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Larval Development of the Chinese Mitten Crab Eriocheir Sinensis H

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Larval Development of the Chinese Mitten Crab Eriocheir Sinensis H HELGOL,~NDER MEERESUNTERSUCHUNGEN Helgolilnder Meeresunters. 50, 223-252 (1996) Larval development of the Chinese mitten crab Eriocheir sinensis H. Milne-Edwards (Decapoda: Grapsidae) reared in the laboratory M. Montfi 1, K. Anger 2 & C. de Bakker 3 1Fundag~o Universidade do Rio Grande, Base Oceanogr~fica Atl~ntica; 96.200 Rio Grande, RS, Brazil 2Biologische Anstalt Helgoland, Meeresstation; 27483 Helgoland, Germany 3Centro de Biologia Marinha, Universidade Federal do Paran~; 80. 000 Curitiba, PR, Brazil ABSTRACT: Larvae of the Chinese mitten crab Eriocheir sinensis were reared in the laboratory from the time of hatching and through metamorphosis. Development normally consists of a Prezoea, 5 Zoea stages, and a Megalopa. Occasionally, an additional (stage VI) Zoea and, in one case, an additional Megalopa (transitional to the first crab stage) were observed. Detailed morphological descriptions of all larval and the first two juvenile instars are given, and larval morphology is compared with that of two closely related species, Eriocheir japonicus and Eriocheir rectus, descriptions of which have recently become available. The zoeal stages of these species can be distinguished by their different number of aesthetascs and setae on the antennules, and different setation of maxillipeds 1 and 2. The Megalopa shows differences in the shape of the rostrum and again in the morphology of the antennule. These and other morphological differences (mainly in setation and spinulation of the zoeal carapace) between E. sinensis and E. japonicus larvae suggest that they may be very closely related but separate species; this contradicts a recent study of adult morphometrics and molecular genetics (Li et al., 1993), suggesting that they are only varieties of a single species. INTRODUCTION The mitten crab Eriocheir sinensis was introduced early this century from China to German rivers, from where it spread over great parts of Europe (Panning, 1933). While the juvenile and adult stages have been investigated in much detail (review of old hterature: Panning, 1938; for recent references see Bianchini & Gilles, 1990; Lee & Yamazaki, 1990), very httle is known about the larvae of E. sinensis. Rough and incomplete descriptions of their morphology were given by Schnakenbeck (1933) and Panning (1939). The latter author suggested the existence of 4 zoeal stages and a Megalopa (Panning, 1936a). Panning (1936b) and Hinrichs & Grell (1937) concluded from field observations that the larvae perform ontogenetic horizontal migrations during their development in estuarine and coastal regions. First, only partially successful rearing experiments were carried out by Buhk (1938). Anger (1991) reared the larvae of E. sinensis under controlled conditions from the time of hatching and through metamorphosis, studying effects of temperature and salinity on ~ Biologische Anstalt Helgoland, Hamburg 224 M. Montfi, K. Anger & C. de Bakker survival and development. In the present paper, the morphology of all larval and the first two juvenile stages is described and compared with that of closely related species, Eriocheir japonicus (Kim & Hwang, 1990) and Eriocheir rectus (Shy & Yu, 1992); however, according to a recent taxonomical revision of adult material (Chan et al., 1995), the larvae described in the latter paper did not belong to E. rectus, but to a new species, Eriocheir formosa. The comparison of larval morphology in E. sinensis and E. japonicus became particularly interesting since Li et al. (1993) concluded from morphometric and biochemical evidence that these two species, which co-exist in some Chinese rivers, should actually be only varieties of a single species. MATERIAL AND METHODS Ovigerous mitten crabs were dredged in early 1989 from the outer Elbe estuary (North Sea) and transported to the Helgoland marine biological station, where they were maintained at constant 12 ~ and 20 %o S until larvae hatched (in May). Larvae were reared both individually in vials (30 cm 3) and communally in bowls (400 cm 3) at 25 different combinations of temperature (6, 9, 12, 15, and 18 ~ and salinity (10, 15, 20, 25, and 32 7~). Water and food (freshly hatched San Francisco Bay Brand TM Artemia spec. nauplii) were either changed daily (at -> 12 ~ or every second day (at ~- 9 ~ Further details of rearing conditions are given in a previous paper by Anger (1991), where effects of temperature and salinity on rates of larval development and survival are described. The exuviae and dead larvae from these experiments were used in the present study, in addition to material obtained from cultivation experiments that were carried out exclu- sively for morphological studies. Specimens of larvae and exuviae were fixed in 4 % seawater-formaldehyde. Later, at least 20 individuals of each larval stage (where available) were measured and dissected under Wild stereo microscopes. Material was also available for the study of exceptionally occurring additional stages (Zoea VI, Megalopa II). Drawings were made with the aid of a camera lucida attached to a Leitz compound microscope. The drawings of whole larvae and single appendages thereof are not grouped by stage but by character (e.g. total lateral view of larva, antenna, telson, etc.), so that each plate shows the development of one character. The following abbreviations are used: CL = carapace length from orbit to posterior carapace border; CW = carapace width measured at the widest part of carapace; DS-RS = distance from tip of dorsal spine (DS) to tip of rostral spine (RS); LS = lateral spine; P1-P5 = pereiopods 1-5; Pll-Pl5 = pleopods 1-5; TL = total length from tip of the rostrum to the posterior border of the telson (only for Megalopa). RESULTS Larval development of Eriocheir sinensis consists normally of a brief, non-feeding Prezoea, five Zoea-stages, and a Megalopa. At 15 ~ m combination with low salinity (15 %0 S), survival rate was poor (Anger, 1991), and five individuals passed a sixth zoeal stage before reaching the Megalopa. In one case (at 5 700 S, after stepwise reduction of salinity) an additional Megalopa was observed; however, this individual died before metamorphosis. Since such unfavourable conditions also occur in the field, namely in Larval development of Er~ocheir sinensis 225 estuarine environments, additional larval stages may exist also there and are thus described here as well. No other effects of rearing conditions on larval morphology were observed. Prezoea Size: TL 1.23-1.48 ram; CL 0.48-0.52 mm. Carapace (Fig. IA): Cephalothorax globose, with rostrum, DS and LS developing; DS pointed toward rostrum. Antennule: Three aesthetascs emerging, 2 spines. Antenna: Protopod and exopod emerging. Mandible, Max/llula, Maxilla: Nonfunctional, all embedded in mass of ceils. Maxillipeds: Biramous, covered with cuticular sheaths. Maxillipeds 1 and 2 with 5 seg- ments. Abdomen: Five somites. Somites 2, 3 and 4 with a knob on each medial lateral surface. Telson (Fig. 14A): Rectangular, furca and setae emerging from Cuticular sheath. Zoea I Size: CL 0.64-0.67 mm; DS-RS 1.25-1.27 mm. Carapace (Fig. 1B): Cepha]othorax punctuated, DS well developed, covered with small spines, longer than RS and lateral spine; RS covered with small spines; LS with small spines on surface; posterolateral region with 6-9 spines on each side. Eyes un- stalked. Antennule (Fig. 3A): Conical and slender with 3 aesthetascs and 2 setae of different size. Antenna (Fig. 4A): Long protopod with short spines along margins; exopod small with terminal and tateral spines. Mandible (Fig. 5A): Symmetrical, molar and incisive parts well defined; incisive part with 2 ventral teeth. Maxillula (Pig 6A): Endopod 2-segmented; proximal article with 1 long seta; distal article with 4 terminal and 1 subterminal plumose setae; basal endite with 4 spines and 1 seta; coxal endite with 5 setose spines. Maxilla (Fig. 8A): Endopod bilobed with 2, 2 terminal setae; basal endite bilobed with 4, 4 sp/setae; coxal endite bilobed with 2, 3 setae; scaphognathite with 3 plumose, 2 smooth setae plus 1 plumose projection. Maxilhped 1 (Fig. t0A): Basis with 6 ventral setaei exopod with 4 natatory setae; endopod 5-segmented with 1, 1, 2, 2, 5 setae. Maxilliped 2 (Fig. 12A): Basis with 3 sparsely plumose setae; exopod with 4 natatory setae; endopod 3-segmented with 0, 1, 5 sparsely plumose setae. Abdomen (Fig. 14B): Five somites; somites 2, 3 and 4 with small hook on each lateral surface; somite 5 with posterolateral spines overlapping the telson. Somites 3-5 with 2 minute dorsal setae. Telson (Fig. 14B): 3 pairs of setae on posterior margins, with spines and minute setae; one dorsal marginal series of spinules and one on internal margin of furca. 226 M. Montfi, K. Anger & C. de Bakker Zoea II Size: CL 0.68-0.71 mm; DS-RS 1.56-1.60 mm. Carapace (Fig. 1C): As in Zoea I, with series of 5 plumose setae on the upper postero- lateral region, marginal part with alternate series of spines and minute setae; small spines on DS and LS reduced as compared to Zoea I. Eyes stalked. Antennule (Fig. 3B): As in Zoea I, but with 3 aesthetascs, 1 short, 1 minute setae. Antenna (Fig. 4B): As in Zoea I, but with exopod more developed. Mandible (Fig. 5B): As in Zoea I, molar part well developed with little teeth, incisive part more developed. Maxillula (Fig. 6B): Endopod 2-segmented, with 1 smooth and 5 sparsely plumose setae; basal endite with 7 serrate spines; coxal endite with 5 serrate spines; protopod with 1 plumose seta. Maxilla (Fig. 8B): Endopod bilobed with 2, 2 sparsely plumose setae; basal endite bilobed with 3 stout sparsely plumose, 1 fine and 5 stout plumose setae; coxal endite with 3 and 4-5 stout sparsely plumose setae; scaphognathite with 8 plumose setae. Maxilliped 1 (Fig. 10B): Basis with sparsely plumose setae arranged in groups of 2 and 3; exopod with 6 plumose setae; endopod 5-segmented with 2, 1, 1, 2, 5 setae.
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