THE RAFFLES BULLETIN OF ZOOLOGY 2005 THE RAFFLES BULLETIN OF ZOOLOGY 2005 Supplement No. 12: 31–37 © National University of Singapore PRESENCE OF THE ALPHEID SHRIMP GENUS POTAMALPHEOPS POWELL, 1979 (CRUSTACEA: DECAPODA: CARIDEA) IN SOUTH ASIA, WITH DESCRIPTION OF A NEW SPECIES FROM SRI LANKA Arthur Anker Department of Biological Sciences, University of Alberta, Edmonton, AB, T6G 2E9, Canada Email: [email protected] ABSTRACT.–A new species of the alpheid shrimp genus Potamalpheops Powell, 1979, is described on the basis of a single male specimen collected in a brackish water stream, 6 km from the sea, in southern Sri Lanka. Potamalpheops galle, new species, differs from all other species of the genus by a combination of several features, and appears to be closest to the South-East Asian species P. amnicus Yeo & Ng, 1997, P. miyai Yeo & Ng, 1997 and P. palawanensis Cai & Anker, 2004. The new species is the first record of Potamalpheops in South Asia and constitutes an important link between two major radiation zones of this genus, West Africa and South-East Asia/Australia. KEY WORDS.–Potamalpheops, Alpheidae, Sri Lanka, South Asia, brackish waters, Tethyan distribution. INTRODUCTION nearest localities being situated in the southern Malaysian state of Johor and Singapore (Yeo & Ng, 1997). Furthermore, The alpheid genus Potamalpheops was established by Powell this specimen differs in several subtle features from all (1979) for three West African species: P. pylorus Powell, 1979, previously known species and represents a new species, the type species, P. haugi (Coutière, 1906) and P. monodi which is described herein. (Sollaud, 1932). The presence of a fourth West African species was mentioned (Powell, 1979: addendum); however, this species remains undescribed. In West Africa the range of MATERIAL AND METHODS Potamalpheops stretches from Senegal to Gabon (Coutière, 1906; Sollaud, 1932; Gordon, 1956; Powell, 1976, 1979; Anker, The carapace length (CL) and total length (TL) are given in 2001), with all four species occurring in the Niger Delta region mm and were measured from the tip of the rostrum to the in Nigeria. Subsequently members of this genus were found posterior margin of the carapace (CL) or the posterior margin in other parts of the world: P. stygicola (Hobbs, 1973) in south- of the telson (TL). Other abbreviations used in the text: P – ern Mexico (Hobbs, 1973); P. hanleyi Bruce, 1991 and P. pereiopod; Mxp – maxilliped. The type specimen is deposited darwiniensis Bruce, 1993 in northern Australia (Bruce, 1991, in the Zoological Reference Collection, Raffles Museum of 1993);P. pininsulae Bruce & Iliffe, 1992 in New Caledonia Biodiversity Research, National University of Singapore (Bruce & Iliffe, 1992; Anker, 2001); P. amnicus Yeo & Ng, 1997 (ZRC). Specimens of the following species were used for in southern peninsular Malaysia and Singapore (Yeo & Ng, comparison and remain deposited in the Muséum National 1997); P. miyai Yeo & Ng, 1997 in western Indonesia and d’Histoire Naturelle, Paris, France (MNHN) and the National Philippines (Yeo & Ng, 1997; Cai, in prep.); P. tigger Yeo & Museum of Natural History, Smithsonian Institution, Ng, 1997 in Singapore and northern Australia (Yeo & Ng, Washington, D.C., U.S.A. (USNM): Potamalpheops pinin- 1997; Anker, 2003); P. johnsoni Anker, 2003 in Singapore sulaeBruce & Iliffe, 1992: 2 specimens, Pindai caves, New (Anker, 2003) and most recently P. palawanensis Cai & Anker, Caledonia (MNHN); P. palawanensis Cai & Anker, 2004: 2 2004 in Palawan (Cai & Anker, 2004). The habitats of Potam- specimens, St. Paul Cave, Palawan (MNHN); P. johnsoni Anker, alpheops species range from freshwater streams and brackish 2003: 1 specimen, Mandai Kechil, Singapore (MNHN); P. tidally influenced portions of rivers to mangroves and peat amnicus Yeo & Ng, 1997: 1 specimen, Sungai Kahang, Johor, flats, and also include freshwater and anchialine caves. S Malaysia (MNHN); P. tigger Yeo & Ng, 1997: 4 specimens, Lim Chu Kang, Singapore (MNHN); P. monodi (Sollaud, 1932): In 1996 M. M. Bahir (Wildlife Heritage Trust, Sri Lanka) 3 specimens, Porto-Novo, Benin (MNHN); P. pylorus Powell, collected a single, complete male specimen of Potamalpheops 1979: 1 specimen, Port Harcourt, Nigeria (MNHN); P. stygicola in a brackish water stream in southern Sri Lanka. This (Hobbs, 1973), 1 specimen (paratype), Oaxaca, Mexico specimen is the first record of this genus in South Asia, the (USNM). 31 Anker: New Potamalpheops from Sri Lanka TAXONOMY First chelipeds (Figs. 1g–i) symmetrical in shape, equal in size, not particularly enlarged compared to second pereiopod; FAMILY ALPHEIDAE RAFINESQUE ischium elongated; merus slightly longer than ischium and carpus; carpus cylindrical, distally slightly widening, ventro- GenusPotamalpheops Powell, 1979 mesially with at least five rows of grooming setae (Fig. 1h); chela subcylindrical, longer than carpus, fingers slightly Potamalpheops galle, new species shorter than palm (Fig. 1i), with scattered tufts of setae; finger (Fig. 1) tips bidentate (Fig. 1i’), cutting edges unarmed. Material examined. – Holotype - male, CL 3.15, TL 11.1, Second pereiopod slender (Fig. 1j); ischium and merus ZRC 2004.0673, SW Sri Lanka: Lelkada near Dodangoda, Galle subequal in length, slender; carpus five-articulated, ratio of District, Gin River basin, 6 km from the sea, altitude 5 m, 06º08’N carpal articles approximately equal to (from proximal to distal): 80º10’E, brackish water, 0.5–1 m deep, stream bank with vegetation 3–1–1.2–1.2–2 (Fig. 1j); chela simple, longer than distal carpal and leaf debris, coll. M.M. Bahir, 21 Mar.1996. article, subequal to first carpal article; fingers longer than palm. Third pereiopod slender (Fig. 1k); ischium armed with Description. – Carapace smooth, glabrous, laterally with one spine; merus armed with two spines (Fig. 1k); carpus two distinct longitudinal-oblique grooves, one anterior and unarmed, distally with strong seta; propodus armed with six one posterior (Fig. 1a). Rostrum triangular, broad at base, minute spines on ventral margin, including one distal spine only slightly exceeding distal margin of cornea (Fig. 1c, d), (Fig. 1k); dactylus about 0.4 length of propodus, simple, distally acute, without setae. Extra-corneal teeth well slender, gradually and slightly curving distally, furnished with developed, acute, infra-corneal region broadly rounded, fine setae (Fig. 1k). Fourth pereiopod similar to third pereiopod. produced anteriorly (Figs. 1a, b). Pterygostomial angle very Fifth pereiopod generally similar to third and fourth slightly projecting anteriorly, rounded, with a few simple pereiopods, but with different proportions of articles (Fig. (non-plumose) setae (Fig. 1a). Eyes largely exposed in 1l); ischium and merus each armed with one spine; propodus dorsal and lateral views; cornea well pigmented, occupying with two-three minute spines on ventral margin and at least most of eyestalk except for mesial and antero-mesial rows of grooming setae, distal setae longest (Fig. 1l). portions, antero-mesial margin with small angular projection (Fig. 1d), without setae. Ocellar beak visible in Abdominal segments with postero-ventral angles rounded, dorsal view, with black spot (Fig. 1c). fifth segment with postero-ventral angle angular, not acutely produced (Fig. 1n); sixth segment postero-ventrally with Antennular peduncles slender, second article slightly longer triangular articulated flap (Fig. 1n); posterior margin sub- than visible portion of first in dorsal view (Fig. 1c); first rectangular with minute subacute tooth (Fig. 1n’). Second segment with distal margin not toothed, dorsal margin with pleopod with appendix masculina twice as long as appendix small spines (Figs. 1b, e); stylocerite acute, reaching slightly interna (Fig. 1m), distally with four slender spines (Fig. 1m’). beyond distal margin of first segment (Figs. 1c, e); ventro- Uropodal exopod with diaeresis finely toothed for about 3/4 mesial carina with well developed, subacute tooth as of its length between insertion of lateral spine and abrupt illustrated (Fig. 1f); lateral flagellum lacking on both sides. incision (Fig. 1o), with around 19–20 very small triangular Antenna with basicerite bearing acute ventro-lateral tooth acute or subacute teeth (Fig. 1o’); lateral spine strong. Telson (Fig. 1b); scaphocerite not reaching distal margin of antennular relatively broad, sub-rectangular, slightly tapering distally (Fig. peduncle, disto-lateral spine robust (Fig. 1c), anterior margin 1p), with two pairs of dorsal spines situated as illustrated convex, not exceeding disto-lateral spine; carpocerite short, (Fig. 1p); posterior margin of telson medially rounded, with at with distal margin reaching to about 4/5 of scaphocerite (Fig. each angle two strong postero-lateral spines, mesial slightly 1b); flagellum (broken) slender. longer than lateral (Fig. 1p); anal tubercles absent. Mouthparts typical for Potamalpheops. Mandible with incisor Gill formula typical for genus: 5 pleurobranchs (P1–P5); 0 process bearing six triangular acute teeth, two central teeth podobranch; 1 arthrobranch (Mxp3); 2 lobe-shaped epipods largest; molar process with circular rows of setae and (Mxp1 and Mxp2); 5 strap-like epipods (mastigobranchs, conspicuous tubercle. Maxillule with endopod bilobed, dorsal Mxp3 + P1-P4); 5 sets of setobranchs (P1–P5), 3 exopods lobe without setae, ventral lobe with single very thick seta. (Mxp1–Mxp3). Colour in life unknown. Maxilla with scaphognathite rather narrow; dorsal lacinia with small incision; endopod (palp) slender, not setose. First Etymology. – The new species is named after the old Sri maxilliped with caridean lobe on exopod moderately Lankan port city of Galle situated near the type locality. developed; endopod (palp) bearing distally long, thick plumose setae; epipod rounded-triangular. Second maxilliped Habitat. – The single specimen was collected 0.5–1 m deep without specific features, with epipod elongated. Third in a stream, along a bank with vegetation and leaf debris. The maxilliped slender; coxal lateral plate ear-shaped, distally water condition was noted as brackish (M. M. Bahir, pers. subacute, covering base of exopod; tip of ultimate segment comm.). The stream belongs to the Gin River basin system, distally with at least two small spines; arthrobranch present, and is situated 6 km from the sea at 5 m above sea level. The well developed. only other decapod inhabiting this stream was the atyid shrimp 32 THE RAFFLES BULLETIN OF ZOOLOGY 2005 Fig. 1. Potamalpheops galle, new species, a – carapace, lateral view; b – frontal region, lateral view; c – same, dorsal view; d – detail of rostrum, orbital teeth and eyestalks; e – antennule, basal segments, dorsal view; f – same, mesio-ventral tooth of first segment; g – first pereiopod (cheliped); h – same, mesial view of chela and carpus; i – same, chela, i’ - detail of bidentate tips; j – second pereiopod; k – third pereiopod; l – fifth pereiopod; m – second pleopod, m’ – detail of appendix masclina; n - posterior fifth and sixth abdominal segments, n’ – detail of posterior margin of sixth segment; o – uropod, o’ – detail of diaeresis; p – telson. Scales: 1 mm (a-c, m), 0.5 mm (d-l, n, o); figures i’, m’, n’, o’ drawn without scale. 33 Anker: New Potamalpheops from Sri Lanka Table 1. All presently known records of Potamalpheops with brief summary of ecology. Species Locality Habitat Reference P. amnicus - Singapore: Central Catchment Area Freshwater streams free of tidal influence - Yeo & Ng, 1997 - S Malaysia: S Johor: Kota Tinggi P. darwiniensis - Australia: Northern Territory: Darwin Mangrove creeks, marine water - Bruce, 1993 P. galle, new species - Sri Lanka: Galle District Streams with tidal influence, brackish - Present study water P. hanleyi - Australia: Northern Territory: Darwin Mangrove creeks, marine water - Bruce, 1991 P. haugi - Gabon: Ogôoué River Rivers and streams upstream of tidal limit - Coutière, 1906 - Nigeria: Niger Delta - Powell, 1979 P. johnsoni - Singapore: Sungei Buloh, Lim Chu Kang Mangroves, marine and brackish water - Anker, 2003 P. miyai - W Indonesia: Pulau Bintan Mangroves, marine and brackish water - Yeo & Ng, 1997 - Philippines: Mindanao - Cai &Anker, 2004 P. monodi - Senegal: Néma Mangrove creeks and peat forests, - Gordon, 1956 - Sierra Leone: Murray Town brackish and tidal influenced freshwater - Powell, 1979 - Benin: Porto-Novo streams - Anker, 2001 - Nigeria: Niger Delta - Powell, 1976, 1979 - Cameroon: Manoka Bay - Sollaud, 1932 P. palawanensis - Philippines: Palawan: St. Paul caves Anchialine limestone caves, brackish or - Cai & Anker, 2004 oligohaline water, tidal influence P. pininsulae - New Caledonia: Pindai Peninsula Anchialine limestone caves, brackish - Anker, 2001; Cai water, tidal influence & Anker, 2004 - New Caledonia: Isle of Pines - Bruce & Iliffe, 1992 P. pylorus - Nigeria: Port Harcourt Mangrove creeks and mangrove peat flats,- Powell, 1979 also in transition zone between oligohaline and freshwater P. stygicola - S Mexico: Oaxaca: Cueva del Nacimiento Freshwater caves - Hobbs, 1973, 1983; and Cueva Gabriel Mejía-Ortíz et al., 2003 P. tigger - Singapore: Sungei Buloh, Lim Chu Kang Mangroves, marine or brackish water - Yeo & Ng, 1997; Anker, 2003 - Australia: Northern Territory: Darwin - Anker, 2003 P.sp. - Nigeria: New Calabar River Streams, transition zone between - Powell, 1979 (addendum) oligohaline and freshwater Fig. 2. Potamalpheops amnicus Yeo & Ng, 1997 (a, b) and P. palawanensis Cai & Anker, 2004 (c, d): a, b - male from Sungai Kahang, Johor, Malaysia (MNHN), frontal region in dorsal (a) and lateral (b) view showing a relatively long rostrum (cf. Yeo & Ng, 1997: fig. 1a, b); c - ovigerous female from St. Paul cave, Palawan (MNHN), frontal region showing a relatively short rostrum (cf. Cai & Anker, 2004: fig. 15A); d – male from the same locality (MNHN), second pleopod, d’ – detail of appendix masculina. Scales: left - 1 mm (a-c), right: 0.5 mm (d); figure d’ drawn without scale. 34 THE RAFFLES BULLETIN OF ZOOLOGY 2005 Caridina costai De Silva (M. M. Bahir, pers. comm.), which is segments of the second pereiopod (cf. idem.: fig. 4C) and believed to be a synonym of Caridina simoni simoni Bouvier, the more robust walking legs (cf. idem.: fig. 4F). Finally, P. 1904 (see Benzie & De Silva, 1984). Ovigerous females of this monodi differs from the Sri Lankan species by the very species have relatively large eggs and live mostly in different proportions of the carpal segments in the second freshwater habitats. The presence of C. costai (C. s. simoni) pereiopod (cf. Gordon, 1956: fig. 14), the lower number of could indicate indirectly that the stream where P. galle, new teeth in the uropodal diaeresis (cf. idem.: fig. 20) and the species, was collected was only feebly brackish. presence of numerous conspicuous setae on the anterior margin of the eyestalks (cf. Sollaud, 1932: fig. 1; Gordon, Distribution. – Presently known only from the type locality, 1956: fig. 1). The fourth, still undescribed West African Lelkada, Galle District, southwestern Sri Lanka. species was briefly described as “morphologically less specialized” than any of the other three African species Remarks. – Potamalpheops galle, new species, belongs (Powell, 1979: addendum, p. 150); this species also could toP. monodi (Sollaud, 1932) group, characterized by two be related to P. galle, new species. pairs of disto-lateral spines on the posterior margin of the telson. Within this group it can be usefully contrasted with The remaining six species of Potamalpheops, namely P. species with a medium to long rostrum; slender, pylorus and P. haugi from West Africa, P. darwiniensis from unspecialized chelipeds; and normally developed northern Australia, P. tigger from Singapore and northern pigmented cornea: P. miyai from Indonesia and the Australia,P. johnsoni from Singapore, and P. pininsulae from Philippines, P. amnicus from southern Malaysia and New Caledonia, are more distantly related to P. galle, new Singapore,P. palawanensis from Palawan, P. hanleyi from species, differing from the latter in a number of characters, northern Australia and P. monodi from West Africa. The including the development of the chelipeds (P. pylorus, P. new species differs from all these species by the presence haugi, P. darwiniensis) or in the shape and length of the of conspicuous grooves on the lateral sides of the carapace, rostrum (P. tigger,P. johnsoni and P. pininsulae). The minute although these grooves are indicated or feebly marked in dorsal spines on the first segment of the antennular peduncle some species, e.g., P. pininsulae or P. palawanensis (Anker, (Fig. 1e) are not characteristic of P. galle, new species. These pers. obs.). The only other species bearing well marked spines or spine-like setae are present in some other species, (although slightly differently arranged) grooves is P. e.g.,P. johnsoni and P. pininsulae (pers. obs.). However, as stygicola from southern Mexico (Hobbs, 1973, 1983). they are not very conspicuous, these spines may be easily However, this stygobitic species can be easily separated overlooked and have not been illustrated or mentioned in by the poor pigmentation of the eyes (cf. Hobbs, 1973: fig. previous descriptions. 1b) and the uniquely incised (V-shaped) diaeresis (Hobbs, 1983: fig. 1a). The rostrum of P. amnicus and P. palawanensis shows some variation in length (Fig. 2a–c), approaching the condition Besides the absence of well developed grooves on the found in P. galle, new species. A more significant variation of carapace all above-mentioned species differ from P. galle, the rostrum length and shape exists in P. pininsulae (Bruce & new species, by a combination of several other features. Iliffe, 1992), P. tigger and P. johnsoni (Anker, 2003). Potamalpheops miyai appears to be closely related to P. Furthermore, both P. pininsulae and P. johnsoni appear to be galle, new species, but differs from the new species in the highly variable in the development of the ventral rostral teeth, absence of a spine on the ischium of the fifth pereiopod which may be present in number of one to three or absent (cf. Yeo & Ng, 1997: fig. 4i), the non-protruding mesio- (Bruce & Iliffe, 1992; Anker, 2003). The previously unnoted anterior margin of the eyestalks (cf. idem.: fig. 3b) and the variation in P. amnicus and P. palawanensis appears to be shorter and stouter carpal segments of the second slight, but demonstrates once more that the rostrum in pereiopod (cf. idem.: fig. 4c). Potamalpheops amnicus, Potamalpheops species is not always a reliable taxonomic differs from P. galle, new species, by the shape of the character. uropodal diaeresis (cf. Yeo & Ng, 1997: fig. 1e), the presence of small setae on the anterior margin of the The appendix masculina of P. palawanensis was previously eyestalks (cf. idem.: fig. 1b), the absence of a spine on the unknown, as both type specimens of this species are merus and the ischium of the fifth pereiopod (cf. idem.: fig. females (Cai & Anker, 2004). Therefore, I take the 2i) and especially by the conspicuously elongated opportunity to illustrate the second pleopod and the appendix masculina (cf. idem.: fig. 1h). The stygophilic P. appendix masculina of the first known male of P. palawanensis can be separated from the Sri Lankan species palawanensis (Figs. 2d, d’). The appendix masculina of by its much longer rostrum (cf. Cai & Anker, 2004: fig.15A, most species of the genus, including P. galle, new species, B), slightly narrower telson (cf. idem.: fig. 15C), much more and P. palawanensis, is furnished with spines on the tip slender dactyli on the walking legs (cf. idem.: fig. 16D, E) (Figs. 1m’, 2d’). The only exception is P. pininsulae, in and the antero-mesial margin of the eyestalks non- which the spines are present on the tip but also along the protruding beyond the cornea (cf. idem.: fig. 15B). margins of the appendix masculina (Bruce & Iliffe, 1992: Potamalpheops hanleyi can be easily distinguished from fig. 31). In P. amnicus the appendix masculina is the new species by its much more robust chelipeds (cf. conspicuously elongated, reaching far beyond the distal Bruce, 1991: fig. 4A, B), the proportions of the carpal margin of the endopod (Yeo & Ng, 1997: fig. 1h). 35 Anker: New Potamalpheops from Sri Lanka Fig. 3. Distribution of 14 presently known Potamalpheops species: Pam – P. amnicus;Pda – P. darwiniensis;Pga – P. galle, n. sp.; Pha – P. haugi;Phn – P. hanleyi;Pjo – P. johnsoni;Pmi – P. miyai;Pmo – P. monodi;Ppa – P. palawanensis;Ppi – P. pininsulae;Ppy – P. pylorus;Psp –Potamalpheops sp. (Powell, 1979); Pst – P. stygicola;Pti – P. tigger. * indicates regions with the presently highest diversity of Potamalpheops (at least 3 species): the Niger delta in Nigeria, Singapore, and Darwin in Northern Territory, Australia. daile, are restricted to “refugial” habitats, such as caves and DISCUSSION land-locked anchialine pools. In contrast to the modern species of these presumably very old genera, most recent With the present new species the genus Potamalpheops species of Potamalpheops are epigean, although confined to contains 14 species and appears to have a disjunct worldwide “stressful” habitats (e.g., mangroves, streams with or without distribution (Table 1, Fig. 3). The finding of P. galle, new tidal influence, and caves). species, in Sri Lanka extends the distribution of this genus from Southeast Asia to South Asia. The genus appears to All presently known records of Potamalpheops species and have radiated in two major zones: in tropical South-East Asia the ecology of each species are summarized in the Table 1. It and Australia with seven species, and in tropical West Africa can safely be assumed that the present knowledge of with four species, one of which remains undescribed. Bruce Potamalpheops is far from being complete. The major reason (1991) suggested a former Tethyan distribution and an Upper is that Potamalpheops spp., are rarely if ever collected. Most Cretaceous origin (80–90 my) of the genus. Since P. galle, species are very small (less than 15 mm TL), inconspicuous in new species, appears to be morphologically (and possibly colour and cryptic in habits, and therefore, may easily be phylogenetically) closer to some South-Asian species, it is overlooked even by an experienced collector. Some of the possible that it originated from the same ancient stock as the described species presently restricted to the type locality modern Southeast Asian and Australian species. With respect may have a much wider distribution range. Furthermore, new to its ecology, P. galle, new species, belongs to a group of species may be discovered in other parts of the Indo-Pacific species living in tidally influenced freshwater or brackish- in the future. water habitats, which also includes P. monodi and the stygophilic P. palawanensis. The ancestors of the present species were possibly shallow water marine or brackish water ACKNOWLEDGEMENTS species, from the same lineage as today’s mangrove species P. miyai, P. tigger or P. hanleyi. A phylogenetic hypothesis I would like to thank Mr. M. M. Bahir (Wildlife Heritage Trust, for the genus Potamalpheops (Anker, in prep.) would enable Colombo, Sri Lanka) and Dr. Yixiong Cai (Department of a more solid evolutionary interpretation of biogeographical Biological Sciences, National University of Singapore, and ecological features. Singapore) for collecting and making the specimen available for study, respectively. This study was supported by a NSERC Other caridean genera with disjunct worldwide distributions, research grant (A7245) to Dr. A. Richard Palmer (Department e.g., the atyid genera Typhlatya Creaser and Antecaridina of Biological Sciences, University of Alberta, Edmonton, Edmondson and the barbouriid genus Parhippolyte Borra- Canada). Dr. Sammy De Grave (Oxford University Museum of 36 THE RAFFLES BULLETIN OF ZOOLOGY 2005 Natural History, Oxford, U.K.) reviewed the first draft of the Proceedings of the Biological Society of Washington,106: 698– manuscript and made valuable suggestions and corrections. 704. Dr. Rafael Lemaitre (National Museum of Natural History, Bruce, A. J. & T. M. Iliffe, 1992. Potamalpheops pininsulae sp. nov., Smithsonian Institution, Washington, D.C., U.S.A.) arranged a new stygiophilic shrimp from New Caledonia (Crustacea: the loan of the USNM paratype of P. stygicola, used for Decapoda: Alpheidae). Stygologia,87: 231–242. comparison with the new species. Mr. Hendrik Freitag (Institut Cai, Y. & A. Anker, 2004. A collection of freshwater shrimps für Zoologie - Tierökologie, Martin-Luther-Universität Halle- (Crustacea: Decapoda: Caridea) from the Philippines, with Wittenberg, Halle, Germany) and Dr. Darren C. J. Yeo descriptions of five new species. Tropical Zoology,17: 233–266. (Department of Biological Sciences, National University of Coutière, H., 1906. Sur une nouvelle espèce d’Alpheopsis,A. Haugi, Singapore, Singapore) collected and donated specimens of P. provenant d’un lac d’eau douce du bassin de l’Ogoué (Vouyage palawanensis and P. amnicus, respectively, which were also de M. Haug). Bulletin du Muséum d’Histoire Naturelle,12: 376– used for comparison. 380. Gordon, I., 1956. Redescription of Alpheopsis monodi Sollaud, a rare freshwater prawn from Senegal. Bulletin de l’Institut Français de LITERATURE CITED l’Afrique Noire,série A, 18: 1110–1117. Hobbs, H. H. Jr., 1973. Two new troglobitic shrimps (Decapoda: Anker A., 2001. Taxonomie et évolution des Alpheidae (Crustacea, Alpheidae and Palaemonidae) from Oaxaca, Mexico. Bulletin of Decapoda). Thèse de Doctorat, Muséum National d’Histoire the Association for Mexican Cave Studies,5: 73–80. Naturelle, Paris. 2 vols : 547 + 331 pp. Hobbs, H. H. Jr., 1983. 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