenile crab was observed. bud present. no megalops were observed in the laboratory Maxillule (Fig. 3D): coxal and basial LarVaL DeVeLoPMent oF CHIROSTYLUS STELLARIS 5557

Fig. 2. Chirostylus stellaris Osawa, 2007, first zoea. A, lateral; B, carapace, abdomen, and telson, dorsal; C, rostrum, dorsal; D, posterior margin of telson, dorsal. scale bars: a, b = 0.5 mm; c, d = 0.1 mm.

endites lacking setae, with some minute with that of maxilliped 1, unsegmented, protuberances; endopod unsegmented, devoid of setae; exopod 2-segmented with 4 without setae. long plumose natatory setae. Maxilla (Fig. 3e): coxal and basial endites third maxilliped (Fig. 3h): biramous; weakly bilobed, without setae; endopod endopod unsegmented, without setae; exopod not segmented, without setae; exopod unsegmented, without setae. (scaphognathite) margin with 19–22 setae of Pereopods (Fig. 3I): pereopods 1–5 various lengths. present, with some incomplete segmentation; First maxilliped (Fig. 3F): coxa and basis pereopods 1–4 with simple pair of gill without setae; endopod arising from midpoint buds; pereopod 1 chelate; pereopod 5 small, of basis, relatively short compared with that markedly shorter than pereopod 1–4, not of maxilliped 2, unsegmented, devoid of chelate, but weakly bifid distally. setae; exopod 2-segmented, terminal segment abdomen (Figs. 2a, B): 5 somites and with 4 long plumose natatory setae. telson; somite 1 unarmed; somite 2 with second maxilliped (Fig. 3G): coxa and 1 posteromedian spine; somites 3–5 each basis without setae; endopod arising from with 1 posteromedian spine and pair of midpoint of basis, relatively long compared posterolateral spines; somite 6 fused to telson, 5856 Y. FuJIta & P. F. cLarK

Fig. 3. Chirostylus stellaris Osawa, 2007, first zoea. A, antennule; B, antenna; C, mandibles (r, right side; l, left side); D, maxillule; E, maxilla; F, first maxilliped; G, second maxilliped; H, third maxilliped; I, pereopods; J, pleopod 2; K, pleopod 3; L, pleopod 4; M, pleopod 5; n, uropod. scale bars = 0.1mm.

with 2 posterolateral spines; all these spines biramous uropod present on undifferentiated perpendicular to dorsal surface; biramous somites 6 (Fig. 3n), endopod and exopod pleopods present on somites 2–5 (Fig. 3J- without marginal setae, but with some minute M), endopods and exopods both naked; protuberances present. LarVaL DeVeLoPMent oF CHIROSTYLUS STELLARIS 5759

Fig. 4. Chirostylus stellaris osawa, 2007, second zoea. a, lateral; B, antennule; c, antenna; D, mandibles (r, right side; l, left side); e, maxillule; F, maxilla; G, first maxilliped; h, second maxilliped; I, third maxilliped; J, pereopods; K, pleopod 2; L, pleopod 3; M, pleopod 4; n, pleopod 5; o, uropod. scale bars: a = 0.5mm; B-o = 0.1mm.

telson (Fig. 2B, 2D): broadened posteriorly; lateral margin with 3 pairs of second zoea strong spines; posterior margin with 6 pairs Material examined: 2 specimens (including of spinulose spines, 4 lateral strong, 2 mesial one exuvia). fine; anomuran seta (hair) absent. size: cL 1.25 mm. carapace (Fig. 4a): eyes stalked; 6058 Y. FuJIta & P. F. cLarK

Fig. 5. Chirostylus stellaris osawa, 2007, megalop. a, dorsal; B, thoracic sternites, ventral; c, abdomens, lateral; D, pereopod 1, chela; e, pereopod 2, propodus, dactylus; F, pereopod 5, propodus, dactylus; G, pleopod 2; h, pleopod 3; I, pleopod 4; J, pleopod 5; K, uropod. scale bars: a = 0.5mm; B-K = 0.1mm. LarVaL DeVeLoPMent oF CHIROSTYLUS STELLARIS 5961

Fig. 6. Chirostylus stellaris osawa, 2007, megalop. a, antennule; B, antenna; c, mandibles; D, maxillule; e, maxilla; F, first maxilliped; G, second maxilliped; H, third maxilliped. Scale bars = 0.1mm.

otherwise unchanged. (approximately 50%) of protopodal process; a ntennule (Fig. 4B): peduncle otherwise unchanged. 3-segmented, proximal segment bearing Mandible: (Fig. 4D): unchanged. a small opening of statocyst; second and Maxillule (Fig. 4e): bud of seta present third segments without setae; endopod on proximal part of basial endite; otherwise incompletely articulated with peduncle; unchanged. exopod 3-segmented, with 3 proximal, 3 (1+2) Maxilla (Fig. 4F): exopod (scaphognathite) median and 3 terminal aesthetascs; otherwise margin now with 20–24 setae, posterior unchanged. lobe with small protuberances; otherwise antenna (Fig. 4c): endopod half length unchanged. 6260 Y. FuJIta & P. F. cLarK

First maxilliped (Fig. 4G): exopod with with very long distal seta. 8 long plumose natatory setae; otherwise Mandible: (Fig. 6c): nearly symmetrical; unchanged. incisor margins distinctly dentate; palp second maxilliped (Fig. 4h): exopod with 2-segmented, proximal segment naked, distal 8 long plumose natatory setae; otherwise segment with 1 terminal seta. unchanged. Maxillule (Fig. 6D): coxal endite with 7 third maxilliped (Fig. 4I): endopod now setae; basial endite with 19 (11 cuspidate and 5-segmented; otherwise unchanged. 8 plumodenticulate setae) processes distally, Pereopods (Fig. 4J): all pereopods and 2 plumose setae on proximal lateral distinctly segmented; pereopod 5 weakly bifid margin; endopod unsegmented, with 1 or 2 distally, subchelate; otherwise unchanged. subterminal setae. abdomen (Fig. 4a): minute protuberances Maxilla (Fig. 6e): coxal endite weakly (= setal buds) present on exopods of pleopods bilobed, with 8 (5+3) setae; basial endite 2–5 and exopod and endopod of uropod; weakly bilobed with 17 (6–7+10–11) setae; otherwise unchanged. endopod unsegmented, with 1 proximal seta; telson (Fig. 4a): unchanged. exopod (scaphognathite) margin with 33–35 setae. Megalop First maxilliped (Fig. 6F): coxa and basis Material examined: 2 specimens (including well flattened, with 4–6 and 5–6 marginal one exuvia). setae, respectively; endopod small, without size: cL 1.16 mm. setae; exopod unsegmented, with 2–4 short carapace (Fig. 5a): similar to adult in setae distally; epipod not visible. general shape; anterolateral and epigastric second maxilliped (Fig. 6G): coxa spines present; gastric region with 1 small and basis undifferentiated, without setae; spine; cardiac region with 1 small spine; endopod relatively long compared with that branchial region with 4 or 5 small spines; of maxilliped 1, 5-segmented with 0, 3–4, 2, 5, posterior margin concaved. rostrum short, 5–7 (1–2 plumodenticulate and 5–6 serrate) rounded, without spine. setae, respectively; exopod 2-segmented, sternum (Fig. 5B): sternite 3 laterally much longer than endopod, with 5 or 6 setae produced into blunt spine, anterior margin on distal segment; epipod not visible. nearly transverse, with 2 simple setae and 3 or third maxilliped (Fig. 6h): coxa with 2 4 small blunt spines; sternites 4–6 each with setae and 2 arthrobranchs; basis with 2 setae; blunt spine on each proximal lateral margin; endopod extremely long relative to those of sternite 7 without lateral spine. maxillipeds 1–2, 5-segmented, with 4, 6–5, a ntennule (Fig. 6 a ) : p e d u n c l e 8–9, 15–17, 12–13 setae, respectively, first 3-segmented, proximal segment broad, segment (ischium) with 13–15 teeth on crista dorsal surface with rows of setae in crescent dentata, remaining segments (merus, carpus, arrangement suggesting opening of statocyst, propodus, and dactylus) without spines; median segment without setae, distal exopod 2-segmented, proximal segment with segment with 1 short terminal seta; endopod 1 subterminal seta, distal segment with 6 2-segmented, proximal segment with 2 setae, terminal plumose setae; epipod absent. distal segment with 6 or 7 (3 subterminal, 3 Pereopods (Fig. 5a, D-F): distinctly or 4 terminal) setae; exopod 3-segmented, segmented, well formed; pereopods 1–4 proximal segment with 3 aesthetascs, median exceedingly long, slender, subcylindrical, segment with 3 (1+2) aesthetascs and 2 setae, surfaces with sparsely setose and spinose distal segment with 3 aesthetascs and 1 seta. as illustrated; pereopod 1 longer than other antenna (Fig. 6B): 8-segmented, pereopods, opposable margins of fingers proximal first to third segments incompletely each with prominent acute tooth proximally articulated, third segment proximally fused and moderately small teeth distally, movable to second; fifth segment with 1 distal spine; finger distally ending in sharp spine fitting short setae arranged 0,0,0,1,1,0,3,3 from between 2 terminal spines of fixed finger proximal to distal segments, terminal segment when closed; pereopods 2–4 with 5 flexor LarVaL DeVeLoPMent oF CHIROSTYLUS STELLARIS 6163 spines on dactylus, ultimate as long as the case, because C. dolichopus ranges from and somewhat narrower than penultimate; Japan to the eastern coast of africa, via pereopod 5 reduced in size, distally chelae Western australia (Baba et al., 2008). In and setose, fingers short relative to propodus, contrast to the zoeal phase, the megalopal especially fixed finger. phase of C. stellaris survived about three abdomen (Fig. 5a, c, G-K): 6 somites, weeks before metamorphosing to the first sparsely setose as figured, unarmed (long juvenile stage. therefore, it can be assumed spines as in zoeal stages disappeared; that the megalop, which is transitional biramous pleopods present on somites 2–5 between planktonic and the benthic stages, (Fig. 5G–J), endopods with 3 or 4 terminal may be the main dispersal phase for cincinnuli, exopods with 7–8, 7–8, 8, 8 Chirostylus. long marginal setae on pleopods 2, 3, 4, 5 clark & ng (2008) noted that the respectively; uropods biramous, protopod pleopodal exopods in zoeas bear short with 1 plumose seta, endopod with 11–13 marginal setae in C. ortmanni, instead of long marginal setae and 1 short subterminal setal “buds” in C. sandyi. these setae are seta, exopod with 16–18 long marginal setae. barely discernible in C. stellaris. other than telson (Fig. 5K): elongate rectangular, the above differences, the first zoeas of these dorsal surface with 2 pairs of median setae. three species are morphologically similar and no significant distinguishing characters are found. Discussion In the chirostylidae, zoeal morphology Chirostylus stellaris has two zoeal and a described on the basis of the laboratory- megalopal phase before metamorphosing to reared materials is known for six species first juvenile crab stage. clark & ng (2008) in four genera: Chirostylus ortmann, 1892 presumed that Chirostylus sandyi (under (two species), Gastroptychus caullery, 1896 C. ortmanni) zoeas may be lecithotrophic, (one species), Uroptychus henderson, 1888 based on the morphological and behavioural (one species), and Eumunida smith, 1883 characters displayed by the first zoeas: (1) the (two species) (see Pike & Wear, 1969; de absence of setal armature from the mouthparts saint Laurent & Macpherson, 1990; ogawa (maxillule, maxilla, first maxilliped and & Matsuzaki, 1992; Guerao et al., 2006; second maxilliped); (2) yellow yolk stored clark & ng, 2008). First zoeas of Eumunida under the carapace; and (3) the hatched larvae capillata de saint Laurent & Macpherson, were not active and remained on the bottom 1990 and Eumunida annulosa de saint of the rearing container. they also believed Laurent & Macpherson, 1990 are quite that lecithotrophic larvae are associated with different from those of other chirostylids abbreviated zoeal development. these are (including Chirostylus) and close to galatheid fully supported by the present study showing zoeas in the “caridean-like” rather than “crab- that the C. stellaris zoea are apparently like” carapace, and the telson, in particular (de lecithotrophic and the megalop, with well saint Laurent & Macpherson, 1990; Guerao developed setal armature on the mouthparts, et al., 2006). the Chirostylus zoeas can be commenced feeding. It is conceivable that distinguished from the zoea of Gastroptychus the dispersal of decapod species with a novaezelandiae Baba, 1974 by the carapace lecithotrophic phase is localized and not that is armed with large, denticulate spinulose over a wide area, as suggested by clark & spines instead of two rows of slender lateral ng (2008). actually, C. stellaris has been spines and the telson that is armed with 6 or reported solely from the shallow reef slope (at 7 instead of 14 pairs of posterior marginal depths of 14.9–30 m) of the ryukyu Islands, processes. the Chirostylus zoeas are also southern Japan (osawa, 2007). Given that differentiated from the zoea of Uroptychus the entire zoeal phase of C. stellaris lasted tomentosus Baba, 1974 by the carapace with only two days in the laboratory, the dispersal more numerous spines (> 47 vs. 12 spines) during the zoeal phase of Chirostylus may be on the surface, abdominal somites 2–5 each restricted. however, this may not be always bearing, instead of lacking, a posterodorsal 6462 Y. FuJIta & P. F. cLarK

spine, and the absence of antennal exopod side. In the adult morphology, osawa (2007) (scaphocerite). noted that the armature of the male pereopod the anomuran seta (hair) as known 1 is one of the characters to distinguish C. in most of galatheoideans is absent in the stellaris from C. ortmanni. the merus of the chirostylid Chirostylus (see clark & ng, 2008 pereopod 1 of C. stellaris has a row of strong, and present study) as well as Gastroptychus closely-set spines on the proximal half of the and Uroptychus (see Pike & Wear, 1969). ventral surface. however, such spines are not these chirostylid larvae are presumed to be found in the megalop. lecithotrophic (Van Dover & Williams, 1991; the antenna of megalop described here clark & ng, 2008). the zoeas of the other for the first time for the chirostylidae is chirostylid Eumunida, which are possibly different from what has been expected. In planktotrophic, possess the anomuran hair the Galatheidae this appendage is similar (de saint Laurent & Macpherson, 1990; to that of the adult, being composed of Guerao et al., 2006). according to clark 4-segmented peduncle and a multi-segmented & ng (2008), the lack of the anomuran flagellum. however, the megalopal antenna hair in Chirostylus may be explained by of C. stellaris is simply 8-segmented, without the fact that lecithotrophic zoeas hatch clear distinction between the peduncle and in an advanced state such that the seta is flagellum. the structure of this appendage lost during the development. however, in may be explained by the fact that the antennal Galathea Fabricius, 1793, Munida Leach, peduncle in chirostylids consists of five 1820, Pleuroncodes stimpson, 1860, and segments. the distal fourth segment may Sadayoshia Baba, 1969, all belonging to correspond to the ultimate segment of the the Galatheidae, the anomuran hair persists antennal peduncle because it bears a distinct throughout all zoeal stages (huus, 1934; distal spine that is consistent in the adult of all Boyd, 1960; Fagetti & campodonico, 1971; known species of Chirostylus. the three distal roberts, 1973; Gore, 1979; christiansen segments that are relatively broad and not & anger, 1990; Fujita et al., 2001, 2003; clearly different in size from the fourth may Fujita & shokita, 2005). In addition, the correspond to the flagellum. In the adult, the zoea of Munidopsis serricornis (Lovén, flagellum is 4- or 5-segmented but somewhat 1852), which are lecithotrophic and show an narrower than the ultimate segment of the abbreviated development, have the anomuran peduncle (osawa, 2007). hair (samuelsen, 1972). although more extensive studies, including embryogenetic acknowledgements development, are required for the other species producing large eggs, it is likely that We are indebted to Prof. K. Baba, Dr. M. the presence or absence of the anomuran hair osawa and an anonymous reviewer for their corresponds to the galatheoidean systematics. valuable comments and suggestions on the the present study contributes to the first manuscript. thanks are extended to akane description of the megalop of a Chirostylus Ito of the Faculty of Medicine and university species. although the megalop of C. sterallis hospital, university of the ryukyus, for her is similar to the adult in the general shape, kind help in the animal collection. the following characters clearly distinguish it from the adult: (1) the gastric region of the Literature cited carapace is armed with a small submedian spine instead of being unarmed; (2) the Baba, K., 1969. Four new genera with their rostrum is unarmed, instead of bearing a representatives and six new species of median spine; (3) the merus and carpus of the Galatheidae in the collection of the the third maxilliped are unarmed, instead Zoological Laboratory, Kyushu university, of bearing a distolateral spine; and (4) the with redefinition of the genus Galathea. opposable margins of the pereopod 1 fingers ohMu, occasional Papers of Zoological are unarmed, instead of bearing a large Laboratory Faculty of agriculture, Kyushu subtriangular tooth on the distal third of each university, 2: 1–32. LarVaL DeVeLoPMent oF CHIROSTYLUS STELLARIS 6365

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