Spatial Distributions of Phyllosoma Larvae (Crustacea: Decapoda: Palinuridae and Scyllaridae) in Taiwanese Waters
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Journal of Oceanography, Vol. 57, pp. 535 to 548, 2001 Spatial Distributions of Phyllosoma Larvae (Crustacea: Decapoda: Palinuridae and Scyllaridae) in Taiwanese Waters 1 1 2 NARIAKI INOUE , HIDEO SEKIGUCHI * and SHINN-PYNG YEH 1Faculty of Bioresources, Mie University, 1515 Kamihama-cho, Tsu, Mie 514-8507, Japan 2Department of Aquaculture, National Pingtung University of Science and Technology, Nei-Pu Hsiang, Pingtung, Taiwan 91207 (Received 20 November 2000; in revised form 19 March 2001; accepted 20 March 2001) Distributions and taxonomy of phyllosoma larvae were examined in Taiwanese wa- Keywords: ters, based on ichthyoplankton samples collected from May 1990 to July 1995. ⋅ Phyllosoma, Phyllosoma larvae belonged to the two families Scyllaridae and Palinuridae repre- ⋅ Palinuridae, ⋅ senting 6 genera and 13 species. Of the collected phyllosoma larvae, those of Scyllarus Scyllaridae, ⋅ and Panulirus species were most abundant, forming 90% of total numbers. Early spatial distribu- tions, stage Scyllarus and Panulirus phyllosoma larvae were abundant in Taiwanese waters. ⋅ Taiwanese waters. Middle to late stages (except the final stage) of Panulirus phyllosoma larvae were absent from the waters throughout the year, while those of Scyllarus phyllosoma lar- vae were collected in the waters. This suggests that all stages of Scyllarus phyllosoma larvae may be retained in the northern part of the waters around northern Taiwan while middle to late stages of Panulirus phyllosoma larvae may be flushed out from the waters, the sub-final and final stages then possibly returning to the waters. An anticlockwise eddy existed in the waters off northeastern Taiwan, which may be closely related to flushing out and returning of Panulirus phyllosoma larvae through a much longer planktonic period. 1. Introduction et al., 1988). However, few studies have been conducted Phyllosoma larvae, confined to palinurid and on the population dynamics of the lobsters in Taiwan, scyllarid lobsters, are plankton with transparent, leaf-like particularly on the larval recruitment processes by which forms (Phillips and McWilliam, 1986). The palinurid the populations may form and be maintained in Taiwan- phyllosoma larvae have a long life span of over 6 months ese waters. (Booth and Phillips, 1994), while those of scyllarids have Phyllosoma samples used in the present study were a comparatively short life span of 2 or 3 months collected during the ichthyoplankton surveys conducted (Robertson, 1968; Ito and Lucas, 1990). Seventeen by the Kuroshio Edge Exchange Processes Project palinurid and 16 scyllarid species have been reported in (KEEP). This multidisciplinary study of the Taiwan (Chan and Yu, 1993, 1995; Poupin, 1994; Chan biogeochemical cycle of carbon and associated elements and Chu, 1996). Of the above lobsters, 21 species have in the East China Sea revealed many of the sea’s been described for phyllosoma larvae (Sekiguchi, 1986a). hydrographic features since 1989 (Chuang et al., 1995). Three Panulirus species (P. japonicus, P. longipes For example, Tang et al. (1999) reported that an anticlock- and P. stimpsoni) are of economic importance in Taiwan wise eddy with a diameter of <70 km exists in the waters (Chan and Yu, 1993). Phyllosoma larvae of the first two off northeastern Taiwan (Fig. 1). have been described (Sekiguchi, 1986a). Previous stud- The goal of our future research is to clarify the lar- ies of the lobsters in Taiwan dealt with the aspects of ecol- val recruitment processes of commercially important spe- ogy (Deng, 1963), aquaculture (Jong and Lin, 1981), tax- cies of palinurid and scyllarid lobsters in Taiwan, par- onomy (Chan and Yu, 1993) and fishery biology (Huang ticularly of Panulirus species, in relation to the hydrographic features of Taiwanese waters. To our knowledges, however, phyllosoma and puerulus (or nisto) * Corresponding author. E-mail: [email protected] larvae in Taiwanese waters have not yet been dealt with, Copyright © The Oceanographic Society of Japan. except by Sekiguchi and Saisho (1994). So before begin- 535 Fig. 2. Map of the study area and location of sampling sta- tions. a, b: Sampling stations for surface and oblique tow- Fig. 1. Mean circulation pattern at depths of 16, 104 and 200 ing, respectively; solid circles: sampling stations in the m in August 1994 (modified from Tang et al., 1999). The waters around northern Taiwan; open circles: sampling sta- bathymetry is shown with thin depth contours at intervals tions in the waters around southwestern Taiwan; dotted lines: of 200 m from 200 to 1000 m. isopleths of 200 m depth. ning work on the larval recruitment processes, it is first ern Taiwan from May 1990 to July 1993, covering all of all necessary to identify to the species level phyllosoma months (Table 1, Fig. 2a). The second runs involved a and puerulus (or nisto) larvae contained in total of 195 oblique towings at different depths (15–430 ichthyoplankton samples collected in Taiwanese waters, m) from the surface with a ship speed of 2–4 kt for 5 and then to examine their spatial distribution. min., samples being collected at 41 and 11 stations in the northern part of the waters around northern Taiwan and 2. Materials and Methods southwestern Taiwan, respectively, from May 1990 to Ichthyoplankton samples were collected using two May 1995 (Table 2, Fig. 2b). All samples were fixed in types of ring nets (diameter: 1.3 m and 1.5 m, length: 4.5 5% formalin seawater immediately after sampling and m, mesh: 0.5 mm). Samples were collected by surface preserved in 70% ethanol. and oblique towing during the day or night. The first sam- Phyllosoma larvae were sorted from the samples in pling runs involved a total of 89 surface towings with a the laboratory. Stages of these phyllosoma larvae were ship speed of 2 kt for 10 min., samples being collected at determined according to Braine et al. (1979) and Phillips 57 stations in the northern part of the waters around north- and McWilliam (1986). The late stage phyllosoma larvae 536 N. Inoue et al. Table 1. Data for sampling phyllosoma larvae by surface towing. Numerals in the upper table: number of towings; horizontal line: no data. Numerals in the lower table: number of collected phyllosoma; horizontal line: no data. were then identified to the species level according to of the cephalon, and widths of the cephalic and thorax Sekiguchi (1986a) and Inoue et al. (2000). Based on were measured to the nearest 0.1 mm. All figures were morphological features of the late stage phyllosoma lar- drawn with the aid of a drawing tube. vae identified, we identified the middle stage ones, but The individual number of phyllosoma larvae col- the early stage ones were not identified to the species lected per net haul was very low, as indicated in Tables 1 level. Body length (BL) from the anterior margin of the and 2. Accordingly, we examined the spatial distributions cephalon (forebody) to the posterior end of the hindbody, of phyllosoma larvae of different species by pooling the cephalic length from the anterior to the posterior margins data for the phyllosoma larvae collected by the two types Spatial Distributions of Phyllosoma Larvae off Northern Taiwan 537 Table 2. Data for sampling phyllosoma larvae by oblique towing. Numerals in the upper table: number of towings; horizontal line: no data. Numerals in the lower table: number of collected phyllosoma; horizontal line: no data. 538 N. Inoue et al. of nets, and the phyllosoma larvae were counted as indi- vidual numbers per tow for both types of net. Spatial dis- tributions of phyllosoma larvae of the two genera Scyllarus and Panulirus were examined on the basis of samples collected by surface towing. Identification and taxonomic descriptions were based on samples collected by surface and oblique towings. 3. Results A total of 312 phyllosoma larvae were collected, of which 123 and 189 specimens were collected by surface and oblique towing, respectively (Tables 1 and 2). These phyllosoma larvae belonged to the two families Scyllaridae and Palinuridae, representing 6 genera and 13 species. Of the family Scyllaridae, 3 genera and 10 species were identified (Tables 1 and 2): 7 species of the genus Scyllarus (S. bicuspidatus, S. cultrifer, S. kitanoviriosus, S. martensii, Scyllarus sp. c, Scyllarus sp. d, Scyllarus sp. e), 2 species of the genus Ibacus (I. novemdentatus, I. ciliatus) and 1 species of the genus Scyllarides (Scyllarides sp.). Of the family Palinuridae, 3 genera and 3 species were identified (Tables 1 and 2): Panulirus longipes or P. japonicus, Linuparus sordidus and Puerulus angulatus. The phyllosoma larvae in stages I to IV of the Scyllarus and in stages I to VI of the Panulirus were not identified to the species level. 3.1 Spatial distributions All stages of the Scyllarus phyllosoma larvae were collected by surface towing (Table 1). The phyllosoma larvae in the stages I to IV, which are probably identical Fig. 3. Distributions and abundance of phyllosoma larvae of with those of S. kitanoviriosus, and also in the stages V the Scyllarus found by surface towing. a: Early to middle to VIII stages of S. kitanoviriosus, were mainly found in stage phyllosoma larvae of the Scyllarus; numerals with the northern part of the waters around northern Taiwan open squares and triangles: individual number of stages I (Figs. 3a and b). No changes of spatial distributions ac- to II and stages III to IV, respectively. b: Middle to late stage phyllosoma larvae of Scyllarus kitanoviriosus; numer- cording to larval stages were detected for Scyllarus als with open squares and triangles: individual number of phyllosoma larvae. This also holds for those of the other stages V to VI and stages VII to VIII, respectively. Solid Scyllarus species. circles: sampling stations. Dotted lines: isopleths of 200 m In contrast to Scyllarus phyllosoma larvae, no stage depth. VII and VIII phyllosoma larvae were collected of Panulirus (Table 1).