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The Complete Larval Development of Sadayoshia Edwardsii (Miers, 1884) (Decapoda: Anomura: Galatheidae) Described from Laboratory-Reared Material

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The Complete Larval Development of Sadayoshia Edwardsii (Miers, 1884) (Decapoda: Anomura: Galatheidae) Described from Laboratory-Reared Material Journal of Natural History, 2005; 39(12): 865-886 (T^\ Taylor & Francis \^ J Taylor & Francis Group The complete larval development of Sadayoshia edwardsii (Miers, 1884) (Decapoda: Anomura: Galatheidae) described from laboratory-reared material YOSHIHISA FUJITA & SHIGEMITSU SHOKITA Department of Chemistry, Biology and Marine Science, University of the Ryukyus, 1 Senbaru, Nishihara-cho, Okinawa 903-0213, Japan (Accepted 29 January 2004) Abstract The complete larval development of Sadayoshia edwardsii (Miers, 1884) is described and illustrated from laboratory-reared material. The development comprises four zoeal and one megalopal stages. Diagnostic zoeal characters of Sadayoshia are provided and these are compared with other galatheid genera for which the larval morphology is known. Zoeas of 5. edwardsii are readily distinguished from those of other galatheid species by the setation of the maxillular endopod together with the basis and endopod of the first maxilliped. The megalop of 5. edzvardsii has a flattened, triangular-shaped rostrum, which differs remarkably from that of the adult. Although the rostral shape resembles that of Galathea megalops, the armature of the lateral margins is different between megalops of the two genera. The present larval study suggests that Sadayoshia is more closely allied to Galathea than to Munida. Keywords: Crustacea, Decapoda, Anomura, Galatheidae, Sadayoshia edwardsii, larval development Introduction The family Galatheidae contains 28 genera (see Tirmizi and Javed 1980; Baba 1988, 1991, 1993; Baba and de Saint Laurent 1996; Baba and Wicksten 1997; Baba and Williams 1998; Macpherson 1998; Macpherson and Machordom 2000). However, the larvae have been described for only 12 species belonging to six genera as follows: Agononida Baba and de Saint Laurent, 1996, Cervimunida Benedict, 1902, Galathea Fabricius, 1793, Munida Leach, 1820, Munidopsis Whiteaves, 1874 and Pleuroncodes Stimpson, 1960 (see Gore 1979; Konishi and Saito 2000; Fujita et al. 2001, 2003). Recently, we obtained an ovigerous female of Sadayoshia edwardsii (Miers, 1884) from Okinawa Island of the Ryukyu Islands. This species is common on the coral reefs of the Ryukyu Islands, and inhabits dead coral rubble (Kamezaki et al. 1988, as 5. miyakei Baba, Correspondence: Yoshihisa Fujita, Department of Chemistry, Biology and Marine Science, University of the Ryukyus, 1 Senbaru, Nishihara-cho, Okinawa 903-0213, Japan ISSN 0022-2933 print/ISSN 1464-5262 online c 2005 Taylor & Francis Ltd DOI: 10.1080/00222930410001671264 866 Y. Fujita & S. Shokita 1969). Sadayoshia edwardsii had been referred to as S. miyakei Baba, 1969 or 5". acroporae Baba, 1972, until Baba (1990) considered that 5. edwardsii was the senior synonym of these two species. The purpose of this study is to describe and illustrate the complete larval development of Sadayoshia edwardsii, provide diagnostic larval characters for the genus and to compare it with the known larvae of other genera within the Galatheidae. Materials and Methods An ovigerous specimen of 5. edwardsii was collected in dead coral branches, at Cape Zanpa of Okinawa Is., the Ryukyu Islands, at a depth of 19.2 m, on 26 May 2001. The female was maintained in 5-litre circular plastic aquaria until hatching occurred. The hatched larvae were mass-cultured in circular plastic tanks containing 8 litres of filtered sea water. In order to determine the normal number of zoeal stages, 20 fourth-stage zoeas were cultured individually in 50-ml glass beakers. The salinity and water temperatures of the sea water were 35.0%o and 24.0-27.0°C, respectively. Approximately one-third of the water in the tanks and the beakers was changed daily. Food given throughout was newly hatched Anemia nauplii. The zoeal and megalopal specimens were stored in 50% ethylene glycol. Five larvae of each stage were measured and dissected for setal observations. Methods for mounting, drawing and measurement of carapace length (CL), total length (TL) and postorbital carapace length (PCL; measured only in the megalopal stage) followed those of Fujita et al. (2001). The long, plumose natatory setae on the exopods of maxillipeds and pleopodal exopods are drawn truncated. Terminology generally followed that of Gore (1979) and Ingle (1991). Usage of the terms of'zoeas (the plural)', 'megalop (the singular)', 'megalops (the plural)' and 'basial endite' of appendages followed Clark et al. (1998). The spent female and undissected larvae are deposited in the Coastal Branch of the Natural History Museum and Institute, Chiba, Japan (CMNH) under the following registration numbers: CMNH-ZC 1146 for the spent female, CMNH-ZC 1147-1150 for first to fourth zoeas and CMNH-ZC 1151 for megalops. Results Sadayoshia edwardsii passed through four zoeal stages and the megalop before attaining the first juvenile stage. The megalopal phase appeared 15-20 days after hatching, and the first juvenile stages 24-28 days after hatching. Minimum durations of the first to fourth zoeal stages were 3, 3, 3 and 5 days, respectively. Five of 20 fourth-stage zoeas that were reared solitarily metamorphosed into the megalop stage. The remaining 15 larvae died, all of them failing to metamorphose. The morphological characters of the larvae are described below. Sadayoshia edwardsii (Miers, 1884) (Figures 1-10) First zoea Size. CL 1.29-1.39 mm (mean 1.34 mm), TL 2.41-2.57 mm (mean 2.46 mm). Larval development of Sadayoshia edwardsii 867 Figure 1. Sadayoshia edwardsii (Miers, 1884 ). (A) First zoea, lateral; (B) second zoea, lateral; (C) third zoea, lateral; (D) fourth zoea, lateral; (E) megalop, carapace, abdomen, telson and pleopods, lateral; (F) megalop, anterior margin of carapace, lateral. Scale bars: 0.5 mm (A, B); 0.1 mm (C-F). 868 Y. Fujita & S. Shokita D, G, J Figure 2. Sadayoshia edwardsii (Miers, 1884 ). (A) First zoea, carapace, dorsal; (B) first zoea, rostrum, dorsal; (C) second zoea, carapace, dorsal; (D) second zoea, rostrum, dorsal; (E) third zoea, carapace, dorsal; (F) fourth zoea, carapace, dorsal; (G) megalop, entire animal, dorsal; (H) megalop, rostrum, dorsal; (I) megalop, lateral margin of carapace, dorsal; (J) thoracic sternites, ventral. Scale bars: 0.5 mm (A, B); 0.1 mm (C-F). Larval development of Sadayoshia edwardsii 869 ,1 \ Lt I Si1 ! ~"l 1 1 B 1 ii >n r > 1 i V / 1 -l"':ir \ v\ VVJi/ / ) X -^ - v -* \ A \ v x.% --^ xi * •' / //. \ .XT *t /. \^ V i_» \> \\ I •*--•—_^ \ v M -=«= \ \ -A,F - \ > f^ ^ -B ^"=- y X. C,E — f\ •i\ D,I / / G,J £•*-" G —H H Figure 3. Sadayoshia edwardsii (Miers, 1884). (A-E) Antennule: (A) First zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) megalop. (F-J) Antenna: (F) first zoea; (G) second zoea; (H) third zoea; (I) fourth zoea; (J) megalop, middle segments omitted. Scale bars: 0.1 mm. 870 Y. Fujita & S. Shokita <S -£ /^\ B D -r f^f -v / .\ \ •B •D -E E Figure 4. Sadayoshia edwardsii (Miers, 1884), mandible (r, right; 1, left). (A) first zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) left mandible of megalop; (F) same, palp. Scale bars: 0.1 mm. Larval development of Sadayoshta edwardsii 871 Figure 5. Sadayoshta edwardsii (Miers, 1884), maxillule. (A) First zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) megalop. Scale bars: 0.1 mm. 872 Y. Fujita & S. Shokita Figure 6. Sadayoshia edwardsii (Miers, 1884 ), maxilla. (A) First zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) megalop. Scale bars: 0.1 mm. Larval development of Sadayoshia edwardsii 873 -B ;/ )m TV / " t ft y%^ -D -E Figure 7. Sadayoshia edwardsii (Miers, 1884), first maxilliped (A) First zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) megalop. Scale bars: 0.1 mm. 874 Y. Fujita & S. Shokita Figure 8. Sadayoshia edwardsii (Miers, 1884), second maxilliped. (A) First zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) megalop. Scale bars: 0.1 mm. Larval development of Sadayoshia edwardsii 875 Figure 9. Sadayoshia edwardsii (Miers, 1884). (A-E) third maxilliped: (A) First zoea; (B) second zoea; (C) third zoea; (D) fourth zoea; (E) megalop (F-K) Pereiopods: (F) Second zoea; (G) third zoea; (H) fourth zoea; (I) megalop, first pereiopod; (J) megalop, second pereiopod, carpus, propodus and dactylus; (K) megalop, fifth pereopod, propodus and dactylus. Scale bars: 0.1 mm. 876 Y. Fujita & S. Shokita a1 nil wm\^I \\M| Figure 10. Sadayoshia edwardsii (Miers, 1884 ), abdomen and telson. (A) First zoea, abdomen and telson, dorsal; (B) first zoea, posterodorsal margins of fourth and fifth segments, dorsal; (C) first zoea, posterolateral margin of telson, dorsal; (D) first zoea, innermost plumose seta, dorsal; (E) second zoea, fourth and fifth abdominal segments and telson, dorsal; (F) third zoea, sixth abdominal segment, uropod and telson, dorsal; (G) fourth zoea, uropod and telson, dorsal; (H) fourth zoea, first pleopod, ventral; (I) megalop, tail fan, dorsal; Q-M) megalop, first to fourth pleopods, ventral, exopodal plumose setae of second to fourth pleopods truncated. Scale bars: 0.1 mm. Larval development of Sadayoshia edwardsii 877 Carapace (Figures 1A, 2A, B). Typical galatheid larval form; anterodorsal setae absent; anterolateral spine absent; pair of produced, acute spines present posteriorly; posterodorsal margin with 17 or 18 small teeth, posteroventral margin with 20-25 small teeth; rostrum elongated and spine-like, extending to level of anterior apex of the antennal scaphocerite, without distinct lateral teeth but covered with minute denticles; eyes sessile. Antennule (Figure 3A). Uniramous; elongated, with three aesthetascs and three setae terminally and one long plumose seta subterminally. Antenna (Figure 3F). Biramous; protopod with one robust serrated spine at distoventral end; endopod fused to protopod, terminally ending in acute spine and with a long plumose seta; scaphocerite with produced, acute spine on distolateral margin, mesial margin with nine plumose setae, ventral and dorsal surfaces with sparse minute denticles. Mandibles (Figure 4A). Asymmetrically dentate; incisor processes each with some strong teeth and few small teeth; molar processes strongly serrate or spinose; palp buds absent.
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