REDESCRIPTIONS AND NEW RECORDS OF ISOPODA BOPYRIDAE (CRUSTACEA) FROM THAILAND

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REDESCRIPTIONS AND NEW RECORDS OF ISOPODA BOPYRIDAE (CRUSTACEA) FROM THAILAND ABSTRACT. - Three species ofbopyrid isopods, parasites of decapod crustaceans, previously known from elsewhere in the western Pacific, are reported for the first time from Thailand. In order to overcome deficiencies in the existing descriptions, all species are figured in detail and redescribed or partly to completely diagnosed. Probopyrus giardi Weber infests Macrobrachium sintangense (de Man), a new host record; Dactylokepon richardsonae Stebbing infests Portunus tuherculosus (A. Milne Edwards), a new host record; and Eophrixus shojii Shiino infests Alpheus sp. The total number of bopyrids how known from Thailand is 34. After the preparation of a report on the bopyrid isopods of Thailand (Markham, 1985b), representatives ofthree more species belonging to that fauna came to light. Though this material contains no undescribed species, each sample constitutes a noteworthy extension of the known range of its species, all of the species needed further description, and two are from species of decapods not previously reported as bopyrid hosts. These discoveries raise the documented bopyrid fauna of Thailand (still mostly from Phuket and environs) to 34 species. Probopyrus giardi Weber, 1892 (Fig. 1) Probopyrus Giardi Weber, 1892: 535, 557,561,571, Text-fig. 2, PI. XXX,Fig. 1 [Type-locality,near Kaja Tanam, Sumatra, Indonesia; infesting Palaemon placidus de Man [= Macrobrachium placidum (de Man)]; Bonnier, 1900: 48, 62, 159, 171,222,342,459,460, PI. 32, Figs. 1-6. Probopyrus giardi - Stebbing, 1893: 416; Chopra, 1923: 507,508,509,510,512; Nierstrasz & Brender a Brandis, 1923: 95, Lemos de Castro & Brasil Lima, 1974: 210.

Fig. I. Prohopyrus giardi Weber, 1892. A-L, female; M-R, male. A, dorsal view; B, ventral view; C, right antennae; D, right side of barbula; E, right maxilliped; F, palp of same; G, plectron of same; H, right oostegite I in external view; I, same in internal view; J, right oostegite 5 in external view; K, right pereopod I; L, right pereopod 7; M, dorsal view; N, ventral view; P, left pereopod I; Q, left pereopod 7; R, end of pleon in ventral view. Scale: 2.2 mm for A, B, H-J; 1.1 mm for D, E; 0.4 mm for F, G, K-N; 0.2 mm for C,O-R.

Material examined. - Infesting Macrobrachium sintangense (de Man), L. B. Holthuis, det. of host. Lumpun Lake, Phatalung Province, S. Thailand, Phaibul Naiyanetr coli., 22.v.l979: I male, 1 female, RMNH Crust 1. Diagnosis. - Female (Fig. IA-L): Head about '/4 of body length, not extending beyond pereonal margin (Fig. 1C); distal articles of antennae greatly reduced (Fig. I C), barbula broad and short (Fig. ld). First oostegite falcately pointed (Fig. I H, I). Pleopods proportionately broad and final one uncleft; uropods obscure (Fig. la, B). Male (Fig. IM-R): Head non-extended (Fig. 1M); second antennae with only two articles (Fig. 10). Pereon broad relative to length, pereomeres not separated. Pleon short (Fig. 1M, N); only four pairs of incompletely tuberculiform pleopods (Fig. IR). Remarks. - Weber's (1892) original description of Probopyrus giardi was extremely brief, and his drawings, of the female with the male attached and the pleon of the male, while good, were too few to permit a reliable identification. Bonnier (1900), evidently using Weber's typespecimens (though he does not explicitly say so), presented little more verbal description but adequate illustrations. On the basis of Bonnier's illustrations, the present material is assignable to P. giardi and thus becomes only the second record for the species. The type female of P. giardi, as depicted by Bonnier (1900), differs from the present material in having a more extended non-articulating maxilliped palp, and the outer process of its barbula laterally dentate; the type male has only five distinct pleomeres. The fauna of southeastern Asia contains several species of Probopyrus, all parasites of palaemonids, especially Macrobrachium spp., and frequently difficult to distinguish (Markham, 1985a). The diagnosis above may help solve this problem. This is the first reported record of infestation of Macrobrachium bopyrid. sintangense (de Man) by a Dactylokepon richardsonae Stebbing, 1910 (Fig. 2) Dactylokepon richardsonae Stebbing, 1910: 85, 113, PI. 11C [Type-locality Seychelle Isla~ds; infesting Trapezicepon cymodoce (Herbst)]; Nierstrasz & Brender a Brandis, 1923: 83; Shiino, 1942: 444,447; Markham, 1975: 61, 64, 66, Table 1. Dactylocepon richardsonae - Bourdon, 1967: 122; 1980: 243; 1983: 855-857,859, Fig. 7 [MarseguIsland, Moluccas; infesting T. cymodoce].

Material examined. - Infesting Portunus tuberculosus (A. Milne Edwards), L. B. Holthuis det. of host. Naklua and Si Racha, Bangkok Bight, ca. 100 km SE of Bangkok, Thailand. A. C. J. Burgers & L. B. Holthuis colis., 14.iii.1985: I male, I female, RMNH Crust I. Redescription. - Female (Fig. 2A-L). Length 8.71 mm, maximal width 5.57 mm, head length 1.93 mm, pleonallength 2.36 mm. Body axis distortion 35. Body nowhere abruptly narrowing (Fig. 3A, B). Head about as broad as long, extending somewhat beyond pereonal margin, faintly bilobate. No eyes visible. Antennae (Fig. 2C) of two and four articles respectively, first antenna much reduced. Barbula (Fig. 2D) with two rather broad truncate projections on each side, each bearing some irregular teeth. Maxilliped (Fig. 2E) nearly triangular, its anterior article much longer; prominent non-articulating falcate palp arising just lateral to anteromedial comer; plectron only slightly developed. Pereon broadest across third pereomere. Pereomeres indistinctly defined. No midventral tubercles. Brood pouch not completely covered by oostegites. Oostegite I (Fig. 2F, G) bluntly rounded anteriorly, produced into sharp extended point posterolaterally; internal ridge deeply digitate. Pereopods (Fig. 2H-K) markedly increasing in length posteriorly; each with all articles distinct; first one (Fig. 2H) with prominent basal carina and (Fig. 21) serrated ridges on end of propodus receiving dactylus; pereopods 4-7 with coxae produced into extended knobs; pereopod 7 (Fig. 2J) elongate, except propodus (Fig. 2K) small in proportion to other articles and minutely setose distally. Pleon (Fig. 2L) completely covered laterally by appendages. Lateral plates and uniramous pleopods extended and deeply digitate, anterior lateral plates extending far forward. Uniramous uropods similar in size and structure to pleopods. Male (Fig. 2M-R). Length 3.13 mm, maximal width 0.89 mm, head length 0.33 mm, pleonal length 1.13 mm. Body broadest across pereomere 4 and tapered evenly both ways from there. Head medially fused with pereomere I, but all other segments distinct (Fig. 2M, N). Head oval, broader than long. Small but conspicuous dark eyespots near posterior edge. Antennae (Fig. 20) of three and five articles respectively, both distally setose. Pereomeres deeply separated by anterolateral notches, bearing scattered pigment spots dorsally. Small midventral tubercle on each pereomere. Pereopods (Fig. 2P, Q) all of about same size, but dactyli of first pair very large and pointed, dactyli of second pair shorter, equal-sized dactyli of pairs 4-7 smaller yet. Pleon elongate, extending to blunt point. Pleomeres less separated than pereomeres, some dorsally spotted. Five pairs of flaplike pleopods extending somewhat beyond posterior margins of all pleomeres. Final pleomere (Fig. 2R) truncate, lacking uropods but with tufts of setae on posterolateral comers. Remarks. - The original description of Dactylokepon richardsonae by Stebbing (1910) was so brief that it became difficult to be certain of the identification of subsequent specimens. The shape of the maxilliped and its palp, however, are especially diagnostic. Bourdon's (1983)

Fig. 2. Dactylokepon richardsonae Stebbing, 1910. A-L, female; M-R, male. A, dorsal view; B, ventral view; C, right antennae; D, right side ofbarbula; E, right maxilliped; F, right oostegite 1 in external view; G, same in internal view; H, right pereopod 1; I,distal region of same; J, right pereopod 7; K, distal region of same; L, pleon in ventral view; M, dorsal view; N, ventral view; 0, left antennae; P, left pereopod 1; Q, left pereopod 6; R, final pleomere in ventral view. Scale: 4.5 mm for A, B; 2.2 mm for E-G, L; 1.1 mm for C, D; 1.0 mm for H, J, M, N; 0.4 mm for O-Q; 0.2 mm for I, K, R.

description and illustrations, though not quite complete, are clearly of the same species as here considered. This is only the second account ofthe male; that reported by Bourdon (1983) differs from the present one in being proportionately narrower, having a much longerpleon, and having the head separated from the pereon. Both males have the same distinctive peculiarly shaped anterior pereopods and midventral tubercles on all pereomeres. The host is a portunid, while previous ones were xanthids. It thus might have been expected that the present material would belong to D. catoptri Stebbing (1910), whose type infested another portunid, Catoptrus nitidusa. Milne Edwards at Amirante Island in the western Indian Ocean. D. catoptri was also inadequately described and illustrated and is known only from its type female, but the shape of its maxilliped palp is very different from that of the present female. Eophrixus shojii Shiino, 1941 (Fig. 3) Eophrixus shojii Shiino, 1941: 154-156, Figs. 1, 2 [Type-locality, Tomioka Bay, Amakusa, Japan; infesting Alpheus japonicus infesting A. rapax de Haan]; Markham, Ortmann]; Shiino, 1958: 71 [Momotori, Ise Bay, Mie Prefecture, Japan; 1982: 375-377, 385, Figs. 28, 29 [Tolo Harbour, Hong Kong; infesting A. lobidens de Haan?]; Page, 1985: 185,205,206-208 [New Plymouth, near Foxton and Foveaux Strait, New Zealand; infesting A. socialis Heller]; Markham, 1990: 555, 563 [Mirs Bay, Hong Kong; infesting A. dispar Randal]. Anisarthrus shojii - Codreanu & Codreanu, 1956a: 119; Codreanu & Codreanu, 1956b: 577; Codreanu, 1961: Fig. 1. Eophryxus [sic] shojii - Shiino, 1972: 9. Material examined. - Infesting Alpheus sp., L. B. Holthuis, del. of host. Ko Libong (= Pulau Talibong) southwest of Ban Kantang, Phuket Province, Thailand, C. Swensen coli., 24.x.1984: 1 male, 1 female, RMNH Crust I. Remarks. - The present material, especially the female (Fig. 3A-J), matches the typespecimens (Shiino, 1941) more closely than those from Hong Kong (Markham, 1982, 1990). Unlike the Hong Kong females, this one has all seven pereopods on each side fully developed, and it has broad pleonal appendages. The male (Fig. 3K-0), has the head fused with the first pereomere, as do the type male and that from New Zealand (Page, 1985), but, unlike those of other males, its final pereomere is separate from the pleon; its pleon, though of peculiar shape, lacks all indications of segmentation. The present host, like all the others, is a species of Alpheus, whose damaged condition prevented specific identification. Acknowledgements. - Lipke B. Holthuis of the Rijksmuseum van Natuurlijke Historie Leiden (designated RMNH) made the material available for study, identified the hosts and provided valuable curatorial services. Mrs W. A. Markham provided facilities at the Arch Cape Marine Laboratory, of which this is contribution number 23.

Fig. 3. Eophrixus shojii Shiino, 1941. A-J, female; K-O, male. A, dorsal view; B, ventral view; C, head and pereomeres 1, 2 in dorsal view; D, barbula and pereomere 2 in ventral view; E, left maxilliped in external view; F, right pereopod 3; G, left pereopods; H, left oostegite 1 in external view; I, same in internal view; J, end of pleon in dorsal view; K, dorsal view; L, left antennae; M, left pereopod 1; N, left pereopod 7; 0,end of pie on in ventral view. Scale: 2.2 mm for A, B, H, I; 1.1 mm forc-g, J; O.4mm fork; 0.2 mm for M-O; 0.1 mm for L.

Bonnier, J., 1900. Contribution II l' etude des epicarides. Les Bopyridae. Trav. Stat. Zool. Wimereux, 8: 1-476. Bourdon, R., 1980.Aporobopyrus dollfusi n. sp. (Crustacea, Epicaridea, Bopyridae) parasite de porcellanes de la Mer Rouge. Bull. Mus. natn. Hist. Nat., Paris, (4) 2 (Sect. A) (no. 1): 237-244. Bourdon, R., 1983. Expedition Rumphius II (1975). Crustaces parasites, commensaux, etc. (Th. Monod d.). VIII. Crustaces isopods (3e partie: Epicarides Bopyridae). Bull. Mus. natn. Hist., Paris, (4e) S(A3): 845-869. Codreanu, M. & R. Codreanu, 1956a. Sur I'Anisarthrus pelseneeri, Epicaride parasite abdominal de la crevettea thanas nitescens; sa presence dans la Mer Noire et ladispersion du genre Anisarthrus. Bull. bioi. Fr. Belg., 90(2): 111-121. Codreanu, M. & R. Codreanu, 1956b. Un epicaride abdominal nou pentru Marea Neagra, Anisarthrus pelseneeri Giard 1907, parazit pe creveta Athanas nitescens Leach. Revizuirea si raspinderea genului Anisarthrus. Bul. Stiint. Sec. Bioi. Stiinte Agric., 8: 567-584. Codreanu, R., 1961. Crustacei paraziti cu affinitati indo-pacifice in Marea Neagra. Hidrobiologia, Acad. Rep. Pop. Romine, 3: 133-146. Lemos de Castro, A. & I. M. Brasil Lima, 1974. Crustaceos Isopodos Epicarideos do Brasil. IX. Genero Probopyrus Giard e Bonnier. Rev. Brasil. Bioi., 34: 209-217. Markham, J. c., 1975. New records of two species of parasitic isopods of the bopyrid subfamily Ioninae in the western Atlantic. Crustaceana, 29: 55-67. Markham, J. C., 1982. Bopyrid isopods parasitic on decapod crustaceans in Hong Kong and Southern China. In: Proc. First Internatl. Mar. Bioi. Workshop: The Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong. 1980, 1: 325-391. Eds. B. S. Morton & C. K. Tseng, Hong Kong University Press, Hong Kong. Markham, J. C., 1985a. Redescription and systematic remarks on Probopyrus buitendijki (Horst, 1910) (Isopoda, Bopyridae) parasitic on Macrobrachium rosenbergii (de Man) in southeast Asia. J. Crust. Bioi., S: 673-682. Markham, J. c., 1990. Furthernotes on the Isopoda Bopyridae of Hong Kong. In: Proc. Second Internatl. Mar. Bioi. Workshop: The Marine Flora and Fauna of Hong Kong and Southern China. Hong Kong. 1986. 2: 555-568. Ed. B. S. Morton, Hong Kong University Press, Hong Kong Nierstrasz, H. F. & G. A. Brender II Brandis, 1923. Die Isopoden der Siboga-Expedition. II. Isopoda Genuina. I. Epicaridea. Siboga-Expeditie Monogr., 32b: 57-121. Page, R. D. M., 1985. Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea). N. Z. Journ. Zool., 12: 185-212. Shiino,S.M., 1941. Further notes on bopyrids from Kyusyu and Ryukyu. Annot. Zool. Japan., 20(3): 154-158. Shiino, S. M., 1942. Bopyrids from the South Sea Islands with description of a hyperparasitic cryptoniscid. falao Tmp. BioI. Sta. Studies, 2: 437-458.

Shiino, S. M., 1972. The Epicaridea (list of species) from Japan. Kansai Shizenkagaku, 24: 7-10. (In Japanese) Stebbing, T. R. R., 1893. A history of Crustacea: Recent Malacostraca. London: Kegan Paul, Trench, Triibner & Co. xvii+466 pp. The international scientific series. Stebbing, T. R. R., 1910. Isopoda from the Indian Ocean and British East Africa. Reports of the Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership ofmr J. Stanley Gardiner, M.A. Trans. Linn. Soc. London (2 Zoo1.), 14(1): 83-122. Weber, M., 1892. Die Siisswasser-Crustaceen des Indischen Archipe1s, nebst Bemerkungen uber die Siisswasser-Fauna in Allgemeinen. Zool. Ergebn. einer Reise in Niederl. Ost-Indien, 2: 528-571.