THE RAFFLES BULLETIN OF ZOOLOGY 2009

THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 1–4 Date of Publication: 28 Feb.2009 © National University of

LEPTOSERIS KALAYAANENSIS (: AGARICIIDAE), A NEW SPECIES FROM THE

Wilfredo Y. Licuanan Bro Alfred Shields Marine Station and the Biology Department, De La Salle University-Manila 2401 Taft Avenue, Manila 1004, The Philippines Email: [email protected]

Porfi rio M. Aliño The Marine Science Institute, University of the Philippines, Diliman, Quezon City 1101, The Philippines Email: [email protected]

ABSTRACT. – Leptoseris kalayaanensis new species, from the Kalayaan Island Group, the Philippines, is described. It is characterized by unique solid spines all over its encrusting surface.

KEY WORDS. – Scleractinia, Agariciidae, Leptoseris, new species.

INTRODUCTION Philippines. The authors have not seen the species in the Philippines outside the type locality. Mr. Phan Kim Hoang A new species of Leptoseris, L. kalayaanensis is described (Nha Trang Institute of Oceanography, ) has from three coralla found under overhangs in about 15m collected another colony (a photograph of which was sent depth at its type locality, the Northeast Investigator Shoal by Mr. Brian Stockwell of Silliman University, Philippines) of the Kalayaan Island Group, the Philippines. It is unusual at 27–28m depth at the south reef of North Danger Reef among Leptoseris in that it bears solid spines (up to 10 mm in the Spratly Islands. Like the holotype, it was found in long and three mm thick at the base) all over its encrusting indents on a steep reef slope. surface. These spines are different from the hollow tubes sometimes found in L. scabra Vaughan, 1907 and L. tubulifera Vaughan, 1907. The spines are three to fi ve mm MATERIALS AND METHODS apart and are usually found either on concentric ridges near the middle of the corallum, or on radiating ridges near were collected at the Investigator Shoal (9.172°N the colony margin. Corallite centers are larger and more 116.467°E) on 6 May 1999 on the way back from the Kalayaan recognizable near the colony center, where a slightly larger Islands Group Inter-Agency Marine Scientifi c expeditions central corallite is found. to the Spratly Islands on board the Training and Research Vessel Sardinella, of the University of the Philippines in Leptoseris kalayaanensis, new species, was fi rst mentioned the Visayas, and funded by the Philippine Department of in Licuanan (2003) but since no holotype or syntypes were Science and Technology. The site of the collection is a steep designated then, this name as originally used should be wall at 15 m depth. Paratypes were located approximately considered as a nomen nudum under ICZN Code Article 2 m from the holotype at the same depth. After soaking in 16.4.1. This new species is thus described in this paper and sodium hypochlorite solution, the corals were examined using a holotype (P1L001932, deposited at the Coral Collection of a Zeiss Stemi DV400 stereomicroscope. A stage micrometer the Marine Science Institute, University of the Philippines, for the latter microscope was used for the measurements. Quezon City, the Philippines) is designated. Macro photographs of the corolla were made using Nikon D70S with 60 mm Micro Nikkor and Sigma EM-140 DG Type locality is the Northeast Investigator Shoal of the while photomicrographs were made using an Olympus Kalayaan Group of Islands west of Palawan, western SZX7 dissecting scope with an Evolution 5.0 MP digital camera attachment.

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The holotype and paratypes are deposited at the Coral Septo-costae are equal, closely packed, with non-dentate Collection of the Marine Science Institute, University of edges and an acute keeled profi le. Fine granulations are the Philippines, Quezon City, the Philippines. found at regular intervals at the edges of lateral ridges running parallel to the septo-costae giving them a saw-like appearance. The septo-costae are fairly straight, also on the TAXONOMIC ACCOUNT spines. About 12–23 reach the corallite center but some fuse before reaching the centre while some others form interrupted Scleractinia Bourne, 1905 plates near the center.

Fungiina Verril, 1943 The columella, usually present in the larger corallites, is usually a solid mass with an uneven surface that may be Agariciidae Gray, 1847 connected to septo-costae. The encrusting unifacial laminae and fi ne septo-costal structure suggests this species is better Leptoseris Milne-Edwards & Haime, 1849 placed in Leptoseris than in any other of the attached agariciid genera. Leptoseris kalayaanensis, new species Paratypes. – Both paratypes have central corallites with Type material. – Holotype P1L001932 (94 mm in diameter). slightly raised margins (Fig. 6). P1L001933 is dome-shaped Paratypes P1L001933 (84 mm in diameter) and attached to but clearly distinct from P1L001934 (103 mm long). All collected from the NE Investigator Shoal (9.172°N 116.467°E), 15 m depth, coll. W. Licuanan, 6 May 1999. The paratypes are attached to the same rock and have no skeletal connection.

Holotype. – Corallum small, encrusting (but with margins mostly raised from the substratum) to unifacial laminate and slightly dome-shaped. Margin entire or lobate (Fig. 1). Thickness at the margins is about 0.5 mm and increasing towards the center. Living colony light brown with whitish margin [photographs can be seen in Licuanan (2003) and in Pg. 51 of the same book). Newly cleaned coralla had greenish tinge in the center.

Corallites circular to elliptical, 1.8 to 3.6 mm in diameter measuring from where septo-costae descend into the shallow fossa (Fig. 2). Central (primary) corallite deeper but barely distinct from lateral ones, which may be of similar size (Fig. 2, see also Fig. 3 of a paratype). Corallites are located mainly within the inner zone of the corallum, from the centre until halfway the periphery, somewhat in a regular pattern three to eight mm apart from each other, but rarely about 10 mm from the margin. These are not outwardly inclined but are Fig. 1. Leptoseris kalayaanensis, new species: Holotype (diameter immersed to sub-immersed. 94 mm), NE Investigator Shoal, Kalayaan Islands (P1L01932). Photo: W. Licuanan. Distinctive spines are distributed over the upper surface; up to 10 mm tall and elliptical in cross-section, diameter about 2.7 mm at the base and 1.1 mm at the blunt tips. The spines are unbranched and are scattered between corallites in concentric rows. Most spines initially point out towards the corallum periphery then curve upwards (Fig. 4). Some spines are found on top of folds of laminae near the corallum margins, whereas others emanate from a mound-like common base. Spines are solid, as can be seen in a broken spine, (Fig. 5), which suggests that they are not homologue to the hollow tubes in L. scabra Vaughan, 1907, and L. tubulifera Vaughan, 1907. The spines are much taller than the mound- like nodules described for L. solida (Quelch, 1886), L. hawaiiensis Vaughan, 1907, and L. scabra (see Dinesen, 1980). About 10–15 individual septo-costae run continuously up to the tips and, except for the ones at the sharp ends of the elliptical base, down to the other side. Fig. 2. Leptoseris kalayaanensis, new species. Holotype: Inset showing the slightly larger central corallite and its columella. Photo: K. Luzon.

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and attached only at the centre. The distinctive spines, when folds or collines of L. mycetoseroides, nor the deep rounded examined from the underside of the corallum (where the corallites of L. hawaiiensis. Hydnophora-like projections have underlying substratum was cleared), are clearly solid as been described for L. incrustans, and mound-like nodules well. It partially covers P1L01934 and is attached to the for L. solida, L. hawaiiensis, and L. scabra but these are same rock. P1L01934 has shorter spines but remain distinct shorter and do not have the shape of L. kalayaanesis, new from other Leptoseris. species, spines.

Etymology. – The species is named after the Kalayaan Island group, near Palawan province in the western Philippines, in DISCUSSION which the type locality is found. With the addition of L. kalayaanensis, new species, the Affi nities. Ð The spines of L. kalayaanensis, new species, number of Leptoseris species in the Philippines has increased clearly distinguishes it from the other Leptoseris. It differs to 14 (as based on Veron, 2000), next to L. gardineri, L. from L. gardineri Horst, 1921, and L. papyracea (Dana, 1846) papyracea, L. tubulifera, L. amitoriensis, L. explanata, in that there it shows no tendency to form branching fronds L. scabra, L. solida, L. striata, L. mycetoseroides, L. nor does it form tiered or whorled fronds like L. amitoriensis hawaiianensis, L. incrustans, L. foliosa, and L. yabei. Veron Veron, 1990. Among the encrusting to laminate forms, L. kalayaanensis, new species, differs from L. explanata Yabe & Sugiyama, 1941, L. scabra, L. solida, L. striata Fenner & Veron, 2000, and L. yabei (Pillai & Scheer, 1971) in that the septo-costae do not alternate. Leptoseris hawaiiensis, L. incrustans (Quelch, 1886), L. mycetoseroides Wells, 1954, and L. foliosa Dinesen, 1980, also have even septo-costae but L. kalayaanensis, new species, does not have the irregular

Fig. 5. Leptoseris kalayaanensis, new species. Holotype: Detail and inset showing the base of a broken spine demonstrating it is solid, not hollow. Photo: K. Luzon.

Fig. 3. Leptoseris kalayaanensis, new species. Paratype (P1L01933): Detail showing the larger central corallite on the upper right. Photo: K. Luzon.

Fig. 6. Leptoseris kalayaanensis, new species. Paratypes: ex (diameter 84 and 103 mm), NE Investigator Shoal, Kalayaan Islands (P1L01933 is the lower part). P1L01934 is partially covered Fig. 4. Leptoseris kalayaanensis, new species Holotype: oblique by P1L01933 and is attached to the same rock. The two have no photo to show spines. Photo: K. Luzon. skeletal connection. Photo: W. Licuanan.

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& Fenner (2000) however did not include the Philippines LITERATURE CITED in the species ranges of L. tubulifera and L. striata (a new species then) unlike in Veron (2000). Dinesen, Z. D., 1980. A revision of the coral Leptoseris (Scleractinia: Fungiina: Agariciidae). Memoirs of the Queensland All colonies of L. kalayaanensis, new species, that have Museum 20(1):181–235. been reported so far all came from the same reef complex. Licuanan, W. Y., 2003. A ray of light: A new species of coral None have been seen in the other biogeographic sections of – Leptoseris kalayaanensis. In Aliño PM & MCC Quibilan, the Philippines despite the intensive search for the species. The Kalayaan Islands. Our Natural Heritage. U.P. Marine Like the type specimens, the one collected by Dr. Phan Science Institute, Diliman, Quezon City. 60 pp. Kim Hoang was also in deep water (27–28 m) at the base Veron J. E. N. 2000. Corals of the World. Australian Institute of of the reef slope. Marine Science, Townsville, Australia. Vol. 2: 202–220. Veron J. E. N. & D. Fenner. 2000. Coral species recorded from the Philippines or likely to occur there. In: Werner, T. B. & G. ACKNOWLEDGEMENTS R. Allen (eds.), 2000. A Rapid Marine Biodiversity Assessment of the Calamianes Islands, Palawan Province, Philippines. We wish to thank the editors and reviewers for helping RAP Bulletin of Biological Assessment 17. Washington, D.C. Conservation International: 66–74. improve the manuscript. We also thank the Philippine Department of Science and Technology for funding the Kalayaan Island Group Project, the partner institutions of the project, and the RV Sardinella and its crew. This work was also supported by the GEF/WB Coral Reef Targeted Research and Capacity Building for Management Program. This is contribution number 378 of the U.P. Marine Science Institute.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 5Ð15 Date of Publication: 28 Feb.2009 © National University of Singapore

HELMINTHS OF MELOMYS RUFESCENS AND MELOMYS SPP. (MURIDAE: HYDROMYINAE) FROM PAPUA NEW WITH THE DESCRIPTIONS OF A NEW GENUS AND FIVE NEW SPECIES IN THE HELIGMONELLIDAE (NEMATODA: TRICHOSTRONGYLOIDEA)

L. R. Smales Parasitology Section, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia Email: [email protected]

ABSTRACT. – A cestode, a trematode and 14 nematodes, including fi ve new species in the Heligmonellidae (Trichostrongyloidea), were collected from the digestive tracts of 25 Melomys rufescens (Rodentia: Hydromyinae) from Papua New Guinea. Of these Heligmonoides mirzai, new species, is nearest to H. bulbosus in the form of the synlophe but is distinguished by males with longer spicules and females with a praepuce and a spiked tail tip. Melomystrongylus sepikensis, new genus, new species, differs from all other heligmonellids in having a single hypertrophied ventral ridge extending from the excretory pore along the anterior third of the body. Odilia similis, new species, is allied to O. maxomyos, O. praeputialis, O. polyrhabdote, and O. uromyos in the number of ridges in the midbody but differs in having a spicule to body length ratio of 1:6.5 and a blunt conical female tail. Odilia species 1, tiny male worms with 16Ð18 ridges in the synlophe and spicules 260–280 µm long, could not be characterised further because there was insuffi cient material available. Paraheligmonelloides singauwaensis, new species, resembles P. triangulus but has longer spicules and smaller eggs. Overall the composition of the helminth assemblage found in M. rufescens resembled neither that of M. lutillus (also from Papua New Guinea) nor those of M. burtoni or M. cf burtoni from Australia but was indicative of closer relationships with the helminths found in Indonesian and Malaysian rodent hosts.

KEY WORDS. – Papua New Guinea, Muridae, Melomys, Heligmonellidae, Odilia, Paraheligmonelloides, Melomystrongylus.

INTRODUCTION Melomys rufescens is widespread across lowland New Guinea and is also found in areas of disturbed vegetation The endemic Australasian rodent tribe, the Uromyini at higher altitudes (Flannery, 1995). Little is known about (Muridae: Hydromyinae), is a heterogeneous clade of the helminth fauna of Melomys species in New Guinea, with primarily herbivorous murids with medium to large eyes four species of nematode and one cestode reported from and sparsely haired tails (Flannery, 1995). Assumed to have Melomys lutillus (Thomas, 1913) (see Smales, 2005a) but been derived from the earliest murid invasion of the island none from M. rufescens. of New Guinea, uromyin ancestors may have arrived, from Southeast Asia, during the Miocene Epoch (10Ð15 million This study was undertaken to report on the helminth fauna of years ago (mya) when the region consisted of no more than the digestive tract of M. rufescens and to compare it with that an island archipelago (Holloway & Hall, 1988). Subsequently, of M. lutillus and with that of the closely related Australian representatives of at least one genus, Melomys (Thomas, representatives of the genus, Melomys burtoni (Ramsay, 1922), may have travelled to Australia during the Pleistocene 1887) and Melomys cf. burtoni (see Menzies, 1996), whose 0.02Ð0.04 mya across the land-bridges formed during that helminths have been documented (Smales, 2005a). time (Flannery, 1995; Godthelp, 2001). Following revision by Menzies (1996) the genus now comprises four species from New Guinea, including Melomys rufescens (Alston, MATERIALS AND METHODS 1877) (the first rodent species from New Guinea to be described), four species from Australia and 13 extralimital The digestive tracts from the bodies of 23 M. rufescens, and species (Duff & Lawson, 2004). 10 Melomys, not identifi ed to species level and not including M. lutillus, previously collected from Papua New Guinea,

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fi xed in 5Ð10% formalin and stored in 70% ethanol, were RESULTS examined for helminths. Additional specimens dissected from two individuals of M. rufescens were also studied. Two platyhelminths, a cestode and trematode, and 14 nematodes were identifi ed, to species level where possible, The following list is of 16 localities where hosts (with from 22 of 25 M. rufescens (Table 1). The trematode the number of individuals in parentheses) were collected. fragments, the capillariid nematode, Capillaria sensu lato, the Latitude and longitude are given where they were recorded larval heligmonellids and a single female heligmonellid could or where they can reasonably be estimated from the locality not be identifi ed further, nor was there suffi cient material to data available. fully characterise one of the two species of Odilia described below. All are new host records. Melomys rufescens. – Yapsiei area, 4¡ 35'S 141¡5'E (1); 2km S Mt Samoro, West Sepik (1); Sanduan Province: The cestode, Raillietina celebensis, occurs across the region Sideia Mission, 10¡32'S 150¡48' E (2); Milne Bay Province: from , Papua, Papua New Guinea to Australia, in NNW slopes Mt. Boravi, 6¡35'S 142¡52'E (1); Southern a range of murid hosts. Highlands Province: Jumbora Plantation, Popondetta, 8°46’S 148¡14'E (1); Oro Province: Aribaai Village, 13 miles (21 Of the nematodes, the spirurid Protospirura kaindiensis, km) E Chuave, 6¡06'S 144¡56'E (1); Doido Village, Mt and the oxyurid Syphacia longaecauda have been described Karimui, 6¡32' S 142¡52'E (1); Yuro Village (1); Chimbu from New Guinean hosts, the heterakid, Heterakis spumosa Province: Western Highlands Province, no further data is a cosmopolitan species that has previously been recorded (1); 12 km NW Tep Tep, 5¡57'S 146¡05'E (3); Wanuma, from the Island of New Guinea and the chabertiid, 4¡54'S 145¡19'E (2); Madang Province: Kalolo (4); 10 km Cyclodontostomum purvisi, although previously noted from W Bulolo, 7¡12'S 146¡39' E (3); Minava Village, Watut, Indonesia and Australia, is a new record for New Guinea. 6¡50'S 146¡2'E (1); 14 miles (23 km) NW Nauti Village, 7¡15'S 146¡13'E (1); Singauwa River, Lae, 6¡07'S 146¡59'E The dominant nematode group, the trichostrongyloids, (1); Morobe Province. were represented by the heligmonellids Nippostrongylus brasiliensis, a cosmopolitan species but a new record for Melomys species. – Boulder Camp, Goodenough Island New Guinea, Odilia mackerrasae previously recorded 9¡21'S 150¡16'E (2); Munimun Village, Aguan, 9¡53'S from Australia and New Guinea and Macrostrongylus 149¡23'E (4); Sanduan Province: Dokfuma Star Mountains ingens, known only from New Guinean hosts as well as 5¼01'S 141¡07'E (2); Milne Bay Province: Aru Island, 6¡10'S the trichostrongylid Peramelistrongylus sp. previously 134¡30'E (2). recorded from Australia and New Guinea. New species of the Heligmonellidae from the genera Heligmonoides, Odilia All helminths recovered from the digestive tracts were and Paraheligmonelloides were found and together with a stored in 70% ethanol. Prior to microscopical examination new genus, Melomystrongylus, are described below. nematodes were cleared in lactophenol and examined as temporary wet mounts; cestodes were stained in celestine One Melomys sp. was infected with the new species of blue, dehydrated in ethanol, cleared in methyl salicylate Heligmonoides, one with the new species of Odilia and fi ve and mounted in Canada balsam. Transverse sections were with the new genus, Melomystrongylus. prepared by cutting by hand using a cataract scalpel and mounting in polyvinyl lactophenol and spicules were mounted in Berlese’s medium, for detailed study. Figures were prepared with the aid of a drawing tube and measurements, in micrometres unless otherwise stated, given as the range Heligmonoides mirzai, new species followed by the mean in parentheses, were taken using an (Figs. 1Ð14) ocular micrometer. Terminology of hosts follows Flannery (1995) and Menzies (1996) and parasites Durette-Desset Material examined. – Holotype male and allotype female: BBM- (1983, 1985). NG-99529, from the small intestine of Melomys rufescens (Muridae: Hydromyinae), type host, Nauti Village, 7¡15'S 146¡13'E, Morobe All specimens have been returned to their home institutions, Province, Papua New Guinea, coll. A. B. Mirza, 10 Oct.1970. the Bernice Bishop Museum, Honolulu, Hawaii (BBM) or the Paratypes: 12 males, 12 females, SAM AHC34783; collection Australian Museum, Sydney, Australia (AM), or the South data as above. Australian Museum, Adelaide, Australia (SAM). Voucher specimens: BBM NG- 97928 1 male, 2 females All measurements are in micrometres (µm) except for lengths from the small intestine of Melomys rufescens, Minava which are given in milimetres (mm). Village, Watut, 6¡50'S 146¡21'E, Morobe Province, coll. A. B. Mirza, 23 Oct.1969; AM W32620, 2 males, 3 females from small intestine of Melomys rufescens, Yapsiei area, 4¡35'S 141¡5'E, Sanduan Province, coll. T. Flannery, 17 Jan.1984; 1 male, 2 females, AM W32625 from the small intestine of Melomys species, Dokfuma Star Mountains

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Table 1. Helminth parasites from 25 Melomys rufescens from Papua New Guinea.

Taxon Site in host Prevalence % Cestoda Davaineidae Raillietina celebensis (Janicki, 1902) Small intestine 4.0 Trematoda fragments Small intestine 4.0 Nematoda Capillariidae Capillaria sensu lato Small intestine 4.0 Chabertiidae Cyclodontostomum purvisi Adams, 1933 Caecum 16.0 Heligmonellidae Heligmonoides mirzai, new species Small intestine 20.0 Macrostrongylus ingens Smales, 2008 Small intestine 24.0 Melomyostrongylus sepikensis, new species Small intestine 20.0 Nippostrongylus brasiliensis (Travassos, 1919) Small intestine 4.0 Odilia mackerrasae (Mawson, 1961) Small intestine 16.0 Odilia similis, new species Small intestine 16.0 Odilia sp. Small intestine 4.0 Paraheligmonoides singauwaensis, new species Small intestine 4.0 heligmonellid larvae Small intestine 12.0 heligmonellid species Small intestine 4.0 Heterakiidae Heterakis spumosa (Schneider, 1866) Caecum 4.0 Oxyuridae Syphacia longaecauda Smales, 2001 Caecum 20.0 Spiruridae Protospirura kaindiensis Smales, 2001 Stomach 4.0 Trichostrongylidae Peramelistrongylus sp. Small intestine 24.0

5¼01'S 141¡07'E, West Sepik, Sanduan Province, coll. T. Male (holotype and 9 paratypes): Length 2.2Ð3.3 (2.8) Flannery, 5 Apr.1984. mm, maximum width 83Ð107 (90). Cephalic vesicle 36Ð46 (39.25) long. Oesophagus 350Ð550 (420) long; nerve ring Prevalence. – A total of 5 of 25 M. rufescens and 1 of 10 145 from cephalic end. Bursa asymmetrical, left lobe larger, Melomys spp. rays of left lobe more robust; without dorsal median notch. Dorsal ray symmetrical; divided at about half its length, each Etymology. – The species name is in recognition of the branch dividing again at distal tip; terminal divisions, rays extensive collecting done by A. B. Mirza in the 1960s and 9, 10 symmetrical; rays 8 arising at same level proximally 1970s in New Guinea. to division of dorsal ray, left ray 8 slightly more robust than right. Rays 4, 5, 6 with common stem, reaching margin of Description. – Small coiled worms; prominent cephalic bursa, rays 6 more slender than rays 4, 5. Rays 2, 3 with vesicle present; buccal capsule vestigial. Mouth opening common stem, widely divergent distally, rays 3 longer than triangular with rudimentary lips; labial and cephalic papillae rays 2. Genital cone short, lightly sclerotized, ventral lobe not observed. Oesophagus claviform. Nerve ring surrounds with unpaired papillae 0, dorsal lip bifi d, each lobe with oesophagus at about middle third; deirids and excretory single papilla 7. Spicules equal, fi liform, proximal ends pore at about same level, posterior to nerve ring. Synlophe rounded, distal tips simple, straight, 250Ð320 (275) long. of pointed longitudinal cuticular ridges extends from the Gubernaculum 27 long. posterior margin of the cephalic vesicle to level of spicules or vulva; 16Ð18 ridges in anterior, 21Ð24 in midbody. Axis Female (allotype and 9 paratypes): Length 2.95Ð3.7 (3.4) of orientation of ridges passing from ventral right to dorsal mm, maximum width 99Ð107 (104). Cephalic vesicle 42.0 left side, inclined about 75¡ from sagittal axis; 10Ð12 ridges Ð49.5 (45) long. Oesophagus 350Ð400 (380) long; nerve ring dorsal side, 11Ð12 ridges ventral side in midbody. Ridges 190, 220, excretory pore 250, 280 deirids 250, 280 from 1, 1’ large but not thickened basally, together with ridges cephalic end. Posterior end twisted and fl exed ventrally, with 2, 2’ form a carene of type B. Ridges 3–7 decreasing in praepuce in mature worms; vulva opening about 100 from size, ridges 8Ð10 increasing in size, ridge 11 smaller; ridges tail tip. Monodelphic; ovejector with sphincter, 95, longer 3’– 8’ increasing in size, ridges 9’–11’ decreasing in size. than vestibule 40, and infundibulum, 45. Tail conical, tip with Posterior region of body with 19Ð20 (male) 20Ð21 (female) spike 39.5Ð55 (48.5) long. Eggs thin- shelled, ellipsoidal, in ridges reduced in size; 10Ð11 dorsal side, 9Ð10 ventral side utero 51Ð69 (57) by 30Ð36 (33.8). ridges.

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Remarks. – Despite the fact that much of the material had species, difers from H. bulbosus in spicule length (250Ð320 been fi xed in the hosts in situ, making it diffi cult to discern compared with 92Ð100), the proportions of the dorsal ray, the morphology of the anterior end of the worms, it could the length of rays 3 and the shape of the female posterior be seen that Heligmonoides mirzai, new species, has most end, with a praepuce and spiked tail tip. of the morphological features of the genus Heligmonoides and resembles H. bulbosus Ow Yang et al., 1983, in the Hasegawa & Syafruddin (1994a) originally described number and arrangement of the ridges on the synlophe (Ow Maxomystrongylus musseri as Heligmonoides musseri, noting Yang et al., 1983, Fig 11A, B). Heligmonoides mirzai, new that it most closely resembled H. bulbosus, particularly in

Figs. 1–14. Heligmonoides mirzai, new species, from Melomys rufescens and Melomys sp. from Papua New Guinea: 1, female anterior end, lateral view; 2, male cross section, anterior body; 3, female cross section, anterior body; 4, male cross section, midbody; 5, female cross section, midbody; 6, spicule tips; 7, male cross section, posterior body; 8, female cross section, posterior body; 9, genital cone, dorsal view; 10, ovejector, lateral view; 11, gubernaculum, lateral view; 12, female posterior end, lateral view; 13, dorsal ray; 14, bursa, left lateral view. Scale bars: 1, 10, 11, 14 = 50 μm; 2–5, 7, 8, 12 = 25 μm; 6 = 10 μm; 9, 13 = 12.5 μm.

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the features of the carene; supported by 3Ð4 slender ridges Melomys sp., Munimun Village, Aguan, 9¡53'S 149¡23'E, lacking thickened roots. Subsequently these authors erected coll P. German, 8 Aug.1992; 1 female, AM W32558, from the new genus Maxomystrongylus Hasegawa & Syafruddin, the caecum of Melomys sp., Boulder Camp, Goodenough 1997, and transferred H. musseri into it on the basis of Island, 9¡21'S 150¡16'E, Sanduan Province, coll. T. Ennis, 25 the characters of the synlophe discussed above, having Aug.1987; 2 females, AM W32559 from the small intestine the division of the dorsal ray distal to the derivation of of Melomys sp., Boulder Camp, Goodenough Island, 9¡21'S rays 8 and a vagina vera with a unilateral diverticulum 150¡16'E, Sanduan Province, coll. T. Ennis, 25 Aug.1987; (Hasegawa & Syafruddin, 1994a). Further they commented AM W32626, 5 males, from the small intestine of Melomys that the systematic relationship between H. bulbosus and sp., Dokfuma Star Mountains 5¼01'S 141¡07'E, Milne Bay Maxomystrongylus remained unclear. Although H. mirzai, Province, coll. T. Flannery, 5 May 1987. new species, has characters of the synlophe and a form of the dorsal ray consistent with Maxomystrongylus, it does not Prevalence. – A total of 5 of 25 M. rufescens and 3 of 10 have a unilateral diverticulum of the vagina. Consequently it Melomys spp. has been placed in the genus Heligmonoides together with the other Southeast Asian representatives of the genus, H. Etymology. – The species name refl ects the locality in which bulbosus and H. lanceolatus Ow Yang et al., 1983, pro tem. this nematode is found. A revision of the heligmonellid genera from the Oriental and Australasian regions is needed to resolve the relationship Description. – Relatively robust worms; prominent cephalic between Heligmonoides and Maxomystrongylus and the vesicle present; buccal capsule vestigial. Mouth opening systematic position of the Australasian species. triangular with rudimentary lips; surrounded by 4 cephalic papillae and 2 amphids; labial papillae not observed. Oesophagus claviform. Nerve ring surrounds oesophagus Melomystrongylus, new genus at about a third its length, digitiform deirids and excretory pore at about same level, posterior to nerve ring. Synlophe Diagnosis. – Trichostrongyloidea: Heligmonellidae: of continuous longitudinal pointed cuticular ridges extends Nippostrongylinae. Synlophe well developed with pointed from the posterior margin of the cephalic vesicle to just ridges; in midbody axis of orientation of ridges passing anterior to the bursa or vulva; 8Ð14 ridges in anterior, 14Ð16 through ventral right and dorsal left sides inclined about 65¡ in midbody. Axis of orientation passing from ventral right from sagittal axis in anterior body, lacking clear orientation to dorsal left sides, inclined about 65¡ to sagittal axis in in mid and hind body. Ventral ridge 5’ hypertrophied anterior body; 4Ð8 ridges dorsal side, 4Ð8 ridges ventral anteriorly. Bursa asymmetrical with larger left lobe. Dorsal side. Ridges 1Ð6 decreasing in size; ridges 1'Ð4' decreasing ray divided distal to level of branching of rays 8 from dorsal in size, ridge 5' largest, ridges 6'Ð8' decreasing in size. trunk. Parasites of hydromyine murids. Posterior region of body with 15 (male) 10Ð14 (female) ridges reduced in size with no clear axis of orientation; Type species. – Melomystrongylus sepikensis, new 7Ð8 (male), 6Ð7 (female) ridges dorsal side, 8 (male), 7Ð9 species. (female) ridges ventral side.

Male (holotype, 2 paratypes and 7 vouchers): Length Melomystrongylus sepikensis, new species 3.2Ð4.3 (3.8) mm, maximum width 82.5Ð99 (90.5). Cephalic (Figs. 15Ð28) vesicle 20.5Ð42.5 (33) long, 34Ð39 (37) wide. Oesophagus 280Ð355 (322) long; excretory pore 169Ð221 (186), deirids Material examined. – Holotype male and allotype female: BBM- 204 from cephalic end. Bursa slightly asymmetrical, left NG-104657B, from the small intestine of Melomys rufescens lobe larger; dorsal lobe with median dorsal notch. Dorsal (Muridae: Hydromyinae) type host, 2 km south of Mt Samoro, ray symmetrical, divided at about half its length, each West Sepik, Sanduan Province, Papua New Guinea, coll. A. B. branch dividing again at distal tip; terminal divisions, rays Mirza, 10 May 1975. 9, 10 symmetrical; rays 8 arising at same level proximal to Paratypes: 2 males, 5 females BBM-NG-105128B, from small division of dorsal ray, right ray 8 slightly longer than left. intestine Melomys rufescens, 12 km northwest of Tep Tep, 5¡57'S Rays 4, 5, 6 with common stem, reaching margin of bursa, 146¡05'E, Madang Province, coll. A. B. Mirza, 2 Dec.1975. rays 6, more slender, curving anteriorly, rays 4, 5 curving posteriorly. Rays 2, 3 with common stem, dividing into 2 Voucher specimens: 4 males, 1 female BBM-NG-105155B, widely divergent branches, curving posteriorly, reaching 5 females, 1 male, BBM-NG-105156B, from the small margin of bursa. Genital cone slightly extended, lightly intestine of Melomys rufescens, 12 km northwest of Tep sclerotized, ventral lobe with unpaired papilla 0 and dorsal Tep, 5¡57'S 146¡05'E, Madang Province, coll. A. B. Mirza, lip with paired papillae 7. Spicules equal, filiform, tips 2 Dec.1975; 1 female, BBM-NG-103717, Wanuma, 4¡54'S pointed, 450Ð660 (610) long. Gubernaculum forming thin 145¡19'E, Madang Province, coll A. B. Mirza, 3 Feb.1974; plate 38Ð56 (43) long. 1 male, 2 females, AM W32617, from the small intestine of Melomys rufescens, Sideia Mission, 10¡32'S 150¡48'E, Female (allotype, 4 paratypes and 5 vouchers): Length Milne Bay Province, coll. G. Hangay 31 Dec.1988; 3 3.9Ð5.5 (4.6) mm, maximum width 80Ð114 (100). Cephalic males, 8 females, AM W32557, from the small intestine of vesicle 29.5Ð39.5 (30) long, 33Ð46 (35.5) wide. Oesophagus

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Figs. 15–28. Melomystrongylus sepikensis, new genus, new species, from Melomys rufescens and Melomys spp. from Papua New Guinea: 15, female anterior end, lateral view; 16, male cross section, anterior body; 17, female cross section, anterior body; 18, female cephalic end, lateral view showing synlophe; 19, male cross section, midbody; 20, female cross section, midbody; 21, female posterior end, lateral view showing ovejector; 22, dorsal ray; 23, female cross section, posterior end; 24, female posterior end, ventral view showing well developed praepuce; 25, bursa, left lobe, lateral view; 26, genital cone, lateral view; 27, gubernaculum, lateral view; 28, spicule tips. Scale bars: 15, 24 = 100 μm; 16, 18–20, 22, 23, 25–28 = 25 μm; 17 = 20 μm; 21 = 50 μm.

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310Ð390 (345) long; nerve ring 150, excretory pore 145Ð248 vulva; 35Ð37 ridges in anterior, 35 in midbody of male. Axis (216), deirids 145Ð248 (216) from cephalic end. Vulva with of orientation from ventral right to dorsal left sides, inclined prominent lips, opens 60Ð115 (95) from tail tip. Body wall about 60¡ from sagittal axis. Ridges 1Ð16 increasing slightly extends over vulva and tail as praepuce in mature specimens. in size; ridges 1’–10’ increasing in size, ridges 11’–21’ Monodelphic; ovejector with infundibulum, 90Ð100, longer smaller. Posterior region of body with 28Ð30 ridges reduced than vestibule, 60Ð80, sphincter 30, 50, shortest element. in size; 20 ridges dorsal side, 8Ð10 ridges ventral side. Tail conical, tip rounded, refl ected ventrally, 29Ð42.5 (35) long. Eggs thin-shelled, ellipsoidal, in utero 49Ð66 (57) by Male (holotype and 9 paratypes): Length 2.1Ð3.2 (2.5) mm, 30Ð39 (34). maximum width 67Ð100 (84). Cephalic vesicle 39.5Ð49.5 (43) long, 23Ð29.5 (27) wide. Oesophagus 320Ð520 (390) long; Remarks. – Melomystrongylus new genus has all the nerve ring 130, excretory pore 180, deirids 180 from cephalic characteristics of the subfamily Nippostrongylinae (see end. Bursa slightly asymmetrical, right lobe larger, rays of Durette-Desset, 1985). Lacking a carene and with a right lobe more robust; without dorsal median dorsal notch. hypertrophied ventral ridge Melomystrongylus resembles Dorsal ray symmetrical, divided at about half its length, each Hasanuddinia Hasegawa & Syafruddin, 1994. It can be branch dividing again at distal tip; terminal divisions, rays 9, readily distinguished from Hasanuddinia in that it has a single 10 symmetrical; rays 8 arising at same level proximally to hypertrophied ventral ridge beginning from the excretory pore division of dorsal ray, right ray 8 slightly more robust than and ending in the anterior third of the body. Melomystrongylus left. Rays 4, 5, 6 with common stem, reaching margin of has smaller similar sized ridges showing no clear axis of bursa, rays 6 curving anteriorly, rays 4, 5 curving posteriorly. orientation in the mid and posterior body while the axis of Rays 2, 3 with common stem, diverging distally, curving orientation of the ridges is maintained along the whole length posteriorly, reaching margins of bursa. Genital cone short, of the body in Hasanuddinia (see Hasegawa & Syafruddin, lightly sclerotized, with papillae 0 and 7. Spicules equal, 1994a). The gradient of ridge size in the anterior body in fi liform, tips simple, sharply curved, 340-450 (390) long. Melomystrongylus is from right to left (excepting ridge 5), Gubernaculum 37Ð41 (39) long. but in Hasanuddinia it is from left to right. Female (allotype and 9 paratypes): Length 2.4Ð3.4 (2.8) mm, maximum width 66Ð87 (81). Cephalic vesicle 36Ð52 Odilia similis, new species (43) long, 23Ð33 (28) wide. Oesophagus 250Ð520 (370) (Figs. 29Ð38) long; nerve ring 200, excretory pore 250, deirids 250 from cephalic end. Vulva opens 66Ð72.5 (71) from tail tip. Material examined. – Holotype male and allotype female: BBM- Monodelphic; ovejector with sphincter 40, 40, and vestibule NG-53130, from the small intestine of Melomys rufescens (Muridae: 40, 50, about same length, infundibulum 30, 40, and vagina, Hydromyinae), type host, Kalolo, Morobe Province, Papua New 25, 30, shorter. Tail blunt, conical, tip rounded, 16.5Ð24 Guinea, coll. O. R. & J. W. Wilkes, 28 Aug.1966 (19) long. Eggs thin- shelled, ellipsoidal, in utero 53Ð63 (57) by 32Ð33 (32.8). Paratypes: 1 male, 2 females, SAM AHC34782; collecting data as above. Remarks. – Although the fi xation of this material, in situ in the intestine of the host, was not ideal it was suffi cient Voucher specimens: 2 males, 3 females, BBM-NG-54020 to allow adequate examination of the specimens. Odilia & 20 males, 18 females, BBM-NG- 53691, from the small similis, new species, has every morphological character intestine of Melomys rufescens, 10 km west of Bulolo, 7¡12'S of the genus Odilia (Mawson, 1961) (see Durette-Desset, 146¡39'E, Morobe Province, coll. A. C. Zeigler, 7 Aug.1967; 1983). Following the key to the species of Odilia of Smales 3 males, 2 females, AM W31470, from the small intestine (2005b), O. similis, new species, with 35 ridges in the of Melomys rufescens, Munimun Village, Aguan, 9¡53'S midbody of the male, falls between the group of species 149¡23'E Sanduan Province, coll. P. German, 8 Aug.1992. with up to 35 ridges, O. maxomyos Hasegawa et al. 1999, and O. praeputialis Gibbons & Spratt, 1995, and the group Prevalence. – Four of 25 M. rufescens and one of 10 with more than 35 ridges in the midbody, O. polyrhabdote Melomys spp. (Mawson, 1961) and O. uromyos (Mawson, 1961). Odilia similis, new species, further differs from each of these species Etymology. – The species name indicates that whilst having a in having a spicule to body length ratio of 1: 6.5, simple unique set of characters there are no singular morphological pointed spicule tips and a female tail blunt, conical, neither features to highlight. twisted nor with a praepuce (Smales, 2005b).

Description. – Small coiled worms; prominent cephalic vesicle present; buccal capsule vestigial. Mouth opening Odilia sp. triangular with rudimentary lips; labial and cephalic papillae (Figs. 39Ð43) not observed. Oesophagus claviform. Nerve ring surrounds oesophagus in middle third, deirids and excretory pore Material examined. – 6 males, AM W32552, from the small at about same level, posterior to nerve ring. Synlophe of intestine of Melomys rufescens (Muridae: Hydromyinae), Yapsiei longitudinal cuticular ridges extends from the posterior area, 4¡35'S 141¡5'E, Sanduan Province, Papua New Guinea, coll margin of the cephalic vesicle to just anterior to the bursa or T. Flannery, 17 Jan.1984.

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Figs. 29–38. Odilia similis, new species, from Melomys rufescens and Melomys spp. from Papau New Guinea: 29, female anterior end, lateral view; 30, male cross section, anterior body; 31, bursa, right lateral view; 32, male cross section, midbody; 33, spicule tips; 34, dorsal ray; 35, female posterior end, lateral view; 36, male cross section, posterior body; 37, gubernaculum, lateral view; 38, genital cone, lateral view. Figs. 39–43. Odilia species from Melomys rufescens and Melomys sp. from Papau New Guinea: 39, male cross section, anterior body; 40, dorsal ray; 41, bursa, left lateral view; 42, male cross section, midbody; 43, spicule tips. Scale bars: 29 = 100 μm; 30–34, 37–43 = 25 μm; 35 = 200 μm; 36 = 10 μm.

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Description. – Tiny, coiled worms; synlophe of 16Ð18 Male (holotype and 4 paratypes): Length 2.35Ð2.75 (2.6) mm, continuous ridges in the anterior and midbody. Axis of maximum width 99Ð100 (99). Cephalic vesicle 37.5Ð42.5 (40) orientation of ridges passing through ventral right and dorsal long, 30.5Ð34 (32.5) wide. Oesophagus 320ÐÐ350 (345) long; left sides inclined at about 65¡ from the sagittal axis; gradient nerve ring 165, excretory pore 240 from cephalic end. Bursa of size right to left on dorsal side, left to right ventrally. asymmetrical, right lobe larger, rays of left lobe more robust; with small median dorsal notch. Dorsal ray symmetrical, Male (6 specimens): Length 1.9Ð2.35 (2.08) mm; maximum divided at about 2 thirds its length, each branch dividing again width 47Ð53 (50). Cervical vesicle 23Ð29.5 (26) long; at distal tip; terminal divisions, rays 9, 10 symmetrical; rays oesophagus 280Ð350 (303) long. Bursa asymmetric, right 8 arising at slightly different levels proximally to division lobe larger, dorsal lobe shorter than lateral lobes, with dorsal of dorsal ray. Rays 4, 5, 6 with common stem, reaching notch; dorsal ray divided in distal half; rays 8 arising at margin of bursa, rays 6 more slender than rays 4, 5. Rays same level in dorsal trunk, proximal to branching of dorsal 2, 3 with common stem, robust, diverging distally, curving ray. Spicules fi liform, tips pointed, curved but not hooked, posteriorly, reaching margin of bursa. Genital cone short, 260Ð280 (275) long; gubernaculum13.6Ð15.3 (14.2) long. lightly sclerotized; ventral lobe with unpaired papillae 0, dorsal lip bifi d, each lobe with single papilla 7. Spicules Remarks. – The morphology of the synlophe and bursa of equal, fi liform, proximal ends rounded, tips simple pointed, the present species was congruent with the characteristics of 230Ð290 (265) long. Gubernaculum 34Ð35.5 (35) long. the genus Odilia (Mawson, 1961) (see Durette-Desset, 1983). The suite of measurements and the form of the synlophe, Female (allotype): Length 2.9 mm, maximum width 130. were not however, consistent with any known species. The Cephalic vesicle 42.5 long, 34 wide. Oesophagus 390 long. available specimens were diffi cult to study, with the details Vulva opens 95 from tail tip. Posterior end refl ected dorsally; of the anterior end and bursa concealed by the coiling of the body wall extended to form praepuce. Monodelphic; ovejector body and the infolding of the bursa. And since no females with sphincter, 80, longest, infundibulum, 50, vestibule, 30, were obtained the proposal of a new species is withheld. smallest element. Tail conical tip with spike. Eggs thin- shelled, ellipsoidal, in utero 45 by 30.

Paraheligmonelloides singauwaensis, new species Remarks. – Paraheligmonelloides singauwaensis, new (Figs. 44Ð53) species, is characterized by having a synlophe with ridge 1« much more developed than ridge 1, and with the relative Material examined. – Holotype male, allotype female: BBM-NG- sizes and orientation of the ridges as described for the 24762 from the small intestine of Melomys rufescens (Muridae: southeast Asian members of the genus (Ow Yang et al., Hydromyinae), type host, Singauwa River, Lae, 6¡07'S 146¡59'E, 1983; Hasegawa et al., 1999). Of these P. singauwaensis, Morobe Province, Papua New Guinea, coll. P. J. Shanahan, 7 new species, most closely resembles Paraheligmonelloides Apr.1966. triangulus Ow Yang et al., 1983 in the number of ridges on Paratypes: 3 males, BBM-NG-24762, collecting data as above. the synlophe. It differs from P. triangulus in having more ridges in the midbody (20Ð22 ridges compared with 19Ð21 Prevalence. – Two of 25 M. rufescens. ridges), as well as in the length of the spicules (230Ð290 compared with 61Ð85) the form of the spicule tips and the Etymology. – The species name is taken from the type size of the eggs (45 by 30 compared with 67 by 36) (Ow locality. Yang et al., 1983).

Description. – Small coiled worms; prominent cephalic vesicle present; buccal capsule vestigial. Mouth opening DISCUSSION triangular with rudimentary lips; labial and cephalic papillae not observed. Oesophagus claviform. Nerve ring surrounds Although this study presents information on the helminths oesophagus in middle third; deirids and excretory pore at occurring in M. rufescens, it is limited by the small number about same level, in posterior third of oesophagus length. of hosts collected at each locality and by the examination Synlophe of pointed longitudinal cuticular ridges extends of digestive tracts only from hosts held in museum from the posterior margin of the cephalic vesicle to just collections. These data therefore give conservative estimates anterior to bursa or vulva; 17 ridges in anterior, 20Ð22 in of presence and prevalence of helminth species, with taxa midbody. Axis of orientation of ridges passing from ventral such as Strongyloides being overlooked. In some instances right to dorsal left sides, inclined about 80¡ from sagittal there was insufficient material (the capillariid, and the axis; 8Ð11 ridges dorsal side, 9Ð13 ridges ventral side. heligmonellid species), or all the specimens were immature Ridge 1’ more developed than ridge 1 in anterior; ridges (the helligmonellid larvae) such that identifi cation to species 1’, 2’ more developed than ridges 1, 2 in midbody. Ridges level was not possible. In the case of Peramelistrongylus 1–11 increasing in size; ridges 3’–5’ increasing in size, sp. only females were found. Peramelistrongylus is a ridges 7’–11’ decreasing in size. Posterior region of body monospecifi c genus, described by Mawson (1960) from with 22Ð23 (male) 20Ð21 (female) ridges reducing in size, bandicoots (Peramelidae) in Australia, which has also been retaining axis of orientation. recorded as an occasional infection in Australian rodents,

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including Melomys spp. (Smales et al., 2004; Smales, 2005) (Smales, 2005a) but more than the subset of fi ve species and Hydromys chrysogaster Geoffroy, 1804 (see Mawson, recorded from 20 M. lutillus. When compared, the helminth 1961). There are, however, no records of the genus from assemblages of the two hosts from New Guinea, M. lutillus either bandicoots or rodents in New Guinea, so it cannot be and M. rufescens had in common only two species, the assumed that the female Peramelistrongylus reported here cestode R. celebensis and the trichostrongyloid nematode were Peramelistrongylus skedastos Mawson, 1960. O. mackerrasae, both previously reported in Australia and Papua New Guinea (Smales, 1997, 2005a; Smales et The total number of species recorded from M. rufescens, al., 2004). The other species of helminths occurring in M. 16 from 25 hosts is less than the 22 species recorded from rufescens included the cosmopolitan species H. spumosa 112 grassland melomys; Melomys burtoni and M. cf. burtoni and N. brasiliensis, both usually found in the cosmopolitan from Australia and M. lutillus from Papua New Guinea murine rats Rattus norvegicus (Berkenhout, 1769) and Rattus

Figs. 44–53. Paraheligmonelloides singauwaensis, new species from Melomys rufescens and Melomys spp. from Papau New Guinea: 44, male anterior end, lateral view; 45, male cross section, anterior body; 46, bursa left lateral view; 47, male cross section, midbody; 48, genital cone, ventral view; 49, male cross section, posterior body; 50,genital cone, lateral view; 51, female posterior end, lateral view; 52, spicule tips; 53, dorsal ray. Scale bars: 44 = 50 μm; 45–50, 52, 53 = 25 μm; 51 = 100 μm.

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rattus (Linnaeus, 1758) (see Smales, 1992; Durette-Desset, Hasegawa, H. & Syafruddin, 1994a. Hasanuddinia maxomyos 1970). Their presence in a hydromyine host is indicative of n.gen., n.sp. and Heligmonoides musseri n.sp. (Nematoda: occasional infections acquired from the normal host species. Heligmonellidae) collected from endemic murines of , Species widespread from to Australia were represented Indonesia. Journal of Parasitology, 80: 781Ð788. by C. purvisi (see Hasegawa & Syafruddin, 1994b) and Hasegawa, H. & Syafruddin, 1994b. Cyclodontostomum purvisi species known only from New Guinea by P. kaindiensis (Syn Ancistronema coronatum) (Nematoda: Chabertiidae) and S. longaecauda (see Smales, 2001a, b). from rats of Kalimantan and Sulawesi, Indonesia. Journal of Parasitology, 80: 657Ð660. Apart from O. mackerrasae and N. brasiliensis the Hasegawa, H. & Syafruddin, 1997. Maxomystrongylus yasumai heligmonellid fauna of M. rufescens comprised species or gen. et sp. n. (Nematoda: Trichostrongylina: Heligmonellidae) genera only found in New Guinea (O. similis, new species, collected from murid rodents in Kalimantan, Indonesia. Journal of the Helminthological Society of Washington, 64: 263Ð268. Odilia sp. and M. sepikensis, new species) and species or genera also known from Indonesia or Malayasia, (M. ingens, Hasegawa, H., Miyata, A. & Syafruddin, 1999. Six new nematodes H. mirzai, new species, P. singauwaensis, new species) (this of the Helligmonellidae (Trichostrongylina) collected from endemic murines of Sulawesi, Indonesia. Journal of study; Smales, 2008). This suggests closer relationships Parasitology, 85: 513Ð524. between the helminth fauna of M. rufescens and that of the endemic murids of Indonesia and than that of Holloway J. D. & R. Hall, 1998. SE Asian geology and biogeography: an introduction. In: Hall, R. & J. D. Holloway the endemic murids of Australia, despite the fact that the (eds), Biogeography and Geological Evolution of SE Asia. genus Melomys is found in both geographical regions. An Backhuys Publishers, Leiden The Netherlands. Pp. 1Ð23. examination of the helminth fauna of the extralimital species of Melomys, including M. rufescens from the Molucca Islands Mawson, P. M., 1960. Nematodes belonging to the Trichostrongylidae, Subuluridae, Rhabdiasidae and Trichuridae from bandicoots. could prove informative. Australian Journal of Zoology, 8: 261Ð284. Mawson, P. M., 1961. Trichostrongyles from rodents in Queensland, with comments on the genus Longistriata (Nematoda: ACKNOWLEDGEMENTS Heligmosomatidae). Australian Journal of Zoology, 9: 791Ð826. Special thanks are due to Ms C. Kisinami of the Bishop Menzies, J. I., 1996. A systematic revision of Melomys (Rodentia: Museum and Prof. T. Flannery, formerly of the Australian Muridae) of New Guinea. Australian Journal of Zoology, 44: Museum, for giving me access to the host bodies, to Mr B. 367Ð426. Heinrich for assistance with sorting the specimens and to Prof I. Beveridge who kindly helped with the identifi cation Ow Yang, C. K., Durette-Desset, M.-C., & M. Ohbayashi, 1983. Sur les nématodes parasites de rongeurs de Malaisie. II. Les of the cestodes. Trichostrongyloidea. Annales de Parasitologie Humaine et Comparée, 58: 467Ð492. Smales, L. R., 1997. A review of the helminth parasites of Australian LITERATURE CITED rodents. Australian Journal of Zoology, 45: 505Ð521. Duff, A. & A. Lawson, 2004. Mammals of the World A Checklist. Smales, L. R., 2001a. Syphacia longaecauda n. sp. (Nematoda: A. & C. Black, London. 312 pp. Oxyuridae) Syphacinea from Melomys spp. (Muridae: Hydromyinae) from Papua New Guinea and Irian Jaya, Durette-Desset, M.-C., 1970. Le genre Nippostrongylus Lane, Indonesia. Parasite, 8: 39Ð43. 1923, (Nématode- Heligmosomatidé ) Annales de Parasitologie Humaine et Comparée, 45: 815Ð821. Smales, L. R., 2001b. Protospirura kaindiensis n. sp. (Spirura: Spiruridae) and other helminths from Pseudohydromys Durette-Desset, M.-C., 1983. Keys to genera of the Superfamily (Muridae: Hydromyinae) from Papua New Guinea. Journal Trichostrongyloidea. In: R. C. Anderson & A. C. Chabaud of Parasitology, 87: 169Ð172. (eds.), CIH Keys to the Nematode Parasites of Vertebrates No 10. Commonwealth Agricultural Bureaux, Farnham Royal, Smales, L. R., 2005a. Helminth parasites of the grassland melomys England. Pp. 1Ð85. (Muridae: Hydromyinae) from Australia and Papua New Guinea. Australian Journal of Zoology, 53: 369Ð374. Durette- Desset, M.-C., 1985. Trichostrongyloid nematodes and their vertebrate hosts: Reconstruction of the phylogeny of a Smales, L. R., 2005b. A redescription of Odilia emanuelae parasitic group. Advances in Parasitology, 24: 239Ð306. (Nematoda: Trichostrongylina: Heligmonellidae) from Australian rodents with a key and comments on the genus Durette-Desset, M.-C., Beveridge, I., & D. M. Spratt, 1994. Odilia. Transactions of the Royal Society of South Australia, The origins and evolutionary expansion of the Strongylida 129: 59Ð64. (Nematoda). International Journal for Parasitology, 24: 1139Ð1165. Smales, L.R., 2008. New species of Heligmonellidae (Nematoda: Trichostrongyloidea) from Melomys, Paramelomys and Uromys Flannery, T., 1995. The Mammals of New Guinea. Reed Books, from Papua, Indonesia and Papua New Guinea. Comparative Sydney, Australia. 568 pp. Parasitology, 75: 52Ð60. Godthelp, H., 2001. The Australian rodent fauna, fl otilla’s, fl otsam or just fl eet footed? In Metcalfe, I., J. B. M. Smith, M. Morwood & I. Davidson (eds.), Faunal and Floral Migrations and Evolution in SE Asia- Australasia. A. A. Balkema Publishers, Lisse. Pp. 319Ð322.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 17–24 Date of Publication: 28 Feb.2009 © National University of Singapore

A NEW SYNDYAS LOEW, 1857 (DIPTERA: : ) FROM MANGROVES IN SINGAPORE, WITH A REVIEW OF THE ORIENTAL AND AUSTRALASIAN SPECIES

Patrick Grootaert Department of Entomology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1000 Brussels, Belgium

Ding Yang Department of Entomology, Agricultural University, Beijing 100094, China (Corresponding author)

ABSTRACT. – Syndyas singaporensis, new species, is described from mangroves in Singapore. Syndyas aterrima de Meijere, 1913 (Ceram, Indonesia), S. brevior de Meijere, 1910, (Krakatau, Indonesia) and S. elongata de Meijere, 1910, (Java, Indonesia) are redescribed. Lectotypes are designated for S. elongata and S. brevior. A key, as well as a checklist, are given for all Oriental and Australasian species.

KEY WORDS. – Diptera, Hybotidae, Syndyas new species, Oriental, Australasian, mangrove.

INTRODUCTION holotype female from Ceram (Australasia), Syndyas brevior de Meijere, 1910, described from two females, from Krakatau The subfamily Hybotinae is rare in Singapore. This subfamily and a lectotype is designated for the specimen bearing the radiated in regions with a temperate climate such as the label “type”. Finally, Syndyas elongata de Meijere, 1910, Holarctic and it is far less common in tropical Oriental described from several specimens from Krakatau and conditions. On a total of 544 samples recently taken in Jakarta is also redescribed and a lectotype is designated. It Singapore, only 17 samples yielded Hybotinae (Shamshev was not necessary to redescribe the other Oriental species & Grootaert, 2007) comprising two records of Syneches since they have good recognizable characters as can be seen Walker, fi ve records of Hybos Meigen, and ten records of in the new key that is presented here for all Oriental and Syndyas Loew. The Tachydromiinae on the other hand, Australasian species. another hybotid subfamily, is more common and also more diverse. For example, during the same sampling campaign It is remarkable that the new Syndyas species was only found 987 individuals were recorded from the tachydromiine genus in mangrove. As demonstrated elsewhere, the mangroves in Elaphropeza Macquart, belonging to 52 species (Shamshev Singapore have an unexpected high diversity of empidoid & Grootaert, 2007). fauna (Evenhuis & Grootaert, 2002; Grootaert, 2006a, 2006b; Grootaert & Meuffels, 2001; Zhang et al., 2006, 2007) and Syndyas is a genus of rather small Hybotinae that look merit a special attention from a faunistical point of view as quite fragile in comparison to the stouter Hybos, Parahybos well for conservation. Adult Syndyas are predators and the Kertész, Chilcottomyia Saigusa, and Syneches. Syndyas is larval stages are unknown. cosmopolitan in distribution. It is rare in the Palaearctic where only three species are known. It is most diverse in the Afrotropical Realm with 14 species, followed by the Oriental MATERIALS AND METHODS Realm with nine species. The Australasian Realm has three, the Nearctic six and the Neotropical only one species. For sampling of the material in Singapore and the administrative codes used, we refer to the papers of Shamshev In the present paper, we describe a new species of Syndyas & Grootaert (2007) and Zhang et al. (2008). The holotype from Singapore and in order to confi rm the identity of the of the new species is housed in the collections of the Raffl es new species it was necessary to revise the type material of Museum, National University of Singapore (ZRC) and some the closest related species. Here, we redescribe Syndyas paratypes are deposited in the collections of the Royal aterrima de Meijere, 1913, originally described from the Belgian Institute of Natural Sciences in Brussels (RBINS).

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De Meijere’s material is stored in the collections of the – Veins R4+5 and M converging towards wing tip, but turning up Zoölogisch Museum Amsterdam (ZMAN). The information just before meeting the costa. Hind femur ventrally with a row on the labels is placed between square brackets […]. of long spiny bristles ...... 10 10 Hind femur strongly swollen (club-shaped). Tip of right epandrial lamella bifurcate, outer lobe bearing broad black bristly hairs (Singapore) ...... S. singaporensis, new species SYSTEMATIC ACCOUNT – Hind femur weakly swollen. Tip of right epandrial lamella bifurcate, outer lobe with only ordinary hairs (North Palaearctic, Syndyas Loew, 1857 Oriental China – Guizhou, Hainan) ...... S. nigripes (Zetterstedt) Genus diagnosis and redescription in Chvála, 1983: 99.

Species with holoptic eyes. Third antennal segment elongate Syndyas aterrima de Meijere, 1913 ovoid with apical arista. Proboscis strong, directed forward. (Figs. 3, 6, 11) Hind femur generally thickened, club-shaped hind tibia and thickened hind fi rst tarsomere. Basal section of vein M, the Syndyas aterrima de Meijere, 1913: 53, Fig. 12 (wing). vein separating the two basal cells scarcely visible (distinct in both Hybos and Syneches). Material examined. – Holotype female, INDONESIA, [Ceram, 25.II] [Mevr. De Beaufort leg. 1910] [Syndyas aterrima det. de The species resemble one another in leg setation, general body Meijere Type] Red label [Syndyas aterrima de Meijere, 1913 ZMAN type DIPT.0473.1]. Female on pin, in bad condition: head and leg colour. Colour of wing (clear, yellowish or brownish missing, wings missing on thorax, but mid leg and one wing in a tinged), presence or absence of a wing stigma, length of the dry slide attached to pin. discal cell in relation to length of basal cells are reliable characters. The shape of the fore tibia (thickening caused Diagnosis. – Species with a brownish wing with a large by the development of the tibial gland), hind femur, hind brown stigma. Vein R4+5 and M straight. Hind femur spindle- tibia, hind tarsomere 1 are also useful characters. The male shaped with a ventral row of interspaced bristles that are genitalia are quite distinct, but the genitalia of few species not spiny. have been fi gured and several species are only known from the holotype female. Redescription. – Female: length: 2 mm; wing: 2.5 mm.

Head (missing, description after de Meijere, 1913) Ocellar KEY TO ORIENTAL AND AUSTRALASIAN tubercle with 2 short black bristles. Eyes touching on frons, SYNDYAS face very narrow, linear. Antenna black, medium-sized; third segment elongate oval, with a black, somewhat subdorsal 1 Discal cell shorter than basal cells ...... 2 arista that is much longer than the three antennal segments – Discal cell as long as basal cells ...... 4 2 Discal cell very short, basal cells nearly 3 times longer. (Sri combined. Proboscis and palp shining black. Lanka, Papua New Guinea) ...... S. parvicellulata Bezzi – Discal cell longer ...... 3 Thorax shining black, with a green metallic tinge when seen 3 Basal cells about 1.5 times as long as discal cell. Mesonotum from behind. Hairs and bristles darkened but with a pale black, without blue shine. Abdomen not elongated (Indonesia: tinge. Posterior part of mesonotum covered with a brown Krakatau) ...... S. brevior de Meijere dusting. Pleura also brown dusted. Dorsocentrals not distinct – Basal cells almost as long as discal cell. Mesonotum black in from other pilosity. Scutellum with a pair of black apical ground-colour, with a blue shine. Abdomen long and slender bristles and at least 4 black marginal hairs half as long as S. elongata (Indonesia: Krakatau) ...... de Meijere apicals, on the side (rubbed off on one side). 4 Tarsomeres 1 and 2 of all legs yellow (Taiwan) ...... S. orientalis Frey – All tarsomeres black ...... 5 Legs shining black. Fore femur without ventral bristles. Fore 5 Wing clear (India, ) ...... S. vitripennis Brunetti tibia spindle-shaped (Fig. 6), much swollen at basal third – Wing not clear, but yellowish to brownish tinged ...... 6 (opening of tibial gland not on a tubercle); a double row of 6 Wing yellowish tinged, no stigma (Philippines) ...... long pale posteroventral bristles, twice as long as tibia is wide...... S. dapana Frey The rows continue on the fi rst tarsomere. Hind femur (Fig. – Wing brownish tinged, usually with stigma (except S. 11) slender in basal half, thickened in apical half, ventrally eumera) ...... 7 with a row of black interspaced bristly hairs, in basal half 7 Halter dark yellow, wing brown with dark brown stigma (China: the bristles are a little longer than femur is wide; in apical Zheijang) ...... S. sinensis Yang & Yang – Halter black ...... 8 half the bristles are a little shorter than femur is wide. A 8 Brown stigma present. Hind femur club-shaped (less distinct few long pale dorsal hairs near base. Hind tibia thickened in S. nigripes) ...... 9 on apical third. First tarsomere on hind leg swollen, shorter – No stigma present. Hind femur very thickened (thickest near than apical four tarsomeres together. middle) (Papua New Guinea) ...... S. eumera Bezzi

9 Veins R4+5 and M are straight running parallel toward costa. Hind Wing (Fig. 3) brownish, a large stigma present (see femur ventrally with a row of bristles (not spiny). (Indonesia: discussion). Discal cell longer than basal cells, elongate Ceram) ...... S. aterrima de Meijere rectangular, nearly 3 times as long as vein closing the discal

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cell (dm-cu). Vein R3+4 and M straight, ending parallel in Postoculars short, black, with tips proclinate over the eye. costa. Halter black. Antenna black, third segment elliptical 3 times as long as wide. Arista missing. Palp and proboscis hidden. Abdomen shining black, with paler hairs than on mesonotum, from some point of view appearing whitish. Thorax black in ground-colour, with thin pale brownish pubescence. Mesonotum shining black on anterior 2/3, Comments. – As can be seen on Fig. 3 the stigma is very posterior third thinly covered (subshining) with brownish

large, but it is not clear if the area before the tip of R1 is dusting. Pleura grey dusted. Large thoracic bristles brown. also darkened. De Meijere (1913) indicates only a long oval Acrostichals long biseriate, the rows widely separated;

stigma on his fi gure between R1 and R2+3. dorsocentrals uniseriate but with numerous hairs at sides, ending in a long prescutellar. A pair of long brown apical Frey (1938) reports this species from the Philippines (3 scutellars and some shorter marginal hairs on both sides. females from Luzon and a male and female from Mindoro). These specimens should be re-examined to see if they are Legs brownish-black. Fore femur without ventral bristles, a really conspecifi c with the female holotype of S. aterrima. few fi ne black posteroventrals near tip. Fore tibia (Fig. 8) somewhat curved in dorsal view, slightly dilated, especially around the opening of the tibial gland, opening of the tibial Syndyas brevior de Meijere, 1910 gland not distinct; tibia with 2 short dorsal bristles and with (Figs. 2, 8, 10) a row of long posteroventrals, a little longer than tibia is wide, the row continuing on tarsomere 1. Tarsomere 1 with Syndyas brevior de Meijere, 1910: 68, Fig. 5 (wing). some long dorsal bristles, especially a basal one and a pair of apicals; following tarsomeres also with distinct dorsal Material examined. – Lectotype female INDONESIA [E. Jacoboson hairs. Mid femur ventrally bare. Mid tibia with 4-5 brown Lang Eiland Krakatau Mei 1908] [Syndyas brevior det. de Meijere dorsal bristles, twice as long as tibia is wide and a very type] red label [Syndyas brevior de Meijere 1910 ZMAN type long ventral preapical; a row of long brown posteroventrals. DIPT.0476.1]; 1 female [E. Jacobson. Verlaten Eiland Krakatau Mei 1908] red label [Syndyas brevior de Meijere 1910 ZMAN Mid tarsomere 1 with two long dorsals and a row of long type DIPT.0476.2] posteroventrals and a long ventral near base. Hind femur club-shaped, slender on basal third, gently widening towards Diagnosis. – A small species with clear wing membrane, tip, near tip about twice as wide as at base, ventrally with no stigma and discal cell much shorter than basal cells. a row of equally long spine-like bristles. Hind tibia club- Hind femur thickened on apical 2/3 with a ventral row of shaped, narrow at base but quickly widening towards tip, at spiny bristles, about half as long as femur is wide. Hind its widest part, 3 times as wide as at base (Fig. 10) dorsally tarsomere slightly thickened and much shorter than following with 3-4 long black bristles, in addition a pair of preapical tarsomeres together. dorsal bristles. Hind tarsomere 1 thickened, about half as long as following four tarsomeres together; with a pair of Redescription. – Female: length: 2 mm; wing: 1.75 mm. long preapical bristles.

Head. Eyes holoptic, frons and face linear. A pair of Wing (Fig. 2) clear, without stigma. Discal cell short; basal proclinate ocellars as long as all antennal segments together. cells about 1.5 times as long as discal cell. Vein R4+5 and

Figs. 1–4: Syndyas wings: 1, S. singaporensis, new species; 2, S. brevior de Meijere; 3, S. aterrima de Meijere; 4, S. elongata de Meijere.

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M slightly converging near wing tip and ending parallel Redescription. – Male: length: 3.5 mm; wing: 2.5 mm. into costa. Head. Eyes holoptic, frons and face linear. A pair of Abdomen shining black. Tergites dorsally with short pale proclinate ocellars as long as all antennal segments together. brownish bristles; basal tergite with long brown bristles at Postoculars long above, black, with tips proclinate over the sides. Apical two segments and cerci brown dusted. eye. Antenna black, third segment elliptical 2.5 times as long as wide. Arista at least 3 times as long as all antennal Comments. – Frey (1938) reports a male from the Philippines, segments combined. Palp and proboscis hidden. unfortunately not giving any details. It is clear that in the future these specimens should be examined in order to Fore tibia (Fig. 7) dorsoventrally curved, only a little try to establish conspecifi city with the holotype female of thickened; gland opening a little prominent, with a row of very long pale posteroventral bristles. Hind femur very Krakatau. slender on basal 2/3, only swollen on apical third (Fig. 12). Ventrally a row of fi ne bristles a little longer than femur is wide becoming spine-like on apical third (the club-shaped Syndyas elongata de Meijere, 1910 part). Hind tibia thickened on apical half, with at least 3 pairs (Figs. 4, 7, 12) of long fi ne dorsal bristles, the preapical pair the longest. Hind tarsomere 1 thickened and a little longer than following Syndyas elongata de Meijere, 1910: 67, Fig. 4 (wing). tarsomeres together. Material examined. – Lectotype male (designated here) INDONESIA Mesonotum shining black, viewed from behind with a [E. Jacobson. Verlaten Eiland Krakatau Mei 1908] white label in long hand [Syndyas elongata det. de Meijere Type] red label greenish blue metallic shine. Posterior third of mesonotum [Syndyas elongata de Meijere 1910 ZMAN type DIPT.0478.1]; covered by thin brown dusting. Pleura with the usual thin 3 females, same provenance as lectotype. 1 female [E. Jacobson brown dusting and so only subshining. Scutellum with a pair Batavia Sept 1908]; 1 male, 1 female [Java Jacobson]. The latter of brown apical bristles and at least 2 hairs at each side. specimens are probably from Tandjong Priok near Batavia, November, Jacobson leg as mentioned in de Meijere (1910), but Wing (Fig. 4) clear with a faint yellowish stigma. Veins there is nothing additional on the label. brown. Discal cell a little shorter than basal cells. Squama black with very long, pale cilia. Halter black. Diagnosis. Slender species with long slender abdomen and wings. Wing clear with a faint yellowish stigma. Discal cell Abdomen long and slender, covered with short black a little shorter than basal cells. Mesonotum shining black hairs, even at base of basal segments. Genitalia as wide as with a bluish tinge. abdomen.

Figs. 5–8. Syndyas fore tibia posteriorly: 5, S. singaporensis, new species; 6, S. aterrima de Meijere; 7, S. elongata de Meijere; 8, S. brevior de Meijere.

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Female: length: 3.2–3.7 mm; wing: 2.4–2.5 mm. Identical Paratypes: SINGAPORE, 1 female same provenance as holotype. in most respects to male. Sungei Buloh, mangrove: 1 male, 1 female, 9 Dec.2002 (22057, Si301); 1 male, 1 Jun.2005 (25165, Si1639); 1 male, 22 Jun.2005 Comments. – The male from Krakatau was selected as (25171, Si823; on pin, genitalia dissected); 1 male, 1 female, 6 Jul.2005 (25199, Si881); 1 male, 1 female, 20 Jul.2005, (25264, lectotype not only because its locality was mentioned fi rst Si1056); 1 male, 27 Jul.2005 (25272, Si289); 1 female, 7 Sep.2005 in the description, but also because it is a more precise site (25264, Si1056); 1 female, 27 Nov.2003, sweeping (23092, Si92); than Jakarta (Batavia). 1 female, 13 Jan.2006 (26015, Si1581, RBINS). Chek Jawa, Pulau Ubin, mangrove, 1 male, 1 female, 19 Nov.2002, sweeping (leg. P. Grootaert, RBINS). Syndyas singaporensis, new species (Figs. 1, 5, 9, 13) Diagnosis. – Species with brown wing and brown stigma. Discal cell almost as long as basal cells. Mesonotum Material examined. – Holotype male: SINGAPORE: Sungei Buloh, anteriorly shining black. Hind femur club-shaped, ventrally 7 Sep.2005, Malaise trap 1 (25336, leg. P. Grootaert, Si1055, in with spine-like bristles. ZRC of Raffl es Museum, NUS, Singapore).

Figs. 9–12. Syndyas hind leg: 9, S. singaporensis new species; 10, S. brevior de Meijere; 11, S. aterrima de Meijere; 12, S. elongata de Meijere.

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Description. – Male: Length: 3.0–3.2 mm, wing 3.0–3.2 basal one and a pair of apicals. Following tarsomeres also mm. with distinct dorsal hairs. Mid femur ventrally bare. Mid tibia with 4–5 black dorsal bristles, 3 times as long as tibia Head. Eyes holoptic, frons and face linear. A pair of proclinate is wide and a very long ventral preapical; a row of long ocellars as long as all antennal segments together. Postoculars brown posteroventrals. Mid basal tarsomere with two long short, black, with tips proclinate over the eye. Antenna black, dorsals and a row of long posteroventrals and a very long third segment elliptical 3 times as long as wide. Arista bare, ventral near base. Hind femur (Fig. 9) club-shaped, slender 3 times as long as antennal segments together. Palp elongate, on basal half, gently widening towards tip, near tip about black with a long black dorso-apical bristle. 1.5 times as wide as at base. Ventrally with a row of equally long spine-like bristles, in basal half the spines are a little Thorax black in ground-colour. Mesonotum shining black longer than femur is wide; in apical half about as long as femur is wide. Hind tibia club-shaped, narrow at base but on apical half, posterior half thinly covered (subshining) abruptly widening towards tip, at its widest part, 3 times with thin brownish dusting. Humeri brownish (paler than as wide as at base (Fig. 9), dorsally with 3-4 long black rest of mesonotum). Pleura grey dusted. Large thoracic bristles, in addition a pair of preapical dorsal bristles. Hind bristles brown. Acrostichals long, biseriate; dorsocentrals tarsomere 1 as long as following four tarsomeres together; uniseriate but with numerous hairs at sides. A pair of long much widened; with a few long bristles near tip. brown apical scutellars and a series of shorter marginal hairs on both sides. Wing (Fig. 1) brownish clouded, stigma dark brown occupying the whole apex of cell R (between tip of R and Legs. Anterior four legs shining brownish black; posterior 1 1 R2+3. The vein separating basal cells hardly prominent. Discal pair intensively black. Fore coxa with short black hairs cell almost as long as basal cells, elongate rectangular (almost anteriorly. Fore femur without ventral bristles, a few fi ne three times as long as wide). Apical portion of vein Cu, 1.5 black posteroventrals near tip. Fore tibia (Fig. 5) slightly times as long as vein closing the discal cell (dm-cu). Veins spindle shaped with a very prominent opening of the tibial R and M slightly diverging near middle, then converging gland in the form of a tubercle, bearing a hair-like structure 4+5 and diverging again before meeting costa. Halter black. at tip; with a row of long posteroventrals, a little longer than tibia is wide; the row continues on fore tarsomere 1. Fore Squama and ciliation brown. basal tarsomere with some long dorsal bristles, especially a

Fig. 13. Syndyas singaporensis, new species. Paratype male genitalia. REL, right epandrial lamella; ph, phallus; LEL, left epandrial lamella.

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Abdomen shining black. Tergites dorsally with short pale Grootaert, P., 2006a. Dolichopodidae in mangroves of Southeast brownish bristles; basal three tergites with very long pale Asia: diversity, community structure, zonation and phenology: bristles at sides. Sternites narrow, shining brownish black, a case study in Singapore. 6th International Congress of with few moderately long dark hairs. Genitalia (Fig. 13) with Dipterology, Fukuoka, 23–28 September 2006: 91–92. tip of right epandrial lamella bifurcate; outer lobe bearing Grootaert, P. 2006b. The genus Teuchophorus (Diptera, fl attened black bristly hairs, inner lobe with a row of ordinary Dolichopodidae) in Singapore. The Raffl es Bulletin of Zoology, hairs. Left epandrial lamella at tip also with fl attened black 54: 59–82. bristly hairs. Cerci rather short. Tip of phallus very broad. Grootaert, P. & H. Meuffels, 2001. A note on the marine dolichopodid fl ies from (Insecta, Diptera, Dolichopodidae) The Female: length: 2.9–3.1mm; wing: 3 mm. In most respects Raffl es Bulletin of Zoology, 49: 339–353. identical to male. De Meijere, J.C.H., 1910. Studien über südostasiatische Dipteren. IV. Die neuen Dipterenfauna von Krakatau. Tijdschrift voor Comments. – Syndyas singaporensis, new species superfi cially, Entomologie, 53: 58–194; Taf. 4–8. resembles S. aterrima de Meijere from Ceram. The wing De Meijere, J.C.H., 1913. Praeda Itineris a L.F. de Beaufort in

in the latter species is also brownish tinged, but veins R4+5 Archipelaga indico facti annis 1909-1910. VI. Dipteren I. and M are straight and run parallel toward costa. The hind Bijdragen tot de Dierkunde, 19: 45–69. femur bears a row of long ventral hairs and no spine-like Shamshev, I., & P. Grootaert, 2007. Revision of the genus bristles as in the new species. Elaphropeza Macquart (Diptera: Hybotidae) from the Oriental Region, with a special attention to the fauna of Singapore. S. singaporensis, new species, also resembles S. nigripes Zootaxa, 1488: 164 pp, 261 Figs. (Zetterstedt) that has however a slender hind femur with Yang, D., 2004. One new species of Syndyas with key to species little dilated tip while the hind femur is more distinctly from China (Diptera: Empidoidea). Transactions of the American club-shaped in S. singaporensis, new species. The tip of Entomological Society, 130: 91–94. the right surstylus is bifurcate in both species, but in S. Zhang, L., D. Yang & P. Grootaert, 2007. Paraclius (Diptera: singaporensis, new species, it bears fl attened black bristly Dolichopodidae: Dolichopodinae) of Singapore, with new hairs while only ordinary hairs in S. nigripes. The shape species from mangroves. The Raffl es of Bulletin of Zoology, of epandrial lamellae, length of cerci and shape of phallus 55: 49–62. are completely different in both species. Nevertheless we ZHANG, L., YANG, D. & P. Grootaert, 2007. Mangrove Hercostomus suppose that both species are closely related. sensu lato (Diptera: Dolichopodidae) of Singapore. The Raffl es Bulletin of Zoology, 56: 17–28.

ACKNOWLEDGEMENTS Checklist of Oriental and Australasian Syndyas The fi rst author thanks Prof. Dr. Peter Ng and Prof. Dr. Rudolf Meier for hosting him during his sabbatical stay in Syndyas aterrima De Meijere, 1913. Oriental: Philippines? Singapore in 2005. He also thanks NParks for issuing the Australasian: Indonesia: Ceram. research permits and their interest in the study. We thank Syndyas aterrima De Meijere, 1913, Bijdr. Dierk. 19: 53. Type Mr. Ben Brugge (Amsterdam) for sending us de Meijere’s locality: Indonesia: Ceram. type material. Dr. Igor Shamshev helped with some literature. Andreas Stark and Adrian Plant gave many useful comments Syndyas brevior De Meijere, 1910. Oriental: Indonesia, on the manuscript. Finally PG thanks Dr. Luca Bartolozzi for Philippines?. Syndyas brevior De Meijere, 1910. Tijdschr. Ent. 53: 68. Type his help with the translation of the Italian text of Bezzi. localities: Indonesia: Krakatau, “Lang Eiland” and “Verlaten Eiland”.

LITERATURE CITED Syndyas dapana Frey, 1938. Oriental: Philippines. Syndyas dapana Frey, 1938. Not. Ent. 18: 62. Type locality: Bezzi, M., 1904. Empididi Indo-Australiani raccolti dal signor L. Philippines: Dapa, Siargao. Biró, Annales Historico-Naturales Musei Nationalis Hungarici, 2: 320–361. Syndyas elongata De Meijere, 1910. Oriental: Indonesia. Chvála, M., 1983. The Empidoidea of Fennoscandia and Denmark. Syndyas elongata De Meijere, 1910. Tijdschr. Ent. 53: 67. Type II. General Part. The families Hybotidae, Atelestidae and localities: Indonesia: Krakatau, Verlaten Eiland; Batavia [Jakarta], Microphoridae. Fauna entomologica Scandinavica, 12: 279pp. Tjandjong Priok near Batavia (Jakarta), Java. Scandinavian Science Press, Copenhagen. Syndyas eumera Bezzi, 1904. Australasian: Papua New Evenhuis, N. & P. Grootaert, 2002. Annotated checklist of the Guinea. Dolichopodidae (Diptera) of Singapore, with new records and Syndyas eumera Bezzi, 1904. Ann. Hist. Nat. Mus. Natl. Hung. 2: descriptions of new species. The Raffl es Bulletin of Zoology, 323. Type locality: Papua New Guinea: Sattelberg. 50: 301–316. Frey, R., 1938. Hybotinen (Dipt, Empididae) von Formosa und den Syndyas nigripes (Zetterstedt, 1842). Palaearctic: Austria, Philippines Notulae Entomologicae, 18: 52–62. Belgium, China, Czech, England, Finland, Germany, Hungary, Italy, Netherlands, Poland, Russia, Sweden, Switzerland? Oriental: China.

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Ocydromia nigripes Zetterstedt, 1842. Dipt. Scand. 1; 240. Type Syndyas sinensis Yang et Yang, 1995. Oriental: China. locality: Sweden: Suecia media, Ölandia [= Öland]. Syndyas sinensis Yang et Yang, 1995 of Baishanzu Mountain, Eastern China: 503. Type locality: China: Zhejiang: Baishanzu. Syndyas orientalis Frey, 1938. Oriental: China (Taiwan). Syndyas orientalis Frey, 1938. Not. Ent. 18: 61. Type locality: Syndyas singaporensis, new species. Oriental: Singapore. Type China: Taiwan, Chosokei. locality: Singapore, Sungei Buloh.

Syndyas parvicellulata Bezzi, 1904. Oriental: Philippines, Sri Syndyas vitripennis (Brunetti, 1913). Oriental: India, Myanmar. Lanka? Australasian: Papua New Guinea. Leptopeza vitripennis Brunetti, 1913. Rec. Ind. Mus. 9: 31. Type Syndyas parvicellulata Bezzi, 1904. Ann. Hist. Nat. Mus. Natl. locality: India: Oncha Gaon, Naini Tal; Myanmar: Rangoon. Hung. 2: 321. Type localities: : Colombo; Papua New Guinea: Sattelberg.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 25–27 Date of Publication: 28 Feb.2009 © National University of Singapore

ECTOPOMYIA HANCOCKI, A NEW SPECIES OF ACANTHONEVRINE (DIPTERA: TEPHRITIDAE: PHYTALMIINAE) FROM PENINSULAR MALAYSIA

Chua Tock Hing Monash University, Sunway Campus, Jalan Lagoon Selatan, 461150 Bandar Sunway, Selangor Darul Ehsan, Malaysia Email: [email protected]

ABSTRACT. – A new species of acanthonevrine fl y from Peninsular Malaysia is described and illustrated. Ectopomyia hancocki, new species, is the second species of the genus to be described. A key to Ectopomyia spp. is given. E. hancocki, new species, may be separated from E. baculigera Hardy, by the wing pattern and body markings.

KEY WORDS. – Acanthonevrini, Phytalmiinae, Ectopomyia hancocki, new species, Peninsular Malaysia.

INTRODUCTION SYSTEMATICS

The subfamily Phytalmiinae, considered to be monophyletic Tephritidae Newman, 1834 (Korneyev, 2001), consists of four tribes: Acanthonevrini, Phytalmiini, Phascini and Epacrocerini. Acanthonevrini Phytalmiinae Hendel, 1914 is equivalent to the subtribe Acanthonevrina sensu Hardy (1986). Acanthonevrini Hardy, 1970

The Acanthonevra subgroup of genera (e.g. Ectopomyia, Ectopomyia is morphologically similar to Hexacinia Hendel, Hexacinia and Rioxa) in Acanthonevrini are characterized by based on two pairs of inferior fronto-orbital bristles that are a strong anepimeral bristle, two orbital bristles, long and broad very close together, with the lower bristle incurved and upper surstyli, very long and wrinkled preglans areas of distiphallus, bristle reclinate. Hexacinia is represented by four species: oval spermathecae with long and bulbous necks and apical H. radiosa (Rondani), H. stellipennis (Walker), H. pellucens nipples, and yellow head bristles (Korneyev, 2001). Hardy, and H. punctifera (Walker) (Hardy, 1986). Only the fi rst two species occur in Malaysia (Chua, 1999; Hancock & Species grouped under Acanthonevrini infest damaged or Drew, 1994, 1995). These two genera and the two species decaying plant matter, including bamboo shoots, bamboo of Ectopomyia may be separated by the following key. stems, tree bark and, rarely, fruit. Although many species have been described, the detailed biology of most species remains unknown. This includes the species described here. KEY TO HEXACINIA AND There is no host information available for Ectopomyia or ECTOPOMYIA SPECIES Hexacinia but, given their apparent relationship with Rioxa Walker, it is likely that they breed beneath tree bark. 1. Wing shape broad, about 2× longer than wide (Platensina-like); numerous small hyaline spots around margin and in the fi eld; The genus Ectopomyia was created by Hardy (1973) and with a distinct bristle in lower median portion of anepisternum; has only one previously recorded species, viz. Ectopomyia male genitalia normal; male fore-femur not modifi ed ...... Hexacinia baculigera from . A female Ectopomyia specimen was – Wings normal shape with large hyaline spots at apex and on collected from Pahang, Peninsular Malaysia and found to hind margin, fewer hyaline spots in the fi eld; may lack a distinct belong to a new species as confi rmed by Dr. David Hancock bristle in lower median portion of anepisternum; male genitalia (Australia). Here, the new species is described and a key to has unusual development, with the surstyli folding under venter Hexacinia and Ectopomyia spp. is presented. of abdomen and plainly visible in situ; male fore femur has a subbasal appendage (at least recorded in baculigera Hardy) ...... 2

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2. General body colour pale yellow; Scutum with 2 lateral yellow of frons and two pairs of orbital bristles; ocellar bristles longitudinal streaks; wings: hyaline area in sc crossing R1 into rudimentary, reduced to small hairs scarcely larger than the r1, r1 with most 1 hyaline area, hyaline areas in cells r2+3 and setae in interfrontal area; the frontals close together with m large and rounded; 1 large hyaline area in m; r2+3 with a the lower pair incurved and the upper pair reclinate. Face rounded hyaline area just apical to the crossvein r-m. Laos short, slightly concave in median portion in lateral view, ...... E. baculigera Hardy – General body colour light brown; scutum with 3 brown with epistomal margin produced rather prominently. Lower longitudinal streaks; wings: hyaline area in sc not crossing 2/3 of occiput swollen, upper portion sharply narrowed as

R1, r1 with 2 hyaline areas, hyaline areas in cell r2+3 and r4+5 seen in direct lateral view. Oral margin with a few yellow-

elongate, 2 small hyaline areas in m; r2+3 without hyaline area brown setae in front and rather densely yellow setose along in the middle. Malaysia ...... E. hancocki, new species hind portion over genae and lower occiput. Posteroventral margin of occiput with a dense band of mixed yellow and brown hairs; also with a dense clump of dark brown hairs on Ectopomyia hancocki, new species posterodorsal margin of occiput immediately above cervix. (Figs. 1–3) Antennae as long as face, brown with median portion of 3rd

Material examined. – Holotype. 1 female (ZRC), collected in bamboo grove at Fraser’s Hill, Pahang, Peninsular Malaysia, coll. T. H. Chua, 3 Mar.1996.

Distribution. – Peninsular Malaysia.

Description. – Female: length of body including oviscape 6.5 mm; wings. 5.0 mm (Fig. 1).

Head generally light brown (Fig. 2), with a dark brown narrowing mark extending from ocellar triangle down median portion of frons reaching the edge of lunule, remainder of frons golden pollinose, clear yellow along eye orbits. Face entirely light brown. Genae and lower 2/3 of occiput light brown, upper occiput (except the edge) shining dark brown to black on sides, light brown in median portion. Head length slightly shorter than high (7:8). Eyes oval. Frons fl at, slightly sloping as seen in lateral view, 5.6 wide: 4 long and wider than an eye (3:2.5) as seen from direct dorsal view. Head with two pairs of frontal bristles on lower 1/4

Fig. 2. Ectopomyia hancocki, new species, lateral view of female head

Fig. 3. Ectopomyia hancocki, new species, lateral view of female Fig. 1. Ectopomyia hancocki, new species, female. thorax.

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segment orange brown. Third antennal segment rounded at slightly wider than long, concave on posteromedian portion, apex, arista plumose; the longest plumes are slightly less with 2 strong and at least 2 thinner bristles on hind margin. than the width of 3rd segment. Palpi pale brown, sparsely Oviscape dark brown, as long as tergites III–VI, eversible brown setose. Labella fl eshy, with numerous rather long membrane brown with black lateral margins. pale hairs on mentum. Male: unknown. Thorax largely sub-shining brown with dark brown markings on dorsum and with brown to black fi ne setae. Anterior margin Remarks. – As the male of E. hancocki, new species, is of scutum and postpronotal lobe light yellow. Scutum with unknown, it is not known if male front legs have a subbasal 3 brown longitudinal streaks: one median streak extending appendage on front femur as seen in E. baculigera. The from anterior margin to dorsocentral bristle line, two two species can be differentiated by differences as noted lateral brown streaks each extending from area above the in the key to species. In addition, the wing is less broadly

postpronotal lobe to almost the posterior margin of scutum. rounded apically in E. hancocki, new species,vein R4+5 does Two brown markings also present on posterior part of scutum not intersect the white apical spot at the wing margin and

on both sides of median marking and extend posteriorly to veins R2+3 and R4+5 are not distinctly divergent over apical both sides of scutellum. Posteromedian portion of scutum third. Although it was caught in the bamboo groves, we yellow between acrostichal bristles; this yellow stripe is have no information on its host. continuous over middle part of scutellum. Hind margin and venter of scutellum pale yellow. Pleura light brown (Fig. 3) Etymology. – This species is named after Dr. David Hancock except for darker brown spots in proepimeron, anepimeron, who has contributed greatly to the study of Tephritids in anepisternum, katepisternum, and metapleuron. Mediotergite the Asian region. and postscutellum brown except for a yellow to rufous mark down the median portion. Bristles black and well developed: 1 pair scapulars, 1 postpronotal, 2 notopleurals, 1 ACKNOWLEDGEMENTS presutural supra-alar, 1 postsutural supra-alar, 1 dorsocentral, 1 acrostichal, 1 intra-alar, 3 pairs scutellars; Acrostichals in I thank Monash University, Sunway Campus for the general line with intra-alars; median scutellars shorter than other research facilities, and Miss Magdalene Chong for the scutellar bristles. A strong anepimeral bristle present. illustration, and Dr. David Hancock (Australia) for helping with the identifi cation of the specimen and giving constructive Legs entirely pale brown. Front femur also with strong comments on an earlier draft. anteroventral bristles. Middle tibia with 1 strong apical spur. Tarsi densely setose ventrally with 2nd tarsomere short. LITERATURE CITED Wings entirely covered with microtrichia and predominately brown, with hyaline marks along the margin and in the Chua, T. H., 1999. Fruit fl ies of the subfamilies Ceratitidinae, Tephritinae and Trypetinae in Malaysia, Serangga, 4: fi eld. Vein R4+5 setose over most of its length, the setae extending well beyond a level with m crossvein. Subcostal 189–219.

cell about ¾ as long as costal cell. Vein R2+3 almost straight. Hancock, D. L. & R. A. L. Drew, 1994. New species and records The r-m crossvein situated at about apical 3/4 of cell dm, of Asian Trypetinae (Diptera: Tephritidae). Raffl es Buletin of and lobe of posterior cubital cell (cup) rather short, 1/7 to Zoology, 42: 555–591.

1/8 as long as vein CuA1. Wing markings are as shown in Hancock, D. L. & R. A. L. Drew, 1995. New genus, species and Fig. 1. Hyaline spots present as follows: bc with apical part synonyms of Asian Trypetinae (Diptera: Tephritidae). Malaysian hyaline, c with 2 hyaline spots separated by brown area, sc Journal of Science, 16A: 45–59.

with a marginal hyaline spot; r1 with a large squarish spot Hardy, D. E., 1973. The fruit (Tephritidae – Diptera) of and a smaller rounded spot touching anterior wing margin, Thailand and bordering countries. Pacifi c Monograph, a large hyaline spot at wing apex and 5 rounded to oblong 31: 1–353. spots on posterior margin; approximately 7 small hyaline Hardy, D. E., 1986. Fruit fl ies of the subtribe Acanthonevrina of spots isolated in the fi eld. Indonesia, New Guinea, and the Bismarck and Solomon Islands (Diptera: Tephritidae: Trypetinae: Acanthonevrini). Pacifi c Abdomen mostly shining dark brown to black with median Insect Monograph, 42: 1–191. portion rufous extending from base to 6th tergum; the rufous Korneyev, V. A., 2001. Phylogenetic relationships among higher region extends laterally at hind margin on each segment. groups of Tephritidae. In: Aluja, M. & Norrbom, A. L. (eds.), Sixth tergum about as long 5th. The sterna are light brown Fruit fl ies (Tephritidae): phylogeny and evolution of behavior. except for brown lateral margins. The katepisternum is just CRC Press. Pp. 73–113.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 29–37 Date of Publication: 28 Feb.2009 © National University of Singapore

THE GENUS EOTRECHUS (HETEROPTERA: GERRIDAE) IN VIETNAM, WITH DESCRIPTIONS OF TWO NEW SPECIES

John T. Polhemus Colorado Entomological Institute, 3115 S. York St., Englewood, CO 80113 USA Email: [email protected]

A. D. Tran Systematics and Ecology Laboratory, Department of Biological Sciences National University of Singapore, 14 Science Drive 4, Singapore 117543 Email: [email protected]

Dan A. Polhemus Department of Natural Sciences, Bishop Museum, 1525 Bernice St., Honolulu, HI 96817 USA Email: [email protected]

ABSTRACT. – Two new species of hygropetric water striders in the genus Eotrechus are described from Vietnam: E. fansipan, new species, from the Sa Pa area of northern Vietnam, and E. pumat, new species, from the Pumat Nature Reserve of north-central Vietnam. Figures of key morphological characters are given for both species. Discussion is also provided of the unusual habits of E. brevipes Andersen, which is recorded from Vietnam for the fi rst time and shown to inhabit dry bedrock exposures with scattered clumps of grasses and sedges. Photographs are provided of this habitat and of the adjacent wet rheocrene type locality for E. fansipan.

KEYWORDS. – Eotrechus, new species, Vietnam, taxonomy.

INTRODUCTION described. The four known species from Vietnam are keyed and discussed below. The genus Eotrechus was described by Kirkaldy (1902) to contain a single species, E. kalidasa, based on an Eotrechus species are generally considered to inhabit indeterminate number of specimens collected by L. Fea rheocrenes or waterfall splash zones, but are also known at Carin Cheba, Burma (east of modern Toungoo). These from wet trails or damp ground. During our collecting specimens were said to be deposited in the Museo di Storia around the waterfalls in the Sa Pa area of northern Vietnam Naturale “Giacomo Doria” in Genoa, but possibly never we even found E. brevipes running on perfectly dry, steep returned there (Andersen, 1982). We have seen a male rock faces and hiding in clumps of grass amid the crevices lectotype of this species in the Snow Entomological Museum on such cliffs, an unexpected discovery described in greater in Lawrence, Kansas, and a female in the Distant Collection detail below. Although once thought to be the most primitive at the British Museum (these types were studied by Andersen, of the Gerridae because of their large terminal claws, the 1982). This species, and many others in the genus, have Eotrechinae are now considered to be among the most been rare in collections up to the present time, probably evolved of all water striders, representing a remarkable because collectors did not choose to scramble over slippery, evolutionary reversal from aquatic to terrestrial existence vertical rock faces after creatures of unknown importance. (Andersen, 1982, 1995). As the ecology of Eotrechus species has become better understood, however, a number of substantial collections To date, all captures of Eotrechus in Vietnam have come have been reported over the last decade, with new species from the northern half of the country. In particular, we have described by Andersen (1982, 1998), Tran & Yang (2006), no records of this genus from the Central Highlands of Tran & Zettel (2006), and Vitheepradit & Sites (2007) from Vietnam, despite the presence of many suitable habitats in various parts of southeast Asia and China, bringing the total that area. We independently made extensive collections of number of known species to 14. With the addition of the aquatic Heteroptera throughout the Central Highlands in 2001 two new species reported here, a total of 16 species are now (by J. T.Polhemus & D. A. Polhemus) and in 2003 & 2005

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(by Tran A. D.) and found only the genus Onychotrechus, Eotrechus brevipes Andersen, 1982 a smaller member of the Eotrechinae, inhabiting rheocrene (Figs. 1–3) sites where Eotrechus might reasonably have been expected to occur. Therefore, we believe the absence of the latter Eotrechus brevipes Andersen, 1982: 17–19, Figs. 13, 19, 25–27, genus in the central and southern sections of Vietnam may 32–33 (type locality: Goom, Darjeeling, India); Andersen, 1998: represent a true distributional anomaly, rather than an artifact 4–5 (descriptive notes). of insuffi cient sampling. Material examined. – INDIA: 1 apterous male, “N. India: Darjeeling, 7000 ft., 11–20.iii.1924, Maj. R.W.G. Hingston / Everest Exp. Brit. Mus. 1924-386” (paratype, BMNH); 1 macropterous MATERIALS AND METHODS female, “Gopaldhara, Darjeeling, 4720 [not 4729 as in Andersen, 1982]–6100 ft., on wet ground, 24-7-17, H. Stevens, Brit. Mus. The material studied consisted of dry-mounted and 1922-307” (paratype, BMNH); 1 apterous female, NW Bengal, alcohol-preserved specimens deposited in the following Rimbik-Srikhola, 2,650m, coll. E. Kucera,ˇ 15–27 May 2006 collections: Natural History Museum, London (BMNH), (NHMW). VIETNAM, Lao Cai Prov.: 5 apterous males, 13 apterous Bernice Bishop Museum, Honolulu, Hawaii (BPBM), J. T. females, dry steep rockface, 17 km northwest of Sa Pa, 1,935 m. Polhemus collection, Englewood, Colorado (JTPC), Natural [6,350 ft.], 22°21'17"N 103°46'28.3"W, 8 Apr.2000, 1115–1300 hrs., CL 4401, coll. J.T. Polhemus, D.A. Polhemus & P. Nguyen History Museum Vienna (NHMW), United States National (USNM, JTPC, ZRC). Museum, Washington, DC (USNM), Zoological Museum, Hanoi University of Science (ZMHU), Zoological Reference Diagnosis. – Size: apterous male, length 7.35, width 2.28; Collection, National University of Singapore (ZRC). apterous female, length 6.80, width 2.51. Colour as in Andersen (1982). Measurements of new species refer to the holotype and the allotype. Body size measurements (length and width) are Body mainly brown, covered with green refl ective pubescence. given as ranges based on randomly selected specimens. Antennae about 0.7× body length, fi rst antennal segment Length of the body was measured from the anterior margin shorter than head width across eyes. Male fore femur strongly of the head to the posterior tip of the last abdominal tergum. incrassate (slightly less in female) and not spinose; fore tibia Width of the body was measured across the meso-acetabula. slightly curved with a small projection on apex. Abdomen All measurements are given in millimetres. Illustrations relatively short. Male sternum VII about 1.66× length of two were made using a camera lucida attached to a binocular preceding sterna combined, posterior margin emarginated microscope. with a broad medial notch (about 1/3 of sternum VII length). Male pygophore with a pair of posterolateral projections, each projection with a fi nger-like process. Female genital TAXONOMY segments concealed, proctiger covered by tergum VIII.

Eotrechus Kirkaldy, 1902 Remarks. – The specimens listed above represent the fi rst record of this species for Vietnam, and represent a range Eotrechus Kirkaldy, 1902: 137 (type species: Eotrechus extension of 1,700 kilometres from the original type locality kalidasa Kirkaldy, 1902, by monotypy); Distant, 1903: 182; at Darjeeling, India (Andersen, 1982). This, in combination Matsuda, 1960: 249–251; Andersen, 1982: 5–8. with other records (see below) implies that the species is distributed along the entire southern margin of the Himalayan uplift from Assam, through Burma and Yunnan, to Vietnam. KEY TO SPECIES OF EOTRECHUS OF VIETNAM It has likely been overlooked because of its odd ecological habits, as described in the following section. 1. Male pygophore with posterolateral projections bearing a fi nger-like process at apex. Female genitalia totally concealed by sternum VII and tergum VIII ...... E. brevipes Andersen Ecological notes. – Eotrechus brevipes is notable among – Male pygophore with posterolateral projections (if present) members of the family Gerridae for exhibiting an essentially without finger-like process at apex. Female genitalia not terrestrial mode of existence. At the Vietnamese locality 17 concealed by sternum VII and tergum VII ...... 2 km northwest of Sa Pa, this species was found inhabiting 2. Pronotum and mesonotum with a prominent light median clumps of grasses and sedges clinging to an exposure of dry, longitudinal stripe, laterally with brownish markings, but not brown bedrock adjacent to a waterfall and associated lateral extensive light markings. Male genitalia without prominent rheocrenes (Figs. 1–3). Specimens of E. brevipes were found posterolateral projections (Figs. 8, 10) ...... only in this dry setting, where they sheltered at the bases of ...... E. fansipan, new species the vegetation clumps (Fig. 3). If disturbed, they were able to – Pronotum and mesonotum with extensive light markings medially and laterally. Male genitalia with prominent move across the steeply sloping rock face with great rapidity posterolateral projections (Figs. 14, 16) ...... 3 to another patch of sedges or grasses, and were therefore 3. Male pygophore with a pair of prominent posterolateral digitate most easily collected by application of a light pyrethrin fog, and fl at projections ...... E. vietnamensis Tran & Yang which slowed their movements and caused them to emerge – Male pygophore with a pair of prominent posterolateral from their hiding places. The striking color pattern of this broad foliate processes, directed upward in lateral view (Figs. species, consisting of longitudinally parallel orange and 14–16) ...... E. pumat, new species green stripes, although conspicuous in a collection tray, in

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Figs. 1–3. Type locality for E. fansipan, new species, at a waterfall 17 km. northwest of Sa Pa, Vietnam. Both E. brevipes and E. fansipan, new species, occurred at this location. 1, general view of waterfall and surroundings, showing wet rheocrenes adjacent to the fall itself, on the left side of the photo, which provided habitat for E. fansipan, new species, and dry slopes to the right side of the photo, which provided habitat for E. brevipes. The senior author, standing to the right of the fall, provides scale; 2, lateral view of fall, showing dry bedrock faces in the foreground on which E. brevipes occurred; the photo was taken from near the position of the senior author in Fig. 1; 3, detail of E. brevipes habitat, showing clumps of vegetation scattered on open bedrock. Individuals of E. brevipes sheltered within the vegetation clumps.

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fact provides a remarkably effective camoufl age amid the and metasternopleura usually dark brown; abdominal venter green stems and brownish earth of the vegetation clumps mainly light brown. in which this species hides. A search of the adjacent wet rheocrenes in closer proximity to the waterfall produced no Structural characteristics. Apterous male (holotype): Head further captures of E. brevipes; instead, these wet rheocrenes width across eyes 1.80; interocular width 1.00; eye kidney- were inhabited by E. fansipan, new species (see subsequent shaped in dorsal view, length 0.87. Antennae about 0.77× description). Therefore, it appears that at this single locality, body length (6.55: 8.50), lengths of segments I–IV: 1.92: two distinct Eotrechus species were segregating rock face 1.56: 1.21:1.86; first segment with 5–6 black spines on habitats on the basis of moisture regime. The unusual apical part. Pronotum broader than long, shorter than head preference of E. brevipes for dry, partially vegetated rock length (1.08: 1.49); mesonotum length 1.80. Lengths of faces may also explain the relative dearth of captures for mesosternum and metasternum: 2.28 and 0.71. Lengths of this species, given that most collectors would not search for leg segments (femur: tibia: tarsal segment I: tarsal segment water striders in such settings. II) as follows, fore leg: 2.93: 2.71: 0.27: 0.52; middle leg: 7.30: 6.80: 0.67: 0.60; hind leg: 7.50: 7.70: 0.69: 0.67. Fore Distribution. – India (Sikkim and West Bengal), China femur (Fig. 4) incrassate, length about 4.65× maximum width (Fukien), and Vietnam (north-western area). (2.93: 0.63), slightly constricted before apex, basal part with broad tubercle bearing patch of minute dark hairs; ventral surface of femur with a row of about 10 spine-like hairs; Eotrechus fansipan, new species fore tibia slightly curved in the middle, long spine-like hairs (Figs. 1, 4–10, 17) scattered on external side and denser at apical part (Fig. 4). Mid- and hind femora slender and slightly shorter than Material examined. – Holotype (apterous male) and allotype body. Mid- and hind trochanters without spines, mid- and (apterous female): VIETNAM, Lao Cai Prov., waterfall, 17 hind femora with scattered black spines. Lengths of fore, km northwest of Sa Pa, 1,935 m. [6,350 ft.], 22°21'17.0"N mid, and hind claws: 0.16: 0.25: 0.25. Abdomen relatively 103°46'28.3"W, water temp. 18.5°C., 8 Apr.2000, 1115–1300 short, venter medially depressed from segment III to VI and hrs., CL 4401, coll. J.T. Polhemus, D.A. Polhemus & P. Nguyen more broadly depressed on segment VII, length of abdominal (USNM). venter from sternum II to sternum VII: 1.96; sternum VII Paratypes: VIETNAM, Lao Cai Prov.: 32 apterous males, 35 about 1.48× length of two preceding sterna together (0.71: apterous females, same locality data as holotype (USNM, JTPC, 0.52), posterior margin deeply emarginate, around 2/5 of BPBM, ZRC); 5 apterous males, 2 apterous females, 1 macropterous sternum VII length, margin with long, soft, brown hairs (Fig. female, rocky streamlet with cascades, 16 km northwest of Sa Pa, 8). Abdominal segment VIII long (dorsal length 0.87, ventral 1,890 m. [6,200 ft.], 22°21'28.4"N 103°46'36.8"W, water temp. length 0.83), with posteroventral margin almost straight; 17°C., 8 Apr.2000, 1330–1430 hrs., CL 4402, coll. J.T. Polhemus, abdominal spiracle of segment VIII not produced into minute D.A. Polhemus & P. Nguyen; 9 apterous males, 6 apterous females, tubercle. Genital segments relatively large, modified as 1 macropterous female, Bac Waterfall and smaller side waterfall, follows: Proctiger trilobate (Fig. 7), posterolateral lobes small, 15 km northwest of Sa Pa, 1,829 m. [6,000 ft.], 22°21'39.7"N 103°46'39.5"W, water temp. 19°C., 8 Apr.2000, 1500–1600 hrs., ventrocaudal surface of proctiger with dense patch of long, CL 4403, coll. J.T. Polhemus, D.A. Polhemus & P. Nguyen. brushlike hairs (Figs. 6, 7) (not visible in situ). Pygophore moderately long in ventral view and constricted on central Description. – Size: apterous male, length 7.66–8.50 section (Fig. 8), two small tubercles near the narrowest part (holotype 8.50), width 2.48–2.74 (holotype 2.74); apterous bearing long soft hairs (more visible in dorsal view) (Fig. female, length 8.55–9.49 (allotype 9.20), width 2.77–2.94 10); ventral surface convex, round in lateral view (Fig. 9); (allotype 2.87); macropterous female, length 9.54–9.60, posterior part produced into short plate-like structure, with width 2.61–2.77. dense patch of long soft hairs on ventrolateral sides, apical margin almost straight (Fig. 10). Paramere very short, blunt, Colour. Apterous form: Dorsum mainly brown, covered with some soft hairs on apex. with golden pubescence, with yellow markings on head, pronotum, and mesonotum as follows: head with yellow Apterous female (allotype): Head width across eyes 1.78; markings on anterolateral corners, next to inner margin of interocular width 1.02; eye kidney-shaped in dorsal view, eyes, and a yellow marking on posterior margin; pronotum length 0.84. Antennae about 0.70× of body length (6.41: with yellow median longitudinal stripe, in lateral view 9.20), lengths of segments 1-4: 1.82: 1.58: 1.19: 1.82; fi rst with two yellow markings, one longer in upper part and segment with about 7 black spines on apical part. Pronotum one shorter on lower part; mesonotum mainly dark brown, broader than long, shorter than head length (1.03: 1.40), with yellow median stripe running from anterior to posterior mesonotum length 2.03. Lengths of mesosternum and margin and tapering posteriorly, anterior part expanded metasternum: 2.33 and 0.79. Lengths of leg segments (femur: laterally. Pro-, meso-, metasternopleura, meso- and meta- tibia: tarsal segment I: tarsal segment II) as follows, fore leg: acetabula with dense patches of refl ective silvery pubescence. 2.94: 2.48: 0.37: 0.59; middle leg: 7.20: 6.60: 0.68: 0.67; Metanotum, abdominal tergites mainly dark brown, tergites hind leg: 7.20: 7.25: 0.70: 0.70. Fore femur slender, length II–VII usually with narrow yellowish median stripe. All about 6.68× maximum width (2.94: 0.44), slightly tapering leg segments (including coxae) yellowish, except tarsi dark towards apex, base without tubercle; ventral surface with brown. Ventral body with prosternopleuron yellow; meso- about 12 spine-like hairs in a row; fore tibia almost straight,

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with many long black spine-like hairs along external side, 2006, especially in the structure of fore leg and genitalia. particularly on distal part. Mid- and hind femora slender, However, closer examination shows signifi cant differences slightly shorter than body. Mid- and hind trochanters without that separate the former from the other two (Table 1). spines, mid- and hind femora with black spines scattered distally. Lengths of fore: mid: hind claws: 0.20: 0.24: 0.24. Etymology. – The name of this species is a noun in apposition Abdomen (Fig. 5) moderate in length, length of abdominal and refers to Mt. Fansipan, the highest mountain in Vietnam, venter from sternum II to sternum VII: 3.38; sternum VII which is located near the type locality. large, slightly tapering towards apex, length about 1.10× length of two preceding sterna combined (1.10: 1.00), Ecological notes. – The type series of Eotrechus fansipan, posterior margin smooth. Genital segments not concealed new species, was taken on wet bedrock faces adjacent to a by sternum VII; proctiger acute. wide, unshaded waterfall along the road from Sa Pa to Lai Chau. The waterfall was set slightly back from the road on the Remarks. – Eotrechus fansipan, new species, is closely upstream side, and the majority of specimens were taken on related to E. pingae Andersen, 1998 (E. pingae is known from the wet rheocrenes to the right side of the fall when looking a single male specimen only) and E. luaae Tran & Zettel, upstream (Fig. 1). The surrounding vegetation consisted of

Figs. 4–10. Eotrechus fansipan, new species, structural details: 4, male foreleg; 5, female abdomen, left lateral view; 6, male proctiger, left lateral view; 7, male proctiger, dorsal view; 8, male terminal abdomen, ventral view; 9, male pygophore, left lateral view; 10, male pygophore, dorsal view.

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Table 1. Key characters separating E. fansipan, new species, E. pingae and E. luaae

Male E. fansipan, new species E. pingae E. luaae Fore tibia - Internal side curved at middle - Almost straight - Almost straight Abdominal venter - Median groove from sternum III - Median groove from sternum II - Median groove from sternum III to VI, less conspicuous to VII, narrow and deep to VII, less conspicuous Abdominal - Posterior margin straight, - Without median process - With small acute median segment 8, venter without median process process Proctiger - Posterolateral lobes small - Posterolateral lobes large - Posterolateral lobes large - With patch of long hairs on - Without such hair patch - With patch of long hairs similar ventrocaudal surface to E. fansipan Pygophore - Medium length - Short - Long and slender - Lateral tubercles small - Tubercles large - Without lateral tubercles - Posterior plate short and broad - Posterior plate short and broad - Posterior plate narrow, tapering towards apex - With ventrolateral dense hair patches - Without ventrolateral hair - With less dense ventrolateral patches hair patches

disturbed montane subtropical forest, intermixed with dry Structural characteristics. Apterous male (holotype): Head bedrock exposures and grassy slopes. The stream below the width across eyes 1.36; interocular width 0.83; eye kidney- falls dropped steeply downhill through a boulder-strewn shaped in dorsal view, length of eye 0.65. Antennae about channel and passed under the road bridge via a series of 1.03× body length (6.06: 5.90), lengths of segments I–IV: shaded pools, which harbored the unusual veliid Velia tonkina; 1.88: 1.49: 1.17: 1.52; fi rst segment with 6 black spine- see Polhemus & Polhemus (2003) for further discussion of like hairs in apical part. Pronotum broader than long, this interesting locality. slightly shorter than head length (0.79: 1.19). Lengths of mesosternum and metasternum: 1.92 and 0.43. Posterior margin of metasternum with a median fringe of short black Eotrechus pumat, new species hairs (Fig. 14). Lengths of leg segments (femur: tibia: tarsal (Figs. 11–17) segment I: tarsal segment II) as follows: fore leg: 2.33: 1.88: 0.22: 0.37; middle leg: 6.35: 4.75: 0.67: 0.57; hind leg: 6.60: Material examined. – Holotype (apterous male) and allotype 5.30: 0.79: 0.59. Fore femur (Fig. 11) slightly concave at (apterous female): VIETNAM, Nghe An Prov., Kem waterfall, the middle part of ventral side and constricted at apical part, Pu Mat Nature Reserve, 400 m [1,310 ft.], coll. J.T. Polhemus, length about 6.13× maximum width (2.33: 0.38), ventral 01 Apr.2000, CL 4385 (USNM). surface with three black stout hairs on basal half (two hairs in other paratypes); fore tibia straight, with few long black Paratypes: VIETNAM, Nghe An Prov.: 8 apterous males, 8 apterous females, same data as holotype (USNM, JTPC, ZRC). spine-like hairs on apical margin, fore tarsus covered with long, soft yellowish hairs. Middle and hind femur slender Description. – Size: apterous male, length 5.72–6.10 and slightly longer than the body, with scattered small brown (holotype 5.9), width 1.99–2.19 (holotype 2.13); apterous spines (more on middle femur). Claws stout, lengths of fore, female, length 6.49–8.44, width 2.33–2.55. mid- and hind claws: 0.19: 0.22: 0.22. Abdomen (Fig. 14) short, length of abdominal venter from sternum II to sternum Colour. Apterous form, body mainly brown (ventral side VII = 1.11, with patch of long yellow hairs on median sterna usually paler) with yellowish brown markings dorsally; dorsal II–VI. Sternum VII about 2.13× length of two preceding surface covered with silvery or greenish pubescence. Head sterna combined (0.51: 0.24), posterior margin with very with one narrow median yellow stripe, anterolateral corners deep rectangular median notch (about half the length of and posterior margin yellow. Antennae yellow to brown. sternum VII) (Fig. 14). Genital segments large, pygophore Pronotum with fi ve longitudinal yellow stripes on dorsal and with a pair of broad and thick posterolateral processes, as in lateral sides. Mesonotum mainly brownish yellow with six Figs. 14–16, these processes directed upward in lateral view longitudinal markings: two slender dark brown lateral stripes, (Fig. 15); on each side of the process (dorsal and ventral) two slender dark brown sub-lateral stripes and two brown with a prominent tubercle (visible in lateral and dorsal view, markings in the centre. Metanotum and abdominal tergum Figs. 15, 16); in dorsal and ventral view, each process with mainly dark brown. Pro-, meso- and metasternopleura with a small fi nger-like process directed inwards (Fig. 14, 16). dense silvery and refl ective patches of pubescence. Legs with Proctiger simple, elongate, unmodifi ed (Fig. 13). Parameres all coxae, trochanters and femora yellow (slightly darker in small and blunt. females), all tibiae and tarsi brown.

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Apterous female (allotype): slightly bigger than the male. Lengths of mesosternum and metasternum: 2.21 and 0.56. Head width across eyes 1.40; interocular width 0.84; eye Lengths of leg segments (femur: tibia: tarsal segment I: kidney-shaped, length 0.63. Antennae about 0.8× of body tarsal segment II): fore leg: 2.16: 1.86: 0.27: 0.40; middle length (5.41: 6.80), lengths of segments I–IV: 1.55: 1.29: leg: 6.70: 4.80: 0.75: 0.63; hind leg: 6.90: 5.30: 0.90: 0.62. 1.06: 1.51; fi rst segment with 2–3 black spines sub-apically. Fore femur simple and slender, length about 6.8× maximum Pronotum slightly shorter than head length (0.79: 1.11). width (2.16: 0.32), ventral surface with 3 to 5 black spine-

Figs. 11–16. Eotrechus pumat, new species, structural details: 11, male foreleg; 12, female abdomen, left lateral view; 13, male proctiger, dorsal view; 14, male terminal abdomen, ventral view; 15, male pygophore, left lateral view; 16, male pygophore, dorsal view.

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Fig. 17. Distributions of new species of Eotrechus in Vietnam: square = E. fansipan, new species; circle = E. pumat, new species.

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like hairs; fore tibia straight with some long black spines on Chau, Dien Bien, and Lao Cai provinces (Tran & Yang, apical margin, fore tarsus covered with long, soft yellowish 2006). hairs. Middle and hind femur slender and subequal to body length, scattered with small brown spines (more on middle Habitats. – This species occurs on wet bedrock faces and femur). Claws stout, lengths of fore, mid- and hind claws: rheocrenes; see discussion in Tran & Yang (2006). 0.19: 0.24: 0.24. Length of abdominal venter from sternum II to sternum VII about 0.29× body length (1.98: 6.80). Venter of sterna II–VII without patch of long hairs. Sternum VII about ACKNOWLEDGMENTS 1.63× length of two preceding sterna together (0.83: 0.51), posterior margin straight. Genital segments not concealed, We thank Dr. Mick Webb (Natural History Museum, London) proctiger round and slightly defl ected (Fig. 12). and Dr. Herbert Zettel (Natural History Museum Vienna) for kindly providing access to specimens held under their Remarks. – Eotrechus pumat is closely related to E. care. D. A. Polhemus and J. T. Polhemus also wish to give vietnamensis Tran & Yang, 2006. They share similar special thanks to the late Dr. Cao Van Sung of the Institute characteristics, including the dorsal colour pattern, the of Ecology and Biological Resources, Hanoi, and to Phong structure of the male fore femur (Fig. 11), and modifi cations Nguyen, formerly of Nam Cat Tien National Park, both of of the male and female pregenital abdomen (Figs. 12, 14). whom greatly facilitated the logistics and permitting of our Females of these two species are indistinguishable. These fi eld work in Vietnam. The faunal survey work in Vietnam similarities suggest that they may be sister species. However, by D. A. Polhemus and J. T. Polhemus was also supported by E. pumat can be easily separated from the latter by the unique the Drake Fund of the National Museum of Natural History, structures of male genitalia (Figs. 14–16). Smithsonian Institution, Washington, D.C. Tran Anh Duc sincerely thanks Prof. Peter K. L. Ng and Mrs. Yang Chang Etymology. – The name of this species is a noun in apposition Man (Department of Biological Sciences, National University and refers to the type locality, the Pu Mat Nature Reserve of Singapore) who greatly supported his research project on in north-central Vietnam. systematics of semi-aquatic bugs of Vietnam. We also thank Dr Herbert Zettel and an anonymous reviewer for their critical Ecological notes. – The type series was collected from Kem comments and corrections on the manuscript. waterfall (Thac Kem) at the end of a trail in the Pu Mat Nature Reserve (Fig. 17). This waterfall fl ows down over a rather irregular rock face into a large plunge pool below; an LITERATURE CITED excellent color photo of this waterfall is provided in Sites & Vitheepradit (2007, pg. 18). The Eotrechus specimens Andersen, N. M., 1982. Semiterrestrial water striders of the genera moved rapidly over the slippery face of the waterfall, and Eotrechus Kirkaldy and Chimarrhometra Bianchi (Insecta, were quite diffi cult to capture. Hemiptera, Gerridae). Steenstrupia, 9: 1–25. Andersen, N. M., 1995. Cladistic inference and evolutionary scenarios: locomotory structure, function, and perfomance in Eotrechus vietnamensis Tran & Yang, 2006 water striders. Cladistics, 11(3): 279–295. Andersen, N. M., 1998. Notes on the genus Eotrechus Kirkaldy Eotrechus vietnamensis Tran & Yang, 2006: 11–20, Figs. 1–6, 26 (Hemiptera, Gerridae), with a cladistic analysis and a new (type locality: Tam Dao, Vinh Phuc, Vietnam). species from China. Steenstrupia, 24: 1–8. Kirkaldy, G. W., 1902. Miscellanea Rhynchotalia No. 3. Material examined. – VIETNAM, Vinh Phuc Prov.: 1 apterous Entomologist, 35: 136–138. male, 1 apterous female, Tam Dao National Park, Suoi Thac Bac (near Doi Che), coll. A.D. Tran & Q.K. Hoang, 17 Jun.2003, Polhemus, D. A. & J. T. Polhemus, 2003. A review of the Veliinae TAD0351 (holotype and allotype, ZMHU). For paratypes, see of Vietnam (Heteroptera: Veliidae) with description of a new Tran & Yang (2006). Velia species. Journal of the New York Entomological Society, 111(1): 29–40. Diagnosis. – Dorsal body with yellow markings on brown Sites, R. W. & A. Vitheepradit, 2007. Namtokocoris Sites, a new background (see Tran & Yang, 2006: fi g. 26). Antennae genus of Naucoridae (Hemiptera: Heteroptera) in waterfalls longer than body length, fi rst antennal segment longer than of Indochina, with descriptions of six new species. Zootaxa, 1588: 1–29. width of head across eyes. Abdomen relatively short in both sexes. Male sternites II–VI with patch of long hairs on Tran, A. D. & C. M. Yang, 2006. New species of the water strider median line, sternum VII with a deep rectangular median genera Eotrechus Kirkaldy and Rhyacobates Esaki (Heteroptera: notch. Male genital segments large, pygophore broadly Gerridae) from Vietnam. Raffl es Bulletin of Zoology, 54(1): 11–20. suboval, with a pair of disto-lateral projections, each directed obliquely upwards, plate like, and with slightly pointed apex Tran, A. D. & H. Zettel, 2006. Notes on Eotrechus Kirkaldy, 1902 in lateral view. (Heteroptera: Gerridae), with descriptions of two new species. Zootaxa, 1353: 39–51. Remarks. – See remarks for E. pumat. Vitheepradit, A. & R. W. Sites, 2007. A review of Eotrechus Kirkaldy (Hemiptera: Heteroptera: Gerridae) of Thailand with descriptions of three new species. Zootaxa, 1478: 1–19. Distribution. – Northern Vietnam: Vinh Phuc, Ba Vi, Lai

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 39Ð50 Date of Publication: 28 Feb.2009 © National University of Singapore

SIX CRAB OF THE SUBFAMILY STEPHANOPINAE FROM SOUTHEAST ASIA (ARANEAE: )

Guo Tang, Changmin Yin and Xianjin Peng College of Life Sciences, Hunan Normal University, Changsha, Hunan Province, 410081, P. R. China Email: [email protected] (Corresponding author)

Charles Griswold California Academy of Sciences, 55 Concourse Drive, San Francisco, California 94118, U.S.A.

ABSTRACT. – Six species of crab spiders belonging to three genera are reported from China, Myanmar and India, including two new species: Sanmenia nigra Tang, Griswold & Peng, new species (from Myanmar) and Sanmenia tengchong Tang, Griswold & Yin, new species (from Yunnan, China); three new combinations: Epidius bazarus (Tikader, 1970) new combination (from India and China, transferred from Platythomisus), Epidius ganxiensis (Yin, Peng & Kim, 1999) new combination (transferred from Philodromus, Philodromidae), and Epidius gongi (Song & Kim, 1992) new combination (transferred from Cupa); and one new record to China: Cebrenninus rugosus Simon, 1887 (from Yunnan, China). The genus Cebrenninus is recorded from China for the fi rst time. Two species are synonymized: Cupa kalawitana Barrion & Litsinger, 1995 = Cebrenninus rugosus Simon, 1887; Philodromus longitibiacus Yin, Peng & Kim, 1999 = Epidius gongi (Song & Kim, 1992). Detailed morphological characters, the distribution map, photos and illustrations of genital organs are presented.

KEY WORDS. – Taxonomy, Araneae, Thomisidae, Stephanopinae, new species, synonymy.

INTRODUCTION 2008, Tang, Peng, Griswold, Ubick & Yin, 2008, Tang, Yin, Ubick & Peng, 2008). The family Thomisidae is one of the largest families of spiders, including 2,070 species in 171 genera and 7 Crab spiders of the subfamily Stephanopinae, which have subfamilies in the world, of which about 500 species in 63 teeth on the chelicerae, are typically striking in general genera are known from Southeast Asia (Murphy & Murphy, appearance and structure of genital organs. They are also 2000; Platnick, 2008). Simon (1880, 1903, 1909) described among the most poorly studied Asian Thomisidae. The some thomisids from China, Vietnam and other areas; Thorell genus Cebrenninus is poorly known and there have been no recorded 31 species of thomisids from Burma (Myanmar) reports for about one century since Simon (1887) created (Thorell, 1895). However, the type specimens of most exotic this genus with the type species: C. rugosus, and only 5 genera described by Simon, Thorell, O. P. Pickard-Cambridge species of this genus have been recorded from South Asia. and some others were not studied by earlier authors (Lehtinen, The genus Epidius is also poorly studied and only 3 out of 9 2004). Furthermore, the fi gures of genital organs are not described species have been illustrated. The genus Sanmenia exact enough in the references published by Tikader (1963, was created by Song & Kim for the species: Cupa zhengi 1965, 1966, 1970, 1971, 1980) and his coworkers from Ono & Song, 1986, and 3 species with detailed fi gures so India and Barrion & Litsinger (1995) from the Philippines. far have been reported from China, and Singapore Lehtinen (2004) made a substantial revision of Misumenini (Song & Kim, 1992; Ono, 1995; Yang, Zhu & Song, 2006). and some species of other subfamilies from the Palaearctic This study is ready to report some poorly known crab and Oriental regions, and clarifi ed many wrong generic species of the subfamily Stephanopinae from Asia, and to placements: four new genera were erected, two genera and clarify some wrong generic placements. 28 species were synonymized and 28 new combinations were proposed. Since then, Tang and his colleagues have reported In this paper, we describe two new species: Sanmenia nigra 32 species of crab spiders including 1 new genus and 20 Tang, Griswold & Peng, new species (from Myanmar) and new species from Yunnan, China (Tang, Yin, Griswold & Sanmenia tengchong Tang, Griswold & Yin, new species Peng, 2006, Tang, Yin, Peng, Ubick & Griswold, 2007, (from Yunnan, China).

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MATERIALS AND METHODS retromargin of chelicera), but can be separated from the latter by: ratio of width of ocular area / head area = 1.0 Observed specimens were from the collection of the CaAS (0.5 in Epidius); male palp has a spatulate VTA and a row and HNU; the type specimens of Epidius gongi (Song of macrosetae (with VTA, RTA, sometimes with ITA in & Kim, 1992) were loaned from IZCAS. Specimens Epidius). were described and illustrated under an Olympus SZ11 dissecting stereomicroscope. Photos were taken using the Syncroscopy Auto Montage¨ software with a Leica M420 Cebrenninus rugosus Simon, 1887 (New to China) stereomicroscope or only by Nikon stereomicroscope. (Figs. 1AÐF, 7) Measurements of leg lengths are given as: total length (femur, patella + tibia, metatarsus, tarsus). Spination of legs mainly Cebrenninus rugosus Simon, 1887: 468; Simon, 1897: 9, Figs. 1Ð2 follows Ono (1988), and in some species only the ventral (types not examined); Benjamin, 2008: 4 (no description) spination of tibiae and metatarsi of legs I and II are given. Libania armillata Thorell, 1890: 149. (synonymized by Simon, Left structures (e. g., palp, legs and chelicera) are measured 1897: 9) Cupa kalawitana Barrion & Litsinger, 1995: 208, Figs. 119aÐf and described. The following abbreviations are used: anterior (new synonymy, types not examined) eye row (AER); anterior lateral eyes (ALE); anterior median eyes (AME); apical tegular apophysis (ATA); California Material examined. – 1 male (HNU-Hu 060901), Qinglangdang Academy of Sciences (CaAS), San Francisco, USA; The (27¼41'N 98¼17'E), 1,309 m, Dulongjiang Township, Gongshan George Washington University (GWU) Washington D.C., County, Yunnan Province, China, coll. P. Hu, 1 Sep.2006. USA; Hunan Normal University (HNU), Changsha, China; Institute of Zoology of the Chinese Academy of Sciences Diagnosis. – The species is the type species of the genus (IZCAS), Beijing, China; Kunming Institute of Botany (KIB), Cebrenninus, which can be separated from the allied species Kunming, China; median ocular area (MOA); posterior eye by: the tibia only with one large RTA, the bulb fl at, and the row (PER); posterior lateral eyes (PLE); posterior median embolus and median apophysis both long and spatulate, the eyes (PME); retrolateral tibial apophysis (RTA); ventral apex of median apophysis curved. tibial apophysis (VTA); Zoological Survey of India (ZSI), Calcutta, India. Description. – Male. Total length 4.64. Carapace 2.25 long, 2.12 wide; abdomen 2.39 long, 1.88 wide. Carapace dark brown, slightly swollen, with some small apophyses and some TAXONOMY short white hairs, lateral margins serrated with curved spines Cervical grooves, radial grooves and fovea distinct. Eye area Stephanopinae O. Pickard-Cambridge, 1871 wide, eye tubercles indistinct, eye sizes and interdistances: ALE (0.16) > PLE (0.14) > PME (0.06) > AME (0.04); Cebrenninus Simon, 1887 (New to China) AME-AME (0.14) < AME-ALE (0.15), PME-PME (0.25) < PME-PLE (0.32), MOA long (0.37), back (0.40) > front Cebrenninus Simon, 1887: 468 (0.29). Sternum brown. Chelicerae, gnathocoxae, labium Libania Thorell, 1890: 148 (synonymized by Simon, 1897: 9) blackish brown. Legs I, II dark brown; legs III, IV brown, Ocyllus Thorell, 1887 (synonymized by Lehtinen, 2002: 316) all femora with annuli and 2 dorsal spines; femora I slightly Cupa Strand, in Bösenberg & Strand, 1906 (Barrion & Litsinger, swollen medially, ventrally and prolaterally with some 1995: 208, misidentifi ed) denticles, prolaterally with 5 spines; tibiae I, II with 5 pairs Type species. – Cebrenninus rugosus Simon, 1887, by original of long ventral spines; metatarsi I, II with 2 pairs of spines; designation. tibiae III, IV with 3 pairs of ventral spines; metatarsi III with 2 pairs of ventral spines; metatarsi IV without ventral Diagnosis. – Medium sized thomisids. Cephalothorax: spines. Leg measurements: I 7.90 (2.50, 3.20, 1.20, 1.00), carapace convex, with many small apophyses and covered II 8.20 (2.60, 3.30, 1.30, 1.00), III 5.70 (1.80, 2.10, 1.10, with many white hairs, lateral margins serrated. Chelicera 0.70), IV 6.10 (1.90, 2.20, 1.20, 0.80); formula 2, 1, 4, 3. with 2 promarginal and 2 retromarginal teeth, the larger Abdomen nearly oval with short hairs, dorsum with black retromarginal one bifi d. Abdomen dorsum with small yellow markings and small light brown spots. Its sides with black spots and black markings. Male palp with a long RTA, bulb lines; venter yellow. Palpal tibia with large RTA, the apex fl at with slender embolus and median apophysis. of RTA sharp and slightly curved; bulb fl at, sperm ducts visible within tegulum, both the embolus and the median Remarks. – The genus includes 5 species recorded from Java, apophysis long and spatulate, the apex of median apophysis and , but is poorly known because only the curved in beak-shaped. type species was simply illustrated by Simon (1897). This genus was misidentifi ed by Barrion & Litsinger (1995) as Female. No fresh female specimen was examined. Cupa (judging from his description and fi gures, the male palp of Cupa kalawitana has a long embolus and median Distribution. – Java, Sumatra, Borneo, Thailand, Malaysia, apophysis, which is the typical diagnostic character of the the Philippines; China (Gongshan County, Yunnan genus Cebrenninus). This genus is similar to Epidius (both Province). of them have a small tooth and a large bifi d tooth on the

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Fig. 1. Cebrenninus rugosus Simon, 1887: A, habitus of male, dorsal view; B, habitus of male, front view; C, palp, prolateral view; D, F, palp, ventral view; E, palp, retrolateral view; e, embolus; ma, median apophysis. Scale bars: AÐB = 1.0 mm, CÐF = 0.1 mm.

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Remarks. – This species is widely distributed from South Material examined. 1 female (HNU-Tang 2003102801), Daxinzai Asia to China (including Thailand and Malaysia (Benjamin, Village (24¡32'N 98¡24'E), 1,219 m, Wuhe Township, Tengchong 2008)). After compared the descriptions and figures of County, Yunnan Province, China, coll. G. Tang, 28 Oct.2003. 1 Cebrenninus rugosus Simon, 1887, and Cupa kalawitana female (CaAS-19611021), Lataguri, 110 m, West Bengal, India, coll. E. Ross, 21 May 1961. Barrion & Litsinger, 1995 (types not examined), both male

palps have a long embolus and median apophysis, which Diagnosis. – The species can be easily separated from E. indicates that Cupa kalawitana Barrion & Litsinger, 1995, gongi (Song & Kim, 1992) by: female with larger body should be considered as a junior synonym of Cebrenninus size: 7.36Ð7.62 (6.00Ð6.60 in E. gongi); copulatory ducts rugosus Simon, 1887. curved and tube-shaped (short and straight in E. gongi); spermathecae small and spherical (spermathecae with 2 chambers in E. gongi). Epidius Thorell, 1877

Epidius Thorell, 1877: 492; Simon, 1897: 10; Millot, 1941: 65. Description. – Female. Total length: 7.36Ð7.62. The specimen Cupa Strand, in Bösenberg & Strand, 1906 (Song & Zhu, 1997: of total length 7.62 measured: carapace 3.47 long, 3.14 wide; 23, misidentifi ed) abdomen 4.58 long, 2.76 wide. Cephalothorax: Carapace slightly raised on the median, light yellow, without markings. Type species. – Epidius longipalpis Thorell, 1877, by original Cervical grooves, radial grooves and fovea indistinct. Eye designation. area gray, compact, eye tubercles grayish-black, small, eye sizes and interdistances: ALE 0.17, PLE 0.16, AME 0.06, Diagnosis. – Small to medium size, Head area narrow, PME 0.14; AME-AME 0.17, AME-ALE 0.12, PME-PME eye tubercles of each side continuous. Chelicera with 3 0.28, PME-PLE 0.20, MOA length 0.43 with front width promarginal and 2 retromarginal teeth, the larger retromarginal 0.30 and back width 0.52. Sternum pale yellow. Chelicerae, one bifi d. Legs slender with many spines. Epigyne usually gnathocoxae, labium yellow. Leg femora yellow, metatarsi with an atrium and a pair of posterior sclerotized plates. and tarsi yellowish-brown. Leg measurements: I 19.20 (6.00, Male tibia with VTA and 4Ð6 macrosetae in a row. 7.20, 4.00, 2.00), II 18.60 (5.80, 7.20, 3.80, 1.80), III 9.50 (2.90, 3.80, 1.80, 1.00), IV 10.00 (3.00, 4.00, 2.00, 1.00); Distribution. – Africa, India, Sri Lanka, Java, Sumatra, the formula 1, 2, 4, 3. Legs light yellow. Spination: femur: I Philippines, Indonesia, Vietnam, China. dorsal, weak spine: 0-0-1-0-0, IIÐIV 0-1-0-0-1, prolateral (weak) 0-1-2-1-1 (right), 1-1-2-1-1 (left); tibiae: prolateral Remarks. – This genus is composed of 9 species from Africa IÐIV 0-0-1-1, ventral I-II 2-2-2-2-2, IIIÐIV 2-2-2, metatarsus and Southeast Asia. Epidius is a group of poorly known IÐII prolateral 1-0-0-1ap, ventral 2-2-2-2; IIIÐIV prolateral special crab spiders and differs from other genera greatly by 1-1-1ap, III ventral 0-2-2. Abdomen elongate oval, dorsum their general appearance and genital organs. This genus is with broken silvery white spots; cardiac pattern gray, 2 pairs similar to Sanmenia in the shape of embolus and conductor of muscular depressions visible; laterally silvery white; venter of male palp, but can be separated from the latter by: body light yellow. Epigyne simple, atrium depressed; with a pair usually light yellow to gray without markings (gray to of sclerotized plates posteriorly; copulatory ducts curved in grayish-brown with some black-brown stripes or markings in n-shaped; spermathecae spherical. Sanmenia), epigyne usually with a pair of sclerotized plates posteriorly (without those in Sanmenia) and male palp with Distribution. – China (Tengchong County of Yunnan), only VTA (with both VTA and RTA in Sanmenia). After India. checking the type specimens of Cupa gongi Song & Kim, 1992, (the only representative of the genus Cupa recorded Remarks. – The type specimen of Platythomisus bazarus from China), it is found that Cupa gongi Song & Kim, 1992, was kept in ZSI and is unavailable for study by foreign was misidentifi ed as a representative of the genus Cupa. The arachnologists (Lehtinen, 2004: 178). However, the locality male palp of Cupa gongi has a spatulate VTA and a row of of one of our specimens (Lataguri, West Bengal, India) macrosetae, which are the typical diagnostic characters of is adjacent to that of Platythomisus bazarus (Nayabazar, the genus Epidius. Cupa gongi Song & Kim, 1992, should West Sikkim, India). After comparing the descriptions and be transferred to the genus Epidius as E. gongi (Song & fi gures of Platythomisus bazarus Tikader, 1970, and those Kim, 1992). Therefore, there is no real representative of of our specimens (from India, China), it was found that they Cupa found in China now. should be the same species. Both of our specimens bear the following diagnostic characters of the genus Epidius: tibiae and metatarsi of legs I & II slender with thick ventral spines, Epidius bazarus (Tikader, 1970), new combination epigyne with an atrium and a pairs of posterior sclerotized (Figs. 2AÐF, 7) plates, eye arrangement also as same as that of Epidius. Because of those, Platythomisus bazarus should be transferred Platythomisus bazarus Tikader, 1970: 48, Fig. 27a-c; Tikader, 1971: to the genus Epidius as Epidius bazarus (Tikader, 1970), 65, Fig. 17MÐP; 1980: 171, Figs. 237Ð239. (type specimens new combination. collected from West Sikkim, India and deposited in IZA, India, inaccessible; not examined)

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Fig. 2. Epidius bazarus (Tikader, 1970): A, habitus of female (from Yunnan, China); B, habitus of female (from West Bengal, India); C, E, epigyne; D, F, vulva; sp, sclerotized plate. Scale bars: AÐB = 1.0 mm; CÐF = 0.1 mm.

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Epidius ganxiensis (Yin, Peng & Kim, 1999), expanding for half an hour in lactic acid (Fig. 4C, F). Epigyne new combination with a pair of sclerotized plates posteriorly; copulatory ducts (Figs. 3AÐJ, 7) short; spermathecae with 2 oval chambers.

Philodromus ganxiensis Yin, Peng & Kim, 1999: 356, Fig. 2AÐG Diagnosis. – This species is similar to E. ganxiensis (Yin, (types in HNU, examined) Peng & Kim, 1999), the differences are discussed in the “diagnosis” of E. ganxiensis. The male is similar to E. Material examined. – 2 males, 3 females (HNU), Ganxi Township, longipalpis Thorell, 1877, in the long palpal tibia, but can Tongdao County (26¡03'N 109¡17'E), Hunan Province, China, coll. C. Yin, X. Peng and Y. Zhang, 1 Jun.1996. be separated from the latter by: the tibia with 2 apophyses E. longipalpis (only with one in ). Description. – Male palp with a spatulate VTA and 6 macrosetae in a row at tibial apex, conductor transparent, Distribution. – China (Hunan, Zhejiang, Fujian twisted and concave at apex; embolus slender, apically Provinces). hidden under the conductor. Epigyne with a pair of spatulate sclerotized plates posteriorly; copulatory openings slit like, Remarks. – The male palp of Cupa gongi has a spatulate copulatory ducts complexly twisted; spermathecae curved VTA and a row of macrosetae, the epigyne with a pair of in n-shape. posterior sclerotized plates, which are the typical diagnostic characters of the genus Epidius. Cupa gongi Song & Kim, Diagnosis. – This species can be separated from E. gongi 1992, should be transferred to the genus Epidius as E. (Song & Kim, 1992) by: the atrium slit-shaped laterally (pit gongi (Song & Kim, 1992), new combination. Philodromus like medially); the ratio of length of palpal tibia to palp = longitibiacus Yin, Peng & Kim, 1999, was fi rstly placed under 1.2 (5.0 in E. gongi). the spider family Philodromidae. After examined both type specimens of Philodromus longitibiacus and Cupa gongi, it Distribution. – China (Hunan). is found that Philodromus longitibiacus actually is a junior synonym of Epidius gongi (Song & Kim, 1992). Remarks. – The species was firstly placed under the spider family Philodromidae. However, the male palp has a spatulate VTA and a row of macrosetae, which are the Sanmenia Song & Kim, 1992 typical diagnostic characters of the genus Epidius of family Sanmenia Song & Kim, 1992: 142; Song & Zhu, 1997: 28; Yang Thomisidae. Philodromus ganxiensis Yin, Peng & Kim, 1999, et al., 2006: 42 should be transferred to the genus Epidius as E. ganxiensis (Yin, Peng & Kim, 1999), new combination. Type species. Ð Cupa zhengi Ono & Song, 1986.

Diagnosis. – Small sized thomisids, body covered with Epidius gongi (Song & Kim, 1992), new combination short white hairs. Chelicerae usually with 3Ð4 promarginal (Figs. 4AÐI, 7) teeth and 3 retromarginal teeth. Legs long with many thick spines. Male palp with VTA and RTA, tegulum simple Cupa gongi Song et Kim, 1992: 141Ð142, Figs. 1Ð7; Song & Zhu, without apophysis. Embolus slender fi liform, conductor wide 1997: 23, Fig. AÐG (type in IZCAS, examined) and extend from the tegulum, and shades embolus. Epigyne Philodromus longitibiacus Yin, Peng & Kim, 1999: 356, Fig. 3AÐD (new synonym, types in HNU, examined) usually with a large medial atrium.

Material examined. – 2 males, 1 female (HNU), Nanyue (27¡05'N Remarks. – The type species of this genus was fi rstly placed 112¡19'E), Hunan Province, China, coll. G. Tang, P. Hu and in the genus Cupa (Ono & Song, 1986). Song & Kim (1992) Q. Wang, 3 May 2007; 1 female (HNU), Xiaxiang, Tongdao erected the genus Sanmenia to accommodate Cupa zhengi. County (26¡03'N 109¡17'E), Hunan Province, China, coll. C. This genus only includes 3 species recorded from Asia Yin, 2 Jun.1996; 2 females (HNU), Mujiao Township, Tongdao before this study. This genus is similar to Epidius in the County, coll. X. Peng & Y. Zhang, 4 Jun.1996; 1 female (HNU), structures of the parts of male palpal organ. The differences Linkou Township, Tongdao County, coll. X. Peng & Y. Zhang, are discussed in the Remarks of Epidius. 6 Jun.1996; 1 male, 1 female (IZCAS-Ar9378, Ar9379), Daoxian County (25¡11'N 111¡12'E), Hunan Province, China, coll. L. Gong, May.1984; 1 male, 1 female (IZCAS-Ar9380, Ar9381), Tiantong Park (121¡15'N 29¡17'E), Ningbo City, Zhejiang Province, China, Sanmenia nigra, Tang, Griswold & Peng, new species Jun.1990; 1 male (IZCAS-Ar9382), Yujiaping, Mountain Longxi, (Figs. 5AÐE, 7) Jiangle County (117¡05'N 26¡11'E), Fujian Province, coll. Y. Han, 11 Apr.1991. Material examined. – Holotype Ð female (CaAS9019310), 4.36 km 142¡ east-south-east of Popamyo (20¡33'N 95¡08'E), 2,050 Description. – Male palp with a spatulate VTA and 5 m, deciduous forest, Mt. Popa Wildlife Reservation, Mandalay macrosetae at tibial apex; conductor axe shaped, apically Division, Myanmar, coll. C. Griswold, 26 Sep.2003. curved; sperm ducts visible, embolus needle like, combined with a median apophysis basally, which can be observed after Diagnosis. – This new species can be separated from S. tengchong, Tang, Griswold & Yin, new species by: carapace

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Fig. 3. Epidius ganxiensis (Yin, Peng & Kim, 1999): A, habitus of female; B, F, left palp, C, D, I, epigynum; E, J, vulva; G, embolus; H, VTA and tibial spines. Scale bars: A = 1.0 mm, BÐJ = 0.1 mm.

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Fig. 4. Epidius gongi (Song & Kim, 1992): A, habitus of male; B, C, E, left palp; D, I, epigyne; F, palpal bulb (expanded after storage in lactic acid for half an hour); G, chelicera; H, vulva; c, conductor; e, embolus; ma, median apophysis. sp, sclerotized plate. Scale bars: A = 1.0 mm, BÐI = 0.1 mm.

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with pars cephalica all black (with small sparse brown stripes black markings. Its sides with black markings; venter light in S. tengchong); the epigyne without lateral hoods (with yellow. Epigyne with a median depressed atrium; copulatory a pair of lateral hoods in S. tengchong); the spermathecae ducts short; spermathecae elongate oval. elongate oval (curved in n-shaped in S. tengchong). Male. Unknown. Description. – Female. Holotype total length 3.88. Carapace 1.80 long, 1.66 wide; abdomen 2.02 long, 2.00 wide. Carapace Etymology. – The specifi c name refers to the large black light brown with black margin, pars cephalica all black. markings on carapace and abdominal dorsum. Cervical grooves, radial grooves and fovea indistinct. Eye area with some long stick-like hairs. Tubercles of ALE and Distribution. – Myanmar. PLE continuous. Eye sizes and interdistances: ALE 0.13, PLE 0.12, PME 0.11, AME 0.03; AME-AME 0.07, AME-ALE Remarks. – Thorell published a descriptive catalogue of 31 0.08, PME-PME 0.11, PME-PLE 0.12, MOA 0.21 long, species of thomisids in 20 genera including 12 new species front width 0.24, back width 0.29. Sternum light yellow. from Burma (Myanmar) without fi gures (Thorell, 1895). Chelicerae, gnathocoxae, labium yellow; chelicerae with Up to now, 4 genera recorded by Thorell from Myanmar scopulae on both margins of fang furrows; 3 promarginal were synonymized, but no species recorded by Thorell was and 3 retromarginal teeth (Fig. 5c). Legs yellow with many transferred to the genus Sanmenia. spines, legs I, II darker. Femora I with 5 prolateral spines and 1 dorsal short one. Tibiae I, II with 2 dorsal spines, 3 prolateral ones and 5 pairs of long ventral ones; metatarsi I, Sanmenia tengchong, Tang, Griswold & Yin, II with 2 prolateral spines and 4 pairs of long ventral ones. new species Left legs I, II lost, right leg measurements: I 5.20 (1.50, 2.20, (Figs. 6AÐF, 7) 1.00, 0.50 ), II 6.20 (2.00, 2.60, 1.10, 0.50), III 3.40 (1.20, 1.20, 0.60, 0.40), IV 4.20 (1.50, 1.50, 0.70, 0.50); formula 2, Material examined. – Holotype Ð female (HNU-YHY2202), 1, 4, 3. Abdomen near oval, dorsum light yellow with large Dengma Village (24¡56'N 98¡23'E), 1,113 m, Hehua Township,

Fig. 5. Sanmenia nigra, Tang, Griswold & Peng, new species: A. habitus of female; B, D, epigyne; C, chelicera; E, vulva. Scale bars: A = 1.0 mm, BÐE = 0.1 mm.

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Fig. 6. Sanmenia tengchong, Tang, Griswold & Yin, new species: A, habitus of female, holotype; B, habitus of female, paratype; C, E, epigyne; D, F, vulva. Scale bars: AÐB = 1.0 mm, CÐF = 0.1 mm.

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Tengchong County, Yunnan Province, China, coll. P. Hu, 2 white and continuous. Eye sizes and interdistances: ALE Jun.2006. 0.13, PLE 0.12, PME 0.11, AME 0.05; AME-AME 0.08, AME-ALE 0.08, PME-PME 0.13, PME-PLE 0.14. Sternum Paratypes Ð 1 female (CaAS-YHY2202), same data as holotype. yellow. Chelicerae, gnathocoxae, labium yellowish brown; chelicerae with scopulae on both margins of fang furrows; 3 Diagnosis. – This new species is similar to S. nigra, Tang, promarginal and 3 retromarginal teeth. Legs grayish-yellow Griswold & Peng, new species, the differences are discussed with grayish-black spots. Femora I swollen with 5 prolateral in the “Diagnosis” of S. nigra. The new species can also and 1 dorsal short spines. Tibiae I, II with 3 prolateral and 5 be separated from S. gongshan Yang et al., 2006, by: the pairs of ventral spines; metatarsi I, II with 4 pairs of ventral abdomen with more dark markings and fewer white spots; spines. Leg measurements: I 6.98 (2.32, 2.96, 1.20, 0.50), the abdomen only with short white hairs (short white hairs II 6.66 (2.20, 2.86, 1.15, 0.45), III 4.80 (1.80, 1.90, 0.80, and some long black hairs in S. gongshan); the epigynal 0.30), IV 5.03 (1.80, 1.93, 0.90, 0.40); formula 1, 2, 4, 3. hoods laterally located (posteriorly located in S. gongshan); Abdomen nearly oval, dorsum covered with short white the spermathecae curved, tube-shaped (pyriform in S. hairs, dorsum grayish-brown with paramedian grayish-black gongshan). stripes and some small white spots; laterally grayish-black; venter grayish-brown with some small black spots. Epigyne Description. – Female. Total length 4.80Ð5.00. Holotype with a pair of hoods laterally; copulatory ducts invisible; total length 4.80. Carapace 2.17 long, 1.93 wide; abdomen spermathecae curved, tube-shaped. 2.54 long, 2.19 wide. Cephalothorax elevated moderately, carapace brown with short white hairs, with grayish black Male. Unknown. markings bilaterally and medially. Carapace flat on the head area. Carapace with 3 small black stripes behind the Etymology. – The specific name refers to the type PER and extending backwards, the fovea black. Cervical locality. grooves, radial grooves and fovea indistinct. Eye area with some long hairs. Tubercles of ALE and PLE grayish- Distribution. – China (Yunnan Province, China).

Fig. 7. distribution map of the six crab spiders from South East Asia: Cebrenninus rugosus Simon, 1887, Epidius bazarus (Tikader, 1970), Epidius ganxiensis (Yin, Peng & Kim, 1999), Epidius gongi (Song & Kim, 1992), Sanmenia nigra, Tang, Griswold & Peng, new species, and Sanmenia tengchong, Tang, Griswold & Yin, new species.

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ACKNOWLEDGEMENTS Simon, E., 1880. Etudes arachnologiques. 11e Mémoire. XVII. Arachnides recueilles aux environs de Pékin par M. V. Collin We are grateful to Dr. Suresh Benjamin (GWU) for his help de Plancy. Annales de la Société Entomologique de France, in identifying the species: Cebrenninus rugosus Simon, 1887; (5) 10: 97Ð128. thanks are given to Dr. Shu-Qiang Li (IZCAS) for loaning Simon, E., 1903. Etudes arachnologiques. 33e Mémoire. LIII. the types of Epidius gongi (Song & Kim, 1992); thanks go Arachnides recueillis à Phuc-Son (Annam) par M. H. Fruhstorfer to Dr. Jeremy Miller (CaAS) and Prof. Yun Liu (HNU) for (novÐdec. 1899). Annales de la Société Entomologique de France 71: photographic work; thanks are given to Dr. Peter Fritsch , 725Ð736. (CaAS) and Prof. Heng Li (KIB) for supporting the joint Simon, E., 1897. Histoire naturelle des araignées, Paris, 2: biodiversity survey of the Gaoligong Mountains; thanks go to 1Ð192. Mr. Peng Hu (HNU) for collecting parts of type specimens. Song, D. X. & J. P. Kim, 1992. A new species of crab spider from This work was supported by the grants from the National China, with description of a new genus (Araneae: Thomisidae). Science Foundation of the USA through the grant “Biotic Korean Arachnology, 7: 141Ð144. survey of the Gaoligongshan, a biodiversity hotspot in western Song, D. X. & M. S. Zhu, 1997. Fauna Sinica: Arachnida: Yunnan, China” (No. DEB-0103795); and from the Scientifi c Araneae: Thomisidae, Philodromidae. Science Press, Beijing, Research Fund of Hunan Provincial Education Department 259 pp. (in Chinese) (08C564), and also partly supported by the National Natural Tang, G., C. M. Yin, C. Griswold & X. J. Peng, 2006. Description Sciences Foundation of China (NSFC-30499341, 30670234, of Sinothomisus gen. nov. with a new species from Yunnan 30770255). Province, China (Araneae, Thomisidae). Zootaxa, 1366: 61Ð68. Tang, G., C. M. Yin, X. J. Peng, D. Ubick & C. Griswold, 2007. LITERATURE CITED Five crab spiders of the genus Lysiteles from Yunnan Province, China (Araneae: Thomisidae). Zootaxa, 1480: 57Ð68. Barrion, A. T. & J. A. Litsinger, 1995. Riceland Spiders of South Tang, G., C. M. Yin, X. J. Peng, D. Ubick & C. Griswold, 2008. and Southeast Asia. CAB International, Wallingford, UK, xix The crab spiders of the Genus Lysiteles from Yunnan Province, + 700 pp. China (Araneae: Thomisidae). Zootaxa, 1742: 1Ð41. Benjamin, S. P., 2000. Epidius parvati sp. n., a new species of the Tang, G., Peng, X. J., Ubick, D., Griswold C. & Yin, C. M., genus Epidius from Sri Lanka (Araneae: Thomisidae). Bulletin 2008. Four crab spiders of the family Thomisidae (Arachnida: of the British Arachnological Society, 11: 284Ð288. Araneae) from Yunnan, China. Acta Zootaxonomica Sinica, Benjamin, S. P., D. Dimitrov, R. G. Gillespie & G. Hormiga, 2008. 33(2): 241Ð247. Family ties: molecular phylogeny of crab spiders (Araneae: Tang, G., Yin, C. M., Ubick, D., & Peng, X. J., 2008. Two new Thomisidae). Cladistics, 24: 1Ð15. species of the crab spider genus Talaus (Araneae: Thomisidae) Lehtinen, P. T., 2002. Generic revision of some thomisids related to from Yunnan Province, China. Zootaxa, 1815: 62Ð68 Xysticus C. L. Koch, 1835 and Ozyptila Simon, 1864. European Thorell, T., 1890. Diagnoses aranearum aliquot novarum in Indo- Arachnology, 315Ð327. Malesia inventarum. Annali del Museo Civico di Storia Naturale Lehtinen, P. T., 2004. Taxonomic notes on the Misumenini (Araneae: di Genova, 30: 132Ð172. Thomisidae: Thomisinae), primarily from the Palaearctic and Thorell, T., 1895. Descriptive catalogue of the spiders of Burma. Oriental regions. In Logunov, D. V. & D. Penney (eds.), London, Pp. 1Ð406. European Arachnology 2003 (Proceedings of the 21st European Tikader, B. K., 1963. Studies on interesting south Indian crab-spiders Colloquium of Arachnology, St.-Petersburg, 4Ð9 August 2003). (Family: Thomisidae). Proceedings of the Indian Academy of Arthropoda Selecta, Special Issue, 1: 147Ð184. Sciences, 58(B): 249Ð262. Millot, J., 1941. Les araignées de l’Afrique Occidentale Française Tikader, B. K., 1965. On some new species of spiders of the family - sicariides et pholcides. Mémoires de l’Académie des Sciences Thomisidae from India. Proceedings of the Indian Academy of de l’Institut de France, 65: 1Ð82. Sciences, 61(B): 277Ð289. Murphy, F. & J. Murphy, 2000. An Introduction to the Spiders Tikader, B. K., 1966. Studies on some crab-spider (family: of South East Asia, With Notes on All the Genera. Malaysian Thomisidae) from Khasi and Jaintia hills, Assam, India. Nature Society, Kuala Lumpur, Malaysia, 625 pp. Proceedings of the Indian Academy of Sciences, 64(B): Ono, H. & D. X. Song, 1986. A new Sino-Japanese species of the 53Ð61. genus Cupa (Araneae, Thomisidae) from the coastal areas of Tikader, B. K., 1970. Spider fauna of Sikkim. Records of the the East China Sea. Bulletin of the National Science Museum, Zoological Survey of India, 64: 1Ð83. Tokyo, (A) 12: 25Ð29. Tikader, B. K., 1971. Revision of Indian crab spiders (Araneae: Ono, H., 1988. A revisional study of the spider family Thomisidae Thomisidae). Memoirs of the Zoological Survey of India, (Arachnida, Araneae) of Japan. National Science Museum, 15(8): 1Ð90. Tokyo, 252 pp. Tikader, B. K., 1980. Thomisidae (Crab-spiders). Fauna India Ono, H., 1995. Four East Asian spiders of the families Eresidae, (Araneae), 1: 1Ð247. Araneidae, Thomisidae and Salticidae (Arachnida, Araneae). Bulletin of the National Science Museum, 21(3): 157Ð169. Platnick, N. I., 2008. The world spider catalog, version 9.0. American Museum of Natural History. Available from: http://research. amnh.org/entomology/spiders/catalog/index.html (accessed 1 Sep.2008).

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 51–63 Date of Publication: 28 Feb.2009 © National University of Singapore

CAVE SHRIMPS OF THE GENUS EDONEUS HOLTHUIS, 1978, FROM LUZON, THE PHILIPPINES, WITH DESCRIPTIONS OF THREE NEW SPECIES (CRUSTACEA: DECAPODA: ATYIDAE)

Yixiong Cai National Biodiversity Centre, National Parks Board, 1 Cluny Road, Singapore 259569. Republic of Singapore Email: [email protected]

Daniel Edison M. Husana Institute of Environmental Science and Meteorology, University of the Philippines, Diliman, Quezon City, The Philippines Email: [email protected]

ABSTRACT. – The troglobitic freshwater shrimp genus Edoneus Holthuis, 1978, is revised on the basis of a fresh collection from several caves in Luzon, the Philippines, as well as previously studied collections deposited at the National Museum of Natural History, Leiden. The revised genus is characterized by the absence of an arthrobranch on the fi rst pereiopod, the absence of appendix interna on the endopod of the male fi rst pleopod, the uropodal diaeresis having the outermost spinules distinctly longer than the lateral angle, the short exopodal fl agellum and long endopod of the fi rst maxilliped, the presence of a tiny posteromedian projection on the telson, and the short intermediate setae on the distal end of the telson. Morphological comparisons between Edoneus and several morphologically related genera are discussed. Three new species, namely Edoneus erwini, E. sketi and E. marulas, are described and illustrated in detail.

KEY WORDS. – Freshwater shrimp, cave, Atyidae, Edoneus, new species, Philippines.

INTRODUCTION southwest of Manila. Numerous caves are present in the area but only three were sampled in the present study. The The cave shrimp genus Edoneus was described by Holthuis meandering underground rivers and pools of the caves vary (1978), who emphasized several troglobitic characters, from less than a metre to a few metres wide. Samplings were including the peculiar gill formula, which had no podobranch conducted from near the entrance (twilight zone), at the and arthrobranch present on the pereiopods or maxillipeds. middle, and in the innermost parts of the caves. The number Balete & Holthuis (1992) reported more specimens of the of specimens collected was limited to avoid possible decline type species, Edoneus atheatus Holthuis, 1978, from the of these populations. Aside from shrimps, other aquatic same cave in northern Luzon, with a corrected name for found were fi shes, leeches, crabs, and snails. the type locality. No additional morphological description was added. Specimens examined are deposited in Collection of the National Museum of the Philippines, Manila (NMCR), Examination of a recent collection from several caves in and the Zoological Reference Collection of the Raffles southern Luzon, the Philippines, revealed the presence of Museum of Biodiversity Research, National University of three undescribed species of atyids, which share numerous Singapore, Singapore (ZRC). Type material of Edoneus generic characters with E. atheatus, except for the presence atheatus in the National Museum of Natural History, Leiden, of one arthrobranch on the third maxilliped. Netherlands (RMNH) has also been re-examined. The rostral formula used follows Bouvier (1925). The abbreviation cl In the present study, we revise the defi nition of the genus is used for the carapace length, given in millimetres (mm), Edoneus to include the new species described here. The and measured from the postorbital margin to the posterior sites where the specimens were collected are located in the dorsal margin of the carapace. municipality of Atimonan, Quezon province, about 180 km

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TAXONOMY podobranch at the base of second maxilliped. Holthuis (1978: 220) also described the telson of E. atheatus as follows: Atyidae De Haan, 1849 “short hairs are implanted on the posterior margin of the telson above the spines; the margin itself shows no median Edoneus Holthuis, 1978 tooth.” The examination of topotypical specimens of E. atheatus from the type locality in the present study shows Edoneus Holthuis, 1978: 219; 1993: 51. that there is a very tiny posteromedian projection on the posterior margin of the telson (see Fig. 1A). This structure Type species: Edoneus atheatus Holthuis, 1978, by monotypy; agrees with that of the new species described herein, and is gender: masculine. regarded as diagnositic character for the genus. Edoneus thus includes four species, all of which are endemic to Luzon, Diagnosis. – Body moderately robust, eyes developed or the Philippines. reduced; rostrum short; pterygostomian margin broadly rounded; supraorbital spine absent; ventral margin of pleura Edoneus is most similar to Parisia Holthuis, 1956, from rounded; telson with a very tiny posteromedian projection; Madagascar, Australia (Williams, 1964), and Luzon (Cai & distal margin with intermediate pairs of setae much shorter Anker, 2004). It can only be differentiated from Parisia by than lateral pair of spines; palp of fi rst maxilliped ending in the presence of a very small posteromedian projection on fi nger-like projection, fl agellum of the exopod very short, the telson (vs. posteromedian projection on telson absent), endopod high, as long as or distinctly exceed the fl agellum the lateral angle of the uropodal diaeresis being distinctly of exopod in length; pereiopods without exopods; fi rst and shorter than the outermost spinules (vs. lateral angle of second pereiopods with chelae monomorphic, palm well uropodal diaeresis much longer than outermost spinules), developed, fi ngers tipped with brushes of long setae, carpus of and minor differences in the branchial formula, viz., no or fi rst pereiopod moderately excavated anteriorly, twice longer one arthrobranch on the base of third maxilliped (vs. one or than broad; carpus of second pereiopod much slender than two arthrobranchs). With respect to the short lateral angle of previous one, not excavated anteriorly; branchial formula uropodal diaeresis, and the absence of one arthrobranch on incomplete, with 6 or 7 pairs of gills, no arthrobranch on the fi rst pereiopod, Edoneus resembles to Lancaris, a genus pereiopods; no arthrobranch, or with only one very small that was recently described from highlands of Sri Lanka (Cai arthrobranch on base of third maxilliped; podobranch of & Bahir, 2005). It, however, differs by the absence of an second maxilliped less developed or vestigial; epipods appendix interna on the male fi rst pleopod, which is present present on fi rst 4 pereiopods; fi rst pleopod of male with on species of Lancaris; and the posterior setae on the telson, endopod sub-rectangular, no appendix interna at endopod; in which, the median pairs of setae are much shorter than the second pleopod of male with appendix masculina rod-shaped, lateral pair (vs. much longer in species of Lancaris). slender, spinose on inner surface; Uropodal diaeresis bearing 14–24 movable spinules, with outermost one distinctly longer than lateral angle. Ovigerous females with large eggs, sized Edoneus atheatus Holthuis, 1978 0.90–1.10 × 0.55–0.60 mm. (Fig. 1)

Distribution. – Known only from several caves in Atimonan, Edoneus atheatus Holthuis, 1978: 220, Figs. 5, 6; Balete & Quezon province and the northern part of Luzon, the Holthuis,1992: 99. Philippines. Material examined. – Holotype: Male, cl 4.6 mm, RMNH D 31898, Habitat. – All four species reported here were found in the Philippines, Luzon, Isabela Province Maddela district, Cave the subterranean water in caves of Luzon Island, northern near Santiago [recte: the Philippines, Luzon, Quirino Province, Philippines. Aglipay Municipality, Barrio Palasian, cave (no. 6) near Sitio Disiluad, (see Balete & Holthuis, 1992; Fransen et al., 1997)]; coll. F. Brandsma, 1974. Remarks. – Three undescribed species of atyids from a recent collection from caves in Luzon, the Philippines, share Paratypes: Two females, cl 4.7–4.9 mm, RMNH D 31899, data numerous characters with Edoneus atheatus Holthuis, 1978, same as holotype. except for the presence of one arthrobranch on the third maxilliped in the former three. This difference is attributed Other materials: One male, cl 3.3 mm, 2 females, cl 3.5–3.6 mm, to interspecifi c variation. Interspecifi c variation of the number RMNH D 41183, the Philippines, north Luzon, Quirino Province, and size of arthrobranchs in the third maxilliped have also Aglipay Municipality, Barrio Palasian, Sitio Disiluad. Cave no. 6, been found in the genus Parisia (Holthuis, 1956a, b). We coll. D. S. Balete, 30 Dec.1985. redefine the genus Edoneus here to include the newly discovered species. In the original description, Holthuis Diagnosis – Rostrum short, triangular in lateral view, (1978) described the branchial formula as follows: “In all, unarmed, reaching nearly to end of eyes. Eyes small, fi ve pleurobranchs are present at the bases of the pereiopods; unpigmented; pterygostomian margin broadly rounded; no other pleurobranch were observed, neither were any telson with a very tiny posteromedian projection; distal arthrobranchs or podobranchs.” His fi gures (Holthuis, 1978: margin with intermediate pairs of setae much shorter than Figs. 5-i), however, showed a lamina-shaped vestigial lateral pair of spines; carpus of fi rst pereiopod moderately

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excavated anteriorly, twice as long as high; chela 1.7 times Habitat. – Collected from large fl owing river inside a cave as long as broad; fi ngers as long as palm; carpus of second in northern Luzon. pereipod much slender than previous one, not excavated anteriorly, 6.2 times as long as high; chela 2.3 times as Remarks. – Found only from the type locality (Holthuis, long as broad; fi ngers distinctly longer than palm; branchial 1978; Balete & Holthuis, 1992). Holthuis (1978) described formula incomplete, with 6 pairs of gills; no arthrobranch the new genus and new species Edoneus atheatus, on the basis on pereiopods and maxillipeds; podobranch of second of three specimens. Balete & Holthuis (1992) reported some maxilliped lamina-shaped; uropod with diaeresis bearing more specimens and provided more detailed information on about 15–24 movable spinules, lateral angle much shorter the type locality, as well as detailed habitat information. than movable spinules. Distribution. – Luzon, the Philippines.

Fig. 1. Edoneus atheatus. A, telson; B, second maxilliped; C, fi st pereiopod; D, second pereiopod; E, uropodal diaeresis. Scale bars: A, E = 0.1 mm; B–D = 0.3 mm. (male, cl 2.9 mm, RMNH).

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Edoneus erwini, new species fl agellum of the exopod very short, endopod high, distinctly (Figs. 2, 3) exceed the fl agellum of exopod in length. Second maxilliped (Fig. 2J) typical, except the slightly reduced podobranch. Material examined. – Holotype: Ovigerous female, cl 5.1 mm, Third maxilliped (Fig. 2K) reaching to middle of third eggs 0.92 × 0.55 mm, NMCR, Bantakay Cave station 2, Luzon, segment of antennular peduncle, with ultimate segment as the Philippines, coll. D. E. Husana, 14 Dec.2002. long as penultimate segment. Paratypes: Six males, cl 3.7–4.3 mm, 1 female, cl 5.0 mm, 1 Epipods well developed on first four pereiopods. First ovigerous female, cl 4.4 mm, ZRC 2007.0479, Bantakay Cave station 2, Luzon, the Philippines, coll. D. E. Husana, 14 Dec.2002. pereiopod (Fig. 3A) reaching to anterior end of eye; merus 2.0–2.2 times as long as broad, as long as carpus; carpus Other materials: Five males, cl 4.0–4.5 mm, 4 females, cl 4.0–4.8 excavated anteriorly, shorter than chela, 1.2–1.7 times as mm, 1 ovigerous female, cl 4.4 mm, eggs 0.90 × 0.55 mm, ZRC long as high; chela 1.7–1.9 times as long as broad; fi ngers 2007.0480, Bantakay Cave station 2, Luzon, the Philippines, coll. varying from distinctly shorter than palm to occasionally D. E. Husana, 14 Dec.2002; 3 males, cl 2.8–3.2 mm, 2 females, as long as. Second pereiopod (Fig. 3B) reaching to end cl 2.6–4.3 mm, QM Cave station 3, Luzon, the Philippines, coll. of second segment of antennular peduncle; merus as long D. E. Husana, 14 Mar.2003; 5 males, cl 2.5–3.2 mm, 5 females, as carpus, 5.0 times as long as broad; carpus 1.2 times as cl 3.2–3.5 mm, QM Cave station 2, Luzon, the Philippines, ZRC long as chela, 4.7 times as long as high; chela 2.3 times 2007.0481, coll. D. E. Husana, 14 Mar.2003; 1 male, cl 3.3 mm, 4 females, cl 3.6–4.4 mm, 2 ovigerous females, cl 3.9 mm, eggs as long as broad; fi ngers1.6 times as long as palm. Third 1.1x0.6 mm, ZRC 2007.0482, QM Cave station 1, Luzon, the pereiopod (Figs. 3C, D) reaching beyond scaphocerite by Philippines, coll. D. E. Husana, 14 Mar.2003; 3 males, cl 3.1–3.7 1/3 of propodus, propodus 10 times as long as broad, 6.5 mm, 4 females, cl 3.5–4.3 mm, 1 ovigerous female, cl 3.4 mm, times as long as dactylus; dactylus 2.8 times as long as eggs 0.9 × 0.6 mm, ZRC 2007.0483, QM Cave station 4, Luzon, wide (spines included), terminating in one long claw, 4 the Philippines, coll. D. E. Husana, 14 Mar.2003; 1 male, cl 3.8 or 5 accessory spines on fl exor margin. Fourth pereiopod mm, 9 females, cl 3.3–4.0 mm, ZRC 2007.0484, QM Cave station reaching to end of antennular peduncle, similar in form as 4; Luzon, the Philippines, coll. D. E. Husana, 14 Mar.2003. third one. Fifth pereiopod (Figs. 3E, F) reaching to end of antennular peduncle, propodus 14.4 times as long as broad, Description. – Rostrum short (Fig. 2A), slightly sloping 4.0 times as long as dactylus; dactylus 3.3 times as long down anteriorly, reaching to or slightly beyond end of basal as wide (spinules included), terminating in one claw, with segment of antennular peduncle, to near end of second 47–53 spinules on fl exor margin. segment, rostral formula 0–2+8–13/2–5, ventral teeth closely placed at anterior portion. Inferior orbital angle acute, below Endopod of male fi rst pleopod (Figs. 3G, I) subtriangular, antennal spine; pterygostomian margin sub-rectangular. 2.3 times as long as wide, 1/3 length of exopod, no appendix interna. Appendix masculina of male second pleopod (Figs. Sixth abdominal somite 0.50–0.53 times length of carapace, 3H, J) sub-cylindrical, reaching slightly beyond half length 1.7 times as long as fi fth somite, slightly shorter than telson. of endopod. Appendix interna very small, only about 1/5 Telson (Figs. 2B, C) 3.0 times as long as wide, distal margin length of appendix masculina. rounded, with a very tiny median projection, with 5 pairs of dorsal spinules, and one pair of dorsolateral spinules; Uropodal diaeresis (Fig. 3K) with 16–21 movable spinules, distal end with 3–4 pairs of spines, lateral pair longer than outermost one distinctly longer than lateral angle. intermediate pairs; sublateral pair shortest. Preanal carina (Fig. 2D) low, lacking spine. Ovigerous females with eggs sized 0.90–1.10 × 0.50–0.60 mm. Eyes developed, anterior end reaching to 0.7 times length of basal segment of antennular peduncle. Antennular peduncle Habitat. – This new species were found from two caves, 0.5 times as long as carapace; basal segment of antennular Bantakay Cave and QM (Quezon Mining) Cave. Collection peduncle as long as combined lengths of second and third of specimens from the two localities was conducted on 14 segments, anterolateral angle 0.3 times length of second Dec.2002 and 16 Mar.2003, respectively. QM Cave is located segment, second segment distinctly longer than third segment. at 14º01'0.06"N 121º47'46.5"E, 156 m above sea level and Stylocerite reaching 0.8 times length of basal segment of 8 km away from the sea to its northeast. Its underground antennular peduncle. Scaphocerite (Fig. 2E) 3.2 times as river stretches for more than 750 m. The epigeal stream that long as wide. supplies freshwater to this subterranean river sinks into the cave’s main entrance by dropping a height of about four Incisor process of mandible (Fig. 2F) ending in a row metres before it reaches the fi rst pool, which is about 34 m of small teeth, molar process truncated. Lower lacinia of away from the entrance of the cave. All the specimens coexist maxillula (Fig. 2G) broadly rounded, upper lacinia elongated, in the area and almost evenly distributed from outside to with numerous distinct teeth on inner margin, palp slender. the inside cave except for the light coloured shrimps which Upper endites of maxilla (Fig. 2H) subdivided, palp short, were found in the inside pools only. They usually nibble on scaphognathite tapering posteriorly with numerous long, the skin if you are in the water in contrast to the behaviour curved setae at posterior end. Distal end of palp of fi rst of E. atheatus which “seems not to be attracted to human maxilliped (Fig. 2I) triangular, with a fi nger-like projection; skin…” (Balete & Holthuis, 1992: 100). Bantakay Cave, on

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Fig. 2. Edoneus erwini, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, preanal carina; E, scaphocerite; F, mandible; G, maxillula; H, maxilla; I, fi rst maxilliped; J, second maxilliped; K, third maxilliped. Scale bars: A, E = 1 mm; B, K = 0.5 mm; C = 0.1 mm; D, F–J = 0.3 mm. (male, cl 4.5 mm, paratype, ZRC).

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Fig. 3. Edoneus erwini, new species: A, fi rst pereiopod; B, second pereiopod; C, third pereiopod; D, dactylus of third pereiopod; E, fi fth pereiopod; F, dactylus of fi fth pereiopod; G, male fi rst pleopod; H, male second pleopod; I, edopod of male fi rst pleopod; J, appendix masculina of male second pleopod; K, uropodal diaeresis. Scale bars: A–C, E = 0.5 mm; D, F, K = 0.1 mm; G, H = 0.3 mm; I, J = 0.2 mm (male, cl 4.5 mm, paratype, ZRC).

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the other hand, is probably the most complex and the longest near to or slightly beyond end of basal segment of antennular explored cave in the karstic area of Atimonan-Pagbilao peduncle, rostral formula 0–2(mode 0)+2–8(mode 0+2)/1–4, towns. It has two entrances; one is dry and located at the ventral teeth closely placed at anterior portion. Inferior bank of the main river while the other supplies the water orbital angle acute, indistinctly separated from antennal of the other smaller stream at the southwest. The water that spine; pterygostomian margin sub-rectangular. comes out from the cave entrance drops through a series of waterfalls at the side of the mountain. With a total of more Sixth abdominal somite 0.55 times length of carapace, 1.7 than one kilometre of meandering underground river system, times as long as fi fth somite, as long as or slightly longer its origin is unknown. This cave is located at 14º01'48.01"N than telson. Telson (Figs. 4B, C) 2.5 times as long as wide, 121º47'05.8"E, 100 m above sea level, and about 8 km distal margin rounded, with a very tiny median projection, away from the sea in its northeast. The dry entrance of the with 5 pairs of dorsal spinules, and one pair of dorsolateral cave gives direct and easy access to the small pool (134 spinules; distal end with 3–4 pairs of spines, lateral pair m away) followed by another (164 m away) where most longer than intermediate pairs; sublateral pair shortest. of the unpigmented shrimps (Edoneus sketi, new species) Preanal carina (Fig. 4D) low, lacking spine. were collected. These shrimp are more abundant in these two pools than in the underground streams and were found Eyes strongly reduced, cornea only partially present, pigment outside the cave. reduced to a tiny dot or totally disappear; anterior end reaching to half length of basal segment of antennular peduncle. Colouration. – Dark gray with red spots, grayish-green with Antennular peduncle 0.5 times as long as carapace; basal dark red spots, light coloured with red spots and light gray segment of antennular peduncle as long as combined lengths with red spots. of second and third segments, anterolateral angle 0.3 times length of second segment, second segment distinctly longer Etymology. – The new species is named after Mr. Erwin than third segment. Stylocerite reaching 0.8 times length of Husana, the dedicated field assistant and brother of the basal segment of antennular peduncle. Scaphocerite (Fig. second author. 4E) 3.0 times as long as wide.

Remarks. – Edoneus erwini, new species, can be easily Incisor process of mandible (Fig. 4F) ending in a row of small separated from E. atheatus by the well-developed eyes (vs. teeth, molar process truncated. Lower lacinia of maxillula eye small, unpigmented), the toothed rostrum (vs. rostrum (Fig. 4G) broadly rounded, upper lacinia elongated, with unarmed), and the much stouter carpus of fi rst pereiopod numerous distinct teeth on inner margin, palp slender. Upper (1.2–1.7 times as long as high vs. 2.0 times in E. atheatus). endites of maxilla (Fig. 4H) subdivided, palp elongated, The large egg size of this species suggests that larval scaphognathite tapering posteriorly with numerous long, development of this species is most probably of abbreviated curved setae at posterior end. Distal end of palp of fi rst or directed types (Shokita, 1981). maxilliped (Fig. 4I) triangular, with a fi nger-like projection; fl agellum of the exopod very short, endopod high, as long as fl agellum of exopod in length. Second maxilliped (Fig. Edoneus sketi, new species 4J) typical, except the slightly reduced arthrobranch. Third (Figs. 4, 5) maxilliped (Fig. 4K) reaching to middle of third segment of antennular peduncle, with ultimate segment slightly longer Material examined. – Holotype: Male, cl 3.9 mm, NMCR, small than penultimate segment. pool in Bantakay Cave, Luzon, the Phippines, coll. D. E. Husana, 14 Dec.2002. Epipods well developed on first four pereiopods. First pereiopod (Fig. 5A) reaching to anterior end of eye; merus Paratypes: Seven males, cl 3.6–4.1 mm, 5 females, cl 4.2–4.5 3.1 times as long as broad, as long as carpus; carpus mm, ZRC 2007.0485, small pool in Bantakay Cave, Luzon, the Philippines, coll. D. E. Husana, 14 Dec.2002; 1 ovigerous female, excavated anteriorly, shorter than chela, 2.4 times as long cl 4.7 mm, ZRC 2007.0486, small pool in Bantakay Cave, Luzon, as high; chela 2.3 times as long as broad; fi ngers 1.8 times the Phippines, coll. D. E. Husana, 14 Dec.2002; 2 male, cl 3.8–4.1 as long as palm. Second pereiopod (Fig. 5B) long, reaching mm, ZRC 2007.0487, small pool in Bantakay Cave, Luzon, the to middle of second segment of antennular peduncle; merus Phippines, coll. D. E. Husana, 14 Dec.2002; shorter than carpus, 6.7 times as long as broad; carpus 1.4 times as long as chela, 6.5 times as long as high; chela 2.6 Other materials: Eight males, cl 3.5–4.0 mm, 1 female, cl 3.6 mm, times as long as broad; fi ngers 1.7 times as long as palm. 2 ovigerous females, cl 4.0–4.6 mm, eggs 1.0–1.1 × 0.6 mm, ZRC Third pereiopod (Figs. 5C, D) reaching beyond scaphocerite 2007.0488, small pool in Bantakay Cave, Luzon, the Phippines, coll. by 1/5 of propodus, propodus 11.6 times as long as broad, D. E. Husana, 14 Dec.2002; 7 males, cl 3.7–4.0 mm, 9 females, cl 3.2–4.7 mm, 1 ovigerous female, cl 4.9 mm, eggs 1.0 × 0.6 mm, 4.1 times as long as dactylus; dactylus 3.4 times as long ZRC 2007.0489, QM cave station 5, Luzon, the Phippines, coll. as wide (spines included), terminating in two claws, 4–5 D. E. Husana, 14 Mar.2003; 4 males, cl 3.5–4.4 mm, 11 females, accessory spines on fl exor margin. Fourth pereiopod reaching cl 3.5–4.6 mm, NMCR, QM cave station 7, Luzon, the Phippines, to end of antennular peduncle, similar in form as third one. coll. D. E. Husana, 14 Mar.2003. Fifth pereiopod (Figs. 5E, F) reaching to end of antennular peduncle, propodus 16.8 times as long as broad, 3.7 times Description. – Rostrum short (Fig. 4A), straight, reaching as long as dactylus; dactylus 4.0 times as long as wide

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Fig. 4. Edoneus sketi, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, preanal carina; E, scaphocerite; F, mandible; G, maxillula; H, maxilla; I, fi rst maxilliped; J, second maxilliped; K, third maxilliped. Scale bars: A = 1 mm; B, K, E = 0.5 mm; C = 0.1 mm; D, F–J = 0.3 mm. (male, cl 4.1 mm, paratype, ZRC).

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Fig. 5. Edoneus sketi, new species: A, fi rst pereiopod; B, second pereiopod; C, third pereiopod; D, dactylus of third pereiopod; E, fi fth pereiopod; F, dactylus of fi fth pereiopod; G, male fi rst pleopod; H, male second pleopod; I, uropodal diaeresis. Scale bars: A–C, E = 0.5 mm; D, F, I = 0.1 mm; G, H = 0.2 mm; (male, cl 4.1 mm, paratype, ZRC).

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(spinules included), terminating in one claw, with 39–49 longer than intermediate pairs; sublateral pair shortest. Preanal spinules on fl exor margin. carina (Fig. 6D) low, lacking spine.

Endopod of male fi rst pleopod (Fig. 5G) sub-triangular, 2.3 Eyes strongly reduced, cornea absent, unpigmented or times as long as wide, half length of exopod, no appendix occasionally with very tiny dot; anterior end reaching to half interna. Appendix masculina of male second pleopod (Fig. length of basal segment of antennular peduncle. Antennular 5H) sub-cylindrical, reaching to 2/3 length of endopod. peduncle 0.5 times as long as carapace; basal segment of antennular peduncle as long as combined lengths of second Uropodal diaeresis (Fig. 5I) with 17–20 movable spinules, and third segments, anterolateral angle 0.25 times length outer most one distinctly longer than lateral angle. of second segment, second segment distinctly longer than third segment. Stylocerite reaching 0.7 times length of basal Ovigerous female with eggs sized 1.0–1.1 × 0.6 mm. segment of antennular peduncle. Scaphocerite (Fig. 6E) 3.0 times as long as wide. Habitat. – See habitat under Edoneus erwini, new species. Incisor process of mandible (Fig. 6F) ending in a row of small Colouration. – Shrimps of the present species are found to teeth, molar process truncated. Lower lacinia of maxillula have yellowish head and mostly unpigmented body. (Fig. 6G) broadly rounded, upper lacinia elongated, with numerous distinct teeth on inner margin, palp slender. Upper Etymology. – The new species is named after Professor endites of maxilla (Fig. 6H) subdivided, palp elongated, Boris Sket, University of Ljubljana, Slovenia, for his great scaphognathite tapering posteriorly with numerous long, contributions on the subterranean fauna of the Philippines, curved setae at posterior end. Distal end of palp of fi rst and for his encouragement to the current study. maxilliped (Fig. 6I) triangular, with a fi nger-like projection; fl agellum of the exopod very short, endopod high, as long Remarks. – With regard to the eyes, Edoneus sketi is as fl agellum of exopod in length. Second maxilliped (Fig. morphologically close to E. atheatus. However, it can be 6J) typical, except the slightly reduced podobranch. Third distinguished by the longer rostrum, and more slender carpus maxilliped (Fig. 6K) reaching to end of third segment of of fi rst pereiopod (2.4 times as long as high vs. 2.0 times antennular peduncle, with ultimate segment as long as in E. atheatus). penultimate segment.

Epipods well developed on first four pereiopods. First Edoneus marulas, new species pereiopod (Fig. 7A) reaching to end of basal segment of (Figs. 6, 7) antennular peduncle; merus 2.1 times as long as broad, as long as carpus; carpus excavated anteriorly, shorter than Material examined. – Holotype: Male, cl 4.1 mm, NMCR, at chela, 1.6 times as long as high; chela 1.8 times as long as middle of Marulas Cave, Luzon, the Philippines, coll. D. E. broad; fi ngers slightly shorter than palm. Second pereiopod Husana, 11 Nov.2002. (Fig. 7B) reaching to middle of third segment of antennular peduncle; merus shorter than carpus, 5.3 times as long as Paratypes: Seven males, cl 3.4–4.0 mm, 3 females, cl 4.5 mm, ZRC broad; carpus 1.4 times as long as chela, 4.8 times as long 2007.0490, at middle of Marulas Cave, Luzon, the Philippines, coll. D. E. Husana, 11 Nov.2002; 6 males, cl 3.4–4.0 mm, 11 as high; chela 2.4 times as long as broad; fi ngers1.6 times as females, cl 3.7–4.3 mm, 3 ovigerous females, cl 4.0–4.4 mm, ZRC long as palm. Third pereiopod (Figs. 7C, D) reaching beyond 2007.0491, Marulas Cave, lowest part, Luzon, the Philippines, coll. scaphocerite by length of dactylus, propodus 10.4 times as D. E. Husana, 11 Nov.2002; 12 males, cl 3.4–4.0 mm, 6 females, cl long as broad, 4.1 times as long as dactylus; dactylus 3.3 4.1–4.5 mm, 1 ovigerous female, cl 4.1 mm, eggs 0.90 × 0.55 mm, times as long as wide (spines included), terminating in two ZRC 2007.0492, middle of Marulas Cave, Luzon, the Philippines, claws, 3 or 4 accessory spines on its fl exor margin. Fourth coll. D. E. Husana, 11 Nov.2002; 8 males, cl 3.3–3.8 mm, 1 female, pereiopod reaching to end of scaphocerite, similar in form cl 4.1 mm, 1 ovigerous female, cl 4.5 mm, with hatchings, NMCR, as third one. Fifth pereiopod (Figs. 7E, F) reaching to end Marulas Well, outside the Marulas Cave, Luzon, the Philippines, of scaphocerite, propodus 14 times as long as broad, 3.2 coll. D. E. Husana, 11 Nov.2002. times as long as dactylus; dactylus 4.8 times as long as wide Description. – Rostrum (Fig. 6A) short, straight, pointed, (spinules included), terminating in a single claw, with 42–49 unarmed, reaching near to or slightly beyond end of basal spinules on its fl exor margin. segment of antennular peduncle. Inferior orbital angle acute, fused with antennal spine; pterygostomian margin Endopod of male first pleopod (Fig. 7G) subtriangular, sub-rectangular. 2.1 times as long as wide, 0.4 times length of exopod, no appendix interna. Appendix masculina of male second Sixth abdominal somite 0.54 times length of carapace, 1.3 pleopod (Fig. 7H) sub-cylindrical, spinose, reaching to 2/3 times as long as fi fth somite, as long as or slightly longer length of endopod. than telson. Telson (Figs. 6B, C) 2.8 times as long as wide, distal margin rounded, with a very tiny median projection, Uropodal diaeresis (Fig. 7I) with 14–17 movable spinules, with 4 pairs of dorsal spinules, and one pair of dorsolateral outer most one distinctly longer than lateral angle. spinules; distal end with 3–4 pairs of spines, lateral pair

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Fig. 6. Edoneus marulas, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, preanal carina; E, scaphocerite; F, mandible; G, maxillula; H, maxilla; I, fi rst maxilliped; J, second maxilliped; K, third maxilliped. Scale bars: A = 1 mm; B, E = 0.5 mm; C = 0.1 mm; D, F–K = 0.3 mm. (male, cl 3.9 mm, paratype, ZRC).

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Fig. 7. Edoneus marulas, new species: A, fi rst pereiopod; B, second pereiopod; C, third pereiopod; D, dactylus of third pereiopod; E, fi fth pereiopod; F, dactylus of fi fth pereiopod; G, male fi rst pleopod; H, male second pleopod; I, uropodal diaeresis. Scale bars: A–C, E = 0.5 mm; D, F, I = 0.1 mm; G, H = 0.2 mm; (male, cl 3.9 mm, paratype, ZRC).

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Ovigerous female with eggs sized 0.90 × 0.55 mm. LITERATURE CITED

Habitat. – Marulas Cave is located in Barangay Tinandog, Balete, D. S., & L. B. Holthuis, 1992. Notes on the cave shrimp same area as the previous two caves visited. Of the three Edoneus atheatus Holthuis, 1978, with an account of its type caves studied, this is the shortest one. Its coordinates are locality of habits (Decapoda, Caridea, Atyidae). Crustaceana, 13º59'40.6"N 121º52'23.2"E, 78 m above sea level and 62: 98–101. about 4 km away from the sea at its northeast. There is a Bouvier, E. L., 1925. Recherches sur la morphologie, les variations, dry creek on the surface that sinks to the cave entrance. la distribution géographique des crevettes des la famille des Water is not present at the cave entrance until about 36 m Atyidés. Encyclopédie Entomologique, series A, 4: 1–370, Figs. 1–761. inside where it leaks (spring) from the rock crevices. The stairway-like morphology of the underground stream where Cai, Y. & A. Anker, 2004. On a collection of freshwater shrimps pools could be found in each step provides refuge for the (Crustacea Decapoda Caridea) from the Philippines, with Tropical Zoology 17 aquatic animals. There was also one ovigerous female of descriptions of five new species. , : 233–266. Caridina typus H. Milne Edwards, 1837, with carapace length 8.3 mm, collected from lowest part of this cave in Cai, Y. & M. M. Bahir, 2005. Lancaris, a new genus of freshwater the present collection. Caridina typus is one of most widely shrimp from Sri Lanka (Crustacea: Decapoda: Atyidae). Raffl es Bulletin of Zoology, Supplement No. 12: 229–235. distributed species in islands of the Indo-West Pacifi c and requires seawater to complete its life cycle (Cai, pers. obs.). De Haan, W., 1833–1850, Crustacea, in P. F. von Siebold (ed.), The presence of C. typus thus indicates that the stream in Fauna Japonica sive Descripto Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum Marulas Cave has connection with the sea. in India Batava Imperium Tenent, Suscepto, Annis 1823–1830 Collegit, Notris, Observationibus et Adumbrationibus Illustravit, Etymology. – The species is named after the type locality- i-xxxi, ix-xvi, 1–243, Pls. A–J, l-Q, 1–55, circ. tab. 2. Lugduni- Marulas Cave in Luzon, the Philippines. Name used as an Batavorum. [Leiden]. apposition. Fransen, C. H. J. M., L. B. Holthuis & J. P. H. M. Adema, 1997. Type-catalogue of the decapod Crustacea in the collections of Remarks. – With respect to the rostrum and the eyes, Edoneus the Nationaal Natuurhistorisch Museum, with appendices of marulas is similar to E. atheatus and E. sketi. But the stout pre-1900 collectors and material. Zoologische Verhandelingen, carpus of the fi rst pereiopod (carpus 1.6 times as long as 311: 1–334. high vs. 2.0 times in E. atheatus, and 2.4 times in E. sketi) Holthuis. L. B., 1956a. An enumeration of the Crustacea Decapoda can easily differentiate it from the latter two species. With Natantia inhabiting subterranean waters. Vie et Milieu, 7: regard to the form of the second pereiopod, E. marulas is 43–76. similar to E. erwini, but can be distinguished by its short Holthuis, L. B., 1956b. The troglobitic Atyidae of Madagascar and unarmed rostrum, and the strongly reduced eyes (vs. (Crudtacea, Decapoda, Natantia), Memoirs of the Institute of eyes developed). Science, Madagascar, ser A, 11: 97–110. Holthuis, L. B., 1978. Zoological results of the British Speleological Expedition to Papua New Guinea 1975. 7. Cavernicolous Key to the species of Edoneus shrimps (Crustacea, Decapoda, Natantia) from New Ireland and the Philippines. Zoologische Mededelingen, 53: 209–224. 1. Eyes reduced distinctly ...... 2 Holthuis, L. B., 1993. The recent genera of the caridean and – Eyes developed ...... Edoneus erwini stenopodidean shrimps (Crustacea, Decapoda) with an appendix 2. Rostrum unarmed, carpus of fi rst pereiopods stout, less than on the order Amphionidacea. Nationaal Natuurhistorisch twice as long as high ...... 3 Museum, Leiden: 1–328, Figs.1–312. – Rostrum armed ventrally with 1-4 teeth, carpus of first pereiopods slender, 2.4 times as long as high ...... E. sketi Milne Edwards, H., 1837, Histoire naturelle des Crustacés, 3. Carpus of fi rst pereiopods 1.6 times as long as high ...... comprenant l’anatomie, la physiologie et la classifi cation de ...... E. marulas ces animaux, 2, 1–532, atlas, Pls. 1–14. – Carpus of fi rst pereiopods 2.0 times as long as high ...... Shokita, S., 1981. Life-history of the family Atyidae (Decapoda, ...... E. atheatus Caridea). Aquabiology, 12, 3(1): 15–23, Figs. 1–9 [in Japanese with English abstract]. Williams, W. D., 1964. Subterranean freshwater prawns (Crustacea: ACKNOWLEDGEMENTS Decapoda: Atyidae) in Australia. Australian Journal of Marine and Freshwater Research, 15: 93–106. The authors wish to thank Dr. P. K. L. Ng (ZRC) and Prof. Boris Sket (University of Ljubljana, Slovenia) for their comments and critical review of the manuscript and also for the continuous support and encouragement in the course of the present study. The collection of the specimens is funded partly by the Offi ce of the Vice Chancellor for Research and Development (UP-OVCRD), University of the Philippines in Diliman, Quezon City, Philippines.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 65–89 Date of Publication: 28 Feb.2009 © National University of Singapore

EPIGEAN AND HYPOGEAN FRESHWATER SHRIMPS OF BOHOL ISLAND, CENTRAL PHILIPPINES (CRUSTACEA: DECAPODA: CARIDEA)

Yixiong Cai National Biodiversity Centre, National Parks Board, 1 Cluny Road, Singapore 259569, Republic of Singapore Email: [email protected] (Corresponding author)

Satish Choy Queensland Department of Natural Resources and Water, 120 Meiers Road, Indooroopilly, Queensland 4068, Australia Email: [email protected]

Peter K. L. Ng Tropical Marine Science Institute, National University of Singapore, Republic of Singapore Email: [email protected]

ABSTRACT. – The epigean and hypogean freshwater shrimps from Bohol Island, central Philippines, collected during various speleological expeditions by the Philippine, Slovenian and Singaporean scientists between 1995 and 2000 are reported here. The material contains 25 species of caridean shrimps in two families. The Atyidae are represented by 16 species from four genera: Antecaridina, Halocaridinides and Atyopsis, each with one species; and Caridina, with 13 species, including seven new taxa. The Palaemonidae are represented by two genera, Macrobrachium with eight species and Palaemon with one. Detailed descriptions are provided for new and rarely reported species.

KEY WORDS. – Bohol, Philippine, freshwater shrimp, Atyidae, Palaemonidae, new species.

INTRODUCTION Biodiversity Research, the National University of Singapore, and the Department of Biology, University of San Carlos, The freshwater shrimps of the Philippine Archipelago have Cebu, in December 2000, resulted in the identifi cation of 25 been comparatively well studied (Dana, 1852; Cowles, 1915a, species of caridean shrimps: 16 species of Atyidae, including 1915b; Blanco, 1935, 1939; Holthuis, 1950; 1978; Johnson, seven new species and nine species of Palaemonidae. In this 1962; Chace, 1983; 1997; Balete & Holthuis, 1992; Chace study, detailed descriptions and illustrations are provided & Bruce, 1993; Sket, 1997; Cai & Anker, 2004; Cai & for the new and the rarely reported species, and brief notes Shokita, 2006a). Prior to this study, 52 species of freshwater are provided for the more common and widely distributed caridean shrimps were reported from the Philippines. The species. family Atyidae was represented by seven genera with 28 species: Antecaridina, Halocaridinides, Atyoida, Atyopsis, All the specimens discussed in this study were collected Parisia, Edoneus, with one species each, and Caridina, with from various localities on Bohol Island, the Philippines (Fig. 22 species. The family Palaemonidae was represented by 1). Specimens examined are deposited in the Zoological two genera with 22 species: Macrobrachium with 20 species Reference Collection, the Raffl es Museum of Biodiversity and Palaemon with two species. The family Alpheidae was Research, the National University of Singapore, Singapore represented by two species of Potamalpheops. (ZRC); Department of Biology, University of San Carlos, Cebu, Philippines (USC); and Department of Biotechnology, The examination of the recent collection from the Bohol University of Ljubljana, Slovenia (UOL). The abbreviation cl Island (Fig. 1), central Philippines, collected by Dr. Boris is used for carapace length measured in mm from the post- Sket and his Slovenian colleagues in February 1995 and orbital margin to the posterior margin of the carapace. March 1999, and a joint expedition by the Raffl es Museum of

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SYSTEMATIC ACCOUNT Atyopsis Chace, 1983

Atyidae De Haan, 1849 Atyopsis spinipes (Newport, 1847)

Antecaridina Edmondson, 1954 Atya spinipes Newport, 1847: 159 [type locality: Philipine Islands] Antecaridina lauensis (Edmondson, 1935) Atya moluccensis – De Man, 1902: 893 (not Atya moluccensis De Haan, 1849). Atyopsis spinipes – Chace, 1983: 35, Figs. 20–22; 1997: 4; Cai & Mesocaris lauensis Edmondson, 1935a: 13, Fig. 4 [type locality: Shokita, 2006b: 2133. Namuka and Wangava Island, Lau Archipelago, Fiji]; Edmondson, 1935b: 4. Material examined. – Two females, cl 6.0–7.2 mm, ZRC 2007.0268, Antecaridina lauensis – Edmondson, 1954: 368; Holthuis, 1955: River outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 25, Fig. 8d–f; 1956: 51; 1963: 267; 1965: 4, Figs. 2a–r; 1973: Dec.2000. 19; Shokita, 1975: 118; 1979: 201; Suzuki, 1980: 47, Figs. 1–5; Smith & Williams, 1981: 49; Hayashi, 1989a: 378, Figs. 172–174; Sket, 1997: 62; Leberer & Cai, 2003: 355; Cai & Remarks. – Chace (1997) reported this species from Luzon, Shokita, 2006b: 2124. Catanduanes, Cebu and a location that probably Mindoro. De Man (1902) reported Atya moluccensis from Halmahera, Material examined. – One male, cl 2.8 mm, 16 females, cl 1.8–2.6 for which, Chace (1983) commentd that De Man’s (1902) mm, 12 juveniles, ZRC 2007.0267, Cave Hinaganan, Panglao, coll. specimens “probably represent Atyopsis spinipes, as indicated Y. Cai et al., 17 Dec.2000; 60 juveniles, UOL, Cave Hinaganan, by the fewer (3–5) ventral teeth.” According to Chace Panglao, coll. B. Sket, Feb.1995. (1983), the most reliable characters used to separate Atyopsis moluccensis from A. spinies is the large number of ventral Remarks. – Antecaridina lauensis, an anchialine species rostral teeth as 7–16 (vs. 2–6). Atyopsis spinipes occurs with a wide but disjunct distribution, has been reported from the Philippines and eastern Lesser Sunda Islands (at from the Lau Archipelago, Fiji Islands, (Edmondson, ca.120°00'E) northwards to Taiwan and Tokuno-shima in the 1935a, b), Europa Island, Madagascar (Holthuis, 1965), Ryukyus, and eastwards as far as Samoa (Chace, 1983). Sinai Peninsular, Red Sea (Holthuis, 1963), Hawaii Island (Holthuis, 1973), Solomon Islands (Smith & Williams, 1981), Guam (Leberer & Cai, 2003) and the Ryukyus (Shokita, Caridina H. Milne Edwards, 1837 1975; 1979; Suzuki, 1980). It has previously been reported from Bohol by Sket (1997). Caridina typus H. Milne Edwards, 1837

Caridina typus H. Milne Edwards, 1837: 363 [type locality: Halocaridinides Fujino & Shokita, 1975 unknown]; Holthuis, 1965: 10, Fig. 3; Chace, 1997: 21; Cai & Anker, 2004: 236; Wowor et al., 2004: 341; Fig. 5F; Cai et al., Halocaridinides trigonophthalma 2006: 412, Figs. 13–15; Cai & Shokita, 2006b: 2134. (Fujino & Shokita, 1975) Material examined. – One female, cl 3.2 mm, ZRC 2007.0269, river Halocaidina (Halocaridinides) trigonophthama Fujino & Shokita, outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000; 1975: 106, Figs. 7, 8 [type locality: a well in Okinawa Island, 1 male, cl 4.7 mm, 8 females, cl 2.8–4.0 mm, 2 ovigerous females, Ryukyus, Japan]; Shokita, 1979: 201. cl 6.8–7.2 mm, ZRC 2007.0270, River outside Inabacan Cave, Palauatya dasyomma Hart, 1980: 481, Figs. 1–31 [type locality: Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male, cl 5.7 mm, 3 Anguar Island, Palau, Caroline Islands]. ovigerous females, cl 6.6–7.8 mm, ZRC 2007.0271, Loboc river, Halocaridinides trigonophthalma – Holthuis, 1982: 31, Figs. 3a–u; Loboc, coll. Y. Cai et al, 19 Dec.2000. Gurney, 1984: 591; Hayashi, 1989b: 497, Fig. 177; Shokita, 2003: 249, Fig. 17C; Naruse et al., 2003: 1; Cai & Shokita, Remarks. – Caridina typus is a very common insular or 2006b: 2132. coastal freshwater species, which is widely distributed in the islands of Indo-West Pacifi c region. Material examined. – Eight juveniles, incomplete, UOL, Alijauan Bridge, stream, S. Duero, coll. B. Sket, 5 Mar.1999. Caridina villadolidi Blanco, 1939 Remarks. – The taxonomy of Halocaridinides trogonophthalma has recently been discussed by Naruse et al. (2003) and Cai Caridina typus var. longirostris De Man, 1892: 370, Pl. 22: 22f–i & Shokita (2006b). The present record is based on eight [type locality: Palopo, Sulawesi (Celebes), Indonesia]. incomplete juvenile specimens, all of which were collected Caridina villadolidi Blanco, 1939: 389, Pl. 1 [type locality: Laoag from an interstitial habitat. These specimens, however, clearly River, Luzon, the Philippines] Hung et al., 1993: 485, Fig. 3; show the characteristic of the species, i.e. the extremely short Chace, 1997: 21, Fig. 12; Cai & Ng, 2001: 668, Fig. 4; Cai rostrum, the slender pereiopods, the fewer uropodal teeth & Shokita, 2006a: 248. as well as the short and stout telson. This represents a new Caridina typus – De Silva, 1982: 135. Figs. 4a–h (not Caridina record for the Philippines. typus H. Milne Edwards, 1837).

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Material examined. – Ten females, cl 3.3–3.8 mm, ZRC 2007.0272, mm, ZRC 2007.0276, river outside Inabacan Cave, Antequrra, coll. river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Y. Cai et al., 18 Dec.2000. Dec.2000; 1 male, cl 3.4 mm, 11 females, cl 3.4–4.4 mm, ZRC 2007.0273, river outside Inabacan Cave, Antequrra, coll. Y. Cai Remarks. – Chace (1997) commented that Caridina et al., 18 Dec.2000; 1 male, cl 4.9 mm, ZRC 2007.0274, Loboc laoagensis will eventually fall into synonymy with the River, Loboc, coll. Y. Cai et al., 19 Dec.2000. variable C. weberi De Man, 1892. Cai & Anker (2004) has shown that C. laoagensis differs markedly from all subspecies Remarks. – Caridina villadolidi has been reported from of C. weberi and other congeners, and should be regarded Taiwan (Hung et al., 1993), the Philippines (Blanco, 1939; as a good species. Caridina laoagensis is characterized by Chace, 1997; Cai & Shokita, 2006a), Sulawesi and Halmahera its straight rostrum which is slightly crested basally above in Indonesia (Cai & Ng, 2001), and Sri Lanka (De Silva, the orbit, and the pattern of distal spines and setae on the 1982, misidentifi ed as Caridina typus). telson, on which, the median plumose seatae are inserted betweem two lateral spines. This is one of the most common species of atyids in the Philippines. Caridina laoagensis Blanco, 1939

Caridina laoagensis Blanco, 1939: 390, Pl. 2 [type locality: Laoag Caridina endehensis De Man, 1892 River, Province of Ilocos Norte, Luzon, the Philippines]; Chace, 1997: 12; Cai & Anker, 2004: 237, Fig. 3; Cai & Shokita, 2006a: 248; Cai & Shokita, 2006b: 2141. Caridina brevicarpalis var. endehensis De Man, 1892: 399, Pl. 24 Fig. 30e [type locality: Nuawari, near Ende, Flores, Indonesia]; Material examined . – Two males, cl 2.6–2.9 mm, 1 female, cl Bouvier, 1925: 34; Roux, 1928: 218; Blanco, 1935: 34, Pl. 2 2.9 mm, 2 ovigerous females, cl 5.6–6.0 mm, ZRC 2007.0275, Fig. 25; Chace, 1997: 8, Fig. 3. river outside the Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Caridina brevicarpalis – Holthuis, 1978: 38 (not Caridina Dec.2000; 3 males, cl 3.2–3.7 mm, 2 ovigerous females, cl 6.2–6.3 brevicarpalis De Man, 1892).

Fig. 1. Map of the Bohol Island, Philippines, showing the collection sites.

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Caridina endehensis – Wowor et al., 2004: 341, Fig. 5M; Cai & Remarks. – Cai & Ng (2007) recently reviewed the taxonomy Shokita, 2006a: 248. of the Caridina gracilirostris species group in detail, referring the form with no appendix interna in endopod of male fi rst Material examined. – 13 males, cl 3.0–4.2 mm, 25 females, cl pleopod as C. gracilirostris sensu stricto and the other, 2.7–6.2 mm, 2 ovigerous, females, cl 4.6–6.2 mm, ZRC 2007.0277, with a distinct appendix interna in the endopod of male river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 11 males, cl 3.2–4.6 mm, 13 females, cl 2.9–4.8 mm, 2 fi rst pleopod, as C. appendiculata Jalihal & Shenoy, 1998. ovigerous females, cl 5.3–6.4 mm, ZRC 2007.0278, river outside The specimens from Bohol have no appendix interna on the Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000. endopod of the male fi rst pleopod, and are here referred to C. gracilirostris. It is an insular or coastal freshwater species Remarks. – Holthuis (1978) reported Caridina brevicarpalis commonly distributed in Indo-West Pacifi c region. from Sumba, Lesser Sunda Islands, Indonesia. According to his description, his specimens “can unhesitatingly be brought to ssp. endehensis”. Caridina endehensis has previously Caridina lobocensis, new species been reported from the Philippines by Blanco (1935), (Figs. 2–4) Chace (1997) and Cai & Shokita (2006a). It is distributed from Philippines to Flores and Sumba in Indonesia. The Material examined. – Holotype: male, cl 5.7 mm, USC, tributary prominently elongated rostrum easily separates C. endehensis of Loboc River, Loboc, coll. Y. Cai, 19 Dec.2000. from C. brevicarpalis and does not appear to be variable. Paratypes: Seventy males, cl 3.8–5.3 mm, 19 females, cl 3.9–7.5 We therefore follow Cai & Shokita (2006a) and regard the mm, 83 ovigerous females, cl 5.3–7.2 mm, ZRC 2007.0283, data taxon as a full species. same as holotype.

Description. – Rostrum (Figs. 2A, 3A) straight, reaching Caridina elongapoda Liang & Yan, 1977 slightly beyond end of antennular peduncle; rostral formula: 2–4+10–12/3–7. Antennal spine fused with inferior orbital Caridina nilotica elongapoda Liang & Yan, 1977: 220, Figs. 5–8 angle. Pterygostomian margin subrectangular. [type locality: Xinzai, Gulei village, Zhangpu County, Fujian, southern China]. Caridina aff. brachydactyla - Yeo et al., 1999: 218, Figs. 10–14; Sixth abdominal somite 0.44 times of carapace, 1.4 times Wowor et al., 2004: Fig. 6B; as long as fi fth somite, distinctly shorter than telson. Telson Caridina elongapoda Cai & Shokita, 2006a: 249. (Figs. 3B, 4A) 2.6 times as long as wide, terminating in a projection, with 3 or 4 pairs of dorsal spinules and 1 pair of Material examined. – Two males, cl 4.1–4.5 mm, 3 ovigerous dorsolateral spinules; lateral pair of spines distinctly shorter females, cl 5.6–5.8 mm, ZRC 2007.0279, tributary of Loboc River, than intermediate pairs of setae. Preanal carina (3I) high, Loboc, coll. Y. Cai, 19 Dec.2000; 1 male, cl 4.0 mm, 5 females, cl lacking spine. 2.8–3.4 mm, 3 ovigerous females, cl 5.3–6.4 mm, ZRC 2007.0280, river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Eyes developed, anterior end reaching to 0.8 times length of Dec.2000; 6 males, cl 3.1–4.7 mm, 17 females, cl 2.8–4.3 mm, ZRC 2007.0281, river outside Inabacan Cave, Antequrra, coll. Y. basal segment of antennular peduncle. Antennular peduncle Cai et al., 16 Dec.2000. 0.55 times as long as carapace; basal segment of antennular peduncle as long as both of second and third segment Remarks. – Caridina elongapoda has recently been reported lengths, anterolateral angle reaching 0.20 times length of from the Philippines (Cai & Shokita, 2006a). Cai & Shokita second segment, second segment distinctly longer than third (2006a) reassigned the material identifi ed as Caridina aff. segment. Stylocerite reaching slightly beyond end of basal brachydactyla by Yeo et al. (1999) from Pulau Tioman, segment of antennular peduncle. Scaphocerite (Fig. 2B) 2.5 Malaysia, to this species.Current known distribution is times as long as wide. southern China, Malaysia and the Philippines. Incisor process of mandible (Fig. 2C) ending in irregular teeth, molar process truncated. Lower lacinia of maxillula Caridina gracilirostris De Man, 1892 (Fig. 2D) broadly rounded, upper lacinia elongated, with a number of distinct teeth on inner margin, palp slender. Caridina gracilirostris De Man, 1892: 399, Pl. 25 Fig. 31–31d Upper endites of maxilla (Fig. 2E) subdivided, palp short, [type locality: Balangnipa, Sulawesi (Celebes) Indonesia]; scaphognathite tapering posteriorly with some long, curved Bouvier, 1925: 142, Figs. 305–307; Holthuis, 1965: 23, Fig. setae at posterior end. Palp of fi rst maxilliped (Fig. 2F) 7; Tiwari & Pillai, 1971: 83, Fig. 2a, b; Chace, 1997: 10, Fig. broadly triangular. Second maxilliped (Fig. 2G) typical 4; Wowor et al., 2004, Fig. 5Q; Cai & Shokita, 2006a: 250; of genus. Third maxilliped (Fig. 2H) reaching to end of Cai & Shokita, 2006b: 2135; Cai & Ng, 2007: 1586. antennular peduncle, with ultimate segment shorter than penultimate segment. Material examined. – Two males, cl 2.5–2.6 mm, 3 females, cl 2.0–3.3 mm, 1 ovigerous female, cl 3.7 mm, ZRC 2007.0282, tributary of Loboc River, Loboc, coll. Y. Cai, 19 Dec.2000. Epipods on fi rst 4 pereiopods. First pereiopod (Figs. 3C, 4D) reaching to end of basal segment of antennular peduncle; merus 1.7–2.1 times as long as broad, slightly longer than

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Fig. 2. Caridina lobocensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, scaphocerite; C, mandible; D, maxillula; E, maxilla; F, fi rst maxilliped; G, second maxilliped; H, third maxilliped. Scale bars: A = 2 mm, B–H = 1 mm (female, cl 6.9 mm, Paratype, ZRC, Philippines).

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Fig. 3. Caridina lobocensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, fi rst pereiopod; D, second pereiopod; E, third pereiopod; F, dactylus of third pereiopod; G, fi fth pereiopod; H, dactylus of fi fth pereiopod; I, preanal carina; J, uropodal diaeresis. Scale bars: A = 2 mm, B, F, H, J = 0.2 mm; C–E, G = 1 mm; I = 0.5 mm (male, cl 5.7 mm, paratype, ZRC, Bohol Island, Philippines).

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Fig. 4. Caridina lobocensis, new species: A, telson; B, endopod of male fi rst pleopod; C, appendix masculina and appendix interna of male second pleopod; D, fi rst pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fi fth pereiopod; I, dactylus of fi fth pereiopod. Scale bars: A, D–F, H = 1 mm; B, C, G, I = 0.2 mm (A–C, male, cl 5.7 mm, paratype, ZRC, Bohol Island, Philippines; D–I, female, cl 6.9 mm, paratype, ZRC, Bohol Island, Philippines).

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carpus; carpus excavated anteriorly, shorter than chela, 1.1– of the antennular peduncle vs. does not reaches) (Figs. 2, 1.2 times as long as high; chela 2.2 times as long as broad; 3; cf. Cai & Shokita, 2006b: Figs. 13, 14). fi ngers distinctly shorter than palm. Second pereiopod (Figs. 3D, 4E) reaching to end of second segment of antennular peduncle; merus as long as carpus, 4.2–5.0 times as long as Caridina liaoi, new species broad; carpus 1.1 times as long as chela, 2.3–2.4 times as (Figs. 5, 6) long as high; chela 2.3–2.4 times as long as broad; fi ngers 1.6 times as long as palm. Third pereiopod (Figs. 3E, F, 4F, G) Material examined. – Holotype: Ovigerous female, cl 4.1 mm, reaching slightly beyond end of scaphocerite, propodus 9–11 USC, Bilar River, Bilar, coll. Y. Cai et al., 19 Dec.2000. times as long as broad, 5.2–5.7 times as long as dactylus; dactylus 2.7–3.0 times as long as wide (spines included), Paratypes: 82 males, cl 2.8–4.3 mm, 112 females, cl 3.7–5.0 mm, 26 ovigerous females, cl 4.2–5.0 mm, ZRC 2007.0284, data same terminating in 2 very strong claws, with 4–6 accessory as holotype. spines on its fl exor margin. Fifth pereiopod (Figs. 3G, H, 4H, I) reaching beyond end of second segment of antennular Description. – Rostrum (Figs. 5A, 6A) long, reaching near peduncle, propodus 13–18 times as long as broad, 5.5–5.6 to or slightly beyond end of scaphocerite; rostral formula: times as long as dactylus, dactylus 2.7–3.0 times as long as 3–4+13–14/13–15. Antennal spine fused with inferior orbital wide (spinules included), terminating in 2 large claws, with angle. Pterygostomian margin broadly rounded. 26–30 spinules on its fl exor margin. Sixth abdominal somite 0.51 times of carapace, 1.7 times Endopod of male fi rst pleopod (Fig. 4B) subtriangular, half as long as fi fth somite, distinctly shorter than telson. Telson length of exopod, appendix interna stout, exceeding end of (Figs. 6B, C) 3.5 times as long as wide, not terminating in a endopod. Appendix masculina of male second pleopod (Fig. projection, with 3 or 4 pairs of dorsal spinules and 1 pair of 4C) 2/3 length of endopod, with appendix interna reaching dorsolateral spinules; lateral pair of spines distinctly longer base of distal one-third of appendix masculina. than intermediate pairs of spiniform setae. Preanal carina (Fig. 6K) high, triangular, lacking spine. Uropodal diaeresis (Fig. 3J) with 18 movable spinules. Eyes well developed, anterior end reaching to 0.8 times Ovigerous females with eggs sized 0.45 × 0.25 mm. length of basal segment of antennular peduncle. Antennular peduncle 0.74 times as long as carapace; basal segment Habitat. – Caridina lobocensis, new species, was collected of antennular peduncle longer than both second and third from a tributary of the Loboc River near Loboc town, Bohol segment lengths, anterolateral angle reaching 0.20 times Island in central Philippines. The shrimps were living among length of second segment, second segment distinctly longer submerged grass at edges of the tributary. than third segment. Stylocerite reaching to 0.8 times length of basal segment of antennular peduncle. Scaphocerite (Fig. Etymology. – The new species is named after its type locality- 6D) 3.4 times as long as wide. Loboc River, Bohol Island, the Philippines. Incisor process of mandible ending in irregular teeth, molar Remarks. – Based on the form of the rostrum, the high process truncated. Lower lacinia of maxillula broadly preanal carina, the large number of uropodal teeth and the rounded, upper lacinia elongated, with a number of distinct form of the telson, C. lobocensis, new species, belongs to teeth on inner margin, palp slender. Upper endites of maxilla the C. weberi species group. In the group, it is most similar subdivided, palp short, scaphognathite tapering posteriorly to C. okinawa Cai & Shokita, 2006b, from Okinawa Island with some long, curved setae at posterior end. Palp of fi rst of the Ryukyu Islands, southern Japan, in term of the rostral maxilliped broadly triangular. Second maxilliped typical formula, the long stylocerite, the broad scaphocerite, the stout of genus. Third maxilliped reaching to end of antennular fi rst pereiopod, and its short fi ngers, and the third to fi fth peduncle, with ultimate segment as long as penultimate pereiopods ending in two claws. However, it could be easily segment. separated from C. okinawa by the well developed eyes (vs. eyes less developed, with a short eye stalk and small cornea) Epipods on fi rst 4 pereiopods. First pereiopod (Figs. 5B, 6E) and the longer rostrum which reaches slightly beyond end reaching to end of basal segment of antennular peduncle; of antennular peduncle (vs. reach to end of second segment merus 1.9–2.2 times as long as broad, shorter than carpus; of antennular peduncle) (Figs. 2–4; cf. Cai & Shokita, carpus excavated anteriorly, shorter than chela, 1.3–1.7 times 2006b: Fig. 12). Caridina lobocensis also resembles another as long as high; chela 2.3–3.5 time as long as broad; fi ngers Ryukyuan species, C. macrodentata Cai & Shokita, 2006b, slightly shorter, as long as or distinctly longer than palm. in the form of the dactylus of the third to fi fth pereiopods, Second pereiopod (Figs. 5C, 6F) reaching to end of second Nevertheless, it could be differentiated from the latter by segment of antennular peduncle; merus as long as carpus, the longer rostrum, which reaches slightly beyond the end 4.2–5.0 times as long as broad; carpus 1.1 times as long as of the antennular peduncle (vs. reaches middle of the second chela, 2.3–2.4 times as long as high; chela 2.3–2.4 times segment of the antennular peduncle) and the long stylocerite as long as broad; fi ngers 1.6 times as long as palm. Third (reaches to or slightly beyond the end of the basal segment pereiopod (Figs. 5D, E, 6G, H) reaching to end of antennular

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Fig. 5. Caridina liaoi, new species. A: cephalothorax and cephalic appendages, lateral view; B, fi rst pereiopod; C, second pereiopod; D, third pereiopod; E, dactylus of third pereiopod; F, fi fth pereiopod; G, dactylus of fi fth pereiopod; H, endopod of male fi rst pleopod; I, appendix masculina and appendix interna of male second pleopod. Scale bars: A–D, F = 1 mm; E, G = 0.1 mm; H, I = 0.2 mm (male, cl 4.0 mm, paratype, ZRC, Bilar river).

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Fig. 6. Caridina liaoi, new species, A: cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, telson; D, scaphocerite; E, fi rst pereiopod; F, second pereiopod; G, third pereiopod; H, dactylus of third pereiopod; I, fi fth pereiopod; J, dactylus of fi fth pereiopod; K, preanal carina; L, uropodal diaeresis. Scales: A, C–G, I = 1 mm; B, L = 0.2 mm; H, J = 0.1 mm; K = 0.5 mm. (ovig. females, cl 4.6 mm, paratype, ZRC, Bilar River).

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peduncle, propodus 11–12 times as long as broad, 4.5–4.7 Antequrra, coll. Y. Cai et al., 16 Dec.2000; 41 males, cl 2.6–3.5 times as long as dactylus; dactylus 3.1–3.3 times as long as mm, 38 females, cl 2.9–4.1 mm, ZRC 2007.0287, Cave Bongkawi, wide (spines included), with 4–6 accessory spines on fl exor Canla-As village, Antequrra, coll. Y. Cai.et al., 16 Dec.2000; 3 margin. Fifth pereiopod reaching to end of second segment males, cl 3.0–3.8 mm, 11 females, cl 2.3–3.8 mm, ZRC 2007.0288, spring outside cave Bones, Canla-As village, Antequrra, coll. P. of antennular peduncle, propodus 14–18 times as long as K. L. Ng & L. Liao, 16 Dec.2000; 6 males, cl 2.8–3.4 mm, ZRC broad, 3.6–3.8 times as long as dactylus, dactylus 4.0–4.4 2007.0289, spring outside Cave Inabacan, Antequrra, coll. Y. times as long as wide (spinules included), terminating in 1 Cai et al., 18 Dec.2000; 22 males, cl 3.0–4.0 mm, 37 females, large claw, with 60–65 spinules on fl exor margin. cl 2.2–3.3 mm, 15 juveniles, ZRC 2007.0290, Cave Serpia, Ginguyuran, Dimiao, coll. B. Sket, Feb.1995; 1 juvenile, ZRC Endopod of male fi rst pleopod (Fig. 5H) sub-rectangular, 2007.0291, Cave Camulaon, lower part, Cugon, Jagna, coll. B. two-fi fth length of exopod, no appendix interna. Appendix Sket, Feb.1995; 7 juveniles, ZRC2007.0292 , spring no.1, Roxas masculina of male second pleopod (Fig. 5I) half length Park, Garcia-Hernandez, coll. B. Sket, Feb.1995; 4 juveniles, of endopod, with appendix interna reaching near end of ZRC 2007.0293, spring no. 2, Roxas Park, Garcia-Hernandez, coll. B. Sket, Feb.1995; 6 juveniles, ZRC 2007.0294, spring appendix masculina. Anislaq, outside, Valencia, coll. B. Sket, Feb.1995; 2 juveniles, ZRC 2007.0295, spring below Cave Serpia, Ginguyuran, Dimiao, Uropodal diaeresis (Fig. 6L) with 14–16 movable B. Sket, Feb.1995; 23 juveniles, ZRC 2007.0296, spring Anislaq, spinules. outside, Velencia, coll. B. Sket, Feb.1995.

Ovigerous females with eggs sized 0.92–0.95 × 0.55–0.60 Description. – Rostrum (Figs. 7A, 8A) very short, reaching mm. near end of basal segment of antennular peduncle; unarmed dorsally or armed dorsally with indistinct small teeth, unarmed Habitat. – Caridina liaoi, new species, was collected from ventrally. Antennal spine indistinct, fused with inferior orbital a tributary of the Bilar River near Bilar town, Bohol Island angle. Pterygostomian margin broadly rounded. in central Philippines. Sixth abdominal somite 0.45 times of carapace, 1.8 times Etymology. – The new species is named after Dr. Lawrence as long as fi fth somite, as long as telson. Telson (Figs. Liao, who has been instrumental in helping the fi rst author 7B, C) 2.8 times as long as wide, not terminating in a making collections in the Philippines for the present projection, with 3 or 4 pairs of dorsal spinules and 1 pair of study. dorsolateral spinules; lateral pair of spines distinctly longer than intermediate pairs. Preanal carina high, sub-triangular, Remarks.- Caridina liaoi, new species, is similar to C. lacking spine. buhi Cai & Shokita, 2006a, from Luzon, by the form of pereiopods, but it differs from C. buhi by its longer rostrum Eyes reduced, anterior end reaching to 0.3–0.4 times length (reaches to or slightly beyond the anterior margin of the of basal segment of antennular peduncle. Antennular scaphocerite vs. reaches to the end of the second segment peduncle 0.64 times as long as carapace; basal segment of the antennular peduncle or to the end of the antennular of antennular peduncle shorter than both second and third peduncle); the more ventral rostral teeth (13–15 vs. 3–7); the segment lengths, anterolateral angle reaching 0.20 times telson does not terminate in a projection vs. terminate in a length of second segment, second segment distinctly longer projection; the bigger appendix interna on the male second than third segment. Stylocerite reaching to 0.8 times length pleopod (reaches near the end of the appendix masculina vs. of basal segment of antennular peduncle. Scaphocerite (Fig. reaches the base of the distal two-third length of the appendix 7D) 3.5 times as long as wide. masculina) and the distal spines on the telson (lateral spines much longer than the intermediate pairs of spiniform setae Incisor process of mandible (Fig. 7E) ending in irregular vs. lateral spines subequal to intermediate pairs) (Figs. 5, teeth, molar process truncated. Lower lacinia of maxillula 6; cf. Cai & Shokita, 2006a: Figs. 3, 4). (Fig. 7F) broadly rounded, upper lacinia elongated, with a number of distinct teeth on inner margin, palp slender. Upper endites of maxilla (Fig. 7G) subdivided, palp short, Caridina boholensis, new species scaphognathite tapering posteriorly with some long, curved (Figs. 7, 8) setae at posterior end. Palp of fi rst maxilliped (Fig. 7H) broadly triangular. Second maxilliped (Fig. 7I) typical Material examined. – Holotype: Male cl 3.9 mm, USC, Quila Cave, of genus. Third maxilliped (Fig. 7J) reaching to end of Nueva Vida Norte, Batuan, coll. B. Sket, Feb.1995. antennular peduncle, with ultimate segment slightly shorter than penultimate segment. Paratypes: 87 males, cl 3.0–4.2 mm, 115 females, cl 2.7–4.7 mm, 1 ovigerous females, 3.8 mm, eggs 0.75 x 0.50 mm, UOL, data same as holotype; 10 males, cl 3.0–4.2 mm, 10 females, cl 2.7–4.7 Epipods on fi rst four pereiopods. First pereiopod (Figs. 7K, mm, 1 ovigerous females, 4.6 mm, eggs 0.75 × 0.50 mm, ZRC 8B) reaching beyond end of basal segment of antennular 2007.0285, data same as holotype. peduncle; merus 2.3 times as long as broad, shorter than carpus; carpus excavated anteriorly, shorter than chela, 2.0 Others: Nine males, cl 2.8–3.8 mm, 5 females, cl 37–3.8 mm, times as long as high; chela 2.3 times as long as broad; 11 juveniles, ZRC 2007.0286, Cave Bones, Canla-As village, fi ngers slightly longer than palm. Second pereiopod (Figs.

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Fig. 7. Caridina boholensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, scaphocerite; E, mandible; F, maxillula; G, maxilla; H, fi rst maxilliped; I, second maxilliped; J, third maxilliped; K, fi rst pereiopod; L, second pereiopod. Scale bars: A, B, D = 1 mm, E–L = 0.5 mm, C = 0.1 mm (male, cl 3.7 mm, paratype, ZRC, Cave Bongkawi, Antequrra).

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Fig. 8. Caridina boholensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, fi rst pereiopod; C, second pereiopod; D, third pereiopod; E, dactylus of third pereiopod; F, fi fth pereiopod; G, dactylus of fi fth pereiopod; H, male fi rst pleopod; I, appendix masculina and appendix interna of male second pleopod; J, uropodal diaeresis. Scale bars: A = 1 mm; B–D, F = 0.5 mm; E, G = 0.1 mm; H–J = 0.2 mm (A–C, ovig. females, cl 3.8 mm, paratype, ZRC; D–J, male, cl 3.7 mm, paratype, ZRC, Cave Bongkawi, Antequrra).

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7L, 8C) reaching to end of scaphocerite; merus distinctly basal segment of antennular pedunlcle; unarmed. Antennal shorter than carpus, 4.4 times as long as broad; carpus 1.2 spine fused with inferior orbital angle. Pterygostomian times as long as chela, 6.9 times as long as high; chela 3.7 margin sub-rectangular. times as long as broad; fi ngers 1.6 times as long as palm. Third pereiopod (Figs. 8D, E, 9E, F) reaching beyond end Sixth abdominal somite 0.43 times of carapace, 1.8 times of scaphocerite by length of dactylus, propodus 10 times as as long as fi fth somite, slightly shorter than telson. Telson long as broad, 3.7 times as long as dactylus; dactylus 3.0 (Fig. 9B) 2.7 times as long as wide, not terminating in a times as long as wide (spines included), with 4 or 5 accessory projection, with 4 pairs of dorsal spinules and 1 pair of spines on fl exor margin. Fifth pereiopod (Figs. 8F, G, 9G, dorsolateral spinules; lateral pair of distal spines distinctly H) reaching to end of scaphocerite, propodus 15 times as longer than intermediate pairs. Preanal carina low, sub- long as broad, 3.6 times as long as dactylus, dactylus 3.4 triangular, lacking spine. times as long as wide (spinules included), terminating in 1 large claw, with 62 spinules on fl exor margin. Eyes well developed, anterior end reaching to 0.7 times length of basal segment of antennular peduncle. Antennular Endopod of male fi rst pleopod (Fig. 8H) sub-triangular, a peduncle 0.63 times as long as carapace; basal segment of quarter length of exopod, no appendix interna. Appendix antennular peduncle distinctly longer than both second and masculina of male second pleopod (Fig. 8I) half length of third segment lengths, anterolateral angle reaching 0.20 times endopod, with appendix interna 0.4 times length of appendix length of second segment, second segment distinctly longer masculina. than third segment. Stylocerite reaching to 0.7 times length of basal segment of antennular peduncle. Scaphocerite 3.6 Uropodal diaeresis (Fig. 8J) with 17 movable spinules. times as long as wide.

Ovigerous females with eggs sized 0.75–0.85 × 0.50–0.60 Incisor process of mandible ending in irregular teeth, molar mm. process truncated. Lower lacinia of maxillula broadly rounded, upper lacinia elongated, with a number of distinct Habitat. – Caridina boholensis, new species, was collected teeth on inner margin, palp slender. Upper endites of maxilla from several locations of subterranean water in Bohol, either subdivided, palp short, scaphognathite tapering posteriorly from cave streams or spring water near caves. with some long, curved setae at posterior end. Palp of fi rst maxilliped rounded, exopod with short fl agellum. Endopod Etymology. – The new species is named after its type of second maxilliped slender. Third maxilliped reaching locality-Bohol Island, the Philippines. beyond end of scaphocerite, with ultimate segment slightly longer than penultimate segment. Remarks. – Caridina boholensis, new species, is most similar to C. cebuensis Cai & Shokita, 2006, from Cebu Island, Epipods on fi rst 4 pereiopods. First pereiopod (Fig. 9C) central Philippines. However, it could be differentiated by stout, reaching to end of second segment of antennular the much reduced eyes, the slender carpus of fi rst pereiopods peduncle; merus 1.6 times as long as broad, slightly shorter (2.0 times as long as high vs. 1.6 times in C. cebuensis) and than carpus; carpus excavated anteriorly, shorter than chela, the slender carpus of second pereiopod (6.9 times as long as 1.3 times as long as high; chela 1.6 times as long as broad; high vs. 4.4 times). Caridina boholensis is the most common fi ngers slightly shorter than palm. Second pereiopod (Fig. 9D) subterreanean Caridina species in Bohol Island. reaching beyond end of scaphocerite; merus distinctly shorter than carpus, 3.3 times as long as broad; carpus 1.2 times as long as chela, 4.5 times as long as high; chela 2.5 times Caridina valencia, new species as long as broad; fi ngers 1.5 times as long as palm. Third (Fig. 9) pereiopod (Figs. 9E, F) reaching beyond end of scaphocerite by length of dactylus, propodus 10 times as long as broad, Material examined. – Holotype: Male, cl 3.5 mm, USC, spring 4.3 times as long as dactylus; dactylus 3.0 times as long Anislaq, outside, Valencia, coll. B. Sket, Feb.1995. as wide (spines included), with 5 or 6 accessory spines on fl exor margin. Fifth pereiopod (Figs. 9G, H) reaching beyond Paratypes: 5 males, cl 2.5–3.6 m, 92 juveniles, UOL, data same end of scaphocerite, propodus 15 times as long as broad, as holotype. 3.0 times as long as dactylus, dactylus 3.5 times as long as Others: 1 juvenile, UOL, spring Badyang, cave, Valencia, coll. wide (spinules included), terminating in 1 claw, with 56 B. Sket, Feb.1995; 1 male, cl 3.1 mm, 1 female, cl 2.8 mm, 1 spinules on fl exor margin. female (damaged), 1 juvenile, UOL, Cave Castigio, Batuan, coll. B. Sket, Feb.1995; 4 males, cl 2.2–2.4 mm, 15 females, cl 2.0–3.1 Endopod of male fi rst pleopod (Fig. 9I) triangular, one-third mm, 2 ovigerous females, cl 2.8–3.1 mm, ZRC 2007.0297, spring length of exopod, no appendix interna. Appendix masculina below Cave Serpia, Ginguyuran, Dimiao, coll. B. Sket, Feb.1995; of male second pleopod (Fig. 9J) half length of endopod, 1 male, cl 3.6 mm, 8 juveniles, ZRC 2007.0298, spring Anislaq, with appendix interna half length of appendix masculina, cave, Valencia, coll. B. Sket, Feb.1995. reaching near end of appendix masculina.

Description. – Rostrum (Fig. 9A) short, reaching near end of Uropodal diaeresis (Fig. 9K) with 17 movable spinules.

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Fig. 9. Caridina valencia, new species: A, cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, fi rst pereiopod; D, second pereiopod; E, third pereiopod; F, dactylus of third pereiopod; G, fi fth pereiopod; H, dactylus of fi fth pereiopod; I, endopod of male fi rst pleopod; J, appendix masculina and appendix interna of male second pleopod; K, uropodal diaeresis. Scale bars: A = 1 mm; B, F, H = 0.1 mm; C–E, G = 0.5 mm; I–K = 0.2 mm. (males, cl 3.4 mm, paratype, ZRC, spring Anislaq, outside Cave Anislaq, Valencia).

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Ovigerous females with egg sized 0.90 × 0.65 mm. Incisor process of mandible ending in irregular teeth, molar process truncated. Lower lacinia of maxillula broadly Habitat. – Caridina valencia, new species, was collected rounded, upper lacinia elongated, with a number of distinct from several caves in the southern part of Bohol Island, teeth on inner margin, palp slender. Upper endites of maxilla central Philippines. subdivided, palp short, scaphognathite tapering posteriorly with some long, curved setae at posterior end. Palp of Etymology. – The new species is named after its type locality- fi rst maxilliped rounded. Podobranch of second maxilliped Valencia Town, Bohol Island, Philippines. The name is used reduced to a lamina. Third maxilliped reaching to end of as a noun in apposition. scaphocerite, with ultimate segment as long as penultimate segment. Remarks. – Caridina valencia, new species, morphologically resembles C. cebuensis Cai & Shokita, 2006a, from Cebu Epipods on fi rst 4 pereiopods. First pereiopod (Fig. 10D) Island, central Philippines. But it could be easily differentiated reaching end of antennular peduncle; merus 2.1 times as from C. cebuensis by the much stouter carpus of the fi rst long as broad, slightly shorter than carpus; carpus excavated pereiopod (1.3 times as long as high vs. 1.6 times); the anteriorly, shorter than chela, 1.6 times as long as high; slender scaphocerite (3.6 times as long as broad vs. 3.1 chela 2.1 times as long as broad; fi ngers slightly shorter than times); and more spinules on the dactylus (56 vs. 39) (Fig. palm. Second pereiopod (Fig. 10E) reaching beyond end of 9; cf. Cai & Shokita, 2006a: Figs. 1, 2). scaphocerite by length of chela; merus distinctly shorter than carpus, 4.5 times as long as broad; carpus 1.3 times as long as chela, 5.3 times as long as high; chela 2.6 times as long Caridina batuan, new species as broad; fi ngers 1.3 times as long as palm. Third pereiopod (Fig. 10) (Figs. 10F, G) reaching beyond end of scaphocerite by length of dactylus and half of propodus, propodus 11 times as long Material examined. – Holotype: Ovigerous female, cl 4.2 mm, as broad, 4.3 times as long as dactylus; dactylus 3.3 times as eggs 0.8 × 0.5 mm, USC, Cave Castigio, Batuan, coll. B. Sket, long as wide (spines included), with 4 or 5 accessory spines Feb.1995. on fl exor margin. Fifth pereiopod (Figs. 10H, I) reaching beyond end of scaphocerite by length of dactylus, propodus 15 Paratypes: One male, cl 4.1 mm, ZRC 2007.0299, same data as times as long as broad, 4.4 times as long as dactylus, dactylus holotype; 4 males, cl 2.2–3.7 mm, 5 females, cl 2.5–3.6 mm, 4 3.7 times as long as wide (spinules included), terminating juveniles, UOL, Cave Kalumpan, Behind-the-Clouds, Batuan, coll. in 1 claw, with 44 spinules on fl exor margin. B. Sket, Feb.1995; 1 female, 1 juvenile, UOL, Spring Badyang, cave, Valencia, coll. B. Sket, Feb.1995; 2 females, cl 1.9–2.3 mm, Endopod of male fi rst pleopod sub-triangular, a quarter length ZRC 2007.0300, Cave Bonugan, Batuan, coll. B. Sket, Feb.1995; of exopod, no appendix interna. Appendix masculina of 2 juveniles, UOL, Alijauan Bridge, stream, S. Duero, coll. B. male second pleopod half length of endopod, with appendix Sket, 5 Mar.1999. interna half length of appendix masculina, reaching near end of appendix masculina. Description. – Rostrum (Fig 10A) short, reaching slightly beyond end of basal segment of antennular pedunlcle; Uropodal diaeresis (Fig. 10J) with 20 movable spinules. rostral formula: 0–2+7–12/0–4. Antennal spine distinct, fused with inferior orbital angle. Pterygostomian margin Ovigerous females with eggs sized 0.80 × 0.50 mm. broadly rounded. Habitat. – Caridina batuan, new species, was collected Sixth abdominal somite 0.50 times of carapace, 1.9 times from several caves in southern part of the Bohol Island, as long as fi fth somite, slightly shorter than telson. Telson central Philippines. (Fig. 10B, C) 2.9 times as long as wide, not terminating in a projection, with 4 or 5 pairs of dorsal spinules and 1 pair of Etymology. – The new species is named after its type locality- dorsolateral spinules; lateral pair of spines distinctly longer Batuan, Bohol Island, the Philippines. The name is used as than intermediate pairs. Preanal carina low, sub-triangular, a noun in apposition. lacking spine. Remarks. – Caridina batuan, new species, is morphologically Eyes reduced, anterior end reaching to 0.4 times length of similar to C. minidentata Cai & Anker, 2004, from west basal segment of antennular peduncle. Antennular peduncle Samar, the Philippines. However, it can be distinguished 0.63 times as long as carapace; basal segment of antennular from C. minidentata by the more developed rostral teeth, peduncle shorter than both second and third segment the much reduced eyes, the lower preanal carina and the lengths, anterolateral angle reaching 0.20 times length of larger number of spinules on diaeresis (20 vs. 16) (Fig. 10; second segment, second segment distinctly longer than third cf. Cai & Anker, 2004: Figs. 4, 5). segment. Stylocerite reaching to 0.7 times length of basal segment of antennular peduncle. Scaphocerite 3.4 times as long as wide.

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Fig. 10. Caridina batuan, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, fi rst pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fi fth pereiopod; I, dactylus of fi fth pereiopod; J, uropodal diaeresis. Scale bars: A, B = 1 mm; C = 0.2 mm, D–F, H = 0.5 mm; G, I, J = 0.1 mm (female, cl 3.0 mm, holotype, USC, Cave Kalumpan, Batuan).

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Caridina anislaq, new species to end of antennular peduncle, propodus 12 times as long (Fig. 11) as broad, 3.3 times as long as dactylus, dactylus 3.5 times as long as wide (spinules included), terminating in 1 claw, Material examined. – Holotype: Male, cl 3.4 mm, USC. with 51 spinules on fl exor margin. Spring Anislaq, outside Cave Valencia, Valencia, coll. B. Sket, Feb.1995. Endopod of male fi rst pleopod (Fig. 11J) triangular, one-third Paratypes: 10 males, cl 2.2–3.4 mm, 9 females, cl 2.2–5.4 mm, length of exopod, no appendix interna. Appendix masculina ZRC 2007.0301, data same as holotype; 1 male, cl 2.4 mm, ZRC (Fig. 11K) of male second pleopod half length of endopod, 2007.0302, Spring Anislaq, Cave Valencia, Valencia, coll. B. with appendix interna half length of appendix masculina, Sket, Feb.1995. reaching near end of appendix masculina.

Description. – Rostrum (Fig. 11A) short, reaching slightly Uropodal diaeresis (Fig. 11L) with 17 movable spinules. beyond end of basal segment of antennular pedunlcle; rostral formula: 3–6+6–11/1–6. Antennal spine fused with inferior Habitat. – Caridina anislaq, new species, was collected orbital angle. Pterygostomian margin broadly rounded. from Spring Anislaq, near Cave Valencia, Valencia, Bohol Island, central Philippines. Sixth abdominal somite 0.48 times of carapace, 1.9 times as long as fi fth somite, slightly shorter than telson. Telson Etymology. – The new species is named after its type (Fig. 11B, C) 3.2 times as long as wide, not terminating in locality-Spring Anislaq, Bohol Island, the Philippines, used a projection, with 3 pairs of dorsal spinules and 1 pair of as a noun in apposition. dorsolateral spinules; lateral pair of distal spines distinctly longer than intermediate pairs. Preanal carina low, sub- Remarks. – Caridina anislaq, new species, is morphologically triangular, lacking spine. similar to C. batuan, new species, in the form of rostrum. However, it could be easily separated from C. batuan by Eyes well developed, anterior end reaching to 0.7 times the larger number of the post orbital rostral teeth (3–6 vs. length of basal segment of antennular peduncle. Antennular 0–2), the stouter merus (1.2 times as long as broad vs. 2.1 peduncle 0.70 times as long as carapace; basal segment times) and the carpus of the fi rst pereiopod (as long as high of antennular peduncle as long as both second and third vs. 1.6 times as long as high) and the shorter fl agellum of segment lengths, anterolateral angle reaching 0.20 times exopod of the fi rst maxilliped (Figs. 10, 11). length of second segment, second segment distinctly longer than third segment. Stylocerite reaching to 0.7 times length of basal segment of antennular peduncle. Scaphocerite 3.6 Caridina camaro, new species times as long as wide. (Fig. 12)

Incisor process of mandible ending in irregular teeth, molar Material examined. – Holotype: Female, cl 4.3 mm, USC, Cave process truncated. Lower lacinia of maxillula broadly Camaro, Batuan, coll. B. Sket, Feb.1995. rounded, upper lacinia elongated, with a number of distinct teeth on inner margin, palp slender. Upper endites of maxilla Paratypes: Six more or less damaged specimens, ZRC 2007.0303, data same as holotype. subdivided, palp short, scaphognathite tapering posteriorly

with some long, curved setae at posterior end. Palp of fi rst Description. – Rostrum (Fig. 12A) short, reaching slightly maxilliped rounded, exopod with short fl agellum. Podobranch beyond end of basal segment of antennular pedunlcle; rostral of second maxilliped reduced to a lamina. Third maxilliped formula: 2+10/5. Antennal spine fused with inferior orbital reaching beyond end of scaphocerite, with ultimate segment angle. Pterygostomian margin broadly rounded. slightly longer than penultimate segment. Sixth abdominal somite 0.48 times of carapace, 1.8 times Epipods on fi rst 4 pereiopods. First pereiopod (Fig. 11D) as long as fi fth somite, slightly shorter than telson. Telson stout, reaching to middle of second segment of antennular (Figs. 12B, C) 2.7 times as long as wide, not terminating peduncle; merus 1.2 times as long as broad, slightly shorter in a projection, with 4 or 5 pairs of dorsal spinules and 1 than carpus; carpus strongly excavated anteriorly, much pair of dorsolateral spinules; lateral pair of distal spines shorter than chela, as long as high; chela 2.1 times as long as distinctly longer than intermediate pairs. Preanal carina low, broad; fi ngers shorter than palm. Second pereiopod (Fig. 11E) sub-triangular, lacking spine. reaching to end of scaphocerite; merus distinctly shorter than carpus, 3.3 times as long as broad; carpus slightly shorter or Eyes reduced, anterior end reaching to 0.4 times length of as long as chela, 3.9 times as long as high; chela 2.4 times basal segment of antennular peduncle. Antennular peduncle as long as broad; fi ngers 1.5 times as long as palm. Third 0.60 times as long as carapace; basal segment of antennular pereiopod (Figs. 11F, G) reaching beyond end of scaphocerite peduncle distinctly as long as both second and third segment by length of 1/3 of propodus, propodus 12 times as long lengths, anterolateral angle reaching 0.20 length of second as broad, 4.9 times as long as dactylus; dactylus 2.5 times segment, second segment distinctly longer than third as long as wide (spines included), with 4 accessory spines segment. Stylocerite reaching to 0.7 times length of basal on fl exor margin. Fifth pereiopod (Figs. 11H, I) reaching

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Fig. 11. Caridina anislaq, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, fi rst pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fi fth pereiopod; I, dactylus of fi fth pereiopod; J, endopod of male fi rst pleopod; K, appendix masculina and appendix interna of male second pleopod; L, uropodal diaeresis. Scale bars: A, B = 1 mm; C = 0.2 mm, G, I = 0.1 mm; D–F, H, L = 0.5 mm (males, cl 3.4 mm, paratype, ZRC, spring Anislaq, outside Cave Anislaq, Valencia).

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Fig. 12. Caridina camaro, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, fi rst pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fi fth pereiopod; I, dactylus of fi fth pereiopod; J, uropodal diaeresis. Scale bars: A, B = 1 mm; C = 0.2 mm; D–F, H = 0.5 mm; G, K = 0.1 mm; B, F, H = 0.1 mm; C–E, G = 0.5 mm; I, J = 0.2 mm (females, cl 4.3 mm, holotype, USC, Cave Camaro, Batuan).

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segment of antennular peduncle. Scaphocerite 3.2 times as Palaemonidae Rafi nesque, 1815 long as wide. Macrobrachium Bate, 1868 Incisor process of mandible ending in irregular teeth, molar process truncated. Lower lacinia of maxillula broadly Macrobrachium rosenbergii rosenbergii (De Man, 1879) rounded, upper lacinia elongate, with a number of distinct teeth on inner margin, palp slender. Upper endites of maxilla Palaemon Rosenbergii De Man, 1879: 167 [type locality: Andai, subdivided, palp short, scaphognathite tapering posteriorly New Guinea (Irian Jaya), Indonesia]. with some long, curved setae at posterior end. Palp of fi rst Palamon carcinus rosenbergi – Ortmann, 1891: 701. maxilliped rounded, exopod with short fl agellum. Podobranch Palaemon (Eupalaemon) carcinus – De Man, 1902: 763 (not Cancer carcinus Linnaeus, 1758). of second maxilliped reduced to a lamina. Third maxilliped Macrobrachium rosenbergii – Holthuis, 1950: 111 (part); Johnson, slender, with ultimate segment as long as penultimate 1960: 260, Fig. 1; Chace & Bruce, 1993: 36, Fig. 15; Short, segment. 2004: 44, Fig. 16 (part). Macrobrachium rosenbergii rosenbergii – Johnson, 1973: 277; Epipods on fi rst 4 pereiopods. First pereiopod (Fig. 12D) Holthuis, 1995: 148; Cai & Ng, 2001: 674; Cai & Shokita, slender, merus 2.4 times as long as broad, shorter than 2006a: 263. carpus; carpus excavated anteriorly, shorter than chela, 1.8 to 3.0 times as long as high; chela 2.7 times as long as broad; Material examined. – One dried male, cl 103 mm, ZRC 2000.2407, fi ngers 1.4 times as long as palm. Second pereiopod (Fig. purchased from a fi shermen who collected from the Loboc River in the 1980s Y. Cai, 19 Dec.2000; 1 molted cast, 2 juveniles, coll. 12E) elongated and slender, merus distinctly shorter than Y. Cai et al., Loboc River, Loboc, 19 Dec.2000. carpus, 6.0 times as long as broad; carpus 1.5 times as long as chela, 7.7 times as long as high; chela 3.7 times as long Remarks. – Only one molted cast and two juveniles are as broad; fi ngers 1.9 times as long as palm. Third pereiopod available for the presnt study. Judging by the form of the (Figs. 12F, G) with propodus 16 times as long as broad, 4.4 rostrum of the larger specimen, which is only slightly times as long as dactylus; dactylus 5.6 times as long as wide beyond the end of scaphocerite and has a rostral formula (spines included), with 6 accessory spines on fl exor margin. as 2+11/9. It apparently belongs to the eastern subspecies, Fifth pereiopod (Figs. 12H, I) with propodus 15 times as i.e. Macrobrachium rosenbergii rosenbergii. long as broad, 3.3 times as long as dactylus, dactylus 5 times as long as wide (spinules included), terminating in 1 claw, with 60 spinules on fl exor margin. Macrobrachium latidactylus (Thallwitz, 1891)

Uropodal diaeresis (Fig. 12J) with 17 movable spinules. Palaemon latidactylus Thallwitz, 1891: 97 [type locality: Celebes (=Sulawesi), Indonesia]; Thallwitz, 1892:17, Fig. 3. Habitat. – Caridina camaro, new species, was collected Palaemon (Macrobrachium) latidactylus – De Man: 1902: 805. from the subterranean stream in Cave Camaro, Bohol Island, Palaemon lampropus De Man, 1892: 493, Pl. 29 Fig. 49 [type central Philippines. locality: Celebes and Timor, Indonesia]; Kemp, 1918: 267. Macrobrachium latidactylus – Holthuis, 1950: 239, Fig. 50; 1980: Etymology. – The new species is named after its type 97; Costa, 1979: 57; Naiyanetr, 1980: 17; Yeo et al., 1999: locality—Cave Camaro, Bohol Island, the Philippines. The 236; Cai et al., 2004: 584; Short, 2004: 80, Fig. 31; Cai & Anker, 2004: 258; Wowor et al., 2004: 349, Fig 10E; Cai & name is used as a noun in apposition. Shokita, 2006a: 266.

Remarks.- With regard to the slender pereiopods, Caridina Material examined. – 5 males, cl 7.3–15.1 mm, 5 females, cl camaro, new species, is morphologically similar to C. liaoi, 6.0–11.5 mm, ZRC 2007.0304, River outside Inabacan Cave, new species. It can be easily differentiated from the latter Antequrra, coll. B. Sket, 23 Feb.1999; 1 male, cl 15 mm, ZRC species by the reduced eyes, and the much shorter rostrum. 2007.0305, Alijanan Bridge, stream, S. Duero, coll. B. Sket, 5 Compared with all the other cave species in Bohol, C. camaro Mar.1999; 1 male, cl 10 mm, 1 female, cl 11 mm, ZRC 2007.0306, has more elongated carpus of the fi rst pereiopods, hence, River outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 could readily be distinguished from C. valencia, C. batuan Dec.2000; 2 males, cl 12.8–13.5 mm, 1 female, cl 8.4 mm, ZRC 2007.0307, River outside Inabacan Cave, Antequrra, coll. Y. Cai and C. anislaq (1.8–3.0 times as long as high vs. 1.0–1.6 et al., 18 Dec.2000. times in the latter three species). It can be separated from

C. boholensis by the slender dactylus of third pereiopod (5.6 Remarks . – Macrobrachium latidactylus has been reported times as long as wide vs. 3.0 times) from Sri Lanka (Costa, 1979), Indonesia, the Philippines, Malaysia, southern Thailand (Holthuis, 1950; Johnson, 1963; Chace & Bruce, 1993; Yeo et al., 1999), Hainan (Yu, 1936; Liu et al., 1990), Taiwan (Hwang & Yu, 1982; Shy & Yu, 1998), the Ryukyu Islands (Shokita, 1979) and Australia (Short, 2004).

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Marobrachium australe (Guérin-Méneville, 1838) Macrobrachium horstii (De Man, 1892)

Palaemon australis Guérin-Méneville, 1838: 37 [type locality: Palaemon (Parapalaemon) Horstii De Man 1892: 460, Pl. 27: Fig. Tahiti, French Polynesia]. 39 [type locality: River at Polopo, central Celebes (Sulawesi), Palaemon sundaicus Heller, 1862: 415, Pl. 2 Figs. 38, 39 [type Indonesia]. locality: Java, Indonesia]. Macrobrachium horstii - Holthuis, 1950: 203, Fig. 42; Chace & Palaemon dispar Von Martens 1868: 41 [type locality: Pulau Bruce, 1993: 27; Cai & Ng, 2001: 679, Fig. 10; Wowor et al., Adonara, east of Flores, Indonesia]. 2004: 349, Fig. 9L. Palaemon (Eupalaemon) dispar – De Man, 1902: 766. Macrobrachium australis – Holthuis, 1950: 124, Figs. 27–30; Chace Material examined. – 1 male, cl 13.6 mm, 1 female, cl 16.8 mm, & Bruce, 1993: 23, Fig. 2; Cai & Ng, 2001: 683, Fig. 14a–d; 1 ovigerous female, cl 12.0 mm, ZRC 2007.0311, River outside Cai & Anker, 2004: 258; Cai & Shokita, 2006a: 263. Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male, 3 ovigerous females, cl 13.6 mm, ZRC 2007.0312, River outside Material examined. – 3 males, cl 17.0–24 mm, 6 ovigerous females, Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000; 2 females, 15.7–19.7 mm, ZRC 2007.0308, Loboc River, Loboc, coll Y. Cai cl 13.6, 6 ovigerous females, cl 11.0–13.8 mm, ZRC 2007.0313, et al., 19 Dec.2000. Loboc River, Loboc, coll. Y. Cai et al., 19 Dec.2000.

Remarks. – The species has a wide distribution in the Remarks. – Macrobrachium horstii has previously been Indo-West Pacifi c, from Madagascar to Polynesia (Holthuis, reported from Leyte, the Philippines (Cai & Ng, 2001). 1950; Chace & Bruce, 1993). It is currently known from the Philippines and eastern Indonesia.

Macrobrachium lar (Fabricius, 1798) Macrobrachium placidulum (De Man, 1892) Palaemon Lar Weber, 1795: 94 [nomen nudum] Palaemon Lar Fabricius, 1798: 402 [type locality: in India orientali Palaemon (Macrobrachium) placidulus De Man, 1892: 489, Pl. Dom. Daldorf]; Cowles, 1914: 380, Pl. 2 Fig. 7. 28 Fig. 48 [type localities: Celebes (Sulawesi), Pulau Selajar, Palaemon (Eupalaemon) lar – De Man, 1902: 774. Flores, and Timor]. Macrobrachium lar – Holthuis, 1950: 176, Fig. 37; Chace & Bruce, Macrobrachium placidulum – Holthuis, 1950: 253, Fig. 51c; Chace 1993: 30, Fig. 9; Yeo et al., 1999: 236; Cai & Ng, 2001: 683, & Bruce, 1993: 35, Fig. 14; Cai & Shokita, 2006a: 264. Fig. 14e; Wowor & Choy, 2001: 285; Cai & Ng, 2002: 75; Macrobrachium placidum – Shokita, 1979: 275. Cai & Anker, 2004: 256; Cai et al., 2004: 585; Wowor et al., Macrobrachium horstii – Hwang & Yu, 1982: 164, Fig. 5 (not 2004: 353, Fig. 14; Short, 2004: 75, Fig. 29. Palaemon (Parapalaemon) Horstii De Man 1892). Macrobrachium cf horstii – Shy & Yu, 1998: 30. Material examined. – One male, cl 22.6 mm, ZRC 2007.0309, Loboc River, Loboc, coll. Y. Cai et al., 19 Dec.2000. Material examined. – One male, cl 14.2 m, ZRC 2007.0314, Alijanan Bridge, stream, S. Duero, coll. B. Sket, 5 Mar.1999; 3 Remarks. – Macrobrachium lar is a well known species, males, cl 15.3–17.4 mm, ZRC 2007.0315, river outside Inabacan widely distributed throughout the Indo-West Pacific. Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male, cl 8.4 However, it seems rather rare in Bohol. mm, 1 female, cl 10.3 mm, ZRC 2007.0316, river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000.

Remarks. – Macrobrachium placidulum is a common Macrobrachium scabriculum (Heller, 1862) freshwater species in the Philippines. It is distributed from Ryukyu Islands, Taiwan, the Philippines, Palau, and eastern Palaemon scabriculus Heller, 1862: 527 [type locality: Sri Lanka].- Henderson & Matthai, 1910: 296, Pl.17 Figs. 7a–c, Pl. 18 Indonesia, to New Guinea, as well as Fiji (Chace & Bruce, Figs. 7a–p. 1993; Choy, 1984). Macrobrachium scabriculum – Holthuis, 1950: 224; Chace & Bruce, 1993: 37; Johnson, 1973: 15; Yeo et al., 1999: 231, Figs. 18; 19; Wowor & Choy, 2001: 286; Cai & Ng, 2002: Fig. 16; Wowor Macrobcrachium equidens (Dana, 1852) et al., 2004: 350, Fig. 12I; Cai & Shokita, 2006a: 266. Palaemon equidens Dana, 1852: 26 [type locality: Singapore]. Material examined. – One male, cl 19.5 mm, 1 ovigerous female, Palaemon sundaicus – Cowles, 1914: 355, Pl. 2 Fig. 3 (not Palaemon 24 mm, ZRC 2007.0310, Loboc River, Loboc, coll. Y. Cai et al., sundaicus Heller, 1862). 19 Dec.2000. Macrobrachium equidens – Holthuis, 1950:162, Fig. 36; 1980: 90; Liu et al., 1990: 110, Fig. 8; Chace & Bruce, 1993: 25, Fig. Remarks. – Macrobrachium scabriculum was reported 4; Yeo et al., 1999: 226; Wowor & Choy, 2001: 282; Cai & recently from Mindanao (Cai & Shokita, 2006a). This is the Anker, 2004: 389; Cai et al., 2004: 589; Wowor et al., 2004: second record for the species to occur in the Philippines. 349, Fig. 10H; Short, 2004: 26, Figs. 8, 9; Cai & Shokita, 2006a: 265.

Material examined. – 1 male, cl 7.5 mm, ZRC, River outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male, cl 10.5 mm, 1 female, cl 10.0 mm, ZRC, Loboc River, Loboc, coll. Y. Cai et al., 19 Dec.2000.

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Remarks. – Marcobrachium equidens is a brackish water Bouvier, E. L., 1925. Recherches sur la morphologie, les variations, species, with gravid females and smaller specimens being la distribution géographique des crevettes des la famille des commonly found in mangrove creeks. It is known from a Atyidés. Encyclopédie Entomologique, series A, 4: 1–370, very wide area in the Indo-West Pacifi c, from Madagascar to Figs. 1–761. the Solomon Islands and Fiji (Choy, 1984; Chace & Bruce, Cai, Y. & A. Anker, 2004. On a collection of freshwater shrimps 1993; Short, 2004). However, a recent molecular analysis from the Philippines, with descriptions of fi ve new species. results (Liu et al., 2007) show that the incongruence of Tropical Zoolology, 17: 233–266. non-monophyly of M. equidens, in 16S, COI and 16S+COI Cai, Y., P. Naiyanetr & P. K. L. Ng, 2004. The freshwater analyses, implied the existence of a cryptic species in the prawns of the genus Macrobrachium Bate, 1868, of Thailand Philippines and Taiwan. (Crustacea: Decapoda: Palaemonidae). Journal of Natural History, 38: 581–649. Cai, Y., & P. K. L., Ng, 2001. The freshwater decapod Palaemon Weber, 1795 of Halmahera, Indonesia. Journal of Crustacean Biology, 21: 665–695, Figs. 1–19. Palaemon concinnus Dana, 1852 Cai, Y. & P. K. L. Ng, 2002. The freshwater palaemonid prawns (Crustacea: Decapoda: Caridea) of Myanmar. Hydrobiologia, Palaemon concinnus Dana, 1852: 587 [type locality: Fiji Islands]; 487: 59–83. Holthuis, 1950: 61, Fig. 12; Chace & Bruce, 1993: 40; Cai & Cai, Y. & P. K. L. Ng, 2007. A revision of the Caridina gracilirostris Ng, 2001: 686, Fig. 14f; Cai & Shokita, 2006a: 267. De Man, 1892, species group, with descriptions of two new taxa (Decapoda; Caridea; Atyidae). Journal of Natural History, Material examined. – 2 males, cl 7.0–7.6 mm, 1 female, cl 9.2 41(25–28): 1585–1602. mm, ZRC 2007.0317, tributary of Loboc River, Loboc, coll. Y. Cai, Y. & S. Shokita, 2006a. Report on a collection of freshwater Cai et al., 19 Dec.2000. shrimps (Crustacea: Decapoda: Caridea) from the Philippines, with descriptions of four new species. Raffles Bulletin of Remarks. – Palaemon concinnus has a wide distribution Zoology, 54: 245–270. in the Indo-West Pacifi c, from South Africa, Indonesia, Cai, Y., & S, Shokita, 2006b. Atyid shrimps of the Ryukyu Islands, Philippines to Marshall Islands and Tuamotu Archipelago Southern Japan (Crustacea: Decapoda: Atyidae). Journal of in French Polynesia (Chace & Bruce, 1993). The species Natural History, 40(38–40): 2123–2172. is commonly found in brackish to fresh water in the lower Cai, Y., P. K. L. Ng, S. Shokita & K. Satake, 2006. Revision of reaches of rivers. the Japanese atyid shrimps (Decapoda: Caridea) fi rst described by William Stimpson, 1860. Journal of Crustacean Biology, 26(3): 392–419. ACKNOWLEDGEMENTS Chace, F. A. Jr., 1983. The Atya-like shrimps of the Indo-pacifi c Region (Decapoda: Atyidae). Smithsonian Contributions to The fi rst author would like to thank Boris Sket (UOL) for Zoology, 384: 1–54, 24 fi gures. sending his specimens and the accompanying habitat notes. Chace, F. A., Jr., 1997. The Caridean shrimps (Crustacea: The ZRC-USC Expedition was led by Lawrence Liao (USC), Decapoda) of the Albatross Philippine expedition 1907–1910. and we are grateful to the support cum help of Benito Tan Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, and N. K. Ng (ZRC), and Adonis Porpetcho and Roxie Diaz Bathypalaemonellidae, Processidae, and Hippolytidae. (USC). The Caving Team from the Holy Name University Smithsonian Contributions to Zoology, 58: 1–106, Figs.1–29. at Bohol was instrumental in helping with the speleological Chace, F. A. Jr. & A. J. Bruce, 1993. The Caridean shrimps surveys, and the support of their president, Florentes Camacho (Crustacea: Decapoda) of the Albatross Philippine expedition is gratefully acknowledged. Boris Sket read the manuscript 1907–1910. Part 6: Superfamily Palaemonoidea. Smithsonian and to N. K. Ng kindly prepared the map. Contributions to Zoology, 543: i–vii + 1–152, Figs. 1–23. Choy, S., 1984. On the freshwater palaemonid prawns from the Fiji Islands (Decapoda, Caridea). Crustaceana, 47: 269–277. LITERATURE CITED Costa, H. H., 1979. The Palaemonidae of the inland water of Sri Lanka. Ceylon Journal of Science (Biological Science), new Balete, D. S., & L. B. Holthuis, 1992. Notes on the cave shrimp series, 13: 39–64, 6 text fi gures, 2 plates. Edoneus atheatus Holthuis, 1978, with an account of its type locality of habits (Decapoda, Caridea, Atyidae). Crustaceana, Cowles, R. P., 1914. Palaemons of the Philippines. Philippine 62: 98–101. Journal of Science, (D), 9(4): 319–403, 1 fi gure, Pls. 1–3. Bate, C. S., 1868. On a new genus, with four new species of Cowles, R. P., 1915a. The habitats of some tropical crustacean freshwater prawns. Proceedings of the Zoolological Society II. Philippine Journal of Science, 10: 11–17, Figs. 1, 2, Pls. of London, 1868: 363–368, Pls. 30, 31. 1–3. Blanco, G. J., 1935. The Atyidae of Philipines. Philippine Journal Cowles, R. P., 1915b. Are Atya spinipes Newport and Atya armata of Science, 56: 29–30. Milne Edwards Synonyms for Atya moluccensis De Haan? Philippine Journal of Science, 10: 147–153, Fig. 1. Blanco, G. J., 1939. Four new Philippine species of fresh-water shrimps of the genus Caridina. Philippine Journal of Science, Dana, J. D., 1852. Crustacea, Part 1. In United States Expedition 70: 389–395, Pls. 1–3. during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U. S. N., 13: 1–685. Atlas (1855): 27 pp, 96 plates. Philadelphia.

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De Haan, W., 1833–1850, Crustacea, in P. F. von Siebold (ed.), Henderson, J. R. & G. Matthai, 1910. On some cartain of of Fauna Japonica sive Descripto Animalium, quae in Itinere Palaemon from South India. Records of Indian Museum, 4: per Japoniam, Jussu et Auspiciis Superiorum, qui Summum 277–305, Pls. 15–18. in India Batava Imperium Tenent, Suscepto, Annis 1823-1830 Holthuis, L. B., 1950. Subfamily Palaemonidae. The Palaemonidae Collegit, Notris, Observationibus et Adumbrationibus Illustravit. collected by the Siboga and Snellius Expeditions with remarks Lugduni-Batavorum. [Leiden]: i–xxxi, ix–xvi, 1–243, Pls. A–J, on other species. I. The Decapoda of the Siboga Expedition. l–Q, 1–55, circ. tab. 2. Part X. Siboga Expeditie, 39(a9): 1–268, Figs. 1–52. De Man, J. G., 1879. On some species of the genus Palaemon Holthuis, L. B., 1955. The recent genera of the caridean and Fabr. with descriptions of two new forms. Notes from the Royal stenopodidean shrimps (Class Crustacea: Order Decapoda: Zoological Museum of the Netherlands, Leyden, 1: 165–184. Supersection Natantia) with keys for their determination. De Man, J. G., 1892. Decapoden des Indischen Archipels, in Zoologische Verhandelingen, Leiden, 26: 1–157, Figs. 1–105. Max. Weber (ed.), Zoologische Ergebnisse einer Reise in Holthuis. L. B., 1956. An enumeration of the Crustacea Decapoda Niederländisch Ost-Indien, 2: 265–527, Pls. 15–29. Natantia inhabiting subterranean waters. Vie et Milieu, 7: De Man, J. G., 1902. Die von Herrn Professor Kükenthal im Indischen 43–76. Archipel gesammelten Dekapoden und Stomatopoden. In: W. Holthuis, L. B., 1963. On red coloured shrimps (Decapoda, Caridea) Kükenthal, Ergebnisse einer zoologischen Forschungsreise in den from tropical land-locked salt water pools. Zoolologische Molukken und Borneo. Abhandlungen der Senckenbergischen Mededelingen, Leiden, 38: 261–279. Naturforschenden Gesellschaft, 25: 467–929, Pls. 18–20. Holthuis, L. B., 1965. The Atyidae of Madagasca, Mémoires du De Silva, K. H. G. M., 1982. Studies on Atyidae (Decapoda, Caridea) Muséum national d’Histoire Naturelle, series A (Zoologie), of Sri Lanka. I. On a new species, a new subspecies and two 33: 1–48, Figs. 1–17. species new to Sri Lanka. Crustaceana, 43: 127–141. Holthuis, L. B., 1973. Caridean shrimps found in land-locked Edmondson, C. H., 1935a. New and rare Polynesian Crustacea. saltwater pools at four Indo-West Pacific localities (Sinai Bernice P. Bishop Museum Occasional Papers, Honolulu, 11: Peninsula, Funafuti Atoll, Maui and Hawaii Islands), with the 1–19, Figs. 1–6. description of one new genus and four new species. Zoologische Edmondson, C. H., 1935b. Atyidae of southern Polynesia. Bernice Verhandelingen, Leiden, 128: 1–48. P. Bishop Museum Occasional Papers, Honolulu, 10: 1–40. Holthuis, L. B. 1978. Zoological results of the British Speleological Edmondson, C. H., 1954. Substitute for an invalid generic name Expedition to Papua New Guinea 1975. 7. Cavernicolous in the Crustacea. Pacifi c Science, 8: 368. shrimps (Crustacea, Decapoda, Natantia) from New Ireland Fabricius, J. C. 1798. Supplementum Entomologiae Systematicae. and the Philippines. Zoolologische Mededelingen, Leiden, 53: Hafniae. 572 pp. 209–224. Fujino, T. & S. Shokita, 1975. Report on some atyid shrimps Holthuis, L. B., 1982. Notes on Indo-West Pacific Crustacea (Crustacea, Decapoda, caridea) from the Ryukyus Islands, Decapoda II. The genus Halocaridinides Fujino and Shokita Bulletin of Science and Engineering Division, University of the and its type species H. trigonophthalma (Fujino and Shokita). Ryukyus, (Mathmatics and Natatural Science), 18: 93–113. Crustaceana, 42(1): 31–36. Guérin-Méneville, F. E., 1829–1838. Crustacés, Arachnides et Holthuis, L. B., 1995. Notes on Indo-West Pacific Crustacea Insectes. In L. J. Duperrey (ed.), Voyage autour du monde, Decapoda III to IX. Zoölogische Mededeelingen, Leiden, 69: excuté par ordre du Roi, sur la corvette de Sa Majesté, La 139–151. Coquille, pendant les annes 1822, 1823, 1924 et 1825. Zoologie Hung, M.S., T.Y. Chan, & H.P. Yu, 1993. Atyid shrimps (Decapoda, 2 (no. 2, sect. 1): 1–47 (Crustacés), 48–319 (Arachnides et Caridea) of Taiwan, with descriptions of three new species. Insectes), Pls. 1–5 (Crustaces), 1–21 (Insectes). Paris: Arthus Journal of Crustacean Biology, 13: 481–503, Figs. 1–14. Bertrand. Hwang, J. J. & H. P. Yu, 1982. Studies on the freshwater Gurney, A. R., 1984b. Halocaridinides fowleri (Gordon) comb. shrimps of the genus Macrobrachium (Crustacea, Decapoda, nov.: a redescription of Parisia (?) fowleri Gordon in Gordon Palaemonidae) from Taiwan. Quarterly Journal of Taiwan and Monod 1968 (Decapoda: Caridea: Atyidae) from Zanzibar. Museum, 25: 157–180. Journal of Natural History, 18: 591–598, Figs. 1a–g, 2a–g, Jalihal D. R. & S. Shenoy, 1998. Taxonomic revision of some 3a–f. Indian prawn species of genus Caridina H. Milne Edward, 1837 Hart, C. W., 1980. A new atyid shrimp, Palauatya dasyomma, (Atyidae), Proceedings and abstracts of the Fourth Internatinal from Palau, Caroline Islands, Proceedings of the Biological Crustacean Congress, Melbourne, Australia: 128–129. Society of Washington, 93(2): 481–489. Johnson, D. S., 1960. Sub-specifi c and infra-specifi c variation in Hayashi, K.-I., 1989a. Prawns, shrimps and lobsters from Japan some freshwater prawns of the Indo-Pacifi c region. Proceedings (49), Family Atyidae-Genus Caridina (3) and Antecaridina. of the Centenary and Bicentenary congress of Biology, Aquabiology, 64, 11: 376–379, Figs. 171a–g, 172, 173a–e, Singapore,1958: 259–267, Figs. 1–5. 174. (in Japanese). Johnson, D. S., 1962. On a new species of Macrobrachium Hayashi, K.-I., 1989b. Prawns, shrimps and lobsters from Japan (Decapoda, Caridea). Crustaceana, 4: 307–310, Fig. 1. (50), Family Atyidae-Genus Paratya and Halocaridinides. Johnson, D. S., 1963. Distribution and other notes on some fresh- Aquabiology, 65, 11: 497–501, Figs. 175, 176a–l, 177, 178a–c. water prawns (Atyidae and Palaemonidae) mainly from the (in Japanese). Indo-west Pacifi c region. Bulletin of the National Museum, Heller, C., 1862. Neue Crustaceen gesammelt wahrend der Singapore, 32: 5–30, Fig. 1–5. Weltumseglung der k.k. Fregatte Novara: Zweiter vorläufi ger Johnson, D. S., 1973. Notes on some species of the genus Bericht. Verhandlungen des Kaiserlich-Königlichen Macrobrachium (Crustacea: Decapoda: Caridea: Palaemonidae). Zoologisch-botanischen Gesellschaft in Wien, 12: 519–528.

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Journal of Singapore National Academy of Science, 3: Shokita, S., 1979. The distribution and speciation of the inland 273–291. water shrimps and prawns from the Ryukyu Islands-II. Bulletin Kemp, S., 1918. Zoological results of a tour in the Far East Crustacea of College of Science, University of Ryukyus, 28: 193–278. (In Decapoda Stomatopoda, Memoirs of the Asiatic Socciety of Japanese with English abstract) Bengal, 6: 219–297. Shokita, S., 2003, Atyidae, In: Nishida, M., Shikatani N. & S. Leberer, T. & Y. Cai, 2003. Shrimps of the family Atyidae from Shokita (Eds.), The fl ora and fauna of inland waters in the Guam, Mariana Islands. Micronesica, 35–36: 355–360. Ryukyu Islands, Tokai University Press, Tokyo, Japan. Pp. 249–254. (In Japanese) Liang, X. -Q. & S. -L. Yan, 1977. New species and subspecies of Caridina (Decapoda, Caridea) from Fukien, China. Acta Short, J. W., 2004. A revision of Australian river prawns, Hydrobiologia Sinica, 6: 219–225, Figs. 1–13. (in Chinese Macrobrachium (Crustacea: Decapoda: Palaemonidae). with English abstract) Hydrobiologia, 525: 1–100. Linnaeus, C., 1758. Systema Naturae per Regna Tria Naturae, Shy, J. Y & H. P. Yu, 1998. Freshwater shrimps of Taiwan. Secundum Classes, Ordines, Genera, Species, cum Characteribus, National Museum of Marine Biology and Aquarium. 103 pp. Differentiis, Synonymis, Locis. Holmiae. Edition 10, volume (In Chinese) 1: iii + 824 pp. Sket, B., 1997. Hypogean aquatic fauna in Bohol Island, Philippines. Liu, M. Y., Cai, Y. X. & C. S. Tzeng, 2007. Molecular systematics Our Caves. Bulletin of the Speleology Association of Slovenia, of the freshwater prawn genus Macrobrachium Bate, 1868 39: 62–67. (Crustacea: Decapoda: Palaemonidae) from mtDNA sequences, Smith, M. J. & W. D. Williams, 1981. The occurrence of with emphasis on East Asian species. Zoological Studies, Antecaridina lauensis (Edmondson) (Crustacea, Decapoda, 46(3): 272–289. Atyidae) in the Solomon Islands. Hydrobiologia, 85: 49–58. Liu, R., Liang, X.& S. Yan, 1990. A study of the Palaemonidae Suzuki, H., 1980. An atyid shrimp living in anchialine pool (Crustacea Decapoda) from China I. Macrobrachium, Leander on Kuro-Shima, the Yaeyama group, Okinawa Prefecture. and Leandrites. Transactions of the Chinese Crustacean Society, Proceedings of the Japanese Society of Systematic Zoology, 2: 102–134, Figs. 1–24. (In Chinese with English abstract) 18: 47–53, Figs. 1–5. Milne Edwards, H., 1837, Histoire naturelle des Crustacés, Thallwitz, J., 1891. Ueber einige neue indo-pacifi sche Crustaceen comprenant l’anatomie, la physiologie et la classifi cation de (vorläufi ge mittheilung). Zoologischer Anzeiger, 14: 96–103. ces animaux, 2, 1–532, atlas, Pls. 1–14. Thallwitz, J., 1892. Decapoden-Studien, insbesondere basiert auf A. Naiyanetr, P., 1980. Crustacean fauna of Thailand (Decapoda and B. Meyers Sammlungen im Ostindischen Archipel, nebst einer Stomatopoda). Department of Biology, Faculty of Science, Aufzählung der Decapoden und Stomatopoden des Dresdener , , Thailand, 73 pp. (In Museums. Abhandlungen und Berichte des Königlichen Thai) Zoologischen und Anthropologisch-Ethnographischen Museums Naruse, T., M. Toda & S. Shokita, 2003. A record of Halocaridinides zu Dresden, 1890-91(3): 1–55, Pl. 1. trigonophthealma (Atyidae) from Hatoma Island, Yaeyama Tiwari, K. T. & R. S. Pillai, 1971. Atyid shrimps of the genus Group, Southern Ryukyus, Japan. Cancer, 12: 1–6. (In Japanese Caridina H. Milne Edwards, 1837, from the Andaman Islands with English abstract) (Decapoda, Caridea). Crustaceana, 21: 79–91. Newport, G., 1847. Note on the genus Atya of Leach, with Von Martens, E., 1868. Ueber einige ostasiatische Süsswasserthiere. descriptions of four apparently new species, in the cabinets of Archiv für Naturgeschichte, 34(1): 1–67, Pl. 1. the British Museum. Annals and Magazine of Natural History, Weber, F., 1795. Nomenclator entomologicus secundum 19: 158–160, Pl. 8, Fig. 1. Entomologiam systematicum ill. Fabricii adjectis speciebus Ortmann, A., 1891. Versuch einer Revision der Gattungen recens detectis et varietatibus. Chilonii et Hamburgi: viii+171 Palaemon sens. strict. und Bithynis. Die Decapoden-Krebse des pp. Strassburger Museums, mit besonderer Berücksichtigung der Wowor, D., & S. Choy, 2001. The freshwater prawns of the von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln genus Macrobrachium Bate, 1868 (Crustacea: Decapoda: gesammelten und z. Z. im Strassburger Museum aufbewahrten Palaemonidae) from Darussalam. Raffl es Bulletin of Formen. II. Theil. Zoologische Jahrbücher Abteilung für Zoology, 49 (2): 269–289. Systematik, Geographie und Biologie der Thiere, 5: 693–750, Pl. 47. Wowor, D., Y. Cai & P. K. L. Ng, 2004. Crustacean: Decapoda: Caridea. In: C. Yule & H. S. Yong (editors). The freshwater Rafi nesque, C. S., 1815. Analyse de la Nature Tableau de l’Univers invertebrates of Malaysia and Singapore. Malaysian Academy et corps organizes. 224 pp. of Sciences, Kuala Lumpur: 337–357. Roux, J, 1928. Notes carcinologiques de l’Archipel indo-australien. Yeo, D. C. J., Cai, Y., & P. K. L. Ng, 1999. The freshwater and I. Décapodes Macroures d’eau douce de l’Archipel Indo- terrestrial decapod Crustacea of Pulau Tioman, Peninsular Australien. Treubia, 10: 197–216, Figs. 1–9. Malaysia. Raffles Bulletin of Zoology, Supplement No. 6: Shokita, S., 1975. The distribution and speciation of the inland 190–244, Figs. 1–20. water shrimps and prawns from the Ryukyu Islands-I. Bulletin Yu, S. C., 1936. Report on the Macrurous Crustacea collected of Science & Engineering Division, University of Ryukyus, during the “Hainan Biological Expedition” in 1934. Chinese Mathmatics and Natural Science, 18: 115–136. (In Japanese Journal of Zoology, 2: 85–99, Figs. 1–7. with English abstract)

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 91–107 Date of Publication: 28 Feb.2009 © National University of Singapore

THE ALPHEID SHRIMP GENUS LEPTALPHEUS WILLIAMS, 1965, IN THE TROPICAL WESTERN PACIFIC, WITH DESCRIPTIONS OF TWO NEW SPECIES (CRUSTACEA: DECAPODA: CARIDEA)

Arthur Anker Dickinson Hall, Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800 USA Email: aanker@fl mnh.ufl .edu

Ivan N. Marin Laboratory of Ecology and Morphology of Marine Invertebrates, A.N. Severtzov Institute of Ecology and Evolution, R. A. S., Leninsky prospect, 33, 119071, Moscow, Russia Email: [email protected]

ABSTRACT. – Two new species of the alpheid shrimp genus Leptalpheus Williams, 1965, are described from the tropical western Pacifi c. Leptalpheus denticulatus, new species, is described on the basis of several specimens collected in Nhatrang Bay, Vietnam (type locality); Panglao, off Bohol, the Philippines; and Yaqara Bay, Viti Levu, Fiji. Leptalpheus dworschaki, new species, is described on the basis of two specimens collected in Panglao. All specimens were collected from burrows of callianassid ghost shrimps, including Glypturus cf. armatus (A. Milne-Edwards, 1870) and Neocallichirus calmani (Nobili, 1904). Both L. denticulatus, new species, and L. dworschaki, new species, are closely related to L. pacifi cus Banner & Banner, 1974, from Hawaii, differing from the latter species in subtle morphological details.

KEY WORDS. – Alpheidae, Leptalpheus, Indo-West Pacific, infauna, commensal, new species, Callianassidae.

INTRODUCTION Leptalpheus is particularly well represented in the western Atlantic, with at least seven species (Anker et al., 2006); The alpheid shrimp genus Leptalpheus Williams, 1965, is however, only four of them are described: L. forceps characterised by the strongly asymmetrical chelipeds, carried Williams, 1965 (type species), L. axianassae Dworschak folded, with subcylindrical, peculiarly shaped major and & Coelho, 1999, L. felderi Anker, Vera Caripe & Lira, minor chelae; the presence of an articulated plate at the 2006, and L. pierrenoeli Anker, 2008 (Anker et al., 2006; posteroventral angle of the sixth pleonite; the absence of Anker, 2008). In the eastern Pacifi c, at least four species orbital teeth and rostrum (although a subtriangular rostral are known, but again only one of them is described: L. projection may be present in some species); the third mexicanus Ríos & Carvacho, 1983 (Anker et al., 2006). maxilliped with a produced, often acute lateral plate on Only one species, L. pacifi cus Banner & Banner, 1974, has the coxa; and the uropodal diaeresis bearing a conspicuous been described from the vast Indo-West Pacifi c region, with subacute tooth near the mesial margin of the exopod only two records: Hawaii (type locality) and southwestern (Williams, 1965; Ríos & Carvacho, 1983; Dworschak & Madagascar (Banner & Banner, 1974, 1983). However, Coelho, 1999; Anker et al., 2006). All these features are Anker et al. (2006) pointed out that Banner & Banner’s also present in Fenneralpheus Felder & Manning, 1986, (1983) record of L. pacifi cus from Madagascar, based on which differs from Leptalpheus mainly by the major untraced material, requires confi rmation. cheliped being stouter and bearing unique lamellar teeth on the fingers (Felder and Manning, 1986). All species Recently, thalassinidean burrows were sampled with the of Leptalpheus live as “commensals” in burrows of aid of a bait suction pump at three distant localities in the thalassinideans, mainly mudshrimps (Upogebiidae) and tropical Indo-West Pacifi c: (1) Yaqara Bay on the northern ghostshrimps (Callianassidae); the hosts of Fenneralpheus coast of Viti Levu, Fiji (AA, in 2005); (2) Nhatrang Bay, presently remain unknown (Anker et al., 2006). south-central Vietnam (IM, in 2004); and Panglao Island, southwest off Bohol, the Philippines (P. C. Dworschak, in 2004). These efforts enabled to obtain several specimens of

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Leptalpheus, all preliminarily identifi ed as L. cf. pacifi cus. Type material. – Vietnam: Holotype: male, CL 6.4, TL 19.1 However, a closer inspection of this material revealed the (MNHN-Na 17064), Nhatrang Bay, Dam Bay, intertidal sand fl at, presence of two species in Panglao, Leptalpheus sp. A and fringe mangrove, bait suction pump, from burrow of Glypturus Leptalpheus sp. B, each represented by two specimens, cf. armatus, coll. I. Marin, 17 Jun.2004. Paratypes: 1 female, CL 7.6, TL 23.8 (FLMNH UF Arthropoda 15018), same collection differing from each other by the dentition on the fi ngers data as for holotype; 1 female, CL 10.0, TL not measured (USNM of the major chela. The fi nger armature of Leptalpheus sp. 1110002), same collection data as for holotype [specimen dissected]; A was very similar to that of the Hawaiian type specimen 1 breeding female, CL 14.5, TL 44.7 (MNHN-Na 17065), same of L. pacifi cus (Banner & Banner, 1974), and also to the collection data as for holotype [specimen photographed]. armature observed in Vietnamese and Fijian specimens. However, subtle differences in some other features, e.g., Additional (non-type) material. – Fiji: 1 female, CL 6.0, TL 18.8 on the uropod, were detected between Leptalpheus sp. A (LACM CR 2005-006.1), Viti Levu, Yaqara Bay, ca. 2 km north and L. pacifi cus. On the other hand, the fi nger armature of of Yaqara Pastoral Community, sand beach fringed with mangrove, Leptalpheus sp. B was clearly different from both Leptalpheus bait suction pump, from burrow, depth: 1.0–1.5 m, coll. A. Anker & T. Faught, 10 Jan.2005 [fcn 845; specimen photographed]; 1 sp. A and L. pacifi cus. male, CL 6.7, TL not measured (LACM CR 2005-007.1), Viti Levu, Yaqara Bay, ca. 2 km north of Yaqara Pastoral Community, Nasilau At our request, R. Robles and D. L. Felder (University of Point, sand beach, bait suction pump, from burrow, depth 0.5–1.0 Louisiana, Lafayette, USA) performed DNA sequencing of m, coll. A. Anker & A. Bunouf, 27 Jan.2005 [fcn 594; specimen the Nhatrang, Panglao and Fiji specimens, and then compared photographed, dissected]; 1 male, CL 7.4, TL 19.5 (MNHN-Na the obtained sequences to the already available sequences 15776), same collection data as for previous specimen [fcn 562; from the Hawaiian specimens. They confi rmed the presence specimen photographed]; 1 female, CL 5.3, TL 16.7 (OUMNH-ZC of two genetically distinct forms – one in Panglao and 2005-07-16), same collection data as for previous specimen [fcn another in Fiji, Nhatrang and Panglao – both suffi ciently 574]. Philippines: 1 male, CL 6.2, TL 16.8 (ZRC 2005.0086), Panglao Island, Alona Beach, Sta. M1, 9°32.9’N 123°46.6’E, different from the Hawaiian form to be considered as intertidal and shallow subtidal, depth 0–1 m, bait suction pump, separate species. Thus the above-mentioned morphological from burrow of Neocallichirus calmani, coll. P. C. Dworschak, 3 differences between Leptalpheus sp. A,. Leptalpheus sp. B Jun.2004 [fcn PD-1; specimen photographed]; 1 female, CL 6.5, and L. pacifi cus are supported by genetic data, which will be TL 17.65 (ZRC 2005.0087), same collection data as for previous published elsewhere (Robles & Felder, in prep.). Therefore, specimen [fcn PD-1]. Leptalpheus sp. A and Leptalpheus sp. B are described as new species in this study. Comparative material. – Leptalpheus pacifi cus. Hawaii: 1 breeding female, CL 8.1, TL not measured (FLMNH UF Arthropoda 12473), Maui, Hekili Point, 20°8108’N 156°6214’E, back reef 0–1 m, hand collected, C. Pittman, 28 Sep.2007; 1 breeding female, CL and TL MATERIALS AND METHODS not measured (USNM 205907), Oahu, Honolulu, Waikiki Beach, coll. R. Heard, 7 Jan.1988. All specimens were collected with the aid of a steel bait suction pump (“yabby pump”). Some were photographed Description. – (based on Nhatrang specimens) – Body alive prior to preservation in 70% ethanol. Drawings were moderately slender, rather stout in large individuals, carapace made under the dissection microscope equipped with a and abdomen very slightly compressed laterally, glabrous camera lucida. The material is deposited in the following except for minute pits (Fig. 1a). Carapace with inconspicuous institutions: FLMNH, Florida Museum of Natural History, suture proximal to base of antenna (Fig. 1b). Frontal margin Gainesville, USA; LACM, Natural History Museum of Los protruding and broadly rounded, without rostral projection or Angeles County, Los Angeles, USA; MNHN, Muséum orbital teeth, without orbital crests (Fig. 1a). Pterygostomial national d’Histoire naturelle, Paris, France; NMCR, National angle rounded (Fig. 1b, c); branchiostegial region with Museum, Manila, the Philippines; OUMNH, Oxford pronounced “lip” anteriorly (Figs. 1b, c); cardiac notch deep University Museum of Natural History, Oxford, UK; USNM, (Fig. 1d). Eyes not visible in dorsal view, anterior portion National Museum of Natural History, Washington DC, USA; visible in lateral view (Fig. 1a–c); anteromesial process well ZRC, Zoological Reference Collection, Raffl es Museum marked; cornea small, lateral, pigmented (Fig. 1c, e). Ocellar of Biodiversity Research, NUS, Singapore. Abbreviations beak not conspicuous. used in the text: P, pereiopod; Mxp, maxilliped; fcn, fi eld collection number. Antennular peduncle relatively stout (Fig. 1a), fl attened dorsoventrally; second segment about twice as long as broad, about as long as dorsally visible portion of first SYSTEMATICS segment; stylocerite not reaching distal margin of first segment, subacute distally (Fig. 1a, b); ventromesial carina Leptalpheus Williams, 1965 of fi rst segment with strong tooth (Fig. 1f); lateral fl agellum biramous, with shorter ramus well developed, situated at third Leptalpheus denticulatus, new species segment (Fig. 1g). Antenna with basicerite bearing strong (Figs. 1–6, 7a, b, 8) ventrolateral tooth (Fig. 1b); scaphocerite broadly ovate, anterior margin of blade slightly convex, but not protruding Leptalpheus pacifi cus cf. B – Anker et al, 2006: Table 1. beyond distolateral tooth (Fig. 1a); carpocerite long, stout, Leptalpheus cf. pacifi cus C – Anker et al, 2006: Table 1. reaching far beyond scaphocerite (Fig. 1b).

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Mouthparts and third maxilliped typical for genus (Fig. 2a–f). 1.5 times as long as former and bearing several slender, Mandible with incisor process bearing six teeth, median two spiniform, apical and subapical setae (Fig. 2m). largest (Fig. 2a). Maxillule with bilobed palp, both lobes with one stiff seta (Fig. 2c). First maxilliped with expanded Uropod with lateral lobe of protopod bearing two small caridean lobe on exopod (Fig. 2e). Second maxilliped with subacute teeth distally (Fig. 1h); endopod somewhat longer elongate epipod (Fig. 2f). Third maxilliped relatively slender, than exopod, without specifi c features; exopod with truncate elongate; lateral plate acutely produced (Fig. 2g); ultimate posterior margin and with feebly protruding, blunt distolateral segment with rows of long, distally thickened setae, tip tooth adjacent to robust distolateral spiniform seta; lateral unarmed; arthrobranch very large (Fig. 2g). half of diaeresis mostly straight except for central-most portion curved into small tooth (Fig. 1i); mesial portion Chelipeds strongly asymmetrical in shape, unequal in size deeply incised forming stout subtriangular tooth proximal (Fig. 3), carried folded when not in use (Fig. 8a). Major to mesial margin (Fig. 1i). cheliped (on either left or right side) enlarged, elongate (Fig. 3a, b); ischium short, ventromesial margin without Telson moderately slender, slightly tapering distally, about subtriangular tooth (Fig. 3b); merus long, slender, with 2.5 times as long as wide proximally (Fig. 1j); dorsal surface smooth margins, distally not widening, ventrally depressed, pitted (Fig. 1k), with two pairs of strong spiniform setae distal margin ending bluntly; carpus short, cup-shaped, with inserted at short distance from lateral margin, at about 1/3 blunt distolateral process; chela subcylindrical, palm smooth, and 3/5 length of telson, respectively (Fig. 1j); posterior ventromesially excavated, about three times as long as high; margin broadly rounded, each posterolateral angle with two adhesive discs well developed (Fig. 3a, c); fi ngers about spiniform setae, mesial at least fi ve times longer than lateral 1/3 length of palm; dactylus moderately curved distally, (Fig. 1k); anal tubercles small, feebly sclerotised. with subacute tip, cutting edge with one feebly protruding, distally rounded, median tooth, about 1/3 as high as long Gill/exopod formula typical for genus: 5 pleurobranchs and small, subtriangular, proximal tooth; pollex as long as (above P1–5); 1 arthrobranch (above Mxp3); 0 podobranch; dactylus, abruptly ending, tip truncate, with two subacute 2 lobe-shaped epipods (Mxp1–2); 5 mastigobranchs or strap- teeth, cutting edge with one much larger subtriangular like epipods (Mxp3, P1–4); 5 sets of setobranchs (P1–5); 3 proximal tooth and one smaller truncate distal tooth (Fig. exopods (Mxp1–3). 3c, d). Minor cheliped weaker than major cheliped, slender (Fig. 3e), ischium short, unarmed; merus slender, ventrally Colour pattern. – Semitransparent with more or less depressed (Fig. 3e); carpus short, cup-shaped; chela smooth, pronounced pinkish tinge; carapace dorsally with reddish subcylindrical, with fingers about as long as palm, tips ; each pleonite with diffuse transverse crossing when chela closed; cutting edges of dactylus and band of reddish chromatophores along posterior margin; pollex with small curved teeth proximally and two larger antennular and antennal peduncles, telson and uropod opposing teeth at about 2/3 of fi nger length (Fig. 3f). more intense pink due to higher concentration of reddish chromatophores; walking legs and antennular and antennal Second pereiopod relatively slender (Fig. 2h); ischium about fl agella semitransparent; major chela hyaline-whitish with 3/4 length of merus; carpus fi ve-segmented, segments with scattered reddish chromatopores (Fig. 8). ratio approximately equal to 3.5/1/1.5/1/2.5; chela simple, slightly shorter than fi rst carpal segment; fi ngers as long as Size. – The smallest examined specimen is a female from palm. Third pereiopod moderately slender (Fig. 3i), ischium Fiji with 5.3 mm CL and 16.7 mm TL; the largest is an unarmed; merus fl attened mesially, about 2.5 times as long ovigerous paratype female from Nhatrang with 14.5 mm as ischium, about four times as long as wide, with convex CL and 44.7 mm TL. The diameter of embryos (measured dorsal and ventral margins (Fig. 2i); carpus less than 1/2 in the largest female from Nhatrang) is approximately 0.6 length of merus, with distoventral spiniform seta; propodus × 0.5 mm. longer than carpus, with two ventral spiniform setae and one distoventral spiniform seta proximal to dactylus; dactylus Etymology. – The specifi c name (denticulatus – Latin for simple, conical, about 2/5 length of propodus, curved (Fig. 2j). “bearing a denticle or small tooth”) refers to the presence of Fourth pereiopod generally similar to third. Fifth pereiopod a small tooth on the central portion of the uropodal diaeresis, much more slender than third and fourth pereiopods (Fig. a key feature to distinguish the new species from the closely 2k); ischium and merus not fl attened mesially, unarmed; related L. pacifi cus (see under Remarks below). carpus without distal spiniform seta; propodus as long as merus, without spiniform setae, distally with at least six Type locality. – Nhatrang Bay, Vietnam. rows of setae; dactylus similar to that of third and fourth pereiopods. Distribution. – Western tropical Pacifi c; presently known from three distant localities: Nhatrang Bay in Vietnam (type First to fifth pleomeres with minute pits on surface; locality); Viti Levu in Fiji; and Panglao Island, southwest posteroventral angles rounded; sixth pleonite with large of Bohol, in the Philippine Archipelago. articulated plate posteroventrally, dividing suture appearing somewhat incomplete (Fig. 1l). Male second pleopod (Fig. 2l) Ecology. – All specimens were collected from burrows with appendix interna and appendix masculina, latter about of large callianassid ghostshrimps on intertidal or shallow

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Fig. 1. Leptalpheus denticulatus, new species, female paratype from Nhatrang Bay, Vietnam (USNM 1110002): a, frontal region, dorsal view; b, same, lateral view; c, frontal margin of carapace, lateral view; d, carapace, lateral view; e, right eye, ventrolateral view; f, fi rst segment of right antennular peduncle, lateral view; g, lateral fl agellum of right antennule, lateral view; h, right uropod, dorsal view; i, same, detail of exopod, setae omitted; j, telson, dorsal view; k, same, detail of posterior half; l, sixth pleonite and tail fan, setae omitted, lateral view. Scale bars = 1 mm.

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Fig. 2. Leptalpheus denticulatus, new species, a–k: female paratype from Nhatrang Bay, Vietnam (USNM 1110002); l, m: male holotype from Nhatrang Bay (MNHN-Na 17064): a, mandible, mesial view; b, same, molar process; c, maxillule, lateral view; d, maxilla, lateral view; e, fi rst maxilliped, lateral view; f, second maxilliped, lateral view; g, third maxilliped, lateral view; h, second pereiopod, lateral view; i, third pereiopod, lateral view; j, same, distal carpus, propodus and dactylus; k, fi fth pereiopod, lateral view; l, second pleopod, lateral view; m, same, detail of appendix masculina and appendix interna. Scale bars = 1 mm; m drawn without scale.

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Fig. 3. Leptalpheus denticulatus, new species, female paratype from Nhatrang Bay, Vietnam (USNM 1110002): a, major (left) cheliped, lateral view; b, same, mesial view; c, same, chela, carpus and distal merus, ventrolateral view; d, same, distal portion of chela, mesial view; e, minor (right) cheliped, lateral view; f, same, chela, ventrolateral view. Scale bars = 1 mm.

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Fig. 4. Leptalpheus denticulatus, new species, largest female paratype from Nhatrang Bay, Vietnam (MNHN-Na 17065): a, major (left) cheliped, distal merus, carpus and chela, ventrolateral view; b, same, detail of fi ngers; c, same, ventromesial view. Scale bars = 1 mm.

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Fig. 5. Leptalpheus denticulatus, new species, male from Yaqara Bay, Viti Levu, Fiji (LACM CR 2005-007.1): a, frontal region, dorsal view; b, same, lateral view; c, fi rst segment of right antennular peduncle, lateral view; d, mandible, mesial view; e, maxillule, lateral view; f, maxilla, lateral view; g, fi rst maxilliped, lateral view; h, second maxilliped, lateral view; i, third maxilliped, lateral view; j, posterolateral margin of carapace, detail of cardiac notch; k, endopod of second pleopod; l, same, detail of appendix masculina; m, sixth pleonite and tail fan, setae omitted, lateral view; n, left uropod, dorsal view; o, right uropod, detail of exopod, setae omitted, dorsal view; p, telson, dorsal view; q, same, ventral view (showing anal tubercles). Scale bars = 1 mm; l drawn without scale.

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Fig. 6. Leptalpheus denticulatus, new species, male from Yaqara Bay, Viti Levu, Fiji (LACM CR 2005-007.1): a, major (left) cheliped, lateral view; b, same, ventrolateral view; c, same, distal chela, ventromesial view; d, same, ventrolateral view; e, minor (right) cheliped, lateral view; f, same, chela, ventrolateral view. g, second pereiopod, lateral view; h, third pereiopod, lateral view; i, fi fth pereiopod, lateral view. Scale bars = 1 mm.

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subtidal sand flats, fringed by mangroves. In Nhatrang, to the age-related allometry of the major cheliped, but we most specimens were collected from burrows of Glypturus feel that further studies, involving morphometric studies cf. armatus (A. Milne-Edwards, 1870) [IM, pers. obs.], and DNA sequencing, are needed to elucidate the nature whereas the two Panglao specimens were collected from of this variation. burrows of Neocallichirus calmani (Nobili, 1904) [fi eld notes by P. C. Dworschak]. None of the Fijian specimens was collected together with a host, but one specimen of Leptalpheus dworschaki, new species Neocallichirus sp. was collected at the same locality and (Figs. 9–12) the presence of typical mounds of Glypturus sp. was also noted [AA, pers. obs.]. Leptalpheus sp. 7 aff. pacifi cus – Anker et al, 2006: Table 1.

Remarks. – Leptalpheus denticulatus, new species, is closely Type material. – Philippines: Holotype: male, CL 7.7, TL 21.9 related to L. pacificus. These two species share a great (NMCR 27540), Panglao Island, Sungcolan Bay, Sta. M11, 9°38.3'N 123°49.6'E, intertidal fl at, fringed with mangrove and seagrass, bait number of characters, e.g., they have a very similar frontal suction pump, from burrow of Glypturus sp., coll. P. C. Dworschak, margin of the carapace and a nearly identical dentition on 7 Jun.2004 [fcn PD-37; specimen dissected]. Paratype: 1 female, the major chela. However, L. denticulatus, new species, can CL 7.7, TL 23.9 (ZRC 2005.0085), same collection data as for be separated from L. pacifi cus by the presence of a small holotype [fcn PD-36]. tooth on the central portion of the uropodal diaeresis, just before the diaereis slopes abruptly into the large mesial Description. – Body not particularly slender or stout (Fig. incision (Figs. 1i, 5o), and the slightly more slender second 10a), carapace and abdomen slightly compressed laterally, pereiopod, with a smaller chela and shorter first carpal glabrous. Carapace with inconspicuous suture proximal to segment (cf. Figs 2h, 6g and Fig. 7f). The fi gures of L. base of antenna (Fig. 9b, c). Frontal margin protruding and pacifi cus in Banner & Banner (1974) must be used with broadly rounded, without rostral projection or orbital teeth, some caution for they appear to be slightly inaccurate. For without orbital crests (Fig. 9a, b, k). Pterygostomial angle instance, the stylocerite was illustrated with tip overreaching rounded (Fig. 9c); branchiostegial region with pronounced the distal margin of the first segment of the antennular “lip” anteriorly; cardiac notch fairly deep (Figs. 9c, 10a). peduncle (see Banner & Banner, 1974: Fig. 1A) and the Eyes not visible in dorsal view, anterior portion visible lateral lobe of the uropodal protopod as bluntly rounded in lateral view (Fig. 9a, c); anteromesial process feebly (Banner & Banner, 1974: fi g. 1M). However, examination of marked; cornea small, lateral, pigmented. Ocellar beak not these two features in L. pacifi cus (FLMNH UF12473) shows conspicuous. that the stylocerite actually falls short of the distal margin of the fi rst segment (Fig. 7e), while the lateral lobe of the Antennular peduncle relatively stout, fl attened dorsoventrally; uropodal protopod bears two broadly rounded teeth or lobes second segment about twice as long as broad, about as long (Fig. 7g, h), although the latter appear to be blunter than the as dorsally visible portion of fi rst segment; stylocerite not more distinctly protruding, subacute teeth of L. denticulatus, reaching distal margin of fi rst segment, subacute distally (Fig. new species (compare with Figs. 1h, 5n). More specimens 9a, b, k); ventromesial carina of fi rst segment with strong tooth of L. denticulatus, new species, and L. pacifi cus must be (Fig. 9d); lateral fl agellum biramous, with shorter ramus well examined in order to conclude whether the confi guration of developed, situated at third segment. Antenna with basicerite the lateral lobe of the protopod (blunt vs. subacute teeth) can bearing strong ventrolateral tooth (Fig. 9b); scaphocerite be reliably used to distinguish these two species. broadly ovate, anterior margin of blade convex but not protruding beyond distolateral tooth (Fig. 9a); carpocerite Some intraspecifi c variation was observed among specimens long, stout, reaching far beyond scaphocerite. of L. denticulatus, new species, either from the same or from different localities. For instance, the Nhatrang specimens Mouthparts and third maxilliped as in previous species (Fig. (Figs. 1–4) differ from the Fijian specimens (Figs. 5–6) by the 10b–h; for description see above). proportions of the third pereiopod, particularly in the merus being somewhat broader and having more convex margins Chelipeds strongly asymmetrical in shape, unequal in size (cf. Figs. 2i, 6h). The two illustrated Nhatrang paratypes, (Figs. 11, 12a, b), carried folded when not in use (Fig. a 10 mm CL female and a 14.5 mm CL breeding female, 10a). Major cheliped (on either left or right side) enlarged, differ from each other by the extension of the blunt tooth elongate (Fig. 11); ischium short, ventromesial margin on the cutting edge of the major chela dactylus, which is without subtriangular tooth; merus long, slender, with smooth relatively short and more protruding in the smaller female margins, distally not widening, ventrally fl attened, distally (Fig. 3c), and distinctly longer and more fl attened in the more depressed, distal margin with blunt lobes; carpus short, large female (Fig. 4b); this latter confi guration is similar cup-shaped, with blunt distal process; chela subcylindrical, to that of the Fijian specimens (Fig. 6b, c) and especially palm smooth, ventromesially excavated, about 2.5 times as to the Hawaiian L. pacifi cus (Fig. 7c, d). They also differ long as high (Fig. 11c); adhesive discs well developed (Fig. by the ratio of the fi ngers to the palm, which is closer to 11b, c); fi ngers about 1/3 length of palm; dactylus moderately 1/3 in the smaller female (Fig. 3c) and almost 1/2 in the curved distally, with subacute tip, cutting edge with one larger female (Fig. 4a), approaching the ratio observed in median subtriangular-rounded tooth, about 1/2 as high as L. pacifi cus (Fig. 7c, d). These differences are possibly due long, without proximal tooth; pollex shorter than dactylus,

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Fig. 7. Leptalpheus denticulatus, new species (a, b), male from Panglao, Philippines (ZRC 2005.0086): a, frontal region, dorsal view; b, major (left) cheliped, carpus and chela, lateral view. Leptalpheus pacifi cus Banner & Banner, 1974 (c–h), female from Hawaii (USNM 205907): c, major (right) cheliped, ventrolateral view; d, same, detail of fi ngers; breeding female from Hawaii (FLMNH UF Arthropoda 12473): e, frontal region, dorsal view; f, second pereiopod, lateral view; g, right uropod and telson, dorsolateral view, setae omitted; h, left uropod, detail of protopod. Scale bars: a–g = 1 mm; h = 0.5 mm.

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abruptly ending, with one blunt terminal tooth, cutting edge (Fig. 9f). Male second pleopod (Fig. 9g) with appendix with two teeth: one much larger subtriangular proximal tooth interna and appendix masculina, latter almost twice as long and one smaller truncate distal tooth (Fig. 11c, d). Minor as former, and bearing numerous slender, spiniform apical cheliped shorter and weaker than major cheliped (Fig. 12a); and subapical setae (Fig. 9h). ischium short, unarmed; merus slender, ventrally depressed; carpus very short, cup-shaped; chela smooth, subcylindrical, Uropod with lateral lobe of protopod bearing two small with fi ngers about 1.2 times as long as palm, tips crossing acute teeth distally (Fig. 9i); endopod somewhat longer when chela closed; cutting edges of dactylus and pollex with than exopod, without specifi c features; exopod with truncate small curved teeth proximally and two larger opposing teeth posterior margin and without distinct distolateral tooth at about mid-length (Fig. 12b). adjacent to distolateral spine; lateral portion of diaeresis straight, not curved into tooth centrally; mesial portion Second pereiopod slender; ischium about 3/4 length of merus; deeply incised forming large triangular tooth proximal to carpus fi ve-segmented, segments with ratio approximately mesial margin (Fig. 9i). equal to: 4/1/1/1/2.5 (Fig. 12c); chela simple, about as long as fi rst carpal segment; fi ngers as long as palm. Third Telson moderately slender, more than twice as long as wide pereiopod moderately slender (Fig. 12d); ischium unarmed; proximally (Fig. 9j); dorsal surface pitted, with two pairs of merus fl attened mesially, about 2.5 times as long as ischium, robust spiniform setae inserted at short distance from lateral about fi ve times as long as wide; carpus less than 1/2 length margin, at about 1/3 and 2/3 length of telson, respectively; of merus, with distoventral spiniform seta; propodus longer posterior margin feebly rounded, posterolateral angles each than carpus, with three ventral spiniform setae and one with pair of spiniform setae, mesial at least four times as distoventral spinifom seta proximal to dactylus; dactylus long as lateral; anal tubercles small, feebly sclerotised. simple, conical, about 2/5 length of propodus, curved (Fig. 12d). Fourth pereiopod (Fig. 12e) generally similar to third. Gill/exopod formula as in previous species (see above). Fifth pereiopod (Fig. 12f) much more slender than third and fourth pereiopods; ischium and merus not fl attened mesially, Colour pattern. – Field notes indicated: “Translucent, slightly unarmed; carpus without distal spiniform seta; propodus pink, ovary green”. as long as merus, without spiniform setae, distally with at least seven rows of setae; dactylus similar to that of third Size. – The CL of both specimens is 7.7 mm; the TL ranges and fourth pereiopods. from 21.9 mm in the male holotype to 23.9 mm in the female paratype. First to fifth pleomeres with minute pits on surface; posteroventral angles rounded; sixth pleonite with large Etymology. – The new species is named after Peter C. articulated plate posteroventrally (Fig. 10a). First pleopod Dworschak (Naturhistorisches Museum in Wien, Vienna, woth very short endopod, latter fringed with setae apically Austria), the collector of the present specimens, our estimated friend and colleague, and also a well known specialist of the Thalassinidea.

Type locality. – Panglao Island, southwest of Bohol, the Philippines.

Ecology. – Both specimens were collected from a Glypturus cf. armatus burrow, together with gastropods (Phenacolepadidae) and sentinel crabs, Macrophthalmus sp. (Ocypodidae) [fi eld notes by P. C. Dworschak].

Distribution. – Western Pacifi c: presently known only from the type locality in the Philippines.

Remarks. – Leptalpheus dworschaki, new species, is closely related to L. pacifi cus and the above-described L. denticulatus, new species, but can be separated from both species by the armature on the major chela fi ngers, in particular by the presence of a distinct blunt tooth on the dactylus (absent in L. pacifi cus and L. denticulatus, new species, although larger individuals may have a broad bulge at the same position); the absence of a small proximal tooth on the dactylus (present Fig. 8. Leptalpheus denticulatus, new species, colour pattern in in L. pacifi cus and L. denticulatus, new species); and the life: a, non-type male from Yaqara Bay, Viti Levu, Fiji (MNHN- shorter hiatus between the larger proximal and smaller distal Na 15776); b, large female paratype from Nhatrang Bay, Vietnam (MNHN-Na 17065), major (left) cheliped detached (photograph teeth on the pollex (this hiatus is almost twice as long in by O. Savinkin). L. pacifi cus and L. denticulatus, new species) (compare

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Fig. 9. Leptalpheus dworschaki, new species, male holotype (NMCR 27540): a, frontal region, dorsal view; b, same, lateral view; c, frontal margin of carapace, lateral view; d, fi rst segment of right antennular peduncle, lateral view; e, lateral fl agellum of right antennule, lateral view; f, fi rst pleopod, mesial view; g, second pleopod, mesial view; h, same, appendix masculina and appendix interna; i, right uropod, dorsal view; j, telson, dorsal view; female paratype (ZRC 2005.0085): k, frontal margin, dorsal view. Scale bars = 1 mm.

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Fig. 10. Leptalpheus dworschaki, new species, female paratype (ZRC 2005.0085): a, habitus; male holotype (NMCR 27540): b, mandible, mesial view; c, same, dorsolateral view; d, maxillule, lateral view; e, maxilla, lateral view; f, fi rst maxilliped, lateral view; g, second maxilliped, lateral view; h, third maxilliped, lateral view. Scale bar = 1 mm.

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Figs. 11c, d and Figs. 3c, d; 4 b, c; 7c, d). Leptalpheus ACKNOWLEDGMENTS dworschaki, new species, can be also separated from L. denticulatus, new species, by the absence of a small tooth We are grateful to A. Richard Palmer (University of Alberta, on the central portion of the uropodal diaeresis. We were Edmonton, Canada) for supporting research on the family unable to fi nd any other obvious morphological differences Alpheidae from an NSERC operating grant (A7245). Steve between L. dworschaki, new species, and L. pacifi cus and Dunbar (Loma Linda University, Loma Linda, USA) invited L. denticulatus, new species. one of us (AA) to participate in the Marine Cryptofauna Survey of the Yaqara Bay, Fiji, in January 2005. Leslie The length of the second segment of the antennular peduncle Harris and George Davis (LACM) arranged the loan of and the shape of the frontal margin appear to be somewhat LACM specimens. All Panglao specimens were collected variable in both L. dworschaki, new species (Fig. 9a, k), and by Peter C. Dworschak (Naturhistorisches Museum in Wien, L. denticulatus, new species (Figs. 1a, 5a); the latter species Austria) during the Panglao Biodiversity Project 2004, made is also variable in the proportions of the merus of the third possible through grants from the Total Foundation, the French pereiopod (Figs. 2i, 6h; see also above). Therefore, these Ministry of Foreign Affairs, and the ASEAN Regional features cannot be reliably used to distinguish L. dworschaki, Centre for Biodiversity Conservation, and coordinated by new species from L. denticulatus, new species, or from L. Philippe Bouchet (MNHN) and Danilo Largo (University of pacifi cus. San Carlos, Cebu, the Philippines). Peter K. L. Ng, S. H.

Fig. 11. Leptalpheus dworschaki, new species, male holotype (NMCR 27540): a, major (left) cheliped, mesial view; b, same, lateral view; c, same, chela, carpus and distal merus, ventrolateral view; d, same, distal chela, ventromesial view. Scale bars = 1 mm.

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Fig. 12. Leptalpheus dworschaki, new species, female paratype (ZRC 2005.0085): a, minor (right) cheliped, lateral view; b, same, chela, mesial view; male holotype (NMCR 27540): c, second pereiopod, lateral view; d, third pereiopod, lateral view; e, fourth pereiopod, lateral view; f, fi fth pereiopod, lateral view. Scale bars = 1 mm.

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Tan, Joelle Lai, the late K. L. Yeo (National University of Banner, A. H. & D. M Banner, 1983. An annotated checklist of Singapore, Singapore) and Chia-Wei Lin (National Taiwan the alpheid shrimp from the western Indian Ocean. Travaux Ocean University, Keelung, Taiwan) were helpful during et Documents de l’ORSTOM, 158: 1–164. the sorting of the Panglao 2004 material; S. H. Tan and Dworschak, P. C. & V. R. Coelho, 1999. On two alpheids from late K. L. Yeo arranged the loan of this material. Oleg V. Araça (São Paulo, Brazil) with a description of a new species Savinkin (Moscow, Russia) helped with photography of of Leptalpheus (Decapoda: Caridea: Alpheidae). Annalen des some Nhatrang specimens. Karen Reed (USNM) provided Naturhistorischen Museums in Wien, 101B: 475–488. collection data for the USNM specimen of L. pacificus Felder, D.L. & R. B. Manning, 1986. A new genus and two new and checked a USNM paratype specimen at our request. species of alpheid shrimps (Decapoda: Caridea) from South Two anonymous reviewers made very useful suggestions Florida. Journal of Crustacean Biology, 6: 497–508. improving the quality of the manuscript. Milne-Edwards, A., 1870. Révision du genre Callianassa (Leach) et description de plusieurs espèces nouvelles de ce groupe faisant partie de la collection du Muséum. Nouvelles Archives du LITERATURE CITED Muséum d’Histoire Naturelle de Paris, 6: 75–102, Pls. 1, 2. Nobili, G., 1904. Diagnoses préliminaires de vingt-huit espèces Anker, A., 2008. The shrimp genus Leptalpheus Williams, 1965 nouvelles de stomatopodes et décapodes macroures de la Mer in the southeastern Caribbean Sea, with description of one Rouge. Bulletin du Muséum d’Histoire Naturelle, Paris, 10: new species from Panama (Crustacea, Decapoda, Alpheidae) 228–238. Zoosystema, 30: 781–794. Ríos, R. & A. Carvacho, 1983. Leptalpheus mexicanus, new Anker, A., J. A. Vera Caripe & C. Lira, 2006. Description of a new species (Crustacea, Decapoda, Alpheidae). Caridean shrimps species of commensal alpheid shrimp (Crustacea, Decapoda) of the Gulf of California III. Journal of Crustacean Biology, from the southern Caribbean Sea. Zoosystema, 28: 683–702 3: 306–313. Banner, A. H. & D. M Banner, 1974. Contributions to the knowledge Williams, A. B., 1965. A new genus and species of snapping of the alpheid shrimp of the Pacifi c Ocean, part XVII: additional shrimp (Decapoda, Alpheidae) from the southeastern United notes on the Hawaiian alpheids: new species, subspecies and States. Crustaceana, 9: 192–198. some nomenclatorial changes. Pacifi c Science, 28: 423–437.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 109Ð112 Date of Publication: 28 Feb.2009 © National University of Singapore

DESCRIPTION OF THE PARALARVAE OF WUNDERPUS PHOTOGENICUS HOCHBERG, NORMAN & FINN, 2006 (CEPHALOPODA: )

Christine Louise Huffard Monterey Bay Aquarium Research Institute 7700 Sandholdt Road, Moss Landing, California, USA, 95039 Email: [email protected] (Corresponding author)

Briand Alessandra Gentry Dallas, Texas, USA, 75229 Email: [email protected]

David Wayne Gentry Dallas, Texas, USA, 75229 Email: [email protected]

ABSTRACT. – Recruitment studies of wild animals rely heavily on a means to identify early life history stages. Among shallow-water octopodids (Cephalopoda: Octopodidae), founder patterns are used to identify paralarvae (planktonic hatchlings) to the species level. Here we describe the founder chromatophore patterns and external morphology of paralarval Wunderpus photogenicus, an that is commercially important but remains unmonitored in the wild. Within the brood of a single female Wunderpus photogenicus, we observed variation in key characteristics used to identify paralarvae to species: the number of chromatophores on the ventral and funnel of hatchlings. Chromatophores preserved in various states of relaxation and contraction could generate these different variations of the baseline pattern, even when all chromatophores are anatomically present.

KEYWORDS. – Life history, larvae, octopus, marine ornamentals, plankton, Indo-West Pacifi c.

INTRODUCTION development of cephalopod skin patterns (Packard, 1985). We have limited knowledge of the chromatophore patterns of Wunderpus photogenicus Hochberg, Norman & Finn, 2006 is tropical octopodid paralarvae, especially those from regions a charismatic tropical octopus distributed from the Philippines of extreme biodiversity with pressing marine management to (Norman, 2000; Hochberg et al., 2006). Its distinct needs, such as Indonesia. For example of the more than deimatic display, possible behavior, and apparent ten octopodids that inhabit North Sulawesi, Indonesia rarity help make it an expensive marine ornamental that is also (distributions in Norman, 2000), hatchlings have been popular among underwater photographers and videographers illustrated or photographed for only Hapalochlaena lunulata (Hochberg et al., 2006). Despite the commercial value of Quoy & Gaimard, 1832, Octopus cyanea Gray, 1849, and these animals, life history and population status of wild Abdopus aculeatus (d’Orbigny, 1834) (Overath & Boletzky, stocks remain entirely unstudied. Photoidentifi cation offers a 1974; Young et al., 1989; Huffard, 2007). Here we describe means to estimate the density and home ranges of individual the founder chromatophore patterns and other morphological adults (Huffard et al., 2008). However tools for identifying characteristics of W. photogenicus hatchlings offering a means and monitoring their planktonic hatchlings from larval stocks to identify them from zooplankton samples. are not currently available.

Traditionally, species-specific founder chromatophore MATERIALS AND METHODS patterns are used to identify cephalopod paralarvae, a process that can be verifi ed by collecting hatchlings from confi dently- A mature female Wunderpus photogenicus (20 mm mantle identifi ed brooding females. Founder chromatophores are length post-spawning) was purchased from a live reef fi sh conspicuous dark pigment sacs that appear early in the store by DWG (Dallas North Aquarium, Dallas Texas) on 4

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Sep.2006. The female was reported to have been collected Brooding was monitored from four days after the eggs were from , Indonesia on either 28 or 29 Aug.2006, and laid until hatching. Paralarvae (n = 46, all preserved at autotomized one arm in transit from Los Angeles, California hatching on day 37) and unhatched eggs (n = 35 preserved to Dallas, Texas within the United States. Beginning 13 on day 35; n = 5 preserved on day 37) were preserved in September 2006 she was maintained at 21Ð25¡C in a 100 10% buffered formaldehyde for approximately four weeks L BioCubeª with sand and live rock substrate. She fed and then transferred to 95% isopropyl alcohol. Paralarvae on three to four reef crabs [Varuna litterata (Fabricius, in good condition (n = 22) were measured and drawn under 1798)], carapace width approximately 19Ð32 mm) every a compound microscope with the aid of a camera lucida. three to four days by web-casting and immobilizing prey. Descriptions of paralarvae follow Hochberg et al. (1992). The lighting schedule was initially 13:11 hr daylight and darkness respectively, until 30 Sep.2006 when a lighting system was installed to produce 8 hrs of daylight, preceded RESULTS and followed by one hour of dawn and dusk, with a 14 hr period of blue ‘night’ light. Lighting was provided by 40.6 cm On approximately 22 Sep.2006 the female Wunderpus 36 W Compact Flourescent 10 K Daylight, 40.6 cm Actinic photogenicus stopped eating, hid in live rock, and was 03 Phospor and Lunar Blue- Moon-Glow LED bulbs. believed to have laid eggs. Brooding was considered to

Fig. 1. Hatchlings of Wunderpus photogenicus. (left) Dorsal view. Chromatophores with dashed outlines are situated on the digestive gland. (right) Ventral view. Scale bar = 1 mm.

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have started on this date. The eggs were held in the oral of hatchlings. Some hatchlings did not bear all of the web, near the mouth as described by Miske & Kirchhauser chromatophores in the baseline pattern (Fig. 1). Similar (2006). For an unknown reason, on occasion the female variation has been documented previously for a number dropped small egg strands while crawling along the bottom, of other octopodid populations (Hochberg et al., 1992). In a behavior reported previously for Hapalochlaena sp. by a Loliginid squid, differences in embryonic temperatures Tranter & Augustine (1973). These eggs were transferred are correlated with certain founder chromatophores being to a mesh net basket within the aquarium. By day 33 after missing from the baseline pattern typical of that species spawning the female had crawled into a pump chamber (Gowland et al., 2003). It may be possible that eggs in the (opening 1.5 mm diameter) and abandoned the remaining single brood of W. photogenicus observed here experienced eggs. Hatching took place on day 37. Hatchlings swam in slightly different temperature regimes, but not nearly to the water column rather than crawling on the bottom, and the degree needed to produce such changes in squid. Thus, likely adopt a planktonic lifestyle. The female died on day another mechanism likely led to variation observed in these 38, after more than a month without feeding. preserved specimens. In live octopodid hatchlings (such as in the planktonic paralarvae of Octopus bocki Adam 1941, The female Wunderpus photogenicus laid 86 fertile eggs. CLH pers. obs.) founder chromatophores may not be visible We do not know if additional eggs were laid and either if radial chromatophore muscles are relaxed and the elastic degraded, eaten, or hatched earlier and lost in the tank. At pigment sac is contracted. Chromatophores preserved in death the female’s ovary measured approximately 4 mm in various states of relaxation and contraction could generate diameter and contained immature oocytes. different variations of a baseline pattern, even when all chromatophores are anatomically present. Thus workers Hatchlings measured 1.35 mm ML ± 0.03 S.E. (n = 22; Fig. aiming to identify early stages of W. photogenicus should 1) and exhibited the following morphological characteristics: consider all variations observed here, particularly if using arms short, subequal, and with four suckers each, extending preserved material for which the arms are degraded. approximately two thirds of proximal arm length, distal third of the arms without suckers; approximately 20Ð50 small fi nger-like projections scattered on the distal third of ACKNOWLEDGEMENTS aboral surfaces of all arms; webs subequal, extending down approximately half of arm length; arms with 4 dark brown Maddi Gentry diligently helped care for and observe chromatophores (1 arm base chromatophore plus longitudinal brooding by the W. photogenicus “Wawasan.” Eric Hochberg, row of 1 + 2 chromatophores running from proximal to Janet Voight, Becky Williams, Stephanie Bush, and three distal tip); funnel with 2 + 2 chromatophores, two specimens anonymous reviewers helpfully critiqued this manuscript. with 1 + 2 + 2 confi guration (2 on funnel base, line of 3 on distal funnel); dorsal head with 6 chromatophores (2 + 4); one chromatophore on each side of mantle; ventral mantle LITERATURE CITED with 5Ð7 chromatophores along the lateral and posterior margin; visceral mass with 8Ð13 chromatophores scattered Adam, W., 1941. Notes sur les céphalopods, XIV. –Sur une evenly over surface. Fig. 1 illustrates maximum number of nouvelle espèce d’Octopode (Octopus bocki, sp. nov.) des chromatophores. Chromatophores were missing from this Iles Fidji. Bulletin du Musée royal d’Histoire naturelle de baseline pattern in variable forms. Belgique, 17: 1Ð3. Fabricius J. C., 1798. Supplementum Entomologiae Systematicae. Hafniae, P. 572. DISCUSSION Gowland F. C., P. R. Boyle & L. R. Noble, 2003. Asymmetry in the embryonic chromatophore pattern of the squid Loligo The hatchlings of Wunderpus photogenicus can be identifi ed forbesi (: Cephalopoda): a proxy for developmental by both their unique founder chromatophore patterns and instability. Journal of the Marine Biological Association of the United Kingdom 83 tiny fi nger-like projections on the arm tips. We could fi nd , :1101Ð1105. no reference to arm-tip projections or similar chromatophore Gray, J. E., 1849. ‘Catalogue of the Mollusca in the collection of patterns in previously described octopodid paralarvae. the British Museum. Part 1. Cephalopoda Antepedia’ London. However similar finger-like arm-base projections have Pp. 1Ð164. been described previously for paralarvae of the undescribed Hochberg, F. G., M. D. Norman & J. Finn, 2006. Wunderpus ‘crescent’ octopus from Hawaii (Young et al., 1989). We do photogenicus n. gen. and sp., a new octopus from the shallow not know the degree to which these paralarval characteristics waters of the Indo-Malayan Archipelago (Cephalopoda: Octopodidae). Molluscan Research, 26:128Ð140. may refl ect evolutionary relationships. Because the identity of the ‘crescent’ octopus remains unclear, we are not able Hochberg F. G., M. Nixon & R. B. Toll, 1992. Octopoda. In: to compare similarities in adult forms. Sweeney, M. J., C. F. E. Roper, K. M. Mangold, M. R. Clarke, & S. V. Boletzky, “Larval” and juvenile : a manual for their identifi cation. Smithsonian Contributions to Within the brood of a single female Wunderpus photogenicus, Zoology, 513: 213Ð279. we observed variation in key characteristics used to identify cephalopod paralarvae to species: the number Huffard, C. L., 2007. Ethogram of Abdopus aculeatus (Cephalopoda: Octopodidae): can behavioral characters inform octopodid of chromatophores on the ventral mantle and funnel

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taxonomy and systematics? Journal of Molluscan Studies, Overath H. & S. von Boletzky, 1974. Laboratory observations on 73: 185Ð193. spawning and embryonic development of a blue ringed octopus. Huffard, C. L., R. L. Caldwell, N. DeLoach, D. W. Gentry, P. Marine Biology, 27: 333Ð338. Humann, B. MacDonald, B. Moore, R. Ross, T. Uno & S. Packard, A., 1985. Sizes and distribution of chromatophores during Wong, 2008. Individually unique body color patterns in octopus post-embryonic development in cephalopoda. Vie Milieu, 35: (Wunderpus photogenicus) allow for photoidentifi cation. PloS 285Ð298. One, 3: e3732. Quoy, J. R. & J. P. Gaimard, 1832. Mollusques, Voyage de Miske V. & J. Kirchhauser, 2006. First record of brooding and decouvertes de l’Astrolabe pendant les annees 1826-1827- early life cycle stages in Wunderpus photogenicus Hochberg, 1828-1829. Zoologie, 2:1Ð320. Norman & Finn, 2006 (Cephalopoda: Octopodidae). Molluscan Research, 26:169Ð171. Tranter D. J. & O. Augustine, 1973. Observations on the life history of the blue-ringed octopus Hapalochlaena maculosa. Marine Norman, M. D., 2000. Cephalopods: A world guide. ConchBooks, Biology, 18: 115Ð128. Hackenheim, Pp. 1Ð 318. Young R. E., R. F. Harman & F. H. Hochberg, 1989. Octopodid Orbigny, M. A. d’, 1834. Voyage dans l’Amerique méridionale, t. paralarvae from Hawaiian USA waters. Veliger, 32:152Ð165. V. Mollusques. In ‘Histoire naturelle générale et particulière Cephalopodes Acetabulifères vivants et fossiles’. (Eds M. M. Ferussac and M. A. d’Orbigny.) Pp. ix + 361.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 113Ð118 Date of Publication: 28 Feb.2009 © National University of Singapore

THE FIRST RECORD OF A SHARK OF THE GENUS GLYPHIS IN INDONESIA

Fahmi and Mohammad Adrim Research Centre for Oceanography, Indonesian Institute of Sciences (LIPI) Jl. Pasir Putih I Ancol Timur Jakarta 14430, Indonesia Email: [email protected]

ABSTRACT. – Sharks of the genus Glyphis are one of the elasmobranch fi shes believed to inhabit tropical freshwater habitats. Presently, fi ve species of Glyphis are known: Glyphis gangeticus, G. glyphis, G. siamensis, G. garricki, and G. sp. B. The latter has not been formally described and is known as Borneo River Shark. A shark species of the genus Glyphis was recorded at Sampit Bay, Central Kalimantan, Indonesia in September 2005. It was a mature male (1,660 mm total length) and was caught using hookÐand-line by a local fi sherman. The specimen was not kept due to the large size and lack of facilities in the fi eld. However, this fi nding is the fi rst record of the occurrence of this genus in Indonesia.

KEY WORDS. – Glyphis, shark, freshwater, Kalimantan, Indonesia.

INTRODUCTION Glyphis is G. siamensis that was described from a specimen collected in the mouth of the Irrawaddy River near Rangoon The river shark of the genus Glyphis is known as one of the (now Yangon), Myanmar, by Steindachner in 1896. While few elasmobranch species known to occur and live in tropical three other undescribed Glyphis were named as species A, freshwater habitat. This genus is poorly known compared to B and C based on the specimens collected from Queensland other genera of the carcharhinid family (Compagno, 1988). (Australia), Borneo (Sabah, Malaysia), New Guinea and The rarity, lack of information and diffi culties in identifying Northern Territory (Australia), respectively (Compagno, the genus are the main factors contributing to the poor 1998). Later, Compagno et al. (2005) separated the specimens knowledge of Glyphis biology and taxonomy (Compagno from Bay of Bengal and New Guinea as Glyphis glyphis, et al., 2005). Species of this genus are often confused with while the specimens from the Bizant River, Queensland and the bull shark, Carcharhinus leucas, another member of the Adelaide River, Northern Territory were named as Glyphis family Carcharhinidae, and which is also the more commonly sp. A, with a note that this may be the same species as encountered freshwater shark species inhabiting rivers and Glyphis glyphis. Specimens from Borneo deposited in the estuaries in the Indo-Pacifi c region (Compagno, 2002). Naturhistorisches Museum, Vienna and several juveniles collected from Kinabatangan River, Sabah, were named as Based on Compagno’s (1998; 1999; 2002; 2005) checklists Glyphis sp. B in Compagno (1998; 2002) and Compagno of living elasmobranchs, there are at least six species of et al. (2005), while specimens from Bainuru and Fly River, Glyphis known worldwide, i.e. Glyphis glyphis (Müller & Papua New Guinea, also those from Adelaide and South Henle, 1839), Glyphis gangeticus (Müller & Henle, 1839), Alligator River in the Northern Territory were named as Glyphis siamensis (Steindachner, 1896), Glyphis sp. A [Last Glyphis sp. C (Compagno, 2002; Last, 2002). Lastly, the & Stevens, 1994], Glyphis sp. B and C [Compagno, 1998]. identities of Glyphis sp. A and C have been clarifi ed by In a recent publication, Compagno et al. (2008), G. sp A was Compagno et al. (2008) as G. glyphis and G. garricki, regrouped into G. glyphis and G. sp. C was described as G. respectively. The type species Glyphis glyphis was recently garricki Compagno, White & Last, 2008. The nomenclature redescribed based on neotype specimen (ZMB 5265) and and classifi cation of genus Glyphis were clearly explained in 15 other specimens from Northern Territory, Queensland Compagno (1988). The description of the speartooth shark, and Papua New Guinea, while G. garricki was formally Glyphis glyphis, apparently based on a stuffed specimen named and described based on nine specimens collected from the Zoologisches Museum, Humbolt Universitat, with from Northern Territory, Western Australia and Papua New no information on its locality, and the holotype no longer Guinea (Compagno et al., 2008). exists (Compagno, 1988). Glyphis gangeticus was described from two specimens collected in the Ganges River, India Indonesia is considered to have the highest number of (Compagno, 1988; 1998; 2002). Another species of genus elasmobranch species in the world, due to its extensive

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marine habitat diversity (see Compagno, 1998; Compagno shark found during the study because it was only recorded et al., 2005; White et al., 2006). At least 135 elasmobranch from a single specimen. A mature male specimen of Glyphis species occur in eastern Indonesia of which 78 species of sp. was recorded at a small landing site in Ujung Pandaran them are sharks (White et al., 2006). This number is expected Village, Sampit District, Central Kalimantan Province, on to increase if other areas elsewhere in the country, such as the 3 Sep.2005. The village is located in the western part western Indonesia is considered. Studies of elasmobranch of Sampit Bay, which is connected to the Java Sea (Fig. 1). diversity in Indonesia have been conducted since 2001 by There is a small scale fi shery operation in the waters off a collaboration research team between Indonesian research the village targeting shrimps and crabs. Most local fi shers institutions such as Fisheries and Marine Department catch fi shes only for local consumption using hand lines and (Department Perikanan dan Kelautan, DKP), the Indonesian gillnets. Fishes are also caught as bycatch in the trammel Institute of Sciences (Lembaga Ilmu Pengetahuan Indonesia, nets and lift nets targeting for shrimps. The Glyphis specimen LIPI) and CSIRO Australia under ACIAR project. The was caught by a local fi sherman using a hook and line at research is presently continued by staff of the Research depth of less than 30 m in the waters off the village. Based Centre for Oceanography, Indonesian Institute of Sciences on the diagnostic characteristics of the specimen, the fi sh (RCO-LIPI) itself. clearly belongs to the genus Glyphis (Fig. 2).

There is currently no information and publication on the The primary character distinguishing the genus Glyphis is the occurrence of Glyphis in Indonesia. The extensive Indonesia- non-crescentic (longitudinal) shape of the upper precaudal pit Australia elasmobranch project carried out between 2001 (Fig. 3) as stated by Compagno (1998), and the large relative to early 2006, and which resulted in a guidebook on size of the second dorsal-fi n (Last & Stevens, 1994). Those economically important sharks and rays of Indonesia (White primary characters clearly distinguished between Glyphis and et al., 2006) also did not list any Glyphis species. This the bull shark, Carcharhinus leucas. Due to its large size, paper provides the fi rst information of this genus from the the Glyphis specimen from Sampit Bay was not collected Indonesian Borneo (Central Kalimantan), and extending its but some measurements were taken and the specimen was distribution from other known location on Borneo Island (i.e. photographed. The total length of the specimen was 1,660 Sabah in Malaysian Borneo; see Compagno, 1998; Manjaji, mm and had fully calcifi ed claspers (the outer length of the 2002a & b; Compagno et al., 2005). right clasper was 115 mm).

Morphological features of the Glyphis from Sampit Bay, MATERIALS AND METHODS Kalimantan, are as follows: long and broad pectoral-fi ns with straight posterior margins; snout short and bluntly A study on biodiversity and fisheries of elasmobranchs rounded; mouth long and parabolic, its length is about a was conducted at various landing sites and coastal villages half of its width (Fig 4). Absence of an interdorsal ridge; in Kalimantan, Indonesia from May 2005 to Oct.2006. A fi rst dorsal-fi n large and broadly triangular in shape with total of six fi sh landing sites in Kalimantan were surveyed posterior margin almost straight, origin posterior to insertion (two sites in West and East Kalimantan and one site in of pectoral fi n; second dorsal-fi n high with almost straight Central and South Kalimantan; see Fahmi & Adrim, 2007). edge at posterior margin and has long posterior lobe; teeth All elasmobranch species were identifi ed, recorded, sexed in upper jaw broad and triangular with the outer margins and counted in each location. Measurements such as total serrated; lower teeth erect with narrow cusps and without length, disc width, weight and claspers length were measured prominent cusplets (Fig 5); pectoral-fi n length 0.58 of its if possible. Unidentified and unique specimens were anterior margin length; anterior margin of the pelvic-fi n 0.30 photographed and some measurements and morphometric of pectoral anterior margin; height of the second dorsal-fi n counts were taken following Compagno (1988; 1998). 0.40 of fi rst dorsal fi n, its base 0.42 of fi rst dorsal-fi n base Some of them were kept and preserved if the size was (Fig 6); anal-fi n height 0.77 of second dorsal-fi n height and reasonable for reference collections. Collected specimens its base 0.85 of second dorsal-fi n base; snout length shorter are registered using National Collection of Indonesian Pisces than mouth width (0.5 times), and head 0.2 of its total length. (NCIP) numbers of Research Centre for Oceanography of The measurement ratios comparing to the total length are the Indonesian Institute of Sciences (LIPI). presented in Table 1.

Unfortunately, due to the lack of preservation facility during RESULTS the study, neither tissue sample nor the body part was collected. Instead, the specimen was processed by villagers A total of 2,310 individuals of elasmobranchs were recorded as both smoked and salted meat for local consumption. The from seven fi sh landing sites in Kalimantan representing fi ns were dried, but the head and guts were discarded. 60 species of 13 families (for a list of species see Fahmi & Adrim, 2007). The batoids (Dasyatidae) was the most common chondrichthyan family recorded during the study in DISCUSSION terms of the number of individuals but Carcharhinidae was one of the most speciose groups together with Dasyatidae Glyphis sp. from Sampit Bay, Kalimantan is the largest (Fahmi & Adrim, 2007). The genus Glyphis was the rarest specimen of the genus Glyphis so far recorded in Borneo

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Table 1. Measurement ratio (in %) of total length (TL) for Glyphis sp. from Sampit Bay compared to Glyphis glyphis and G. garricki specimens (Compagno et al., 2008).

Glyphis sp. (1,660 mm TL) Glyphis glyphis Glyphis garricki Head length 21.4 24.0Ð26.0 24.0Ð25.5 Snout length 6.1 2.8Ð7.9 7.0Ð7.7 Mouth width 12.2 9.8Ð10.8 10.4Ð11.2 Pectoral-fi n length 14.9 11.3Ð14.0 12.9Ð13.8 Pectoral anterior margin 25.7 17.5Ð22.2 19.9Ð22.4 First dorsal-fi n length 17.1 16.2Ð19.6 16.7Ð18.7 First dorsal-fi n base 13.6 12.1Ð13.9 11.9Ð13.3 Second dorsal-fi n length 8.6 9.8Ð13.0 10.2Ð12.5 Second dorsal-fi n base 5.7 6.5Ð8.9 6.6Ð8.4 Interdorsal length 22.1 14.5Ð20.8 16.1Ð21.2 Pelvic-fi n length 8.9 7.1Ð10.3 9.2Ð10.6 Pelvic-fi n anterior margin 7.7 7.0Ð9.0 7.6Ð9.1 Pelvic-fi n height 4.3 4.7Ð8.0 6.9Ð8.1 Pelvic-fi n base 5.3 4.6Ð6.6 5.4Ð6.8 Anal-fi n height 3.7 3.0Ð5.4 4.6Ð5.3 Anal-fi n base 4.8 5.5Ð7.3 6.0Ð7.0

Fig. 1. Sampling locality of the reported Glyphis from Ujung Pandaran Village, Sampit District, Central Kalimantan Province.

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Island (including Kalimantan, and Sabah). This (Compagno, 1988; 2002; Manjaji, 2002a). Other specimens is also the fi rst and the only record of this genus found in of this genus were from the Kinabatangan River in East coastal waters of this region. The salinity of Sampit Bay Sabah in 1997 and followed by several fi ndings thereafter varied between 5 ppt in the river mouth and around 20 ppt (Manjaji, 2002a). Later, Compagno et al. (2005) referred to in the bay area (Nursyarifuddin, pers. com.). The previous all specimens from Borneo as Borneo River Shark, Glyphis specimens found from Borneo were known from riverine sp. B (Fig. 7). habitats but also suggests its occurrence in brackish water. The first finding of the genus Glyphis in Borneo was The characteristics of Glyphis sp. B, according to Compagno collected from an unknown location about a century ago et al. (2005), are having broadly rounded short snout, and it is deposited in the Naturhistorisches Museum, Austria

Fig. 2. Lateral view of Glyphis sp. (mature male, 1,660 mm total length) from Sampit bay. Fig. 5. Upper and lower jaws of Glyphis sp. (mature male, 1,660 mm total length).

Fig. 3. Dorsal view of Glyphis sp. (mature male, 1,660 mm total length), indicating absence of the interdorsal ridge, and longitudinally-shaped precaudal pit (arrow).

Fig. 6. Dried fi rst and second dorsal fi ns of Glyphis sp. (mature male, 1,660 mm total length).

Fig. 7. Lateral view of Glyphis sp. B (young female, 595 mm total length) from Kinabatangan River, Sabah. An unregistered collection Fig. 4. Ventral view of the mouth of Glyphis sp. (mature male, of Universiti Malaysia Sabah, collected on 30 Apr.2003 (Photo: 1,660 mm total length). K. Jensen and J.N. Caira)

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absence of an interdorsal ridge, fi rst dorsal origin over rear ACKNOWLEDGEMENTS third of pectoral-fi n bases, second dorsal slightly more than half or two-thirds of fi rst dorsal in height, and anal-fi n with We thank to Dr. M. Kasim Moosa of Coral Reef Management deeply notched posterior margin. In general, the primary Program, Indonesian Institutes of Sciences (COREMAP- characteristics of Glyphis sp. B are similar to Glyphis from LIPI) and Dr. Peter Last (CSIRO Hobart) for their reviews Sampit. However, the fi rst dorsal fi n shape of Glyphis sp. B of the manuscript. We also thank to Dr. Ian Tibbets of the seemed to be lower and broader than the Sampit specimen (see University of Queensland Australia and Dr. William White of Figs. 6 & 7), and the size of second dorsal-fi n of specimen CSIRO Hobart for their comments on this species. Our thanks from Sampit was smaller (0.40 of fi rst dorsal-fi n) comparing also to Dr. J. N. Caira of the University of Connecticut and to Glyphis sp. B (0.50–0.67 of fi rst dorsal-fi n). Moreover, Dr. K. Jensen of the University of Kansas for their permission teeth from the lower jaw of Sampit specimen were more of using their Glyphis sp. B image. Mr. Priyo Agustono erect with unserrated cutting edges comparing to Glyphis (RCO-LIPI), Ir. Tenny Apriliani (PRPP&SEKP-DKP) and sp. B. The shape of its lower teeth was more similar to the Ir. Noorsyarifuddin M.Si of the Palangkaraya University, lower teeth of G. glyphis, but this species differs from G. for their assistance in the fi eld. This study was funded by glyphis in the height of second dorsal-fi n (see Compagno the Indonesian Government through Census of Marine Life et al. 2005; 2008). The above-mentioned morphological Program (Project number: 110.14.06/SK/KP/2004). characters have shown that the specimen from Sampit is probably not conspecifi c with either Glyphis sp. B or G. glyphis. Hence, the species from Kalimantan, Indonesia LITERATURE CITED could be a different species. Unfortunately, due to its large size and lack of preservation facilities, the Sampit specimen Compagno, L. J. V., 1997. Threatened fishes of the world: was not collected and therefore there is presently no way to Glyphis gangeticus (Müller & Henle, 1839) (Carcharhinidae). confi rm the distinctiveness of this species. Environmental Biology of Fishes 49: 400. Compagno, L. J. V., 1988. Sharks of the Order Carcharhiniformes. Nevertheless, the fi nding of this species near the coast in The Blackburn Press, New Jersey. 486 pp. Sampit Bay, Ujung Pandaran Village, Central Kalimantan, Compagno, L. J. V., 1998. Sharks. In; K. E. Carpenter & V. extends the distribution of Glyphis in Borneo and this is H. Niem (eds.), FAO Species identifi cation guide for fi shery the fi rst record of its presence within Indonesian territory. purposes. The living marine resources of the western central Moreover, this fi nding is also evidence that Glyphis from Pacifi c. Cephalopods, crustaceans, holothurians and sharks. Borneo can inhabit coastal and marine habitats. The presence Vol. 2. FAO, Rome. Pp. 1193Ð1366. of this presumed rare species in Sampit Bay is probably Compagno, L. J. V., 1999. Checklist of living elasmobranchs. In: due to the low fi shing pressure at coastal waters in Central W. C. Hamlett (ed.), Sharks, skates, and rays: the biology of Kalimantan, especially in Sampit Bay. The relatively long elasmobranch fi shes. The Johns Hopkins University Press, distance from the nearby city, lack of transportation facilities Baltimore. Pp. 471Ð498. and lack of electricity in the villages along the Sampit Bay Compagno, L. J. V., 2002. Freshwater and estuarine elasmobranch are some factors which have led to low pressure of fi shing surveys in the Indo-Pacifi c region: threats, distribution and and may constrain fi sheries development in this area. speciation. In: S.L. Fowler, T.M. Reed & F.A. Dipper, (eds.), Elasmobranch biodiversity, conservation and management: Proceeding of the International Seminar and Workshop in Sabah, July 1997. IUCN SSC Shark Specialist Group: Gland, CONCLUSION Switzerland and Cambridge, UK. Pp. 168Ð180. Compagno, L. J. V., M. Dando & S. L. Fowler, 2005. Sharks of The fi nding of a Glyphis species at Ujung Pandaran Village, the world. Princeton University Press, New Jersey. 368 pp. Central Kalimantan, is the first record of the genus in Indonesian territory. This record also revealed that members Compagno, L. J. V., W. T. White & P. R. Last, 2008. Glyphis garricki sp. nov., a new species of river shark (Carcharhiniformes: of the genus from Borneo inhabit marine as well as freshwater Carcharhinidae) from northern Australia and Papua New habitats. Moreover, the specimen could belong to a new Guinea, with description of Glyphis glyphis (Müller & Henle, species. This fi nding also revealed that some unexploited 1839). In: P. R. Last, W. T. White & J. J. Pogonoski (eds.), areas in Indonesia, i.e. Sampit Bay, should be maintained Descriptions of new Australian chondrichthyans. CSIRO Marine or further protected, as areas such as these are under low and Atmospheric Research Paper No. 022. CSIRO, Australia. fi shing pressures. The authors realized that due to the lack of Pp. 203Ð225. information on the Indonesian biodiversity, some rare fauna Erdmann, M. V., R. L. Caldwell & M. K. Moosa, 1998. Indonesian could become either extinct or overexploited before they ‘king of the sea’ discovered. Nature, 385(6700): 335. are documented by scientists. An example is the Indonesian Fahmi & M. Adrim, 2007. Elasmobranch diversity of Kalimantan Coelacanth, in which the local community had already waters. Marine Research in Indonesia, 32(2): 97Ð105. known about an “oily grouper” for several decades, but it Last, P. R., 2002. Freshwater and estuarine elasmobranchs of was only scientifi cally described in 1998 (Erdmann et al., Australia. In: S. L. Fowler, T. M. Reed & F. A. Dipper, (eds.), 1998). Therefore, all information on biodiversity covering Elasmobranch biodiversity, conservation and management: marine and freshwater habitats, including the freshwater Proceeding of the International Seminar and Workshop in shark, will contribute to the basic knowledge on their biology Sabah, July 1997. IUCN SSC Shark Specialist Group: Gland, and habitat to support conservation activities. Switzerland and Cambridge, UK. Pp. 185Ð193.

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Last, P. R. & J. D. Stevens, 1994. Sharks and Rays of Australia. Manjaji, B. M, 2002b. New records of elasmobranch species from CSIRO, Australia. 513 pp. Sabah. In: S. L. Fowler, T. M. Reed & F. A. Dipper (eds.), Manjaji, B. M, 2002a. Elasmobranchs recorded from rivers and Elasmobranch biodiversity, conservation and management: estuaries in Sabah. In: S. L. Fowler, T. M. Reed & F. A. Proceeding of the International Seminar and Workshop in Dipper, (eds.), Elasmobranch biodiversity, conservation and Sabah, July 1997. IUCN SSC Shark Specialist Group: Gland, management: Proceeding of the International Seminar and Switzerland and Cambridge, UK. Pp. 70Ð76. Workshop in Sabah, July 1997. IUCN SSC Shark Specialist White, W. T., P. R. Last, J. D. Stevens, G. K. Yearsley, Fahmi & Group: Gland, Switzerland and Cambridge, UK. Pp. 194Ð Dharmadi, 2006. Economically important sharks and rays of 198. Indonesia. ACIAR, Canberra. 329 pp.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 119–125 Date of Publication: 28 Feb.2009 © National University of Singapore

NEMACHEILUS TEBO, A NEW LOACH FROM SANGKULIRANG KARST, EAST KALIMANTAN, INDONESIA (TELEOSTEI: NEMACHEILIDAE)

Renny Kurnia Hadiaty Museum Zoologicum Bogoriense (MZB), Ichthyology Laboratory, Zoology Division of Research Center for Biology, Indonesian Institute of Sciences (LIPI), Jl. Raya Bogor Km 46, Cibinong 16911, Indonesia. Email: [email protected]

Maurice Kottelat Route de la Baroche 12, Case Postale 57, CH-2952 Cornol, Switzerland (permanent address); and Raffl es Museum of Biodiversity Research, National University of Singapore, Kent Ridge, Singapore 119260. Email: [email protected]

ABSTRACT. – Nemacheilus tebo, new species, is described from Lake Tebo drainage, in the Sangkulirang peninsula karst formation, East Kalimantan, Indonesia. It is distinguished from all other species of Nemacheilus by its unique colour pattern: fl ank with 11–16 bars, those in front of dorsal fi n short, close together or fused, forming a kind of large elongate blotch; those below and behind dorsal fi n distinct and usually not continuous across dorsum.

KEY WORDS. – Nemacheilus, Sangkulirang, limestone, East Kalimantan, Indonesia.

INTRODUCTION COLLECTION SITES

The nemacheiline loaches of the Sundaic area were revised Nemacheilus tebo, was collected in Danau Tebo area (danau by Kottelat (1984) and nine species of the genus Nemacheilus means lake in Indonesian). The Tebo limestone is the most recorded in Indonesia, Malaysia and Singapore. Two species isolated hill among the investigated stations (Tabalar, Baai, have been described since, N. elegantissimus from Danum Marang). The expedition team reached this site by helicopter. Valley, Sabah (Chin & Samat, 1992) and N. tuberigum from According to the local people, overland access from the Aceh, Sumatra (Hadiaty & Siebert, 2001). Two species nearest village (Merapun) takes about a three day walk. The earlier treated as synonyms (N. pfeifferae, N. longipinnis) southern part of Tebo burned once during the 1997 fi res, have been re-validated by Kottelat et al. (1993), bringing the number of valid species from this region to 13.

The Sangkulirang Peninsula is a section of 1,000 km2 of karstic formations in East Kalimantan Province, Indonesia, on the border of the districts Kutai Timur and Berau (Fig. 1). This area has been highlighted as a global priority for karst ecosystem conservation. An expedition in Sangkulirang Peninsula coordinated by The Nature Conservancy and the Indonesian Institute of Sciences conducted a 5 weeks biological survey in Jul.—Aug.2004 at four sites, Suatan/ Tabalar, Baai, Marang and Tebo. Several new plant and species were discovered. We describe here a new fi sh species from the Tebo area, Nemacheilus tebo.

Fig. 1. Sangkulirang Peninsula, with the four limestone formations (Tabalar, Baai, Marang, Tebo) surveyed by The Nature Conservancy and the Research Center for Biology-Indonesian Institute of Sciences (LIPI).

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but the area is so remote that other impacts (illegal logging Material for this study is deposited in the following and hunting) are minimal; it currently retains large patches collections: Academy of Natural Sciences, Philadelphia of primary forest (Salas et al., 2005). (ANSP); Natural History Museum, London (BMNH); Collection of second author (CMK); Museum National Nemacheilus tebo was collected at 6 sites in the Danau d’Histoire Naturelle, Paris (MNHN); Museum Zoologicum Tebo drainage (Fig. 2): 1) entrance of cave northwest of Bogoriense, Bogor (MZB); Nationaal Natuurhistorisch camp (17°00'16"N 54°11'59"E); 2) entrance of second Museum, Leiden (RMNH); Florida Museum of Natural cave northwest of camp (17°00'13"N 54°10'54"E); 3) History, University of Florida, Gainesville (UF); United entrance of cave behind camp (16°56'08"N 54°07'35"E); States National Museum, Washington (USNM); Zoologisch 4) stream Tuba-tubaan inside Gua Keluar (gua means cave Museum, Universiteit van Amsterdam, Amsterdam (ZMA); in Indonesian) (16°41'01"N 54°04'43"E); 5) stream Tuba- and Zoological Reference Collections, National University tubaan inside Gua Masuk (16°41'45"N 54°05'23"E); and 6) of Singapore (ZRC). stream Tuba-tubaan between Gua Keluar and Gua Masuk (16°41'20"N 54°05'02"E) at confl uence with a small inlet. Fish were collected with a 12 V electric fi sh shocker. It was SYSTEMATICS not possible to capture fi shes in Danau Tebo itself due to the lack of appropriate equipment, and because the banks Nemacheilus tebo, new species of the lake are very steep. The other fi sh species collected (Fig. 3) from this drainage were Anguilla marmorata (Anguillidae), Cyclocheilichthys sp., Osteochilus sp. 1, Osteochilus sp. Holotype. – MZB 13367, 56.1 mm SL (70.8 mm TL); Indonesia: 2, Rasbora cf. hubbsi (Cyprinidae), Pangio anguillaris Kalimantan Timur: Berau Regency, Kelai District, Merapun Village, (Cobitidae), Hemibagrus cf. nemurus (Bagridae), Channa Lake Tebo area, a pond at mouth of west cave, site 2 (17°00'13"N gachua (Channidae) and Clarias sp. (Clariidae). 54°10'54"E); coll. R. Hadiaty, Sokir & Cai, 31 Aug.2004. Paratypes. – All from Indonesia: Kalimantan Timur: Berau Regency, Kelai District, Merapun Village, Lake Tebo area. BMNH MATERIALS AND METHODS 2007.11.30.1, 1, 43.8 mm SL (58.4 mm TL); CMK 18909, 3, 35.9–45.0 mm SL (46.3–56.2 mm TL); MZB 13380, 2, 37.5–41.1 Measurements were made from point to point on the left side mm SL (48.8–52.5 mm TL); UF 165707, 1, 38.6 mm SL (49.8 of the body whenever possible with a digital caliper and the mm TL); USNM 388744, 2, 40.4–40.4 mm SL (52.6–53.8 mm data recorded to tenth of a millimeter. The measurements TL); ANSP 187006, 1, 42 mm SL (55.1 mm TL); same data as and counts follow Kottelat (1990). holotype. - MZB 13359, 3, 36.1–40.7 mm SL (47.7–52.2 mm TL); cave behind camp, site 3 (16°56'08"N 54°07'35"E); coll. R. Hadiaty, Sokir & Cai, 30 Aug.2004. - MZB 13360, 4, 37.1–44.9 mm SL (48.3–56.4 mm TL); ZRC 50728, 1, 40.6 mm SL (51.1 mm TL); entrance of cave northwest of camp, site 1 (17°00'16"N 54°11'59"E); coll. R. Hadiaty, Sokir & Cai, 31 Aug.2004.

Additional material (non types). – MZB 13341, 43, 18.8–48.2 mm SL (23.6–62.6 mm TL): Tuba-tubaan river, site 6 (16°41'20"N 54°05'02"E); coll. R. Hadiaty, Sokir & Cai, 30 Aug.2004. - MZB 13346, 2, 37.5–41.6 mm SL (48.0–54.2 mm TL); Gua Keluar, site 4 (16°41'01"N 54°04'43"E); coll. R. Hadiaty, Sokir & Cai, 30 Aug.2004. - MZB 13379, 1, 31.8 mm SL (42.3 mm TL); Lake Tebo (16º49'54"N 54º07'43"E); coll. R. Hadiaty, 28 Aug.2004.

Diagnosis. – Nemacheilus tebo belongs to the N. selangoricus group, diagnosed by the presence of longitudinal rows of elongated scales on the caudal peduncle, each scale with a tubercle at posterior extremity (Fig. 4a). It is distinguished from all other species of Nemacheilus by its unique colour pattern: fl ank with 11–16 dark brown bars, those in front of the dorsal fi n short, close together or fused, forming a kind

Fig. 2. Lake Tebo drainage. Nemacheilus tebo, was collected at sites 1–3 at the mouth of the caves, at sites 4 (Gua Keluar) and 5 (Gua Masuk) in a small stream inside the cave, and at site 6 on the same stream at the confl uence with a small inlet. Black: Lake Tebo; white: limestone cliffs; pale gray: valley bottom; dark grey: Fig. 3. Nemacheilus tebo, MZB 13367, holotype, 56.1 mm SL; karst uplands and (in valley) remnants of pinnacles. Indonesia: Kalimantan Timur: Lake Tebo area, site 2.

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of large elongate dark brown blotch; those below and behind Body completely covered by small, deeply embedded scales. the dorsal fi n distinct, usually continuous across dorsum, On a clear and stained specimen, scales on ventral part of irregularly shaped, usually wider along the dorsal midline and head and belly scattered, not embedded as on dorsum and along lateral line, narrower in-between. Nemacheilus tebo is fl ank. Three to fi ve rows of enlarged scales on fl ank, from distinguished from the other species of the N. selangoricus pectoral-fi n base to above pelvic fi n base. One row above group by having smaller tubercles, and by the presence (vs. lateral line, other rows below lateral line, 2–4 rows under absence) of rows of tubercles on the lower half of the fl ank. pectoral fi n, 5–6 rows above pelvic fi n base, 2–3 rows before Further, it is distinguished from N. selangoricus and N. end of pelvic-fi n base. Each scale bearing a small tubercle spiniferus in missing the long, acuminate posterior projection on posterior extremity. On caudal peduncle, 7–10 enlarged on the elongated scales on the caudal peduncle (vs. presence, scales on one row above and one row below lateral line, each with the tubercle located at the tip of the projection) and with a tubercle at posterior extremity (Fig. 4a, b). Largest by the absence of the black spot at the base of the anterior specimen (holotype, 56.1 mm SL) with a single tubercle on dorsal-fi n rays and of the rows of black spot on the rays lateral line scales on caudal peduncle. (vs. presence). Morphometric characters distinguishing N. tebo from N. selangoricus and N. tuberigum are mentioned Lateral line complete, reaching to caudal-fi n base, with under Discussion. 80–85 pored scales. Cephalic lateral line system with 6 supraorbital, 4 + 9–10 infraorbital, 7 preoperculo-mandibular Description. – See Fig. 3 for general appearance and Table and 3 supratemporal pores. 1 for morphometric and meristic data of holotype and 17 paratypes. An elongate nemacheiline with body depth Anterior nostril at tip of a short pointed tube. Mouth gape 11.3–14.8 % SL, slowly increasing from head up to dorsal- about 1.7 times wider than long. Lips thin but fl eshy, with fi n origin. Behind dorsal fi n, body depth decreasing slowly about 5 folds on lower lip (Fig. 5b). A median incision in to caudal-fi n base. Head slightly depressed; body slightly upper lip. A median interruption in lower lip. Processus depressed anteriorly to compressed posteriorly. Pectoral fi n dentiformis present. A median notch in lower jaw. Inner reaching about halfway to pelvic-fi n base. Axillary pelvic rostral barbel reaching to vertical of middle of eye; outer lobe present. Pelvic fi n reaching about halfway of distance one reaching slightly behind preopercle. Maxillary barbel to anal-fi n origin, just reaching anus, which is about one eye reaching to middle of opercle. diameter in front of anal fi n. Origin of anal fi n slightly in front of vertical through dorsal-fi n origin. Caudal fi n forked. Colour pattern. – In life: body background beige, with 11–16 Low ventral and dorsal adipose crests on posterior half of dark olive green bars on fl ank. Bars in front of dorsal fi n caudal peduncle, which is 1.3–2.0 times longer than deep. short, not very distinct, close together or fused, forming a Distal margin of dorsal fi n straight. Largest recorded size kind of large elongate blotch under pectoral fi n. Below and 56.1 mm SL, 70.8 mm TL. behind dorsal fi n, bars distinct, usually continuous across dorsum, irregularly shaped, usually wider along dorsal Dorsal fi n with 4 simple and 9–10½ branched rays. Anal fi n midline and along lateral line, narrower in-between, and with 3 simple and 5½ branched rays. Caudal fi n with 9+8 vertical median area slightly paler than margins. Ventral branched rays. Pectoral fi n with 12 rays; anterior rays often part of body creamy. A conspicuous elliptical black blotch with a row of small tubercle; in a 44.1 mm SL specimen, on lower half of caudal-fi n base, not reaching ventral mid- tubercles on fi rst ray about 30 µm wide and 20 µm high line. A narrow dark olive green bar below middle of eye. (Fig. 4c). Pelvic fi n with 8 rays. Preserved specimens: dark brown bars and dorsum, whitish on belly and between bars. Vertebrae counts: 23+12=35 (2), 24+12=36 (1), predorsal vertebrae 11 (2), 12 (1).

Fig. 4. Nemacheilus tebo: a–b. scanning electron micrographs of caudal peduncle of 34.3 mm SL specimen: a, showing elongated scales above and below lateral line; b, close-up view of tubercle; c, scanning electron micrograph of tubercle on fi rst pectoral-fi n ray of 44.1 mm SL specimen. Scale bars = 10 µm

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Table 1. Morphometric and meristic data of holotype (MZB 13367) and 17 paratypes of Nemacheilus tebo.

Holotype Paratypes Mean Standard Deviation Standard length (mm) 56.1 35.9–45.0 – – Total length (mm) 70.8 46.3–58.4 In percent of standard length Total length 126.2 124.9–133.3 129.8 2.6 Lateral length of head 24.1 21.9–26.5 24.1 1.1 Dorsal length of head 18.4 17.2–21.1 18.8 1.1 Predorsal length 48.5 43.9–49.6 47.0 1.5 Prepelvic length 52.9 48.8–52.4 50.6 0.9 Preanal length 77.5 72.2–78.8 75.7 1.7 Preanus length 69.0 64.2–69.6 66.7 1.4 Head height (at eye) 10.9 9.4–11.7 10.5 0.6 Head height (at nape) 13.4 11.4–14.0 12.3 0.7 Body height (at dorsal origin) 13.7 11.3–14.8 13.2 0.8 Caudal peduncle depth 10.3 8.8–10.0 9.4 0.4 Length of caudal peduncle 16.2 13.1–18.1 16.4 1.4 Snout length 8.7 7.1–8.6 7.9 0.5 Head width at nares 7.3 6.1–8.3 7.3 0.5 Maximal head width 14.4 12.5–14.8 13.4 0.7 Body width (at dorsal–fi n origin) 11.4 7.1–10.9 8.9 0.8 Body width (at anal–fi n origin) 7.3 5.0–6.7 5.8 0.5 Eye diameter 5.5 4.6–6.4 5.7 0.5 Height of dorsal fi n 23.5 20.9–26.9 24.6 1.6 Length of upper caudal fi n lobe 25.3 27.3–35.6 31.5 2.3 Length of lower caudal fi n lobe 23.7 19.0–30.7 27.1 2.5 Length of middle caudal fi n rays 16.2 15.3–30.3 18.7 3.4 Length of anal fi n 18.2 15.9–21.4 19.2 1.2 Length of pelvic fi n 13.9 15.1–19.4 17.0 1.1 Length of pectoral fi n 19.4 18.9–27.4 23.8 2.0 In percent of dorsal HL Lateral length of head 131 118–139 128.4 7.0 Head height (at eye) 59 51–60 55.7 2.7 Head height (at nape) 73 59–78 65.7 4.5 Body height (at dorsal origin) 75 63–82 70.3 5.8 Caudal peduncle depth 56 43–57 50.3 3.7 Length of caudal peduncle 88 66–103 87.5 10.2 Snout length 48 37–49 41.9 2.8 Head width at nares 40 32–46 38.8 3.8 Maximal Headwidth 79 65–78 71.1 3.7 Body width (at dorsal origin) 62 38–57 47.3 4.4 Body width (at anal origin) 40 25–36 30.7 3.2 Eye diameter 30 26–36 30.1 2.7 Interorbital width 33 26–42 31.9 4.3

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Fin membranes hyaline, rays dusky. Dorsal fi n with a short small fi sh dying were also observed there. Some of them faint mark on lower third of last simple ray. were not able to swim, and were preyed upon by swarming insects (water beetles, Dysticidae). Sexual dimorphism. – Male with suborbital fl ap (Fig. 5a) and much more tubercles on fl ank (from pectoral-fi n base to The Tuba-tubaan is named so because some of local peoples above pelvic fi n base) than female. In the largest specimen (who harvest edible ’ nest inside the caves surrounding (holotype) the fl ap is reduced in size. the lake) used tuba (Derris sp., Fabaceae), a plant from which is extracted rotenone, an effective ichthyocide. The stream Distribution and habitat. – Nemacheilus tebo is currently and the lake are the only water sources available for most known only from Lake Tebo drainage, Kalimantan Timur, of the local vertebrates. Indonesia. Lake Tebo area was densely covered by big trees, which made the area very green and lush, but the area was Etymology – Named after the Lake Tebo. A noun in burnt in 1997’s Borneo fi res. The expedition was conducted apposition. during the dry season and the area was very dry. The water level of Lake Tebo was low (depth unknown, reported by locals to be “more than 5 meters”; Fig. 6a) and we could DISCUSSION fi nd only few water bodies inhabited by fi shes, usually in front of the entrance of caves. These apparently are the Nemacheilus tebo belongs to a group of species, which normal dry season conditions, especially after burning. In we call here the N. selangoricus group (comprising N. the rainy season, the water level increases and covers all selangoricus, N. spiniferus and N. tuberigum). The group of the valley area. is characterised by the presence of two rows of elongate scales on the caudal peduncle, one above and one below the All the specimens of N. tebo collected in Tuba-tubaan stream lateral line, each of them bearing a tubercle near the posterior were very emaciated. The stream had very little running extremity. These scales were called acuminate scales by water, the depth was only about 10–20 cm and the width Kottelat (1984, 1990) in N. selangoricus and N. spiniferus, about 1–2 m (Fig. 6b). The fi shes were usually observed in to refer to the long posterior projection (acumen) at whose small groups of about 1–20 individuals in shallow water, but tip the tubercle is located. In N. tebo and N. tuberigum in deeper spots several thousands of fi sh trapped by the low waters formed large dark masses. In both Tuba-tubaan and Lake Tebo we observed many fi sh dying. In Lake Tebo, the fi shes were very crowded in the drying water bodies, and

Fig. 5. Nemacheilus tebo: a, CMK 18909, paratype, male, 45.0 Fig. 6. a. Lake Tebo; b. Tuba-tubaan river, a small stream fl owing mm SL; head showing suborbital fl ap; b, MZB 13367, holotype, out of Gua Keluar (‘Exit Cave’) and entering Gua Masuk (‘Entrance 56.1 mm SL; mouth, ventral view. Cave’; on picture).

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these scales are elongated, with a broad and short extension had to be escorted by policemen. These collectors partly along the posterior edge, but without the acumen, but the rely on local resources for their subsistence including fi sh, structure is clearly homologous. The members of the N. which they obtain by traditional tools (nets, traps) as well selangoricus group are further distinguished by their colour as ichthyocides like Derris roots or potassium. The use of pattern made of numerous dark bars on the body, while poisons affects all fi sh indiscriminately, even those too small most other species of the genus have a colour pattern made to be consumed, and present a very high threat to the local of a more or less regular stripe, or a row of blotches on fi sh populations. the fl ank, usually alternating with saddles along the dorsal mid-line. Nemacheilus tebo is distinguished from all other Comparative material. – Nemacheilus selangoricus: MZB 3551, species of the N. selangoricus group as well of the genus 3, 28.3–29.4 mm SL; Indonesia, Kalimantan Barat, Sungai Kapuas Nemacheilus by its unique colour pattern of many narrow basin, small forest stream fl ows to Sungai Mandai upstream from its bars on the fl ank, those in front of dorsal-fi n origin not confl uence with Kapuas mainstream; T. R. Roberts, 10 Aug.1976. MZB 2395, 3, 29.3–41.4 mm SL; Indonesia, Kalimantan, Lempake, reaching the dorsal midline, and set very closely or fused, Tanah Merah, M. Siluba; 27 Feb.1978. RMNH 28879, 5, 41.1–44.1 forming an elongated blotch. mm SL; Indonesia, Borneo, Sungai Mandai Kecil, 18 km south west of Putussibau; T. R. Roberts, 11 Aug.1975. ZMA 112.829, lectotype Nemacheilus tebo and N. tuberigum are distinguished from of N. kuiperi, 50.2 mm SL; ZMA 102.145, 21 paralectotypes of N. N. selangoricus and N. spiniferus by the absence of the kuiperi, 28.2–65.8 mm SL; Indonesia, Biliton; F. J. Kuiper, 1936. black spot at the base of the anterior dorsal-fi n rays and of RMNH 24995, 1, 47.1 mm SL; Singapore, Sungei Seletar, north the rows of black spot on the rays. of Seletar reservoir; E. R. Alfred, 4 Apr.1963.

Compared to the other three species of the N. selangoricus Nemacheilus spiniferus: all from Indonesia, Kalimantan Tengah, MZB 6807, 6, 32.2–37.5 mm SL; Sungai Tarusan, a tributary of group, in N. tebo the tubercles are much smaller and can be S. Laung, a tributary of Sungai Barito; D. J. Siebert et al., 16 observed only with magnifi cation, while in the others the Jul.1992. MZB 6877, 11, 29.5–38.0 mm SL; Sungai Karingian, tubercles are visible without magnifi cation, even in the live a tributary of Sungai Laung, a tributary of Sungai Barito; D. J. fi sh. A tubercle on the caudal peduncle of a 34.3 mm SL Siebert et al., 7 Jul.1992. MZB 6928, 2, 38.7–40.0 mm SL; Sungai specimen is about 40 µm wide and about 35 µm high (Fig. Laung, a tributary of Sungai Barito, Laung Tuhup, Barito Utara; 4b). In N. tebo there are also 2–5 rows of elongated scales D. J. Siebert et al., 15–18 Jul.1992. MZB 6948, 2, 34.1–34.5 mm on the lower half of the fl ank from behind the pectoral fi n SL; Sungai Mata, a tributary of Sungai Barito below Muara Laung, to the caudal-fi n base; in the other species elongated scales Laung Tuhup, Barito Utara; D. J. Siebert et al., 8 Jul.1992. are present only on the caudal peduncle. There are small Nemacheilus tuberigum: all from Indonesia, Sumatra, Aceh. MZB tubercles at the posterior extremity of these elongated scales 9356, holotype, 48.5 mm SL; MZB 10565, 1 paratype, 43.0 mm SL; on the fl ank; they are much smaller than the tubercles in MZB 9357, 12 paratypes, 39.6–53.2 mm SL; BMNH 2000.4.10.1–5, similar position in the other species (not quantifi ed). 5 paratypes, 42.2–50.5 mm SL; Kecamatan Kluet Selatan, Desa Pucuk Lembang, Gunung Leuser National Park, tributary of Sungai Besides the characters mentioned above, N. tebo is Lembang; R. K. Hadiaty & A. Mun’im, 2 Sep.1997. MZB 9358, 4 distinguished from N. tuberigum by a shorter snout (length paratypes, 44.8–53.4 mm SL; same dataas holotype; 31 Aug.1997. 37–49 % HL, vs. 47–55), a smaller eye (diameter 26–35 % MZB 9359, 1 paratypes, 42.6 mm SL; same data as holotype, HL, vs. 38–46), a shorter predorsal length (43.9–49.6 % SL, 1 Sep.1997. MZB 9360, 4 paratypes, 42.6–49.2 mm SL; Desa vs. 47.4–51.2), and a more slender caudal peduncle (length Pucuk Lembang, Alur Betung, a tributary of Sungai Lembang; R. K. Hadiaty & A. Mun’im, 2 Sep.1997. MZB 9361, 2 paratypes, 13.1–18.1 % SL, vs. 12.0–17.3; depth 8.8–10.0 % SL, vs. 31.9–37.2 mm SL; Suaq Balimbing Research Station, a muddy 10.2–11.9; ratio length/depth 1.3–2.0, vs. 1.1–1.5). forest stream, tributary of Sungai Lembang; R. K. Hadiaty & A. Mun’im, 4 Sep.1997. ZMA 112.875. 1, 49.1 mm SL; ZMA 113.744, Nemacheilus tebo is distinguished from N. selangoricus 4, 34.3–48.7 mm SL; Pageralem; P. A. Ouwens, 22 Nov.1918. by a slightly shorter predorsal length (43.9-49.6 % SL, vs. ZMA 116.645, 1, 46.7 mm SL; no collector and date. 45.7–53.8), and a more slender caudal peduncle (length 13.1–18.1 % SL, vs. 13.2–16.3; depth 8.8–10.0 % SL, vs. Nemacheilus fasciatus: MNHN B 2798, holotype, 54.1 mm SL; 10.0–11.7; ratio length/depth 1.3–2.0, vs. 1.1–1.5). Indonesia, Java; Kuhl & van Hasselt. MNHN 3930, holotype of Cobitis suborbitalis, 59.1 mm SL; Indonesia, Java; no collector and date. ZMA 109.262, 2, 41.9 mm SL, 1 specimen without head; Nemacheilus tebo is known only from Tebo limestone area. Indonesia, Java, Yogya, stream in Gremeng cave, Gunung Sewu; There is no information on the fi sh fauna of the adjacent E. Jacobson, Feb.1911. water bodies and it is not known whether N. tebo is endemic to Lake Tebo area, or might occur outside. The subterranean Nemacheilus chrysolaimos as above: MNHN 3961, lectotype, 1, course of the Tuba-tubaan stream is not known but local 46.7 mm SL; Indonesia, Java; coll. Kuhl & van Hasselt, no date. people report that they threw small oranges in the cave and MNHN B 2972, paralectotype, 1, 46.7 mm SL; Indonesia, Java; that they re-appeared in a stream near the village. Kuhl & van Hasselt, no date.

The lake is quite far from the village, but ’s nest collectors Nemacheilus longipectoralis: RMNH 7641, lectotype, 1, 34.4 mm SL; RMNH 27350, 2 paralectotypes, 28.9–36.7 mm SL; Indonesia, frequently come to this area and there had been occasional Borneo, Mahakam; A. W. Nieuwenhuis, Nov.1898. clashes between different groups, to the point that our team

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ACKNOWLEDGMENTS LITERATURE CITED

The fi rst author thanks the following: The Nature Conservancy Hadiaty, R. K. & D. J. Siebert, 2001. Nemacheilus tuberigum, for funding the fi eldwork in Sangkulirang Peninsula limestone a new species of loach (Teleostei: Balitoridae) from Aceh, area and the use of maps; especially Leo A Salas, expedition northwestern Sumatra, Indonesia. Bulletin Natural History coordinator, for various information and support; the Research Museum London (Zoology), 67(2): 183–189. Center for Biology, LIPI, for permission to participate to Chin, P. K & A. Samat, 1992. A new species of loach, Nemacheilus the expedition; the other team members, Woro Noerdjito, elegantissimus, (Family Balitoridae, subfamily Nemacheilinae) M. Noerdjito, Y. R. Suhardjono, C. Rachmadi, Rofi k, H. from Danum Valley, Sabah, Malaysia. Malayan Nature Journal 46: 25–33. Wiriadinata, for the nice cooperation on site; Tan Heok Hui and Ng Heok Hee (ZRC) for the x-rays and Tan Heok Hui Kottelat, M. 1984. Revision of the Indonesian and Malaysian for photographs in Fig. 5. RH visited European museums noemacheiline loaches. Japanese Journal of Ichthyology, 31: with the support of the All Catfi sh Species Inventory project 115–260. (principal investigator Larry M Page). Ralf Britz (BMNH), Kottelat, M. 1990. Indochinese nemacheilines: A revision of Martien van Oijen (RMNH), Ronald Vonk (ZMA), and Guy nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Duhamel (MNHN) allowed access to material under their Burma, Laos, Cambodia and southern Vietnam. Pfeil, München. 262 pp. care. RH visited the Smithsonian Institution with support of a grant (Schultz Funds) managed by Lynne R Parenti. Kottelat, M., A. J. Whitten, S. N. Kartikasari & S. Wirjoatmodjo, At USNM, RH benefi tted from the help of Scott Whittaker 1993. Freshwater fi shes of western Indonesia and Sulawesi. Periplus, Hong Kong. 293 pp. (SEM manager) and D. Lumbantobing. This work was completed while RH was a RMBR SPRINT visitor and MK Salas, L. A., A. Bedos, L. Deharveng, S. Fryer, R. Hadiaty, Heryanto, a Visiting Senior Research Fellow at the National University Munandar, Nardiyono, M. Noerdjito, W. Noerdjito, C. Rahmadi, A. Riyanto, Rofi k, A. Ruskandi, M. J. Struebig, Y. Suhardjono, of Singapore. A. Suyanto, J. J. Vermeulen, C. Walck, H. Wiriadinata, E. Meijaard & S. Stanley, 2005. Biodiversity, endemism and the conservation of limestone karst in the Sangkulirang Peninsula, Borneo. Biodiversity, 6(2): 12–23.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 127–181 Date of Publication: 28 Feb.2009 © National University of Singapore

REVIEW OF THE GOBIID FISH GENERA EUGNATHOGOBIUS AND PSEUDOGOBIOPSIS (GOBIOIDEI: GOBIIDAE: GOBIONELLINAE), WITH DESCRIPTIONS OF THREE NEW SPECIES

Helen K. Larson Museum and Art Gallery of the Northern Territory P.O. Box 4646, Darwin, NT 0801, Australia

ABSTRACT. – The gobiid fi sh genera Eugnathogobius and Pseudogobiopsis are reviewed and redescribed. Species of these genera are known from shallow estuarine and freshwater habitats in east Africa, the Indo- Malay Archipelago, northern Australia, southern China and Taiwan. Two new species of Eugnathogobius are described, one from Mozambique, Kenya and Seychelles, and one from northern Australia. Mugilogobius polylepis Wu & Ni is placed in Eugnathogobius and redescribed. Pseudogobiopsis includes four species, most of which had previously been placed in several other genera. One new species is described, from Malaysian Borneo, and Gobius tigrellus Nichols is placed in Pseudogobiopsis and redescribed.

KEYWORDS. – Gobioidei, Gobiidae, Gobionellinae, Eugnathogobius, Calamiana, Pseudogobiopsis, new species.

INTRODUCTION the reason for synonymising his genus Pseudogobiopsis with Stigmatogobius. The species usually assigned to the gobiid genera Calamiana, Eugnathogobius and Pseudogobiopsis have been of interest Herre (1945) created Calamiana for a new species (Calamiana to me for some time, as I attempted to discover if they magnoris) that he considered to possess vomerine teeth, formed one or more monophyletic groups, and if so, how mistaking the curved vomer itself for a pair of canine-like many species were included. Many of the species known vomerine teeth. His species C. magnoris also possessed an are represented by few specimens from a range of Indo-west enlarged mouth and “physiognomy peculiar”. Pacifi c and Southeast Asian localities, making identifi cation diffi cult. This paper presents some answers plus some more Smith (1945) considered that Eugnathogobius was related to questions, and I recognise that these may not be the fi nal his genus Gnathogobius (a junior synonym of Calamiana), answers for this group of fi shes. Mahidolia, Pseudogobiopsis and Waitea (the last a synonym of Oligolepis), due to their greatly enlarged jaws. Mahidolia Hugh Smith began it all originally by describing a new genus is a gobiine, while the others are gobionellines (Birdsong et and species of goby, Eugnathogobius microps, from a single al. 1988; Larson 2001). specimen from the lower Bangpakong River, Thailand (Smith, 1931). He characterized the genus by its enormous mouth with The placement of Mugilogobius polylepis Wu & Ni, 1985, elongate maxillary, but did not compare it with any known was uncertain for many years due to insuffi cient material. genus. Koumans (1940) considered that Eugnathogobius This species was described from three specimens: the 34 mm microps was “Probably a good genus and species”. SL holotype (Shanghai Fisheries College S-0001) and two paratypes 22–23 mm SL (Shanghai Fisheries College S-0002, Koumans (1935) created the genus Pseudogobiopsis, to S-0003). The paratypes now appear to be lost in the postal include Gobiopsis oligactis Bleeker, and Gobius römeri system (Wu Han-Lin, pers. comm.). The holotype was made Weber. He stated that Pseudogobiopsis “… in general available for study by I-S. Chen (National Taiwan Ocean appearance is close to Stigmatogobius, but differs in University) and was found to agree with Calamiana. having the maxillary prolonged posteriorly” (Koumans, 1935). Koumans (1953) later placed Pseudogobiopsis Birdsong et al. (1988) placed Calamiana and Pseudogobiopsis oligactis in the genus Stigmatogobius Bleeker, along with in their Gobionellus Group (they also included Gnatholepis, other species now referred to Hemigobius, Redigobius, Mugilogobius, Oligolepis, Oxyurichthys, Pseudogobiopsis, Pseudogobius and Stigmatogobius. Koumans did not discuss Stenogobius and Tamanka). These genera share the characters

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of two epurals, 25–26 vertebrae, dorsal pterygiophore formula specimens with that count, or the range of counts. Vertebral of 3-12210, and having two or three anal pterygiophores counts and other osteological information were obtained by before the first haemal spine. Pezold (1993) assigned radiography and clearing and double-staining. Calamiana and Pseudogobiopsis to the Gobionellinae, but did not mention Eugnathogobius. Cladistic analyses were made using PAUP* v.40b, using both branch and bound and bootstrap methods (branch-and-bound Larson (1999) described a new species of Calamiana and option) (Swofford 2002). Data was acquired from 12 of allocated Apocryptes variegatus Peters, and Vaimosa mindora the 13 nominal species of Calamiana, Eugnathogobius and Herre, to the genus. She retained Calamiana as separate from Pseudogobiopsis. The matrix used is a subset of that in Larson Eugnathogobius, pending further research. (2001), with the addition of data for Pseudogobiopsis tigrellus for which much osteological information was lacking but its Larson (2001) revised Mugilogobius and 13 genera related placement was uncertain (Table 1) and there was insuffi cient to it, and concluded that Pseudogobiopsis (referred to data for an undescribed Calamiana from the Indian Ocean as Eugnathogobius), as it currently stood, was para- or (so it was not included in the analyses). The 41 characters polyphyletic. Larson (1999) considered that Calamiana used are the 40 detailed in Larson (2001: 15–31), with the shared some similarity with Eugnathogobius microps Smith, addition of character 41 (“broken row b”, Fig. 21) possessed 1931, in having headpores absent; sensory papillae in cheek by P. festivus, new species and P. paludosus (papilla row rows c and cp being large and widely spaced, with papillae in b is broken behind the eye into two short sections, with a rows b and d small and close-set; papilla row p composed of wide gap between each section; in all of the other species, widely spaced papillae, and fi ne villi absent from the dorsal this papilla row b is continuous); all characters were run surface of the head. Calamiana was then considered to be unordered and unweighted. Nominal species of Calamiana different from Eugnathogobius in that it had 16 segmented (C. illota, C. kabilia, C. mindora, C. polylepis, C. stictos, new caudal rays (vs. 17), had jaws enlarged in males in one species, and C. variegata), Eugnathogobius (E. microps) and species only (C. kabilia) (vs. jaws are greatly enlarged in all Pseudogobiopsis (P. oligactis, P. paludosus, P. siamensis, P. males), had a low to moderate metapterygoid (vs. a broad tigrellus (this species initially thought to be Redigobius) and metapterygoid) with a process or bridge overlapping the P. festivus, new species), were compared with three species of quadrate (vs. no process reaching the quadrate). The similarity Pseudogobius (P. javanicus, P. melanostictus and P. olorum) of the type species of Eugnathogobius and Calamiana, plus and three species of Redigobius (R. balteatus, R. dispar and the discovery of new species which had features which agreed R. macrostoma) [both these genera are closely related to with those characteristic of one or more of these genera as Calamiana, Eugnathogobius and Pseudogobiopsis and share presently understood, prompted this review. some common characters (Larson 2001)]. Two odontobutids (Micropercops borealis and Sineleotris chalmersi) were used as outgroups. MATERIALS AND METHODS Abbreviations for institutions referred to are: AMS, The Measurements were taken using electronic callipers and Australian Museum, Sydney; BMNH, The Natural History dissecting microscope. Counts and methods generally follow Museum, London; CAS, California Academy of Sciences, Hubbs & Lagler (1970), except as indicated below. Papillae San Francisco; CMK, Collection Maurice Kottelat, Cornol, pattern terminology is based on that of Sanzo (1911), due to Switzerland; FMNH, Field Museum of Natural History, its use in previous literature on this group of gobionellines Chicago; KEW, collection of Dr K. E. Witte, previously by Aurich (1938) and Miller (1987, 1989). Pterygiophore of University of Constance, Konstanz; KUMF, Kasetsart formula follows Birdsong et al. (1988). Transverse scale University Museum of Fisheries, Bangkok; MNHN, Museum counts are taken by counting the number of scale rows National d’Histoire Naturelle, Paris; NIFI, National Inland from the anal fi n origin diagonally upward and back toward Fisheries Institute, Bangkok; NTM, Museum and Art the second dorsal fi n base. The circumpeduncular scale Gallery of the Northern Territory (previously Northern count is taken beginning at the fi rst scale on top of the Territory Museum), Darwin; QM, Queensland Museum, caudal peduncle immediately in front of the caudal fi n, and Brisbane; PMBC, Phuket Marine Biological Laboratory, following the scale rows down and forward to the ventral Thailand; RMNH, Nationaal Naturhistorisches Museum, edge of the peduncle, then around and back to the original Leiden; ROM, Royal Ontario Museum, Toronto; RUSI, now scale. Head length is taken to the upper attachment of the SAIAB, South African Institute for Aquatic Biodiversity, opercular membrane. Interorbital width is the least fl eshy Grahamstown; SFC, Shanghai Fisheries College, Shanghai; width (not least bony width). Although both left and right SMF, Senckenberg Museum, Frankfurt; URM, University of pectoral fi n ray numbers were recorded, the right pectoral the Ryukyus, Naha; USNM, National Museum of Natural fin count is that used in descriptions and tables, unless History, Washington; WAM, Western Australian Museum, otherwise indicated (in cases of damage). The segmented Perth; ZMB, Zoologische Museum, Berlin; ZRC – Raffl es or branched caudal fi n ray pattern (e.g. 9/8 or 9/7) is the Museum of Biodiversity Research, National University of number of segmented caudal fi n rays attaching to the upper Singapore; ZSM, Zoologische Staatsammlung, München. and lower hypural plates respectively. In the descriptions, an asterisk indicates counts of the holotype (or lectotype). Other abbreviations used: HL, head length; SL; standard Numbers in parentheses after counts indicate the number of length; HD, head depth at rear preopercular margin; HW,

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head width at rear preopercular margin; BDA, body depth at 36); ex AMS I.22055-020, C, 3(17-27). Redigobius dispar: anus; BWA, body width above anus; CPL, caudal peduncle syntypes, ZMB 6705, X, 5(31-33.5); syntypes, ZMB 6700, length; CPD, caudal peduncle depth; CL, caudal length. X, 6(40-41); syntypes, ZMB 6703, X, 2(30-36); syntypes, ZMB 6702, X, 4(38-42); ex USNM 263330, C, 2(21.5-25.5). Details of preserved specimens examined are given with Redigobius macrostoma: AMS I.16954-018, X, 9(19-30); ex each species account (descriptions), except for cleared and AMS I.19341-002, C, 3(28-35.5). stained and x-rayed specimens, which are listed below (C = cleared and stained specimen, X = radiograph): SYSTEMATICS ODONTOBUTIDAE. Micropercops borealis: ex AMNH 10441, C, 3(31-33.5). Sineleotris chalmersi: holotype of Results of analyses. – The branch-and-bound analysis Philypnus chalmersi, AMNH 8384, 1(100); ex AMNH resulted in 10 trees, with the best being 78 steps in length, 10456, C, 3(31.5-40.5). GOBIIDAE, GOBIONELLINAE. with consistency and homoplasy indices of 0.50 (consensus Eugnathogobius illotus: holotype of Calamiana illota, ZRC tree in Fig. 1). A bootstrap 50% majority-rule consensus 39268, X, 1(37); paratypes, NTM S.14235-002, X, 7(22.5- tree was the same but for the taxa from Calamiana illota 34); NTM ex S.14235-002, C, 1(30); ZRC 39269, X, 3(37- to Pseudogobiopsis siamensis forming an unresolved 45). Eugnathogobius kabilia: holotype of Vaimosa kabilia, group (Pseudogobius remained as in Fig. 1). Both analyses CAS 32978, X, 1(36.5); paratypes of Vaimosa rambaiae, indicated that the six nominal species of Calamiana and USNM 119647, X, 2(27-29); NTM S.16415-001, C, 1(47); Eugnathogobius microps form a monophyletic grouping. CMK 4789, X, 1(45); NIFI unregistered, X, 3(26-34); NTM Pseudogobiopsis siamensis aligned with this clade in eight S.14302-001, X, 3(21-23); NTM ex S.14302-001, C, 2(22.5- of the trees and with the Pseudogobius clade in two trees by 28). Eugnathogobius microps: holotype, USNM 90316, a reversal (character 1: rear portion of oculoscapular canal X, 1(26); paratype, USNM 11951, X, 1(26); paratypes, over opercle absent). These eight taxa (E. illotus, E. kabilia, USNM 119593, X, 3(20-26.5); KUMF unregistered, X, E. microps, E. mindora, E. polylepis, E. siamensis, E. stictos, 2(17.5-17.5); NTM S.13953-013, X, 6(18-25.5); ex NTM new species, and E. variegatus) are placed together here as S.13953-013, C, 1(24). Eugnathogobius mindora, ex ROM the genus Eugnathogobius Smith, 1931, which has priority 53371, C, 1(24.5); ROM 53371, X, 8 of 20(14-25); CMK over Calamiana Herre, 1945. The species of Eugnathogobius 5366, X, 6(19.5-26). Eugnathogobius polylepis: ex AMS all share the derived condition of character 19 (papilla row I.25523-003, C, 3(17-20.5); QM I.13347, X, 3(18-21). e), which is variably developed in each species. Eugnathogobius siamensis: holotype of Vaimosa mawaia, CAS 29080, X, 1(24); holotype of Vaimosa jurongensis, CAS However, species assigned to Pseudogobiopsis do not all 32982, X, 1(36); paratypes of V. jurongensis, CAS 32983, fall out quite so neatly. In the branch-and-bound analyses, X, 11(19.5-34); ex USNM 119637, C, 3(26.5-32.5); ANSP Pseudogobiopsis siamensis, P. paludosus and P. festivus, new 63126, X, 2(27.5-28); ANSP 87453, X, 13(23-32.5); CMK species, are unresolved in the consensus. Pseudogobiopsis 8485, X, 6(22-31). Eugnathogobius stictos, new species: ex siamensis is best placed with Eugnathogobius (as above). AMS I.32051-032, C, 1(21). Eugnathogobius variegatus: Pseudogobiopsis paludosus and P. festivus, new species, holotype of Tamanka ubinensis, CAS 30964, X, 1(30); ex formed a species-pair in six of the 10 trees, but in four URM P.13341, C, 1(32); URM P.13341, X, 2(29-31.5); ZRC trees they are split (due to headpore characters), P. festivus, 39270, X, 1(33); URM P.13842, X, 1(33). Pseudogobiopsis new species, grouping with the Pseudogobius clade and P. festivus, new species, paratypes, NTM S.16412-001, C, paludosus grouping with the Eugnathogobius-Pseudogobius 1(25); paratypes, ex CMK 8401, X, 26(11-27); ZRC 26026- clade. In two trees they appear next to Pseudogobiopsis 7, X, 2(14.5-19.5). Pseudogobiopsis oligactis: holotype of oligactis and P. tigrellus. Pseudogobiopsis oligactis and P. Vaimosa perakensis, CAS 32975, X, 1(25.5); paratypes of V. tigrellus always appear closer in the 10 trees to Redigobius perakensis, CAS 32977, X, 6(13.5-28.5); NTM S.14239-001, than to any other taxon and form a species-pair in four trees. X, 3(31.5-36); CMK 5385, X, 7(21-36); ex CMK 7422, C, One tree (Fig. 2) is used as the basis for the classifi cation 1(30); NTM S.16413-001, C, 2(28.5-29). Pseudogobiopsis herein. The name Pseudogobiopsis is here retained for four paludosus: holotype of Ctenogobius paludosus, CAS species (P. festivus, new species, P. oligactis, P. paludosus 32998, X, 1(30.5); NTM S.16416-001, C, 2(18-25); CMK and P. tigrellus), based on four characters (4, pectoral girdle 7384, X, 10(12.5-29); ZRC 8411, X, 1(30); CMK 9009, X, lobes; 12, preanal pterygiophores; 21, unossifi ed scapula; 2(13-21); ZRC 14011, X, 1(22). Pseudogobiopsis tigrellus: 32, preopercular bone morphology). Pseudogobiopsis shows holotype and paratypes of Gobius tigrellus, AMNH 18574, some similarity to Redigobius (its 44 putative species are X, 10(15-21). Pseudogobius javanicus: WAM P.30806- presently under revision). 003, X, 10(21-35); ex NTM S.11125-029, C, 2(30-31). Pseudogobius melanostictus: ZMH 19312, X, 3(30.5-36); Several species of Eugnathogobius and Pseudogobiopsis CMK 6286, X, 3(31.5-35.5); USNM 241842, X, 1(35.5); have been described a number of times (e.g. males as one SMF 18199, X, 1(40); ZRC 3521, X, 9(24-38); ex USNM species, females as another) (Table 2). Several museum lots 268186, C, 2(27.5-33). Pseudogobius olorum: ex AMS of specimens exist which may represent an additional one I.20158-001, C, 2(32.5-33); ex AMS I.18478-003, C, or more species, but this cannot be confi rmed until more 2(38-39); NTM unregistered, X, 7(37-53); SAMA F.5123, specimens are obtained. Further work may help clarify X, 7(27-45). Redigobius balteatus: CMK 7184, X, 7(26.5- the relationships of all these taxa. DNA analyses of some

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Fig. 1. Consensus of 10 equally parsimonious trees, resulting from PAUP branch-and-bound search, of Calamiana, Eugnathogobius, Pseudogobiopsis, Pseudogobius and Redigobius (Micropercops and Sineleotris are outgroup).

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Fig. 2. One of the 10 trees from PAUP analyses; preferred tree on which classifi cation is based (CI = 0.4935, RI = 0.6929).

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species of these genera and their relatives are underway nasal pores); preopercular pores absent; sensory papillae in (Jaafar, in prep.; McMahon & Larson, in prep.), but genetic longitudinal pattern; papillae rows a and c consisting of large, samples are not yet available for all the species presently widely spaced papillae, as does row p if present (otherwise under discussion. replaced by headpores), rows b and d always consisting of small close-set papillae, row f of one or two papillae only, As Larson (1999) previously described and illustrated three two or three s rows on snout of one papilla each; modally Eugnathogobius species (as Calamiana: E. illotus, E. mindora two but up to three s papillae rows on snout; papillae rows p, and E. variegatus), these three are not redescribed below, a and c composed of few large widely spaced papillae, other but they do appear in the dichotomous key to species. papillae small and close-set; papilla row c broken under eye, rear portion consisting of one papilla, papilla row e curves upward onto the cheek in advance of the preopercular margin KEY TO GOBIONELLINE GENERA (variably developed); gill opening restricted to pectoral fi n USED IN ANALYSES base or extending past pectoral base to under opercle; smooth bony or fl eshy fl ange or ridge on pectoral girdle, with no 1 16-segmented caudal fi n rays in most species, if 17 fi n rays fl eshy knobs and fl aps; jaws terminal with lower jaw tip present then papilla row e turns upward onto the cheek well usually anteriormost, jaws may be enlarged in males (may in advance of the preopercular margin; preopercular pores be greatly enlarged in some species); inner edges of lips and posterior portion of oculoscapular canal (over preopercle) smooth, without fi mbriae; anterior nostril tubular, placed at always absent ...... 2 or just behind preorbital edge; genital papilla fl attened and – 17-segmented caudal fi n rays; papilla row e follows preopercular margin; preopercular pores and posterior portion oculoscapular usually elongate in males, rounded and bulbous in females; canal usually present (absent in two species) ...... 3 gut simple, S-bend shape. 2 Mouth terminal, lower jaw tip anteriormost; gut short, “S-bend” shape; headpores absent in three species .... Eugnathogobius Pterygiophore formula 3-12210; 26 vertebrae, modally – Mouth subterminal, upper jaw tip anteriormost, with round 10+16; two epurals; two or three (rarely one) anal fin snout overhanging mouth; gut long, spirally coiled about its pterygiophores present before fi rst caudal haemal spine; longitudinal axis; headpores always present ... Pseudogobius anterior end of preopercular bone blunt, rounded or slightly 3 Second dorsal fi n modally with one more ray than in anal fi n; pointed; neural spines of first few vertebrae slender, vertebral pattern 11+15 to 12+14; scapula usually ossifi ed at tip pointed; neural spines on fi rst few vertebrae slender, tips or around foramen; three to four anal pterygiophores before fi rst caudal vertebra; most species stocky, head and body somewhat not broadened (except in some specimens of E. kabilia); compressed; widespread in estuarine to freshwaters across the metapterygoid short, relatively small, may be broadened Indo-Pacifi c ...... Redigobius dorsally or with short dorsal process, but not contacting or – Second dorsal and anal fins with equal numbers of rays; forming bridge to quadrate in three species, metapterygoid vertebral pattern 10+15-17 (usually 16); scapula usually forming distinct low bridge to quadrate in six species; palatine cartilaginous, tip above foramen sometimes ossifi ed; two to and pterygoid slender and nearly equal in length; palatine three anal pterygiophores before fi rst caudal vertebra; head reaching quadrate; quadrate broad, may be somewhat forked; usually depressed; restricted to freshwaters of South-east Asian fi fth ceratobranchial stout, triangular, with high fl ange fi fth region ...... Pseudogobiopsis ceratobranchial narrow and triangular, with distinct fl ange on back (extra crest on back in one species); four to nine ossifi ed gill-rakers; scapula usually unossifi ed but tip above Eugnathogobius Smith, 1931 foramen may be ossifi ed. Eugnathogobius Smith 1931 (Eugnathogobius microps Smith, 1931: 37, Fig. 18, lower Bangpakong River, central Siam, by original designation and monotypy). KEY TO SPECIES OF EUGNATHOGOBIUS Eugathogobius Koumans, 1931: 69 (error for Eugnathogobius). Calamiana Herre, 1945 (Calamiana magnoris Herre, 1945: 80, 1 Segmented caudal fi n rays 16 ...... 3 Calamianes, the Philippines, by original designation and – Segmented caudal fi n rays 17 ...... 2 monotypy). 2 Caudal fin pale, conspicuously barred with dark pigment, Gnathogobius Smith, 1945 (Gnathogobius aliceae Smith, 1945: fi ve elongate black or brown blotches along midside of body; 523, Fig. 104, Bangkok, Thailand, by original designation headpores present; lateral scales 22–24 ...... and monotypy)...... E. siamensis (Fowler, 1934) (Thailand, Malaysia, Singapore, Borneo) Diagnosis. – Distinguished by following combination of – Caudal fi n plain greyish to translucent, darker at base, side characters. Second dorsal fi n rays I,6-9; anal fi n rays I,5-9; of body without brown blotches or bars; no distinct elongate dark blotches along midside of body; headpores absent; lateral unpaired fi n rays modally equal in number; fi rst dorsal spine scales 24–27 ...... E. microps Smith, 1931 (Thailand) may be elongate; pectoral fi n rays 14-20; 16 or17 segmented 3 Upper jaw teeth in single row, teeth compressed and even in caudal fi n rays in 9/7 or 9/8 pattern; predorsal scales variable, height, with tips pointed, indented and bent to one side, lower absent or 2-25; 23-63 scales in lateral series; TRB 7-22; jaw teeth conical, pointed ...... E. variegatus (Peters, 1869) 12-25 circumpeduncular scales; headpores present in three (Thailand, Singapore, Indonesia, Papua New Guinea) species, absent in fi ve; headpores, if present, reduced (no – Upper jaw teeth in at least two rows, all teeth conical and rear part of oculoscapular canal, no preopercular pores, no pointed ...... 4

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4 Lateral scales 27–39; headpores present or absent; fi rst dorsal Description. – Based on 16 specimens, 15–33 mm SL. Counts fi n spines always VI; all fi ns well-developed ...... 6 of female holotype (Fig. 3) indicated by asterisk. – Lateral scales 48–63; headpores always absent; fi rst dorsal fi n spines VI, V or less; fi sh small and slender with very low First dorsal VI; second dorsal I,7–8 (mean I,7*); anal I,6-8 fi ns ...... 5 (mean I,7*), pectoral rays 15–17 (mean 16*), segmented 5 Back and upper sides with small distinct black spots in both sexes, no distinct black spot on upper base of caudal fi n; nape caudal rays 15–17 (mean 16*); caudal ray pattern modally always naked; fi rst dorsal fi n spines V or fewer ...... 9/7*; branched caudal rays 12–16 (mean 14, 15 in holotype); ...... E. stictos, new species (Northern Territory, Australia) unsegmented (procurrent) caudal rays 6/5 (in 1), 6/6 (in – Body indistinctly mottled, males with black spot on upper base holotype), 7/7 (in 2); longitudinal scale count 32–41 (mean of caudal fi n; predorsal scales modally absent, may be 2-16 35*); TRB 10*-13 (mean 11); predorsal scale count 14–20 present; fi rst dorsal fi n spines VI, rarely V ...... (mean 11, 16 in holotype); circumpeduncular scales 12–14* ...... E. polylepis (Wu & Ni, 1985) (mean 13). Gill rakers on outer face of fi rst arch 2+7 (in 1), (China, Taiwan, Malaysia, Singapore, northern Australia) 3+5 (in 1), 3+7 (in 2), 3+8 (in 2), 4+6 (in 1). Pterygiophore 6 Headpores always absent; jaws greatly enlarged in males; head formula 3-12210* (in six). Vertebrae 10+16* (in 8). Neural with at least two dark stripes, each body scale with fi ne brown bar or spot ...... E. kabilia (Herre, 1940) spine of fi rst few vertebrae narrow, pointed. Two* (in 8) (Thailand, Malaysia, the Philippines) epurals. Two* (in 7) or three (in 1) anal pterygiophores – Headpores present or absent; jaw length not greatly dissimilar before haemal spine of fi rst caudal vertebra. between males and females; body with small but distinct brown or blackish spots and mottling ...... 7 Body very slender and compressed, less so anteriorly; head 7 All nape scales small and equal in size, reaching forward as broad and somewhat depressed. Body depth at anal fi n origin far as over preopercular margin, but not reaching to behind 14.4–18.7% (mean 16.5%) in SL. Head rounded, always eyes; headpores always absent; about 10 narrow bars or saddles wider than deep, HL 26.4–31.1% (mean 28.7%) of SL. Depth across dorsum, side of body with dark stripe along midline; at posterior preopercular margin 53.6–61.9% (mean 57.0%) no distinctive bars or stripes on face ...... E. indicus new species (Kenya, Mozambique, Aldabra) of HL. Width at posterior preopercular margin 68.1–81.0% – Nape scales reaching forward to close behind eyes, anteriormost (mean 74.6%) of HL. Mouth terminal, slightly oblique, scale often enlarged; headpores present or absent; dark stripes forming angle of about 15° with body axis; jaws not greatly or broad dusky bars present on snout and/or side of head ..... enlarged in males, reaching to just below anterior half of ...... 8 eye in large adults or to just below anterior margin of eye 8 First three dorsal spines often elongate in males; four to fi ve in male specimens less than 25 mm SL. Lips smooth, with oblique dark brown lines cross side of head; headpores modally fi ne fl eshy fi mbriae present on inner edges of lips; lower absent; anteriormost nape scale enlarged, occasionally all nape lip free at sides, fused across front. Upper jaw 32.6–47.0% scales small, equal in size ...... E. mindora (Herre, 1945) (mean 37.2% in males, 35.1% in females) of HL. Eyes (Fiji, Queensland, the Philippines, Thailand) – No dorsal spines elongate in males, second or third spine relatively small, dorsolateral, high on head, top forming part longest; two or three broad irregular dusky bands cross snout of dorsal profi le, 21.7–30.3% (mean 25.3%) of HL. Snout and cheek; headpores modally present; anteriormost nape scale short, fl attened, rounded in dorsal view, 22.7–30.0% (mean often enlarged ...... E. illotus Larson 1999 26%) of HL. Interorbital moderate to fairly wide, usually (Thailand, Singapore, Brunei, the Philippines) fl at, 14.5–25.0% (mean 20.5%) of HL. Caudal peduncle quite compressed, length 21.9–28.9% (mean 26.7%) of SL. Caudal peduncle depth 11.3–12.9% (mean 12.1%) of SL. Eugnathogobius indicus new species (Figs 3, 4; Tables 3–7) First dorsal fi n low, third spine nearly always longest; spines slightly longer in males than females; tips of spines just Material examined. – HOLOTYPE – AMS I.23623-001, 27 mm reaching past fi rst and second fi n element of second dorsal fi n SL female, Bazaruto Island, off Inhossoro, Mozambique, 4 May (in 33 mm SL specimen); in most specimens, fi rst dorsal fi n 1972. PARATYPES – NTM S. 16251-001, 1(25.0), pools at end spines falling short of second dorsal fi n, or just reach second of mangrove creek, Gazi Bay, Kenya, H. Coehne, Jul.-Aug.1993; dorsal fi n spine base. Third dorsal spine length 11.5–19.7% USNM 316139, 6(13.5–15.5), in well at trail marker 37, Grand (mean 13.8%) of SL. Second dorsal and anal fi ns low, short- Terre, Aldabra Atoll, Seychelles, J. Louton, 3 Jun.1990; RUSI 5461, 8(13.5–33.0), Bazaruto, Mozambique, 4 May 1972; AMS based, posteriormost rays longest, in most specimens, fi n I.23623-002, 1(23), same data as holotype. rays falling short of caudal fi n base when depressed (as in holotype); in 33 mm SL male, rays reaching to caudal fi n Diagnosis. – Small, slender-bodied Eugnathogobius; fi rst procurrent rays. Pectoral fi n moderately broad to slightly dorsal fi n VI; second dorsal rays I,7–8, modally I,7; anal pointed (tips variably damaged in many specimens), central rays I,6–8, modally I,7; pectoral rays 15–17, modally 16; rays longest, 20.3–25.5% (mean 22.4%) of SL; rays usually longitudinal scales 32–41; TRB 10-13; headpores absent; all branched. Pelvic fi ns rounded to somewhat oval, often predorsal scales 14–20; scales on body small, mostly ctenoid; cup-like, nearly reaching anus, 18.4–22.2% (mean 20.3%) gill opening restricted to just under opercle; colour pale of SL. Caudal fi n short, rounded to truncate, 24.6–30.3% yellowish with dark scale margins in network pattern and (mean 27.3%) of SL. variably distinct dark stripe along mid-side of body; known only from Kenya, Mozambique and Seychelles, in estuarine No mental fraenum or fold, chin smooth. Anterior nostril and freshwater habitats. tubular, placed at edge of upper lip, tube short, oriented

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down and forward, preorbital usually curved to accommodate patch of small cycloid scales on upper third to half; sometimes nostril. Posterior nostril small and rounded, placed very close entire operculum scaled. Cheek always naked. Pectoral base to upper front margin of eye. Gill opening usually extending naked. Prepelvic area naked. Belly with cycloid scales; forward to just under opercle. Inner edge of pectoral girdle ctenoid scales may be present anteriorly, close by pelvic smooth with very low, bony ridge on anterior face of fi n base or up to anterior third of belly with ctenoid scales. cleithrum, usually forming distinct thin fl ange of bone, with Scales on body small, those on caudal peduncle tending to two to three small fl eshy knobs. Gill rakers on outer face of be larger. Ctenoid scales on side of body extending anteriorly fi rst arch greatly reduced, very short fl eshy knobs, largest to up behind pectoral fi n; small specimens from Aldabra raker by angle of arch; rakers on inner face of fi rst arch with only few scales anteriorly. also short and stubby; inner rakers on other arches slightly longer and more slender than fi rst arch inner rakers. Tongue Genital papilla in male slender and fl attened, with pointed reduced to nearly absent, tip blunt. Outer teeth across front tip; genital papilla in female short and cylindrical to rounded of upper jaw sharp and curved, larger (but not greatly so) and bulbous. than inner row teeth; behind this row, two to three rows of very small, pointed teeth; one or two rows of small pointed Head pores absent. teeth at side of jaw (outer row teeth not enlarged in males). Lower jaw with three or four rows of small curved sharp Sensory papillae pattern longitudinal. Two s rows present on teeth across front, inner rows all pointing inward; usually snout, of one papilla each. Cheek rows b and d composed one or two rows of teeth at side of jaw. of small closely spaced papillae; rows a, c and cp of few large widely spaced papillae. Mental f rows consisting of Predorsal scales very small and cycloid, reaching to, or just two pairs of papillae. anterior to, posterior preopercular margin. Operculum with

Fig. 3. Eugnathogobius indicus, new species, holotype, 27 mm SL female, AMS I.23623-001, Bazaruto Island, Mozambique.

Fig. 4. Eugnathogobius indicus, new species, paratype, 14 mm SL, male, USNM 316139, Grande Terre, Aldabra Atoll, Seychelles.

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Colouration of fresh material. – No information dorsal and anal fi n ray counts of I,7 (vs. usually I,8) and its available. lower lateral scale counts (32–41 vs. 48–63).

Colouration of preserved material. – Based on holotype and Distribution. – Restricted to the Indian Ocean, to coasts two largest paratypes (Figs. 3, 4); most specimens faded. of Mozambique and Kenya, and Aldabra Atoll in the Head and body light yellowish, with scale margins on most Seychelles. of body, especially upper half, narrowly outlined with light brown, side of body with small cross-hatched brownish Etymology. – From the Latin indicus, referring to India; in blotches, short bars and X-shaped marks, forming lateral this case the Indian Ocean, where this species occurs. stripe or series of blotches linked by dusky midlateral line; blotch at caudal base may be darkest. Holotype most heavily Ecology. – Limited information is available; the species pigmented specimen, with about seven indistinct brownish appears to be marine to estuarine in habitat. The specimens saddles crossing dorsum, blending with dark scale margins from the freshwater well on Aldabra are intriguing in their (Kenya specimen has remnants of these saddles). isolation.

Head and nape speckled with pale brown dorsally, pigment evenly distributed or forming faint spots or diffuse blotches. Eugnathogobius kabilia (Herre, 1940) Side of head with several pale brownish irregular blotches, (Figs. 5–10; Plate 1A; Tables 3–6, 8) opercle with one or two darker brown horizontal bars or blotches, indistinct brownish bars from eye to snout tip and ?Glossogobius mas Hora, 1923: 742–743, Fig. 23 (Chilka Lake: lower jaw visible on Kenyan specimen. Lips plain, pale off Samal Island, Rambha Bay, off Barkul). whitish yellow. Underside of head pale (diffuse brownish Vaimosa kabilia Herre, 1940: 19, Pl. 14 (Kabili River, north Borneo). band crossing isthmus behind chin in Kenyan specimen). – Koumans 1953: 388. Calamiana magnoris Herre, 1945: 80 (Calamianes, the Philippines). Breast pale yellowish-white. Pectoral base pale with short – Herre 1953: 732; Roberts 1989: 168. brown horizontal bar or spot across upper half. Belly Gnathogobius aliceae Smith, 1945: 523–524, Fig. 104 (Bangkok). whitish. – Suvatti 1981: 203; Kottelat 1989: 19. Vaimosa rambaiae (in part) Smith, 1945: 538–540 (Bangkok). First dorsal fi n transparent to light brown, outer half dusky Calamiana kabilia – Roberts, 1989: 168; Kottelat et al. 1993: 146; to brown; or with lower third of fi n translucent and outer Larson 2001: 59–60. portion light to dark brown. Second dorsal fi n translucent Mugilogobius kabilia – Kottelat et al. 1993: 146. with scattered pale brown pigment, which may form row of Calamiana aliceae – Watson & Horsthemke 1995: 91–92. dark blotches. Anal fi n transparent to translucent yellowish. Material examined. – MALAYSIA: Holotype of Vaimosa kabilia, Caudal fi n translucent yellowish, faintly spotted with brown; CAS 32978, 1(36.5), Kabili River, British North Borneo, Sabah, narrow brown vertical bar along hypural crease; just behind A. W. Herre, 1936–1937 Oriental Expedition, Jan.1937. FMNH vertical bar, a pair of faint brownish spots. Pectoral fi ns 51667, 1(15), East Coast Residency, Sungai Gana, tributary of translucent, sometimes with light brown fi n rays. Pelvic Little Kretam River, just above Nypa belt, Kinabatangan, North fi ns plain, translucent. Borneo, Sabah, R. F. Inger, 12 May 1950; NTM S.14302-001, 5(20.5–28.0), Kampong Pangkang Kuap, Sarawak, B. L. Lim, Comparisons. – This species resembles the two other dwarf 1 Jan.1969. THAILAND: Holotype of Gnathogobius aliceae, slender species (E. polylepis and E. stictos, new species) in USNM 119604, 1(38), central canal, Bangkok, Thailand, H. M. its general morphology but differs in usually having second Smith, 2 May 1931. Paratype of Gnathogobius aliceae, USNM

Fig. 5. Eugnathogobius kabilia. Holotype of Vaimosa kabilia Herre, 36.5 mm SL female, CAS 32978, Kabili River, North Borneo [Sabah].

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119605, 1(31), same data as preceding. Paratypes of Vaimosa face of fi rst arch 1+6 to 3+8 (no mode, mean of seven total rambaiae, ex USNM 119647, 2 (27–29), from shallow slough rakers). Pterygiophore formula 3-12210* (in 11), 3-2311 (in behind Department of Fisheries, Bangkok, N. Pongse on 28 May one). Vertebrae 10+16* (in nine); 10+15 (in four). Neural 1931, preserved 2 Dec.1931. CMK 4789, 2(45–47), aquarium spines of fi rst few vertebra relatively straight, pointed (in specimens exported by K. Derwanz, 1985; NIFI uncatalogued, fi ve); tips of fi rst three spines somewhat expanded at tips (in 6(26–34), Bangpakong River, Nongnuch. THE PHILIPPINES: Holotype of Calamiana magnoris, CAS 39881, 1(31), Busuanga, four). Two epurals (in eight)*, one in one specimen. Two Palawan Province, the Philippines, A. W. Herre, 1 Jul.1940. SRI (in 12*) or three (in one) anal pterygiophores before haemal LANKA: SMF 30305, 3(26–33), freshwater, A. Heymer, Dec.1987. spine of fi rst caudal vertebra. Anterior tip of preoperculum NO DEFINITE LOCALITY: AMS I.35823-001, 1(36), Singapore: blunt in males, pointed in females. Metapterygoid broad, probable aquarium specimen, I. Benoit, Sorbonne. forming bridge to quadrate; greatly expanded dorsally and overlapping quadrate in adult males (see Larson 2001: Fig. Diagnosis. – A relatively large Eugnathogobius with second 61) but separate in females (Fig. 6). dorsal rays I,6–8; anal rays I,5–8; pectoral rays 15–18; 15–17 segmented caudal rays (modally 16); longitudinal Body relatively slender, somewhat cylindrical anteriorly, scales 25–34; TRB 8–12; predorsal scales 10–19; shoulder compressed posteriorly. Body depth at anal fin origin girdle with smooth low ridge or fl ange, rarely with fl eshy 16.4–21.5% (mean 18.9%) of SL. Head depressed anteriorly, lobes; mouth terminal, large, greatly enlarged in mature especially in males, always wider than deep, HL 28.1–33.8% males (maxillary sometimes extending past rear edge of (mean 30.8%) of SL. Depth at posterior preopercular margin preoperculum); each scale on side of body with short dusky 44.2–67.1% (mean 57.5%) of HL. Width at posterior bar, two distinct brown stripes extending back from rear preopercular margin 63.2–81.0% (mean 73.6%) of HL, of eye and two from eye to snout tip, fi ns dark, dorsal fi ns preopercular area tending to be infl ated and muscular in usually distinctly spotted with brown; known from estuarine males. Mouth large, terminal, slightly oblique, forming angle to fresh waters of Sri Lanka, Thailand, Malaysian Borneo of about 35–40° with body axis; jaws reaching well past and the Philippines. eye to preopercular edge in males and to below mid-eye in females (jaw in large males often reaching past preopercular Description. – Based on 23 specimens, 15–47 mm SL. An margin). Lips fl eshy, smooth and without fi mbriae; lower lip asterisk indicates counts of holotype of Vaimosa kabilia mostly free, fused near tip of jaw. Upper jaw 31.8–86.6% (Fig. 5). (mean 39.6% in females, 73.9% in males) of HL. Eyes placed dorsolaterally, protruding above dorsal profi le in large First dorsal VI* (in 21), VII (in one); second dorsal I,6–8 specimens, eye width 19.4–31.8% (mean 23.9%) of HL. (mean I,7*); anal I,5–8 (mean I,7*); pectoral rays 15–18* Snout broad, blunt and fl attened, 23.2–33.6% (mean 29.1%) (mean 17); segmented caudal rays 15–17 (mean 16*); of HL; one large male with snout almost square, another with caudal ray pattern 9/6 to 10/7 (modally 9/7*); branched snout almost rectangular (dorsal view). Interorbital broad, caudal rays 13–15 (modally 15*); unsegmented (procurrent) fl at, 17.9–39.5% (mean 26.6%) of HL; wider in males than caudal rays 7/7 to 7/8 (modally 7/7); longitudinal scale females. Caudal peduncle compressed, length 24.2–30.4% count 25–34 (mean 30, 27 in holotype); TRB 8–12 (mean (mean 27.1%) of SL. Caudal peduncle depth 11.2–14.4% 10, 9 in holotype); predorsal scales 10–19 (mean 14, 11 in (mean 12.9%) of SL. holotype); circumpeduncular scales 12*. Gill rakers on outer

Fig. 6. Jaws and suspensorium of Eugnathogobius kabilia, female, ex NTM S.14302-001, Sarawak.

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First dorsal fi n rounded, rather low, usually third (or second) two-thirds of upper jaw; in females, teeth present either on spine longest; fi n not always reaching second dorsal fi n origin all or anterior three-quarters of jaw. Lower jaw with teeth when depressed, otherwise reaching back to base of fi rst mostly across front, with outer row of small pointed curved element of second dorsal fi n. First dorsal spine always shorter teeth and two or three inner rows of slightly smaller sharp than next two. Second dorsal spine sometimes longest (in teeth (inner and outer rows not differing much in size in some males only), spine length 11.1–13.9% (mean 12.7%) of SL. specimens); only outermost row of teeth present at side of Third dorsal spine length 12.5–15.9% (mean 14.1%) of SL. jaw. Males with teeth restricted to front of lower jaw. Second dorsal fi n may be taller than fi rst dorsal, especially anteriorly; posteriormost rays barely longer than anteriormost; Predorsal scales cycloid, mostly evenly sized, reaching fi n not reaching caudal base when depressed. Anal fi n low, forward to behind eyes; often only one scale anteriormost, posteriormost rays longest, rays falling short of caudal fi n in centre of nape close to interorbital space (sometimes this base when depressed. Pectoral fi n oval, central rays longest, scale and one or two scales near it somewhat enlarged). 18.1–26.8% (mean 23.2%) of SL; all rays usually branched. Operculum with small cycloid scales on upper half to upper Pelvic fi ns oval, rays not reaching anus, 14.5–22.2% (mean third only. Cheek always naked. Pectoral base usually covered 19.2%) of SL. Caudal fi n moderate in size, oval, rounded with cycloid scales, naked in one specimen. Prepelvic area posteriorly, 24.5–33.3% (mean 29.6%) of SL. covered with small cycloid scales. Belly scales cycloid, with area of ctenoid scales anteriorly, close to base of Chin smooth, without mental fraenum. Anterior nostril pelvics (ctenoid scales sometimes extending halfway down in short tube, oriented down and forward over upper lip, belly). On side of body, ctenoid scales extending forward preorbital slightly curved to accommodate nostril. Posterior in wedge to close behind pectoral base or at least to below nostril rounded, sometimes with slightly raised rim, placed fi rst dorsal fi n. close to front of eye. Gill opening extending forward to under opercle. Inner edge of pectoral girdle smooth with no Genital papilla in males small and slender, tip slightly ridge or fl ange (in nine), with low fl eshy fl ange (in eight), expanded (pigmented so that it appears bilobed). Genital or knobs and fl aps (in two, including holotype). Gill rakers papilla in females short, bulbous and round, without lobes on outer face of fi rst and second arch short and unspined, at tip. rakers longer and more slender near angle of arch; rakers on inner face of fi rst and second arch stubby and with fi ne Head pores absent. spines at tip; inner and outer rakers on other two arches stubby, spiny at tip and equal in size to the fi rst arch inner Sensory papillae pattern longitudinal, as in Fig. 7. Papillae rakers. Tongue large, broad, concave to rounded at tip. Outer rows g, x, z, b, d, e and opercular series consisting of small, teeth in upper jaw largest, relatively small but sharp and closely spaced papillae. Papillae rows s, p, a, c, cp and i curved, two to three rows of very small sharp teeth behind consisting of widely spaced, relatively large papillae. Three this row; innermost row of few inward-pointing large teeth s rows present, each of one papilla. Row e well separated may be present. In males, teeth present on anterior half or from preopercular margin, running almost vertically, or

Fig. 7. Eugnathogobius kabilia papillae pattern. Holotype of Vaimosa kabilia Herre (CAS 32978). Scale bar = 1 mm.

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curving anteriorly or posteriorly. Row d forming curved arch, edge of eye and running along top of preopercle and opercle, posteriormost end always pointing downward (not rearward). then becoming indistinct. Two short brown streaks extending Two longitudinally oriented f rows, of two papillae each. from front of eye: upper streak from middle of eye curving around top of posterior nostril and running straight to upper Colouration of fresh material. – Herre (1945) gave an lip; lower streak running from front lower edge of eye to just account of the freshly preserved holotype of Calamiana below anterior nostril. Tips of sensory papillae blackish in magnoris: “...very pale tan, each scale stippled with minute large male specimen (Fig. 8); only a few papillae blackish- brown dots; a brown stripe runs from the lower margin of tipped in other specimens. Lips, underside of head and breast the eye across the preopercle and opercle; the dorsals have plain dusky (lips and folds of branchial membranes may be a few dark spots basally, then a white longitudinal bar, the edged with blackish); dusky underside of head may include balance reddish brown; the other fi ns are all more or less pale areas. Belly pale. reddish brown”. Some sexual dichromatism in fi n colour present. In males, Smith (1945) indicated that his type specimens of fi rst dorsal fi n plain dusky, with broad translucent whitish Gnathogobius aliceae were pale yellow with dull brown to margin. In females, fi rst dorsal fi n translucent with about yellowish brown markings; the fi rst dorsal fi n was blackish three rows of dusky rounded spots over rays and membrane; with a white margin, and the second dorsal had reddish- broad translucent margin also present. Second dorsal fi n dusky brown spots. with broad translucent margin (fi n darker in males) and about fi ve irregular rows of dusky spots and streaks (becoming Hans Horsthemke provided several images of captive more irregular toward rear of fi n) in females, in males only adults (for example, Plate 1A), from which the following about three rows of spots on lower half of fi n visible. Anal description is taken. Head and body light pearly grey, paler fi n plain dusky with broad whitish membrane in both sexes. ventrally, with grey-brown edges to scale margins mostly Caudal fi n plain dusky grey with very narrow whitish margin developed on dorsal half of body. Head mostly plain, with in males; fi n whitish to translucent in females, with irregular scattered brownish blotches over nape, diffuse dark grey rows of small rounded brownish spots. Pectorals and pelvics streak from front of eye running below nostrils to upper lip, light dusky in both sexes; pelvic fraenum whitish. and a narrow irregular dark grey horizontal line just ventral to eye, ending near rear preopercular margin. Iris golden Distribution. – Specimens are known only from Malaysian with brown speckling. First dorsal fi n fawn with dark grey Borneo (Sabah and Sarawak), the Philippines, Thailand and to blackish mottling, a indistinct black spot in centre of fi n; Sri Lanka. The species is rarely collected. The specimen outer third of fi n reddish to reddish-orange. Second dorsal from Singapore is in poor condition, and shows every sign of fi n similar in colour to fi rst, fawn with grey to brownish or having died in an aquarium, scales standing out, ulcerations, blackish mottling, with narrow blue-white outer margin and clumps of fungus), and was very likely purchased from broad submarginal red band. Anal fi n whitish to pale grey, an aquarium dealer and not collected from the wild. The becoming darker toward outer margin and with narrow bright species has never been recorded from Singapore (Larson white edge. Pectoral fi n whitish to translucent on distal half, & Lim 2005). basally yellow to pale brownish. Pelvic fi ns translucent bluish grey. Caudal fi n base with grey somewhat square blotches; Ecology. – Apparently restricted to fresh water. Little fi n greyish to brownish, rays reddish to brown, darkening ecological information is available. One specimen (FMNH toward fi n margin, edge of fi n narrowly white. 51667) probably came from slightly brackish water: “just above Nypa belt” according to the label (Nypa grows at upper Colouration of preserved material. – Head and body tidal limits, usually where freshwater infl ow is constant). background colour variable, pale fawn to light brown (Figs. Smith (1945) kept E. kabilia and Mugilogobius rambaiae 5, 8, 9). Top and side of head, back and side of body plain alive in a small tank for seven months, fed on mosquito brown or fi nely speckled with brown; on lower half of side larvae and “entomostracans”. brown spots may be enlarged. Distinct vertical brown line or spot close to margin of each scale on side of body, giving This species fi rst appeared in the German aquarium trade fi sh a fi nely reticulate appearance. On upper half of body in the late seventies but most of the specimens were males and nape, irregular brown mottling and blotches present (Horsthemke, in litt.). Werner (1981) described and illustrated (not always visible). Brown shoulder bar usually present captive behaviour and breeding of this species. He described (distinct on holotype); bar beginning above pectoral base the fi sh as strictly benthic and quite secretive; the males and sloping obliquely backward to midline of body behind constructed nests under fl at stones, shovelling sand with pectoral fi n. At caudal base, three indistinct brown spots their large mouths while pieces of gravel (up to 12 mm forming vague Y-shape. diameter) carried out one by one. Males display to each other with mouths wide open, and may push against each Dark brown streak running almost horizontally from lower other jaw to jaw (Plate 1A); if disturbed by an observer they rear edge of eye, across cheek to opercle; opercle brown to often switched to locking jaws (as if biting). About 1,000 blackish (rounded blotches sometimes present), especially eggs per batch were laid in the spawning burrow; eggs were on rear half, where blackish pigment usually present. Shorter elongate (1.5 × 0.5 mm) and hatched after 44–95 hours at and less distinct brown streak extending from upper rear 25°C. Larvae were 2.4 mm long and fl oated up near the

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water surface until yolk absorbed after 124 hours; they then mouth “... as a shovel in nest construction ...”. Horsthemke moved down into the water column. Werner was not able (in litt.) states that males can open the jaws to an almost to provide suitable food. vertical gape, with the ends of the maxillaries fl ared out at a considerable angle to the body axis; the males then push Watson & Horsthemke (1995) compare the behaviour of against each other with jaws agape. this species (as Calamiana aliceae) with that of Awaous fl avus Valenciennes (adult males of this species have an Remarks. – Glossogobius mas Hora, 1923, is possibly this enlarged mouth which is gaped extremely widely during species. Hora’s syntypes were deposited at ZSI, but were territorial displays). They also report that the fi sh use the reported as lost according to the ZSI type register (Barman

Fig. 8. Eugnathogobius kabilia male, 45 mm SL, CMK 4789, Thailand.

Fig. 9. Eugnathogobius kabilia. Holotype of Calamiana magnoris Herre, 31 mm SL, CAS 39881, Calamianes, the Philippines.

Fig. 10. Eugnathogobius kabilia. Holotype of Gnathogobius aliceae Smith, 38 mm SL, USNM 119604, Bangkok.

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in litt., 19 Aug.1991). Hora’s description was based on six Eugnathogobius microps Smith, 1931 specimens, four female and two male, about 21–25 mm SL. (Figs. 11–13; Tables 3–6, 9) From the description and drawing, Glossogobius mas could be Eugnathogobius microps, Pseudogobiopsis oligactis, E. ?Glossogobius mas Hora, 1923: 742–743, Fig. 23 (Chilka Lake: kabilia or a valid species. Lateral scale counts are given off Samal Island, Rambha Bay, off Barkul). as 24–26, which is within the range for E. kabilia and E. Eugnathogobius microps Smith, 1931: 37, Fig. 18 (Bangpakong microps. Freshly collected specimens from Chilka Lake River, Siam). – Koumans 1931: 68–69; Koumans 1940: 129, 200; Smith 1945: 520; Suvatti 1950: 421 (not seen); Suvatti (the type locality) are required to ascertain what Hora may 1981: 202; Kottelat 1989: 19; Eschmeyer & Bailey 1990: 145; have had. Larson 2001: 68. Eugnathogobius macrops – Herre 1940: 24 (lapsus). Calamiana magnoris Herre, 1945, the type species of the genus Calamiana (Fig. 9), has priority over Gnathogobius Material examined. – THAILAND: Holotype of Eugnathogobius aliceae Smith, 1945, type species of the genus Gnathogobius microps, USNM 90316, 1(26), lower Bangpakong River, central (Fig. 10). The former name was published on the 3.Jun and Thailand, H. M. Smith, 1 Jul.1923. USNM 119591, 1(26), same the latter on the 13 Nov. Herre (1945d) thought that this data as holotype; USNM 119593, 3(20.0–26.5), near Pitrieu, species had two vomerine teeth, placed one behind the other. Bangpakong River, H. M. Smith, 4 Jun.1928; KUMF uncatalogued, 2(17.5–17.5) Bang Nara River, Narathiwat Province, D. Tanwilai, The enlarged, posteriorly directed teeth in the innermost row 23 Dec.1983; NTM S.13953-013, 7(18.0–25.5), small mangrove of the upper jaw (four present in holotype of Calamiana creek, Klong Bang Sai, Phuket Island, H. K. Larson, D. F. Hoese magnoris) may have been mistaken for vomerine teeth. & PMBC staff, 8 Dec.1993. Herre also noticed the “...3 small fl eshy fl aps on the inner margin of the shoulder girdle”. Other material examined (but not used in description). Two specimens, KUMF 1853, near Pitrin, Bangpakong River, H. Roberts (1989) compared scalation and colour pattern of M. Smith, 4 Jun.1928 (fi sh very fragile, heads separated from Calamiana magnoris and C. kabilia (he had examined bodies). the type specimen of C. magnoris but not V. kabilia) and also with specimens he had collected from the Kapuas Diagnosis. – Small, stocky Eugnathogobius with slightly River and identifi ed as Calamiana (these specimens were depressed head; second dorsal and anal rays I,6; pectoral Eugnathogobius paludosus). rays 16–20; longitudinal scales 23–27; TRB 8–9; predorsal scales absent in specimens from central Thailand, 10–12 The Thai name for this species is given by Smith (1945) as predorsal scales present in southern (peninsular) specimens; “pla bu”, a generic name for nearly all small gobies. pectoral girdle smooth or with low fl eshy fl ange; eyes small,

Fig. 11. Jaws and suspensoria of Eugnathogobius microps, male, ex NTM S.13953-013, Phuket, Thailand. Scale bar = 1 mm.

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mostly dorsal and close to snout tip; mouth terminal, large, eye in some males and to below rear half of eye in females may be greatly enlarged in males; tongue deeply bilobed; (nearly to preopercular angle in male holotype). Lips smooth plain grey in colour, with blackish markings along bases and without fi mbriae, not very fl eshy; lower lip mostly free, of dorsal, anal, caudal and pectoral fi ns; known only from fused at tip of jaw. Upper jaw 45.2–69.6% (mean 48.4% in estuarine to fresh waters of Thailand. females, 63.0% in males) of HL. Eyes small, mostly dorsally oriented and close to snout tip and nostrils, 12.0–21.0% (mean Description. – Based on 13 specimens, 17.5–26.5 mm SL. 17.5%) of HL. Snout broad, blunt and fl attened, 22.4–26.6% An asterisk indicates counts of male holotype (illustrated in (mean 25.2%) of HL. Interorbital broad, fl at, 23.9–31.6% Larson 2001: Fig. 60). (mean 27.8%) of HL. Caudal peduncle compressed, length 23.5–28.6% (mean 25.9%) of SL. Caudal peduncle depth First dorsal VI* (in 12), V (in one); second dorsal I,5–6* 12.6–14.6% (mean 13.8%) of SL. (mean I,6); anal always I,6*; pectoral rays 16-20 (mean 19, 18 in holotype); segmented caudal rays 17*; caudal ray pattern First dorsal fi n low, fi rst to fourth spines longest or subequal; 9/8 (11)*; branched caudal rays 8/7* to 9/7 (modally 8/7); spines usually not reaching second dorsal fi n origin when unsegmented (procurrent) caudal rays 7/7 or 8/7 (modally depressed (rays of USNM specimens, including holotype, 8/7); longitudinal scale count 23–27* (mean 24); TRB 8*–9 fl abby and mostly wrinkled); spine tips in some specimens (mean 9); predorsal naked in central Thailand specimens, reaching front part of fi n. First and second dorsal spine length 10–12 in peninsular Thailand specimens (mean 11); 14.3% of SL (in 1). Third dorsal spine length 10.0–13.7% circumpeduncular scales 12*. Gill rakers on outer face of fi rst (mean 12.7%) of SL. Fourth dorsal spine length 10.0–12.2% arch 2+9 (in one), 3+9 (in one), 3+10 (in one). Pterygiophore (in 2) of SL. Second dorsal and anal fi ns low, posteriormost formula 3-12210* (in eight). Vertebrae usually 10+16* (in rays longest, rays falling well short of caudal fi n base when 11); 10+15 (in one), 11+16 (in one). Neural spines of fi rst depressed. Pectoral fi n oval, central rays longest, 21.3–26.7% few vertebra straight, pointed (in seven). Two* epurals (in (mean 24.1%) of SL; rays usually all branched (upper and nine). Two* anal pterygiophores before haemal spine of fi rst lowermost rays sometimes unbranched). Pelvic fi ns oval, caudal vertebra (in nine), three in one specimen. Anterior central rays usually reaching anus, 24.2–27.2% (mean tip of preoperculum blunt (Fig. 11). 25.6%) of SL. Caudal fi n short, round, 22.5–29.1% (mean 25.7%) of SL. Body stout, somewhat depressed anteriorly, compressed posteriorly. Body depth at anal fi n origin 18.5–23.1% (mean Chin with very low bilobed mental fraenum (distinguishable 21.3%) of SL. Head depressed anteriorly, always wider in fresh specimens, but not in types). Nostrils very close than deep, HL 31.3–35.4% (mean 33.9%) of SL; cheeks together, just in front of eye. Anterior nostril with short often considerably infl ated. Depth at posterior preopercular tube, oriented down and forward over upper lip, preorbital margin 53.3–63.2% (mean 59.3%) of HL. Width at posterior slightly infl ated to accommodate nostril. Posterior nostril preopercular margin 63.9–86.4% (mean 76.4%) of HL. oval, without raised rim, placed halfway between eye and Mouth large, terminal, slightly oblique, forming angle of anterior nostril. Gill opening usually extending forward to about 30° with body axis; jaws reaching well past rear of under opercle. Inner edge of shoulder girdle smooth with no

Fig. 12. Eugnathogobius microps papillae pattern. NTM S.13953-013, Klong Bang Sai, Phuket, Thailand. Scale bar = 1 mm.

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ridge or fl ange (in six) or with low fl eshy fl ange on upper small, closely spaced papillae. Papillae rows s, p, a, c, half of girdle (in seven). Gill rakers on outer face of fi rst cp and i include widely spaced, relatively large papillae. arch very short and unspined, longest rakers near angle Three s rows present, each consisting of one papilla. Row of arch; rakers on inner face of fi rst arch also stubby, but e well separated from preopercular margin, running almost slightly bigger than outer rakers; inner and outer rakers on vertically or turning anteriorly. Row d forming curved other arches smooth, stubby, equal in size to the fi rst arch arch, posteriormost end always pointing downward (not inner rakers. Tongue large, deeply bilobed. Outer teeth in rearward). Row f consisting of two papillae only, crowded upper jaw larger than others, relatively small but sharp and behind mental fraenum. curved; behind outer row, two to three rows of very small sharp teeth present; inner rows ending at anterior third of Colouration of fresh material. – Fresh specimens collected jaw. In males, teeth present on anterior half or two-thirds of from Phuket in 1993 were noted as being “plain dark greyish upper jaw; in females, teeth present on anterior two-thirds brown”; unfortunately they were not photographed. to three-quarters of jaw. Lower jaw with teeth mostly across front only, with outer row of small pointed curved teeth Three months after preservation, the fi sh looked very similar and two to four inner rows of very small sharp teeth; only to their remembered live colour, other than the pale underside outermost row of teeth present at side of jaw. Both males appears to have become accentuated. and females with teeth restricted to front of lower jaw; outer row extending halfway along side in females and along one Head and body dark greyish-brown, with belly and underside third of jaw length in males. of head paler (varying among individuals). Thin, very dark brown edge on scales on sides of body (especially rear half Predorsal scales, if present, small, mostly evenly sized, of body) and tiny dense brown spot often present at base usually reaching forward to behind eyes; often only single of each scale. Cheeks paler than interorbital, front of head scale anteriormost, in centre of nape close to interorbital space and preorbital region, which are usually darker brown than (this scale and its immediate neighbours may be somewhat rest of head. Papillae rows dark brown; sometimes distinctly enlarged). Specimens from the Bangpakong River with scales contrasting with rest of head colour. Isthmus whitish to extending only partly over operculum, but predorsal naked. white. Operculum with small cycloid scales in peninsular Thailand specimens, naked in Bangpakong specimens. Cheek always Fins translucent. Dorsals, anal, pectorals and caudal with band naked. Pectoral base covered with cycloid scales (peninsular) of dense dark brown pigment along bases, extending short or naked (Bangpakong). Prepelvic area covered with small distance onto fi n rays and membranes. Pectoral sometimes cycloid scales (peninsular) or naked (Bangpakong). Belly with greyish brown patch on base of fraenum, sometimes scales cycloid (peninsular) or naked (Bangpakong). Ctenoid narrow brownish edge to fraenum. Iris dark gold with dark scales on side of body in wedge, close up to behind or just brown mottling mostly obscuring gold colour. Genital papilla above pectoral base. and area around anus white (usually streak of grey-brown on each side of male papilla). Genital papilla in male slender fl attened and pointed, with one to several tiny lobes at tip; in female, papilla bulbous, Colouration of preserved material. – Available preserved rounded to slightly pointed. specimens have generally lost all markings (other than the specimens from Phuket, described above) (Fig. 13). The two Head pores absent. uncatalogued KUMF specimens from the Bang Nara River have dusky heads, a small spot at the base of each body Sensory papillae pattern longitudinal, as in Fig. 12. Papillae scale and some scattered melanophores on the underside of rows g, x, z, b, d, e and opercular series consisting of the head and breast.

Fig. 13. Eugnathogobius microps, 25.5 mm SL, NTM S.13953-013, Klong Bang Sai, Phuket, Thailand.

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Distribution. – Specimens are known only from central Oligolepis), due to their greatly enlarged jaws. Mahidolia and peninsular Thailand. However, three specimens were is a gobiine, while the others are gobionellines (Birdsong obtained in October 2008 by Koichi Shibukawa (in litt.) from et al. 1988; this paper). Koumans (1940) considered that E. near the mouth of Tien River, drainage system, in microps was “Probably a good genus and species”. southern Vietnam. Glossogobius mas Hora is possibly this species (type lost; Ecology. – The holotype was collected by a pongpang (= see under Remarks for E. kabilia). Hora (1923) states that tidal bag-net) net from “...the swift, tidal [estuarine] part of his species has small eyes and a “deeply notched” tongue, the Bangpakong River, where small gobies abound in both but he does not indicate if there are scales on the nape. If species and individuals” (Smith, 1931). Later Smith collected material from the type locality is shown to be identical to specimens from Petrieu (= Paetriu; also spelled Pitrieu on E. microps, Hora’s name then has priority. one label), a town on the Bangpakong River about 39 miles (62.4 km) east by south of Bangkok where the water was quite fresh (this river enters the Gulf of Thailand at the NE Eugnathogobius polylepis (Wu & Ni, 1985) “corner”) (Smith, 1945). (Figs 14-16; Plate 1B,C; Tables 3-6, 10)

The only recently collected specimens known have all Mugilogobius polylepis Wu & Ni, 1985: 95–98 (Zhonggang, come from peninsular Thailand and south Vietnam. The Fengxian, Shanghai). KUMF specimens came from the Bang Nara River, not far Calamiana polylepis – Larson, in Randall & Lim, 2000: 636; north of the border with Malaysia (unfortunately no habitat Larson et al., 2008: 141. ?Calamiana polylepis – Larson 2001: 57. information was available). The Phuket specimens came from Calamiana sp. nov. 2 – Larson 2001: 61–62. a small (0.1–1.5 m wide) muddy creek draining through tall mangrove forest into Klong Bang Sai (on the other side of Material examined. – CHINA: Holotype of Mugilogobius polylepis, which was an abandoned tin mining area). The substrate SFC S-0001, 31.5 mm SL male, Zhonggang, Fengxian, Shanghai. was sandy-mud to fi ne mud, with leaf litter, sticks and tiny NSMT P.54851, 1(22), mangrove area, Wenchang, east coast of burrows visible, and the creek was occupied by about 20 Hainan Island, K. Matsuura, 10 Mar.1997. TAIWAN: NMMBP-460, species of gobioids. 3(22.5–27.0), Tongshu, Charyi County, I-S. Chen, 5 March 1996. VIETNAM: NTM S.16373-001, 2(21.5–24.5), vicinity of Haiphong, Remarks. – According to the original description, Smith North Vietnam, Pham Dinh Trong, Mar.2000. THAILAND: ZRC 50377, 1(19), Laem Phrao, Changwat Trat, Z. Jaafar et al., Feb.2006. (1931) had only one specimen, the holotype, which was SINGAPORE: ZRC 47532, 1(21), Pulau Tekong, N. Sivasothi, 1 eventually sent to the USNM (USNM 90316). However, Nov.2001; ZRC 50569, 2(20–21), Pasir Ris mangrove, R. Ng et Smith (1945) mentions that other specimens had since been al., 23 Jun.2006. AUSTRALIA, QUEENSLAND: QM I.23883, collected from other parts of the Bangpakong River and he 2(22.5–24.0), in subsurface mangrove mud, Serpentine Creek, lists what he states are paratypes held at USNM (USNM Moreton Bay, 5 Aug.1972; QM I.13374, 3(17.5–21.0), Serpentine 119593, which currently consists of three specimens, and Creek, Moreton Bay, CSIRO prawn survey, 7 Nov.1972; AMS USNM 119591, one specimen). The latter two lots of I.19580-002, 4(18–26), Serpentine Creek, Moreton Bay, CSIRO specimens cannot be paratypes, as Smith (1931) clearly prawn survey, 7 Nov.1972; WAM P.28816-001, 3(18–22), Moreton stated that he was describing his new species and genus “... Bay, V. Wadley & P. Young, 7 Nov.1972; WAM P.28814-002, 9(17–23), Moreton Bay, V. Wadley & P. Young, 1972; AMS from a single specimen...”. I.23262-004, 1(20), The Esplanade, Cairns, D. Hoese & D. Rennis, 2 Oct.1982. AUSTRALIA, NORTHERN TERRITORY: NTM This species is the type of the genus Eugnathogobius. S.14236-001, 19.5 mm SL male, among mud and mangrove roots Smith (1945) considered that Eugnathogobius was on bank of tidal creek, Reichardt Creek, Darwin Harbour, Northern related to Gnathogobius (a junior synonym), Mahidolia, Territory, R. Hanley, 9 February 1993; NTM S.11933-001, 1(19.5), Pseudogobiopsis and Waitea (the last a synonym of mud surface, mouth of Adelaide River, R. Hanley, 23 May 1985;

Fig. 14. Eugnathogobius polylepis. Holotype of Mugilogobius polylepis Wu & Ni, 1985, 31.5 mm SL male, Shanghai Fisheries College S-0001, Zhonggang, Fengxian, Shanghai, China. Photograph by Heok Hui Tan. Specimen has tiny bubbles on dorsal surface.

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NTM S.10419-007, 2(14–19), tiny puddle on mud, East Arm 14, 13 in holotype); unsegmented (procurrent) caudal rays of Darwin Harbour, near Elizabeth River mouth, H. Larson, 29 7/6 to 7/8; longitudinal scale count 37–63 (mean 52, 49 in Mar.1982; NTM S.14075-001, 1(17), on mud surface, Nayarnpi holotype); TRB 12–22 (mean 17, 15 in holotype); predorsal Creek, Roper River, R. Hanley & L. Banks, 7 Sep.1994; NTM scale count 0–20 (mode 0*, mean 3.8); circumpeduncular S.14074-001, 1(21.5), on mud surface, Nayarnpi Creek, Roper scales 13–23 (mean 17, 19 in holotype). Gill rakers on outer River, R. Hanley & L. Banks, 8 September 1994; NTM S.14080- 001, 1(20), on mud surface, Roper River, R. Hanley & L. Banks, face of fi rst arch 2+6 to 4+7 (modally 3+6). Pterygiophore 10 Sep.1994; NTM S.11936-001, 2(18–21), in mud, Rhizophora- formula 3-12210 in two), 3-112100 (in one), 3-12200 (in lined channel, Little Lucky Creek, McArthur River, R. Hanley, 5 one); in fi rst two arrangements, no fi n ray attached to last Aug.1985. WESTERN AUSTRALIA: AMS I.25521-007, 1(22.5), pterygiophore. Vertebrae 10+16 (in six). Neural spine of Crab Creek, Broome, 0–2 m, D. Hoese & D. Rennis, 20 Sep.1985; fi rst few vertebrae narrow, pointed (in fi ve). Two epurals AMS I.25523-003, 25(13.5–23.0), King Sound, NE of Derby, 0–1 (in fi ve). Two (in four) or one (in one) anal pterygiophores m, D. Hoese & D. Rennis, 22 Sep.1985. before haemal spine of fi rst caudal vertebra. Metapterygoid broad, with wide bridge overlapping quadrate; palatine Other material examined (but not used in description). slender; pterygoid relatively short. QUEENSLAND: QM I.13347, 3, Moreton Bay; WAM P.28805- 004, 1, Moreton Bay; AMS I.19580-001, 4, Moreton Bay. NORTHERN TERRITORY: NTM S.14072-001, 1, Roper River; Body very slender and compressed, less so anteriorly; head NTM S.14073-001, 1, Roper River; NTM S.14076-001, 1, Roper relatively broad. Body depth at anal fi n origin 12.4–17.1% River; NTM S.14077-001, 1, Roper River; NTM unregistered, 1, (mean 14.7%) of SL. Head rounded, always wider than deep, Rosie Creek. HL 24.5–28.6% (mean 26.4%) of SL. Depth at posterior preopercular margin 49.1–61.0% (mean 54.8%) of HL. Width Diagnosis. – Small, slender-bodied Eugnathogobius; fi rst at posterior preopercular margin 63.9–87.5% (mean 72.1%) dorsal fi n IV-VI, modally V; second dorsal rays I,6–8; anal of HL. Mouth terminal, slightly oblique, forming angle rays I,7–9; pectoral rays 16–18; longitudinal scales 37–63; of about 20° with body axis; jaws not greatly enlarged in TRB 12–22; headpores absent; predorsal scales modally mature males, reaching to below anterior half of eye in large absent, 2–20 small scales may be present on midline before adults or to just past anterior margin of eye in specimens fi rst dorsal fi n origin; scales on body small, mostly cycloid, less than 20 mm SL. Lips smooth, without fl eshy fi mbriae ctenoid scales on caudal peduncle and in small patch under present on inner edges of lips; lower lip free at sides, fused pectoral fi n; gill opening restricted to just under opercle; across front. Upper jaw 32.7–43.8% (mean 39.1 in males, colour pale yellowish with black speckling, black spot on 36.4% in females) of HL. Eyes relatively small, dorsolateral, upper caudal fi n base in males; known from China, Taiwan, high on head, top usually forming part of dorsal profi le, Vietnam, Singapore and northern Australia, in estuarine and 21.7–31.0% (mean 26.1%) of HL. Snout short, fl attened, mangrove habitats. 21.3–31.3% (mean 26.2%) of HL. Interorbital moderate to narrow, fl at to slightly convex, 14.0–31.3% (mean 23.4%) of Description. – Based on 45 specimens, 14.0–31.5 mm SL. HL. Caudal peduncle very compressed, length 23.5–27.9% Counts of holotype (Fig. 14) indicated by asterisk. (mean 25.9%) of SL. Caudal peduncle depth 10.7–13.6% (mean 12.1%) of SL. First dorsal IV–VI (mean V*); second dorsal I,6–8* (mean I,8); anal I,7–9 (mean I,8*), pectoral rays 16–18* (mean First dorsal fi n reduced, low, second or third spines longest 17), segmented caudal rays 15–17 (mean 16*); caudal ray or subequal; spines slightly longer in males than females; fi n pattern modally 9/7; branched caudal rays 12–16 (mean widely separated from second dorsal fi n origin. Second dorsal

Fig. 15. Eugnathogobius polylepis, sensory papillae. NTM S.10419-007, Elizabeth River, Northern Territory, Australia. Scale bar = 1 mm.

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spine length 6.1–11.0% (mean 8.5%) of SL. Third dorsal anteriorly at least under pelvic fi ns and with cycloid scales spine length 6.5–11.0% (mean 8.8%) of SL. Second dorsal posteriorly. Scales on body small, those on caudal peduncle and anal fi ns low, short-based, posteriormost rays longest, larger. Most of body scales cycloid; ctenoid scales on side rays falling well short of caudal fi n base when depressed, of body usually limited to posterior part of caudal peduncle only reaching about half length of caudal peduncle. Pectoral and small patch behind pectoral fi n; ctenii relatively stout, fi n broad, round, central rays longest, 17.0–22.8% (mean especially on caudal peduncle scales. 19.6%) of SL; rays usually all branched (uppermost ray may be unbranched). Pelvic fi ns short, often cup-like, oval, Genital papilla in male short, slender and fl attened, with reaching about one-third to half distance to anus, 12.5–16.3% pointed tip; genital papilla in female short, rounded and (mean 14.5%) of SL. Caudal fi n short, rounded to truncate, bulbous. 19.1–25.2% (mean 22.0%) of SL. Head pores absent. No mental fraenum or fold, chin smooth. Anterior nostril tubular, placed at edge of upper lip, tube short, oriented Sensory papillae pattern longitudinal, as in Fig. 15. Two or down and forward, preorbital curved to accommodate nostril. three s rows present on snout, of one papilla each. Cheek Posterior nostril small, rounded to oval, placed very close to rows b and d composed of small closely spaced papillae; upper front margin of eye. Gill opening usually extending rows a, c and cp of few large widely spaced papillae. Mental forward to just under opercle. Inner edge of pectoral girdle f rows of two pairs of papillae. smooth with very low, bony ridge on anterior face of cleithrum, usually forming distinct thin fl ange of bone, bent No swim bladder. Gut simple “S-bend”, with one loop; laterally. Gill rakers on outer face of fi rst arch very short intestine looping to left rather than right, tucked behind rest and unspined, longest rakers near angle of arch; rakers on of gut as in E. variegatus. inner face of fi rst arch stubby with tiny pointed papillae resembling spines; inner rakers on other arches similar in Colouration of fresh material. – Freshly collected specimens form and length to fi rst arch inner rakers. Tongue tip blunt, from the Northern Territory were noted in fi eld log by the occasionally slightly concave. Outer teeth across front of author as being similar to preserved specimens in colour. upper jaw largest, sharp and curved; behind this row, two Colour photographs of live male and female specimens from to three rows of very small, pointed teeth; one or two rows Singapore and Taiwan were made available by H. H. Tan of small pointed teeth at side of jaw (teeth similar in males and I-S. Chen respectively (Plate 1B, C). and females). Lower jaw with three or four rows of small pointed teeth across front, outermost row teeth oriented Head and body translucent yellowish grey, with scale nearly upright, inner rows all pointing inward; usually one margins on upper half of body narrowly outlined with light or two rows of teeth at side of jaw. brown, upper half of body with brown to red-brown small spots, cross-hatched blotches, short bars and X-shaped Predorsal scales usually absent, if present, very small, cycloid, marks; markings may extend below mid-side of body on mostly along midline just before fi rst dorsal fi n, occasionally caudal peduncle; most conspicuous mark in males being an additional isolated patch of a few scales or a single row dark brown to black, round spot on upper caudal fi n base along midline nearly to behind eyes. Operculum naked or (Fig. 4, Plate 1B); in females, spot smaller (Fig. 16). Adult with patch of small cycloid scales on upper third to half; sometimes only few scales present. Cheek always naked. male from Singapore (Plate 1B) almost plain yellowish Pectoral base naked. Prepelvic area naked. Belly naked, or grey in colour, slightly darker dorsally, with almost no red- with midline naked and remaining scales cycloid, or naked brown spots or bars visible; very fi ne brown cross-hatched

Fig. 16. Eugnathogobius polylepis, 21 mm SL female, NTM S.14074-001, Nayarnpi Creek, Roper River region, Northern Territory, Australia.

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markings visible; bright yellow stripe along lower part of narrow, beginning below rictus and running just above lower body extending from yellow pectoral fi n base to just above preopercular margin. Lips pale whitish yellow, may be light rear insertion of anal fi n. brown (extension of third cheek streak). Underside of head pale with diffuse brownish band crossing isthmus behind chin, Head and nape speckled with red-brown to grey-brown band joining anterior ends of lowermost cheek streak. Side of dorsally. Side of head with short brown oblique streak body with two rows of 11–13 short, staggered, cross-hatched extending diagonally from lower rear edge of eye, ending brownish blotches or bars; lower third of body mostly pale. anterior to preopercular edge; short blotchy brown to blackish Breast brown, distinctly darker than rest of underside. Pectoral slightly oblique streak just ventral to eye and ending below base pale with short brown horizontal bar across upper half. posterior margin of eye; a narrow dark brown to red-brown Belly whitish, even in heavily pigmented males. Peritoneum narrow stripe commencing just above rictus and running brown dorsally, fading on sides toward pale belly. horizontally along side of head to mid-opercle, extending horizontally onto opercle and ending as indistinct brown First dorsal fi n transparent to light brown, lower half dusky blotch; lowermost brown line on head narrow and less distinct to brown; or with outer third of fi n translucent, central light than others, beginning below rictus and running just above brown stripe, and lower third of fi n translucent whitish. lower preopercular margin. Upper lip concolourous with Second dorsal fin light brownish with transparent outer snout or bright yellow (in live male from Singapore); lower margin, or with scattered pale brown pigment, some forming lip outlined by distinct orange-red to dark pink line ending blotches. Anal fi n translucent yellowish brown to brownish, at rictus. Pectoral fi n base with short brown horizontal bar with transparent margin. Caudal fi n in females and juveniles crossing dorsal half, ending on lower pectoral fi n ray bases; translucent yellowish, faintly spotted with brown; in males, male with yellow bar across bases of fi n rays. caudal fi n dusky brown with broad transparent margin around entire fi n, dark brown edge separating transparent margin from First dorsal fi n transparent with lower third of fi n dusky to dusky centre, and distinct dark brown to blackish round spot translucent brownish; broad yellow margin to fi n in male. on upper base of fi n, just touching hypural crease. Pectoral Second dorsal fi n transparent to translucent yellowish. Anal fi ns with light brown fi n rays, darkest in males. Pelvic fi ns fi n transparent to yellowish brown with diffuse brownish light to dark brown, darkest in males. submarginal band in male. Comparisons. – This species is close to E. stictos, new Caudal fin in female transparent, spotted with brown, species; as stated above, these two grouped together in especially toward base of fi n; in male, caudal fi n translucent cladistic analyses (Figs. 1, 2) and they do not resemble the yellow with broad dusky brown submarginal edging, other Eugnathogobius species. They are small, slender fi sh transparent yellowish margin around entire fin, distinct characterised by a reduction in dorsal fi n spines (usually blackish rounded spot on upper base of fi n. Pectoral fi ns having four or fi ve spines) and a high number of lateral transparent, rays silvery-whitish ventrally. scales (37 to 63). They appear to be more similar to E. indicus than to other species of Eugnathogobius. A photograph by Gianluca Polgar of a captive specimen from Sulawesi shows the typical male caudal fi n colour but Distribution. – Specimens are known from China, Taiwan, there are no stripes on the head as in other specimens of this Vietnam, Singapore and northern Australia (from Broome in species (but this is possibly an artefact of captivity). Western Australia to Moreton Bay in Queensland); possibly also from West Malaysia (see Remarks) and Sulawesi (see Colouration of preserved material. – Head and body light Colouration of fresh material). yellow to yellowish grey, with scale margins on upper half of body narrowly outlined with light brown; side of body with Ecology. – Eugnathogobius polylepis is usually found small spots, cross-hatched blotches, short bars and X-shaped in mangroves, at sites not far from the sea. Much of the marks; most conspicuous mark in males being dark brown Queensland material was collected during beam trawl to black, round spot on upper caudal fi n base; in females, sampling in shallow water. In the Northern Territory, many spot smaller, light brown and not always distinguishable of the specimens were collected by hand at low tide on from other body markings. mangrove mud-banks, in tiny puddles, on the exposed mud surface or partly buried in the mud. A specimen from Rosie Head and nape speckled with pale brown dorsally, pigment Creek in the Gulf of Carpentaria, was found alive among evenly distributed or forming pairs of faint spots or two the bases of dead mangroves Bruguiera exaristata on quite to three narrow bands crossing nape. Side of head with dry, fi rm mud. four short brown streaks, central two often diffused into single dark patch: uppermost (fi rst) forming line extending Remarks. – Mugilogobius polylepis Wu & Ni, 1985, was diagonally from lower rear edge of eye and ending before described from three specimens: the 34 mm SL holotype preopercular edge; second streak short, blotchy, extending (SFC S-0001), and two paratypes 22–23 mm SL (SFC diagonally from ventral edge of eye to centre of cheek S-0002, SFC S-0003). The paratypes now appear to be and intersecting mid-point of third streak; third beginning lost in the postal system (Wu Han-Ling, pers. comm.). just above rictus, extending horizontally onto opercle and My initial impression from Wu & Ni’s illustration (which ending as indistinct brown blotch; fourth (lowermost) streak shows predorsal scales) and the English abstract given in the

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Plate 1. A. Eugnathogobius kabilia. Captive males exhibiting agonistic behaviour. Photograph by Hans Horsthemke. B. Eugnathogobius polylepis, captive male, ZRC 50569, 21 mm SL, Pasir Ris, Singapore. Photograph by Heok Hui Tan. C. Eugnathogobius polylepis, freshly dead female specimen, NMMBP-460, Tongshu, Taiwan. Photograph by I-Shiung Chen.

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original description, was that Mugilogobius polylepis greatly Ng & Tan, 1999: 364; Larson & Lim, 2005: 144; Larson et resembled in colour pattern specimens originally identifi ed al., 2008: 143. as Calamiana (Larson 2001; and originally considered to be Pseudogobiopsis campbellianus – Kottelat et al., 1993: 150, Pl. a new species: Calamiana sp. nov. 2 of Larson (2001)) from 70 (in part). northern Australia, but M. polylepis had VI fi rst dorsal spines Pseudogobius orotai – Kottelat et al., 1993: 150, Fig. 306. Pseudogobiopsis wuhanlini Zhong & Chen, 1997: 79–81 (Min (vs. IV–VI, modally V) and more predorsal scales (25–34 vs. River, Fujian Province, China). 0–16). However, after being able to examine the holotype Eugnathogobius siamensis – Larson, 2001: 71. of Mugilogobius polylepis (brought to Taiwan by Wu), it was found to be a Eugnathogobius almost identical to the Material examined. – THAILAND: Holotype of Vaimosa siamensis, northern Australian specimens, but it possessed VI fi rst dorsal ANSP 60025, 1(31.5), Silom Canal, Bangkok, R.M. de Schauensee, fn spines. Specimens from Taiwan and China (Hainan Island) 18 Dec.1932. ANSP 87453, 13(23.0–32.5), Bangkok, R. de agreed with the holotype of E. polylepis in having the nape Schauensee, 1936; ANSP 63126, 2(27.5–28.0), Bangkok, R. de naked, and in having VI fi rst dorsal fi n spines. However, two Schauensee, May 1934. PENINSULAR MALAYSIA: Holotype specimens from North Vietnam have fi rst dorsal IV–V and of Vaimosa mawaia, CAS 29080, 1(24), Mawai district, Johore, 4–5 predorsal scales. Material from Singapore and Thailand A. W. Herre, 22 Mar.1934. CMK 8485, 11(19.0–30.5), blackwater have V–VI fi rst dorsal fi n spines and 9–20 predorsal scales. stream, 70 km on road to Kuantan-Pekan-Mersing, Pahang, M. Kottelat et al., 24 Jul.1992; ZRC 19323-19324, 2(33.5–34.5), All Australian specimens have IV–VI fi rst dorsal fi n spines stream by Desaru Road, Kota Tinggi, Johor, P. Ng & M. Kottelat, (strongly modally V) and 0–16 predorsal scales (strongly 14 Aug.1991; ZRC 17051-6, 6(23.5–31.0), freshwater stream on modally 0). No consistent colour pattern or other difference Mawai-Tanjung Sedili Road, Johore, P. Ng & M. Kottelat, 14 could be discerned (for example, the Singapore and Taiwan Aug.1991. MALAYSIA, SABAH: Holotype of Vaimosa oratai, populations both have the distinctive orange-red lower lip CAS 32988, 1(18), Tawau, British North Borneo, A. W. Herre, margin). It is diffi cult to make meaningful comparisons of 17 Jan.1937. SINGAPORE: Holotype of Vaimosa jurongensis, the differences in predorsal scale counts and number of CAS 32982, 1(36), Jurong, A.W. Herre, 8 May 1937. Paratypes fi rst dorsal fi n spines (e.g. to determine if there is clinal of Vaimosa jurongensis, CAS 32983, 22(19.5-34), same data as variation or actually several species involved), as there are holotype; BMNH 1938.12.1.215-7, 3(24.5–28.5), same data as 54 northern Australian specimens available, but only 11 previous. BRUNEI: NTM S.14244-001, 8(23.0–28.5), small stream near Kampong Lempong, Temburong Baru, S. Choy, 9 Jan.1992. from all the South-east Asian localities. So I have retained INDONESIA: CMK 7304, 24(11.5–21.0), tributary of Sungei Siak, them all together under E. polylepis, pending further material Riau Province, Sumatra, M. Kottelat, 13 Feb.1991. becoming available. Other material examined (but not used in description). – CHINA: However, there may be undescribed species related to E. ZRC 45961, 1, Lai Chi Wu, New Territories, Hong Kong. polylepis. For example, several specimens (held at NTM) THAILAND: USNM 119638, 37, Bangpakong River; USNM from mangroves at Sementa, Klang, on the west coast of 119637, 23, Bangpakong River; USNM 119648, 29, Bangkok; Peninsular Malaysia, resemble this species but males do USNM 119650, 14, Bangpakong River; USNM 263426, 11, Tale not have the dark caudal fi n spot. Further collections in Sap. CAS 53249, 2, Goh Chang Island, 12°04'30"N 102°22'12"E. appropriate habitat should provide additional specimens PENINSULAR MALAYSIA: CAS 33865, 38, Kota Tinggi; CMK which will help clarify their status. 8538, 38, Johore; CMK 7841, 28, Johore; USNM 258941, 15, Johore; USNM 257131, 53, Muar River; CAS 33169, 4, Mawai, Johore; CAS 39485, 8, Kota Tinggi; USNM 263418, 5, Muar River. MALAYSIA: FMNH 51776, 2, Sandakan; FMNH 44932, 1, Eugnathogobius siamensis (Fowler, 1934) Sandakan, Sabah; CMK 5683, 1, Sungei Sebubut; NTM S.14240- (Figs. 17–20; Tables 3–6, 11) 002, 8, south of , Sarawak; NTM S.14351-002, 4, near Kuching, Sarawak; ZSM/KEW 495, 3, Sungei Stunggang Sarawak. Vaimosa siamensis Fowler, 1934: 157, Fig. 125 (Silom Canal, BRUNEI: NTM S.13051-001, 1, Bukit Patoi; NTM S.14298-001, Bangkok, Thailand). – Fowler, 1935: 161; Smith 1945: 538, 5, Temburong. INDONESIA: CMK 7312, 4, Sungei Siak, Sumatra; 540–541. CMK 9527, 27, Sungei Sebuku, Kalimantan Timur, Borneo. Vaimosa mawaia Herre, 1936: 9, Pl. 6 (Mawai district, Johore, north of Singapore). – Herre & Myers, 1937: 40; Fowler, 1938: Diagnosis. – A robustly built Eugnathogobius with second 267; Koumans 1940: 152. dorsal and anal rays always I,6; pectoral rays 16–19; Vaimosa jurongensis Herre, 1940: 18, Pl. 13 (Jurong, Singapore). longitudinal scales 22–24; TRB 7-9; predorsal scales 6–8, – Koumans, 1953: 386–387. large, reaching close up to behind eyes; preopercular pores Vaimosa oratai Herre, 1940: 20, Pl. 15 (Brook at Tawau, north Borneo). – Koumans, 1953: 389. absent, posterior portion of oculoscapular canal absent; Pseudogobiopsis oligactis – Koumans, 1940: 135. scales on body mostly ctenoid; third spine of dorsal fi n Vaimosa singapurensis – Tweedie, 1940: 75. usually longest, but not elongate; head and body pale with Stigmatogobius oligactis – Koumans, 1953: 116–117; Suvatti, margins of scales outlined in brown, row of brown midlateral 1981: 204. blotches, three diagonal dark streaks on head and vertically Stigmatogobius poicilosoma (in part) –Alfred, 1966: 47. barred caudal fi n; known from China, Thailand and Indo- Calamiana siamensis – Hoese, in Böhlke, 1984: 110. Malayan Archipelago. Pseudogobiopsis jurongensis – Roberts, 1989: 169–170. Mugilogobius jurongensis – Kottelat, 1989: 19. Description. – Based on 36 specimens, 17–36 mm SL. Mugilogobius mawaia – Kottelat, 1989: 19. Pseudogobiopsis siamensis – Kottelat, 1989: 19; Tan & Tan, 1994: An asterisk indicates counts of holotype of Vaimosa 357; Lim & Larson, 1994: 260; Kottelat & Lim, 1995: 247; siamensis.

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First dorsal VI*; second dorsal rays I,6*; anal rays I,6*, fl eshy fi mbriae may be present on inner edges of upper pectoral rays 16–19 (mean 17, 18 in holotype), segmented lip and front of lower lip; lower lip free at sides, narrowly caudal rays 17*; caudal ray pattern 9/8; branched caudal fused across front. Upper jaw 34.2–67.9% (mean 39.0% in rays 6/6 to 8/7 (modally 7/7, 8/7 in holotype); unsegmented females, 54.2% in males) of HL. Eyes dorsolateral, high on (procurrent) caudal rays 8/7 to 8/8; longitudinal scale count head, top forming part of dorsal profi le, 21.6–32.7% (mean 22–24* (mean 23); TRB 7–9 (mean 8*); predorsal scale 27.6%) of HL. Snout slightly pointed, 21.3–35.0% (mean count 5–8 (mean 7*); circumpeduncular scales 12*. Gill 27.3%) of HL. Interorbital moderate to rather narrow, slightly rakers on outer face of fi rst arch 2+5 to 3+8 (modally 3+7). concave, 10.9–31.6% (mean 16.2%) in HL. Caudal peduncle Pterygiophore formula 3-12210 (in 21). Vertebrae 10+15 compressed, length 25.3–31.0% (mean 29.2%) of SL. Caudal (in three), 10+16 (in 29), 10+17 (in one). Neural spines of peduncle depth 11.4–14.9% (mean 13.1%) of SL. fi rst few vertebrae slender, pointed (in 11). Two epurals (in 31). Two (in 16) or three (in 15) anal pterygiophores before First dorsal fi n low, rounded, second to fourth spines longest haemal spine of fi rst caudal vertebra. Scapula unossifi ed. or subequal, third spine modally longest; spines longer in Metapterygoid broad, approximately triangular (apex dorsal), males than females; spines barely reaching second dorsal fi n without process extending toward quadrate (Fig. 17). origin when depressed in males, spines usually falling well short in females. First dorsal spine always shorter than next Body stout, compressed, less so anteriorly. Body relatively three. Second dorsal spine length 13.9–16.9% (mean 16.0%) stocky, depth at anal fi n origin 18.8–23.4% (mean 21.2%) of SL. Third dorsal spine length 14.0–18.3% (mean 15.6%) of SL. Head approximately triangular to rounded, depth of SL. Fourth dorsal spine sometimes longest, in males only, modally greater than width (or subequal), HL 30.8–34.8% length 14.5–16.1% (mean 15.3%) of SL. Second dorsal and (mean 32.9%) of SL; cheeks may somewhat infl ated. Depth anal fi ns short-based, posteriormost rays not much longer than at posterior preopercular margin 55.9–69.3% (mean 61.6%) anterior rays, rays usually falling well short of caudal fi n base of HL. Width at posterior preopercular margin 58.0–86.0% when depressed (just reaching in some large males). Pectoral (mean 70.8%) of HL. Mouth terminal, oblique, forming fi n slender, pointed, central rays longest, 20.3–29.6% (mean angle of about 20–25° with body axis; jaws generally 26.0%) of SL; rays all branched but for uppermost. Pelvic reaching to below rear of eye at least in mature males and fi ns long, rounded to oval, reaching to anus, and to anal fi n to below front half of eye in females. Lips usually smooth, origin in large specimens, 20.7–29.8% (mean 26.0%) of SL.

Fig. 17. Jaws and suspensorium of Eugnathogobius siamensis, male, ex USNM 119637, Bangpakong River, Thailand. Scale bar = 1 mm.

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Caudal fi n approximately rectangular, rounded posteriorly, Predorsal scales large, evenly sized, reaching forward 25.0–35.8% (mean 30.0%) of SL. to behind eyes, anteriormost scale just behind posterior interorbital pore. Operculum with three to fi ve cycloid scales No mental fraenum, chin usually smooth, sometimes slightly on upper half. Cheek always naked. Pectoral base covered infl ated anterior to f papillae row. Anterior nostril in short with relatively large cycloid scales. Prepelvic area with small tube, placed at or just behind preorbital edge, tube oriented cycloid scales. Belly usually covered with ctenoid scales; down and forward, preorbital sometimes slightly curved to some specimens with isolated patch of ctenoid scales under accommodate nostril. Posterior nostril oval, placed halfway pelvics, rest of scales cycloid; occasionally scales along between anterior margin of eye and anterior nostril. Gill midline of belly cycloid. Side of body with ctenoid scales opening usually extending forward to just under opercle. extending at least up to behind pectoral base, sometimes Inner edge of pectoral girdle smooth with no ridge or fl ange extending over top of pectoral base. Anterior scales on sides (in eight); with low bony ridge or fl ange, sometimes angled of body larger than those posteriorly. outward (in 12); or with low irregular fl eshy ridge or knob (in 11). Gill rakers on outer face of fi rst arch very stubby, Genital papilla in male elongate and fl attened, narrowing without spines, longest rakers near angle of arch; rakers on toward tip; in female, short, rounded and bulbous. inner face of fi rst arch also short, about equal to longest raker on outer face; outer and inner rakers on other arches Head pores present. Pair of nasal pores present, pair of same length as inner rakers on fi rst arch. Tongue blunt to anterior interorbital pores (single anterior median pore in one concave, occasionally reduced or absent. Outer teeth in upper specimen), median posterior interorbital pore, and postorbital jaw largest, stout and curved, largest teeth across front of pore and infraorbital pore behind each eye. Rear portion of jaw; behind outer row, about three rows of small, curved, oculoscapular canal absent; preopercular pores absent. Pores sharp teeth; one row of sharp teeth along side of jaw, teeth often at end of short vertical tubes. often absent from rear quarter of jaw in males; teeth slightly smaller in females; tips of teeth may be tinted translucent Sensory papillae pattern longitudinal, as in Fig. 18. Papillae brownish orange. Lower jaw with about four rows of small usually quite tall and conspicuous. Cheek rows a, c and cp curved teeth across front, outermost row oriented upright, composed of few large, widely spaced papillae; rows b and d inner rows all pointing posteriorly; in males, innermost teeth of small, closely spaced papillae. Three s rows on snout, of toward side of jaw large and stout, sometimes three or four one papilla each, anteriormost s papilla just behind upper lip, teeth strongly curved; teeth absent from posterior half of jaw other papillae close together by posterior nostril. Mandibular in males, only one row of teeth at side of jaw in females; papillae widely spaced, with mental f row consisting of one tips of teeth may be tinted translucent brownish orange. papilla on each side of symphysis.

Fig. 18. Eugnathogobius siamensis, papillae pattern. Paratype, Vaimosa jurongensis, CAS 32983, 33.5 mm SL male, Jurong, Singapore. Scale bar = 1 mm.

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Colouration of fresh material. – From colour slides by grey internal blocks of pigment along the lower half of the Maurice Kottelat. Freshly dead specimens very similar to caudal peduncle and diffuse brown saddles and blotches on preserved material. Background colour whitish to greyish- the head and body (see description of preserved material). white with greyish-brown markings on head and body and The caudal and dorsal fins are translucent pale yellow, dark grey to black markings on fi ns. In some slides, three becoming transparent distally, with diffuse grey bands and to fi ve evenly-spaced subcutaneous blackish bars just visible a blackish spot at the rear of the fi rst dorsal fi n. The eyeball through body wall along lower half of caudal peduncle (three is red-brown with pale gold iris. bars) and anal fi n base (two bars); usually only three bars along caudal peduncle visible. Colouration of preserved material. – Head and body whitish yellow to pale brownish (depending upon preservation), Larson & Lim (2005: 144) show a live specimen from Johor, whitish to pale brown on ventral half, with fi ve narrow Malaysia, which is generally translucent golden with dusky brown blotches (resembling short, broad horizontal lines)

Fig. 19. Eugnathogobius siamensis. CMK 7864, 24 and 33 mm SL, Johor, Malaysia. From colour slide by Maurice Kottelat.

Fig. 20. Eugnathogobius siamensis. Holotype of Vaimosa jurongensis Herre, 34.5 mm SL, CAS 32982, Jurong, Malaysia.

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along midside of body; scale margins on most of body (not habitats, including blackwater streams. It has been found ventrally) narrowly outlined with brown, more diffusely syntopically with P. oligactis. outlined on lower half of body; upper half of body duskier than lower half (Figs 19, 20). Five diffuse brown saddles Adult females are about 10 mm smaller than males. Roberts sometimes visible, crossing back and reaching down to (1989) briefl y discussed maximum size at maturity of this midside of body toward midlateral blotches. Posteriormost species, comparing specimens from the Kapuas River with midlateral blotch extending onto caudal fi n base and meeting the type specimens of Vaimosa jurongensis (from Singapore). vertical brownish to blackish bar at fi n base. Ventral midline He noted that a female from the Kapuas was gravid at 20 of caudal peduncle and along anal fi n base with thin black mm SL, with “... numerous eggs about 0.20 mm diameter line and (usually) three evenly spaced blackish blotches. ...”, while three gravid Singapore females were 22.9–24.3 Nape plain brownish to whitish-yellow, scale margins mm SL and had noticeably smaller eggs (0.15 mm diameter) broadly outlined with brown, often with indistinct brownish (Roberts 1989). mottling. Remarks. – Vaimosa oratai Herre, 1940, was based on two Head with three brown markings: fi rst, usually diffuse and specimens, a large female (holotype at CAS) and a smaller indistinct broad bar from front of eye to upper lip (but not male paratype (13 mm long). The latter has not been examined extending onto lip); second, broad diffuse streak or patch from and its whereabouts are uncertain. lower edge of eye to end of jaw; and third, distinct broad diagonal bar from rear of eye extending backward and ending Koumans (1940: 135) considered that Vaimosa siamensis was on middle of cheek, bar often expanded posteriorly; third a synonym of Pseudogobiopsis oligactis, and also observed bar always darkest. Opercle plain brownish or with diffuse that Vaimosa mawaia was probably a Pseudogobiopsis brown blotches, usually darker brown irregular marking in (Koumans 1940: 152). centre. Lips and chin plain dusky to brown. Chin and anterior portion of underside of head with scattered dusky spots or Tweedie’s (1940) record of Vaimosa singapurensis from plain brownish. Pectoral base with blackish to brown spot Singapore does not refer to Stigmatogobius borneensis (of or blotch across bases of uppermost few rays. Breast and which Stigmatogobius singapurensis Bleeker is a junior belly pale to dusky, breast usually darker than belly. synonym), but to P. siamensis, as he refers to paratypes from Jurong in the Raffl es Museum (which Herre took to First dorsal fin translucent with two narrow blackish CAS/SU). It is possible that V. singapurensis may have been horizontal lines crossing middle of fi n, lower line intensifi ed a manuscript name used initially for Vaimosa jurongensis posteriorly as black spot or blotch; broad dusky brown fi n by Herre. margin. Second dorsal fi n with four to six rows of brownish to blackish spots, usually coalescing on posterior third of Smith (1945) fi rst noticed the black line and spots along the fi n at least, forming wavy rows. Anal fi n plain dusky to ventral edge of the caudal peduncle, which are present in dark greyish brown. Caudal fi n translucent to dusky with this species and P. oligactis, as well as in Pseudogobius and four to seven vertically oriented, slightly wavy, brownish to Redigobius. This feature is being investigated further. blackish lines; anteriormost vertical line thickest and darkest, coalescing with posteriormost midlateral blotch in centre Kottelat et al. (1993) combined P. oligactis and P. siamensis of fi n. Pectoral fi ns with light dusky fi n rays, membranes under their account of Pseudogobiopsis campbellianus. The translucent. Pelvic fi ns plain dusky to brownish, including specimens in their Fig. 70 are E. siamensis and show the fraenum. deeper head and barred caudal fi n pattern clearly. Upon examination of their preserved material (deposited at CMK), Comparisons. – This species is most like Pseudogobiopsis it was found that the specimen lot from which the fi sh came oligactis; see under Comparisons for that species. included both species.

Distribution. – Specimens are known from China, Thailand, Material of Pseudogobiopsis wuhanlini Zhong & Chen, Malaysia, Singapore, Brunei and Indonesia. It is probably 1997, was not available for examination. However, it is clear extinct in Bangkok. Ng & Tan (1999) reported the species from the description and illustrations that this is a synonym for the fi rst time from the Endau River system in southern of E. siamensis. The differences cited by Zhong & Chen Peninsular Malaysia. This species was thought to be extinct fall within the defi nition of E. siamensis; they compared it in Singapore until three specimens (ZRC 50271, from against both P. oligactis and E. siamensis. The statements MacRitchie Reservoir) were collected in 2005 (Larson of “shorter jaw length in adult male, extending only to the et al. 2008). Eugnathogobius siamensis was not known vertical line from posterior margin of orbit” (jaws reaching from China until Zhong & Chen’s (1997) description of to below rear edge of eye at least in E. siamensis, not to the Pseudogobiopsis wuhanlini, in which they described material lower corner of the preopercle in both E. siamensis and P. from the Kwangton and Fujian Provinces (Pearl and Min oligactis as stated by Zhong & Chen); “without the head pore Rivers); and in 1999 ZRC acquired a specimen from the (ρ)” (characteristic of E. siamensis); and “predorsal scales New Territories of Hong Kong. always 7” (predorsal scales 5–8, mean 7, in E. siamensis) all agree with E. siamensis. A ZRC specimen from Hong Kong Ecology. – This species is known only from freshwater seems to be a typical specimen of E. siamensis.

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Eugnathogobius stictos new species pierced by relatively large holes, teeth long and robust. (Fig. 21; Tables 3–6, 12) Scapula unossifi ed. Four ossifi ed gill rakers.

Calamiana sp. nov. 3 – Larson 2001: 62. Body compressed, less so anteriorly; ripe females almost cylindrical anteriorly. Body depth at anal fi n origin 15.3– Type Material. – HOLOTYPE – NTM S.14287-001, 22 mm SL 18.0% (mean 16.5%) of SL. Head rounded, somewhat male, mouth of East Alligator River, Northern Territory, Australia, depressed, wider than deep, but not greatly so, HL 25.9– beam trawl, T. Davis, Apr.1979. PARATYPES - AMS I.32051-032, 27.3% (mean 26.6%) of SL. Depth at posterior preopercular 12(12.5–22), same data as holotype; NTM S.14287-002, 2(20–20.5), same data as holotype. margin 53.3–68.8% (mean 61.4%) of HL. Width at posterior preopercular margin 64.4–77.1% (mean 73.4%) of HL. Diagnosis. – Small (to 22 mm SL), slender-bodied Mouth terminal, oblique, forming angle of about 25–35° with Eugnathogobius; fi rst dorsal fi n always V; second dorsal body axis; jaws reaching to below middle of eye in mature rays I,8–9; anal rays I,7–8; pectoral rays 14–18; longitudinal males and to below front half of eye in females and small scales 48–61; TRB 15-22; predorsal scales absent; headpores males (to below mid-eye in holotype). Lips smooth, without absent; scales on body mostly cycloid, weak, ctenoid scales fl eshy fi mbriae present on inner edges of lips; lower lip free present on caudal peduncle; gill opening restricted to pectoral at sides, fused across chin. Upper jaw 37.8–47.2% (mean base; colour pale yellowish with small brown to blackish 35.0% in females, 42.2% in males) of HL. Eyes relatively spots and fl ecks along upper sides of body and caudal fi n small, dorsolateral, high on head, forming part of dorsal dusky with vertical bars; known only from the East Alligator profi le, 22.9–29.1% (mean 25.5%) of HL. Snout fl attened River estuary, Northern Territory. to rounded, blunt when viewed from above, 24.4–30.2% (mean 26.5%) of HL. Interorbital moderate to narrow, fl at, Description. – Based on 14 specimens, 16.5–22 mm SL. 13.3–23.5% (mean 21.9% in females, 18.0% in males) of Counts of holotype (Fig. 21) indicated by asterisk. HL. Caudal peduncle compressed, length 23.5–26.5% (mean 24.5%) of SL. Caudal peduncle depth 10.8–13.7% (mean First dorsal V* (13), VI (1); second dorsal I,8–9 (mean 12.1%) of SL. I,8*); anal I,7–8 (mean I,8*), pectoral rays 14–18 (mean 16, holotype with 18 on right, 17 on left), segmented caudal rays First dorsal fi n very low, rounded, tips of spines free, second 13–16 (mean 16*); caudal ray pattern modally 9/7*; branched or third spines longest or subequal; spines falling short of caudal rays 13–16 (mean 15*); unsegmented (procurrent) second dorsal fi n origin when depressed. First dorsal spine caudal rays 8/7 (in one); longitudinal scale count 48–61 length 10.0% of SL in one specimen. Second dorsal spine (mean 54, 59 in holotype); TRB 15–22 (mean 18*); predorsal length 7.0–11.4% (mean 9.2%) of SL. Third dorsal spine scales absent; circumpeduncular scales 18–25 (mean 22, 21 length 7.0–11.4% (mean 8.5%) of SL. Second dorsal and in holotype). Gill rakers on outer face of fi rst arch 2+6 to anal fi ns low, pointed posteriorly, posteriormost rays longer 3+7 (modally 3+7). Pterygiophore formula 3-12210 (in one). than anterior rays, rays not reaching caudal fi n base when Vertebrae 10+16 (in one). Neural spine of second vertebra depressed. Pectoral fi n broad, rounded, central rays longest, blunted at tip, other spines narrow, pointed (in one). Two 18.2–21.2% (mean 19.8%) of SL; rays all branched but for epurals (in one). Two (in one) anal pterygiophores before uppermost ray (usually unbranched). Pelvic fi ns short, oval, haemal spine of fi rst caudal vertebra. Metapterygoid broad, sometimes cup-like, reaching half or less the distance to anus, with wide bridge overlapping quadrate; palatine slender; 14.1–17.5% (mean 15.1%) of SL. Caudal fi n relatively long, pterygoid short and stout. Upper ceratobranchial broad, rounded posteriorly, 24.1–29.1% (mean 26.7%) of SL.

Fig. 21. Eugnathogobius stictos, new species, holotype, male, 22 mm SL, NTM S.14287-001, East Alligator River, Northern Territory, Australia.

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No mental fraenum, chin smooth, fl at. Anterior nostril in short brown spots and short streaks. Pectoral fi ns pale, rays outlined tube, at edge of upper lip, tube oriented down and forward, in light brown. Pelvic fi ns, including fraenum, pale to light preorbital curved forward to accommodate nostril. Posterior brown. Males more intensely coloured than females; can be nostril oval, placed close to front centre margin of eye. Gill quickly distinguished by dark dorsal and anal fi ns. opening restricted to pectoral base, occasionally extending forward to just under opercle. Inner edge of pectoral girdle Comparisons. – This species is most similar to E. polylepis smooth with thick fl eshy ridge on anterior edge of cleithrum, (see Comparisons for that species). ridge occasionally bumpy, usually smooth. Gill rakers on outer face of fi rst arch very short and unspined, most rakers Distribution. – Specimens are known so far only from one similar in length; rakers on inner face of fi rst arch also stubby; locality, the East Alligator River estuary, Northern Territory. inner rakers on other arches slightly longer than fi rst arch Recent beam trawl surveys in the area have not obtained inner rakers. Tongue tip blunt to bluntly rounded; absent in additional specimens, but it is likely the exact identical habitat two specimens. Outer teeth in upper jaw largest, stout and was not sampled (Larson, 1999a, b, 2000, 2002). curved, one or two rows of small sharp teeth behind this row (outer row teeth slightly larger in males); one row of small Ecology. – Specimens were obtained by 1 mm mesh beam- teeth at side of jaw. Lower jaw with two or three rows of trawl, from muddy substrate. small, curved, pointed teeth across front; usually only one row of teeth at side of jaw. Etymology. – From the Greek stiktos, meaning spotted or dappled, in reference to the fi ne black spots on the body. Predorsal scales absent in all specimens. Body scales mostly cycloid, thin, diffi cult to see, extending anteriorly to over pectoral base. Small patch of weakly ctenoid scales behind Pseudogobiopsis Koumans, 1935 pectoral fi n in seven specimens, remainder with all cycloid scales; patch of distinctly ctenoid scales at base of caudal Pseudogobiopsis Koumans, 1935 (Gobiopsis oligactis Bleeker, fi n, covering approximately posteriormost third (or less) of 1875: 113, Amboina, by original designation). caudal peduncle. Operculum with few small cycloid scales on upper half; scales often obscured by mucous coat. Cheek Diagnosis. – Distinguished by following combination of always naked. Pectoral base, breast and belly naked. characters. Second dorsal fi n rays I,6–10; anal fi n rays I,5–8; fi rst dorsal spine often longest and may be elongate; pectoral Genital papilla in male elongate, slender and flattened, rays 14–20; usually 17 segmented caudal rays in 9/7 or 9/8 with pointed tip; genital papilla in female short, rounded pattern; predorsal scales variable, 2–13; 22–27 scales in lateral and bulbous. series; TRB 6–9; 12–13 circumpeduncular scales; headpores present in three species (Fig. 25), absent in one, headpores Head pores absent. reduced with no nasal pores, and rear part of oculoscapular canal absent in three species, preopercular pores present in Sensory papillae pattern longitudinal, almost identical to two species; sensory papillae in longitudinal pattern; papillae E. polylepis (Fig. 15). Two s rows present on snout, of one rows a and c consisting of large, widely spaced papillae, as papilla each; rows p, c and cp reduced, papillae few and does row p if present (otherwise replaced by headpores), rows widely spaced. Mental f row of two pairs of papillae. b and d always consisting of small close-set papillae, two or three s rows on snout of one papilla each; modally two Colouration of fresh material. – No information but up to three s papillae rows on snout; papillae rows p, a available. and c composed of few large widely spaced papillae, other papillae small and close-set; papilla row c broken under eye, Colouration of preserved material. – Head and body pale rear portion consisting of one papilla; dorsal surface of head yellowish, paler ventrally, with light brownish fi ne speckling; without fi ne villi; gill opening extending past pectoral base cheek with darker speckling (Fig. 21). Underside of head and to under opercle; bony or fl eshy fl ange present on pectoral branchiostegal membranes brown in males; in females, brown girdle, forming fl eshy knobs in some species; jaws terminal, branchiostegal membranes contrasting with pale underside of jaws enlarged in males, reaching posteriorly to behind eye; head. Most conspicuous markings are dark brown to blackish anterior nostril tubular, placed at or just behind preorbital small spots, fl ecks and short streaks scattered over upper edge; genital papilla fl attened and usually elongate in males, half of body, in some specimens, spots roughly forming rounded and bulbous in females. two rows; markings sometimes forming about 12 indistinct blackish bars or blotches crossing dorsal midline. Pterygiophore formula 3-12210; 25-27 vertebrae, strongly modally 10+16; two epurals; two or three (usually two) In males, both dorsal fi ns plain dusky brownish with brown anal fi n pterygiophores present before fi rst caudal haemal streak at base of fi rst few fi n rays; females with dorsal fi ns spine; anterior end of preopercular bone blunt, rounded or translucent with brownish speckling and short streaks. Anal pointed; ridge and/or posteriorly facing groove present along fi n dusky brownish with translucent to whitish margin. Caudal rear edge of preopercle; neural spines of fi rst few vertebrae fi n dusky to light brown with translucent margin on rear edge slender, pointed; metapterygoid short, relatively short and of fi n and four to seven short, vertically oriented rows of broadened dorsally, may have short dorsal process, may

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contact or slightly overlap quadrate or widely separated from Other material examined (but not used in description). quadrate; palatine and pterygoid not particularly slender, SARAWAK: ZRC 40281, 3, Sungei Stok Muda, Lundu; ZRC usually nearly equal in length, palatine falling short of 26026-27, 2, drain off road to Simundjan; ZRC 40282, 4, about quadrate; fi fth ceratobranchial stout, triangular, with distinct 10 km from Kuching; NTM S.14301-001, 3, Sungei Stok Muda, Lundu; NTM S.14300-001, 3, Sungei Cina Matang, Matang; triangular to blunt high fl ange on back; two to 10 ossifi ed ZSM/KEW 1031, 1, fi eld label lost. gill-rakers; scapula usually unossifi ed but tip above foramen may be partly ossifi ed. Diagnosis. – A moderately slender Pseudogobiopsis with second dorsal rays I,7; anal rays modally I,7; pectoral rays Remarks. – There are four valid species of Pseudogobiopsis 17–19; longitudinal scales 28–32; TRB 8-9; predorsal scales recognised here, one of which is described as new below. 8–13, reaching up to behind eyes; jaws enlarged in mature Several species of this genus have been described a number of males; some headpores present, preopercular pores and times (e.g. males as one species, females as another). Several posterior portion of oculoscapular canal absent; fi rst spine museum lots of specimens exist which may represent an of dorsal fi n longest, occasionally fi lamentous in males; light additional one or more species, but this cannot be confi rmed sandy yellow with brown saddles and mottling, fi ns red in until more specimens are obtained. life, dorsal fi ns and caudal fi ns with rows of black or dark brown spots; known only from fresh waters in Sarawak. Key to species of Pseudogobiopsis Description. – Based on 33 specimens, 14.5–34.5 mm SL. An asterisk indicates counts of holotype (Fig. 22). 1 Preopercular pores absent, pores on top of head present or absent ...... 3 – Preopercular pores present, pores on top of head always First dorsal VI*; second dorsal rays always I,7*; anal rays present ...... 2 I,7*–8 (strongly modally I,7); pectoral rays 17*–19 (mean 2 Dorsal rays I,7, and anal rays I,6–7; 15–16 pectoral rays; body 18), segmented caudal rays 16–18 (mean 17*); caudal ray with 12–13 narrow vertical blackish lines; no elongate fi rst pattern 6/6 to 9/6 (modally 7/7*); branched caudal rays dorsal fi n spines ...... P. tigrellus 12–16 (mean 14*); unsegmented (procurrent) caudal rays (Nichols, 1951) (freshwater, Papua New Guinea) 9/9 (in one); longitudinal scale count 24–26 (mean 25*); – Dorsal and anal rays modally I,6; 16–19 pectoral rays; body TRB 8–9 (mean 8*); predorsal scale count 8–13 (mean 10*); with fi ve narrow dark blotches along midside, several short circumpeduncular scales always 12*. Gill rakers on outer dark bars radiate from lower edge of eye; fi rst dorsal fi n spine usually greatly elongate, especially in males ...... P. oligactis face of fi rst arch 2+5 to 3+7 (modally 2+7). Pterygiophore (Bleeker, 1875) (freshwater; Thailand, Malaysia, Singapore, formula 3-12210 (in three). Vertebrae 10+16 (in seven). Borneo, Indonesia) Neural spine of second vertebra blunt or broadened at tip (in 3 Headpores present; pectoral rays 17–19; cheeks, preorbital and one), or slender and pointed (in one). One (in two) or two upper lip crossed with rows of irregular, oblique, distinct black (in four) epurals. Two (in seven) anal pterygiophores before lines ...... P. festivus, new species (freshwater; Sarawak) haemal spine of fi rst caudal vertebra. Palatine not reaching – Head pores absent; pectoral rays 13–16; side of head with quadrate. Metapterygoid relatively low, well separated dark spots and blotches, forming two broad bars, one across from quadrate. Quadrate shallowly forked. Dentary short. preorbital and one to end of jaws ...... P. paludosus Anterior margin of preopercle pointed. Scapula unossifi ed. (Herre, 1940) (freshwater; Kalimantan, Malaysia, Sumatra) Six ossifi ed gill rakers.

Body slender, somewhat compressed posteriorly, less so Pseudogobiopsis festivus new species anteriorly. Body depth at anal fi n origin 15.6–19.2% (mean (Figs. 22–24; Plate 2A; Tables 13–17) 16.9%) of SL. Head rounded, almost cylindrical, wider than deep, but not greatly so, HL 25.3–30.3% (mean 28.4%) of Calamiana sp. nov. – Kottelat & Lim, 1995: 247. Eugnathogobius sp. nov. – Larson, 2001: 66–68. SL; cheeks may be somewhat infl ated in large males. Depth at posterior preopercular margin 50.0–62.1% (mean 55.1%) Material examined. – HOLOTYPE - ZRC 40279, 34.5 mm SL of HL. Width at posterior preopercular margin 56.8–75.5% male, 8.6 km after turnoff to Sungei Cina Matang after entrance to (mean 65.7%) of HL. Mouth terminal, slightly oblique, Matang Reserve, near Kuching, Sarawak, Malaysia, THH 9554, H. forming angle of about 20° with body axis; jaws reaching past H. Tan & D. C. J. Yeo, 4 Sep.1995. PARATYPES - MALAYSIA, rear preopercular margin in large males (such as holotype) SARAWAK: ZRC 40280, 6(20.5–27.0), same data as holotype; and to below mid-eye in females. Lips smooth, without fl eshy NTM S.14299-001, 7(19.5–29.5), same data as holotype; ZRC fi mbriae on edges; lower lip narrowly free at sides, fused 27842-3, 2(14.5–24.4), 7 km on Kuching-Batu-Kawa Road, M. across front. Upper jaw 31.8–66.0% (mean 35.8% in females, Kottelat & K. Lim, 3 Jul.1992; ZRC 29082-3, 2(19.0–20.5), 7 km on Kuching-Batu-Kawa Road, M. Kottelat & K. Lim, 3 Jul.1992; 46.2% in males) of HL. Eyes dorsolateral, high on head, ZRC 50715, 9(18–33), stream 1, Ulu Assam, Bako National Park, forming part of dorsal profi le, 21.6–34.6% (mean 28.3%) Kuching, K. Lim, 30 Jun.1994; ZSM\KEW 538, 5(16–24), NW of HL. Snout short, fl attened, 19.6–27.3% (mean 22.9%) of Bau, Batang Kayan River basin, Sungai Stunggang, C. Kettner, of HL. Interorbital moderately broad to narrow, 9.8–29.4% K.-E. Witte & R. Krumenacher, 24 Mar.1988; CMK 8401 (12 will (mean 16.9%) of HL. Caudal peduncle compressed, length go to ZRC), 26(11–27), Sungei Bejit, road from Balai Ringin to 28.2–33.8% (mean 30.3%) of SL. Caudal peduncle depth Simunjan, M. Kottelat, K. Lim & P. Ng, 2 Jul.1992. 11.7–13.5% (mean 12.4%) of SL.

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First dorsal fi n triangular, fi rst or second spines longest Sensory papillae pattern longitudinal, as in Fig. 23. Papilla or subequal, occasionally fi lamentous, fi rst spine modally row b broken well behind eyes, rear portion with two to four longest; spines slightly longer in males than females; spines papillae. Three s rows, of one papilla each. Mental f row falling short of second dorsal fi n origin when depressed, even consisting of one papilla on each side of symphysis. in large males (unless spines fi lamentous). First dorsal spine length 14.4–24.8% (mean 19.3% in males, 17.3% in females) Colouration of fresh material. – From colour slides by of SL. Second dorsal spine length 14.4–22.7% (mean 17.5% Peter Ng and Heok Hui Tan of fresh specimens. Colouring in males, 16.1% in females) of SL. Second dorsal and anal very similar to preserved specimens, but upper part of head fi ns short-based, lower than fi rst dorsal fi n, posteriormost and body with scattered orange-red spots, dorsal and anal rays longest, rays falling well short of caudal fi n base when fi ns with orange-red to red wash over black and dark brown depressed. Pectoral fi n slender, pointed, central rays longest, spots and streaks; pectoral and caudal fi ns reddish, not as 23.7–29.7% (mean 25.7%) of SL; rays usually all branched. bright as dorsal fi ns (Plate 2A). Pelvic fi ns long, oval, usually reaching to anus, 21.1–24.8% (mean 22.8%) of SL. Caudal fi n slender, pointed to slightly Colouration of preserved material. – Head and body rounded, 27.0–31.9% (mean 29.3%) of SL. yellowish-white to pale yellowish-brown, with scattered dark brown to blackish spots, some scale margins partly No mental fraenum, chin smooth to slightly bulbous. Anterior outlined with dark brown, especially on lower half of body, nostril tubular, placed at preorbital edge, tube oriented down and fi ve oblique to nearly vertical dark brown bars crossing and forward, preorbital often slightly curved to accommodate back and sides, evenly spaced from just before fi rst dorsal nostril. Posterior nostril round to oval, placed close to front fi n origin to just before upper procurrent rays of caudal fi n; centre margin of eye. Gill opening extending forward to occasionally brown blotch on centre of nape (Fig. 24, Plate under opercle. Inner edge of pectoral girdle smooth (in nine) 2A). Oblique bars becoming diffuse and ending on midside or with knobby ridge or bony fl ange, which may support of body; indistinct midlateral brown streak, formed by two to fl aps (in 21), two specimens with distinct fl eshy knobs. Gill three rows of pale scales with brown spots on or near their rakers on outer face of fi rst arch very short and unspined, centres, sometimes present. Brown spots on body of variable longest rakers near angle of arch; rakers on inner face of intensity, usually very fi ne and partly outlining scale margins, fi rst arch not much longer than those on outer face; inner especially on lower half of body. At midbase of caudal fi n, and outer rakers on other arches similar to rakers on fi rst oblique elongate blackish blotch present, extending down and arch. Tongue long, tip usually blunt to slightly concave; back onto lower segmented caudal rays. Pectoral base dusky occasionally folded longitudinally. Outer teeth across front with short dark brown bar extending obliquely from upper of upper jaw largest, sharp and curved, largest teeth toward third of base onto upper ray bases. Ventral midline of caudal side of jaw (considerably larger in males); behind this row, peduncle with thin black line sometimes visible; midventral two or three rows of small sharp teeth; only one or two line of body with fi ve narrow black blotches, evenly spaced, rows at side of jaw; teeth may be absent from posterior from anus to just before lower procurrent rays of caudal fi n; half of jaw; tips of teeth (mostly outer row) often tinted anteriormost black blotch (at anus) sometimes obscure. translucent orange to brown. Lower jaw with four or fi ve rows of small sharp teeth across front; teeth toward sides Side of head with many dark brown to blackish oblique lines, largest and stoutest (enlarged and curved in males); usually often vermiculate or broken into rows of short streaks or only one row of teeth along side of jaw; tips of teeth often spots; opercle usually with rows of spots. Anteriormost two tinted translucent orange or brown. lines darkest, extending from margin of eye across preorbital to front of jaws, crossing jaws to meet counterparts on Predorsal scales moderate, evenly sized, reaching forward lower lip and chin. Upper lip pale, barred with dark brown to behind eyes or at least halfway between rear preopercular (continuation of oblique stripes on head); lower lip pale to margin and eyes. Operculum with two to four cycloid dusky, often with few brown bars or spots anteriorly. Rear scales on upper half. Cheek always naked. Pectoral base of opercle and branchiostegal membranes dusky with small covered with cycloid scales. Prepelvic area covered with brown blackish spots in males; pale with short brown to cycloid scales. Belly with cycloid scales, occasionally naked blackish oblique lines in females. Nape and interorbital with anteriorly, underneath pelvics. Ctenoid scales on side of small brown blotches and small spots. Underside of head body extending up to pectoral base. and belly plain pale whitish to dusky brownish; scales with margins outlined in dark brown forming dark patch on breast Gut simple, S-bend shape. just anterior to pelvic fi ns in males.

Genital papilla in male relatively short, fl attened, tip blunt First dorsal fi n translucent to dusky, with broad dark grey to to rounded, with one or two fi ne lobes present; in female, blackish streak across centre of fi n, diffuse black spot within short, rounded and indented at tip. streak toward rear sometimes present; brownish narrow stripes or dusky streaks along fi n above and below central Headpores present, with pair of anterior interorbital broad dark streak; tips of several dorsal spines with dense scales, single posterior interorbital pore, and infraorbital black spot, usually on fi rst, third and fourth; fi rst dorsal and postorbital pores behind each eye. Rear portion of spine always pale with two to three small dark brown to oculoscapular canal absent. No preopercular pores. black spots on membrane just behind spine. Second dorsal

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fi n whitish to translucent, with fi ve to seven rows of small Pseudogobiopsis oligactis (Bleeker, 1875) dark brown to black round to oval spots; unsegmented spine (Figs. 22, 26; Plate 2B; Tables 13–16, 18) always pale with two black spots, fi n margin plain dusky. Anal fi n plain dusky to brownish. Pectoral fi ns translucent Gobiopsis oligactis Bleeker, 1875: 113-114 (Amboina, Indonesia). dusky to pale brownish, rays becoming darker toward tips; – Koumans, 1931: 67; Bleeker, 1983: Pl. 433, Fig. 4. fi ns distinctly darker in males. Pelvic fi ns plain dusky grey Glossogobius campbellianus Jordan & Seale, 1908: 542, Fig. 2 to brownish. Caudal fi n translucent whitish to pale dusky (Buytenzorg, Java). ?Glossogobius mas Hora, 1923: 742–743, Fig. 23 (Chilka Lake: with many rows of vertically oriented, dark brown to black off Samal Island, Rambha Bay, off Barkul). small spots or short lines, spots becoming smaller distally; Pseudogobius neglectus Koumans, 1931: 102 (nomen nudum; short lines closest to fi n base sometimes joining to form one Bleeker museum name). – Bleeker, 1983: Pl. 438, Fig. 13. or two oblique to vertical dark lines on lower half of fi n; Stigmatogobius neglectus Koumans, 1932: 3, 5 (western Java). lowermost edge of caudal with broad plain dusky to greyish Pseudogobiopsis oligactis – Koumans, 1935: 131–132, Fig. 4; margin; one to three black spots along upper edge of fi n, Fowler, 1937: 251; Koumans, 1940: 126; Smith, 1945: 521–522; crossing dorsal procurrent rays. Kottelat, 1989: 19; Kottelat et al., 1993: 150, Pl.70; Lim & Larson, 1994: 260; Kottelat & Lim ,1995: 247; Ng & Tan, 1999: Males distinguished from females in being darker, on 364; Larson & Lim, 2005: 143; Larson et al., 2008: 143. Vaimosa perakensis Herre, 1940: 21, Pl. 16 (Lake above Chenderoh pectoral fi ns especially, and in having darker and more Dam, Perak, Malay Peninsula). complex vermiculate oblique lines on head (Figs. 23, 24, Stigmatogobius oligactis – Koumans 1953: 110, 116–117, Fig. 27; Plate 2A). Chatterjee, 1980: 229, 230. Stigmatogobius poicilosoma (in part) – Alfred, 1966: 47. Sensory papillae often orange-tipped. Peritoneum pale with Stigmatogobius isognathus - Mohsin & Ambak, 1983: 187, Fig. broad brown dorsal saddle and scattered brown spots (as in 141. colour pattern of surrounding skin). Mugilogobius perakensis – Kottelat, 1989: 19. ?Genus and species undet. – Roberts, 1993: 44, Fig. 51. Comparisons. – Most similar to P. paludosus; most easily Pseudogobiopsis campbellianus – Kottelat et al., 1993: 150, Pl. 70 (in part). distinguished by the presence of headpores and in the oblique Pseudogobiopsis neglectus – Kottelat et al., 1993: 150, Pl. 70. dark lines on the side of the head (vs. headpores absent and Eugnathogobius oligactis – Larson, 2001: 6970. brown blotches on side of head). Material Examined. – INDIA: CAS 61960, 3(15.0–20.5), sandy Distribution. – Specimens are only known so far from beach at Hampi, Tungabahdra River, Bellary District, Karnataka rainforest freshwater streams of Sarawak, Malaysia. State, Mysore, T. Roberts, 3 Feb.1985. THAILAND: CMK 5385, 22(17–36), stream on road from Ranong to Kra Buri, Ranong Ecology. – One specimen lot was recorded as being from a Province, M. Kottelat, 24 Apr.1985; KUMF uncatalogued, slightly turbid blackwater stream, with leaf litter and aquatic 3(37.5–41.0), Tha Lad Waterfall, Samui, Surathani Province, C. Vidthayanon, 22 Jul.1983; NIFI 1230, 3(33.5–41.0), Chiew vegetation present. Larn Dam, Klong Saeng, Surat Thani Province, J. Karnasutra, 20 Mar.1983. CAMBODIA: CMK 4806, 1(35), Stung Tong Hong, Etymology. – From the Latin, festivus, meaning cheerful, 77 km on road from Phnom Penh to Sihanoukville, d’Aubenton, delightful or humorous, all words which relate to the elongate 13 Jan.1963. MALAYSIA: Holotype of Vaimosa perakensis, CAS smiling jaws of mature males. 32975, 1(25.5), Chanderoh Dam, Perak, A.W. Herre, 9 Mar.1937. Paratypes of Vaimosa perakensis, CAS 32977, 17(13.5–28.5), 2 Remarks. – This species and P. paludosus share a miles N of Sauk, Perak, A.W. Herre, 18 Apr.1937; ZRC 255, synapomorphy: cheek papilla row b is broken into two 47(12-24.8), lake above Chanderoh Dam, Perak, A. W. Herre & sections, with a wide gap behind the eyes. M. F. Tweedie, 19 Mar.1937. CMK 7422, 4(24–31), NE foothills

Fig. 22. Pseudogobiopsis festivus, new species, holotype, 34.5 mm SL, male, ZRC 40279, near Kuching, Sarawak, Malaysia.

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of Gunung Panti, N of Kota Tinggi, Johore, M. Kottelat et al., Kalabakan, Tawau, Sabah, R. Inger, 8 Jun.1956; NTM S.14240- 1991; NTM S.16374-001, 1(30.5), cleared and stained, same data 001, 4(23–26), 7 miles (11.2 km) south of Kuching, Kampong as preceding; ZRC 19323–19325, 1 of 3(34), Desaru Road, Kota Pangkalan Kuap, Sarawak, B. L. Lim, 1 Jan.1969. BRUNEI: NTM Tinggi, Johor, P. Ng & M. Kottelat, 14 Aug.1991; ZRC 20457–61, S.14239-001, 3(32–36), Sungai Belalong, KBSFC, S. Choy, 27 5(25.5–37.5), Rembia, Malacca, D. S. Johnson, 26 Feb.1958; ZRC Jul.1992. SINGAPORE: ZRC 1049i–iv, 4(24–28), MacRitchie 1651i–iii, 3(32.0–40.5), Sungei Telok Bahang at 14.5 miles (23.2 Reservoir, S. H. Chung, 4 Nov.1959. INDONESIA: Holotype of km) on Telok Bahang Road, Penang, E. R. Alfred, 21 Oct.1961; Glossogobius oligactis, RMNH 4459, 1(25.5), Java. Holotype of FMNH 68465, 3(27.5–29.0), Sungai Tawan, Sungai Tibas Camp, Glossogobius campbellianus, USNM 61051, 1(26.5), Buitenzorg

Fig. 23. Pseudogobiopsis festivus, new species, papillae pattern. 26 mm SL female, CMK 8401, Sungei Bejit, Sarawak: A, lateral view; B, dorsal view. Scale bar = 1 mm.

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(= Bogor Botanical Gardens), Java, D. H. Campbell. Paratype dark blotches along midside of body with fi ne dark spots of Glossogobius campbellianus, CAS 22498, (23), same data as below and three dark streaks on face; known from fresh holotype. CMK 9621, 6(15–32), clear stream east of Mentok on waters of Indo-Malay Archipelago. road from Pangkalpinang, Banka, Sumatra, M. Kottelat et al., 4 Mar.1993. Description. – Based on 56 specimens, 18.5–46 mm SL. Other material examined (but not used in description). An asterisk indicates counts of holotype of Glossogobius THAILAND: KUMF uncatalogued, 8, Thasaeh Reservoir, oligactis. Chumporn; USNM 263432, 14, Ban Pong; CMK 9765, 2, Koh Chang Island; CMK 5350, 5, Phang Nga; CAS 44347, 4, Goh Chang First dorsal VI*; second dorsal I,6–7 (mean I,6*); anal Island; CAS 53248, 1, Goh Chang Island; CAS 53254, 1, Goh I,5–7 (mean I,6*), pectoral rays 16–19 (mean 18, holotype Chang Island; CAS 51608, 1, Goh Chang Island; NTM S.14350- with 19 on right, 18 on left), segmented caudal rays always 042, 2, Bangpakong River; CMK 10713, 22, Trat Province; SMF 17*; caudal ray pattern 9/8; branched caudal rays 12–15 13346, 4, Lam Pi River, Phuket; USNM 316197, 1, Pungah; NTM (mean 14, modally 7/7, fi n ray tips broken in holotype); S.14241-001, 13, Borapet Swamp, Nakonsawan. CAMBODIA: unsegmented (procurrent) caudal rays 7/7 to 10/10 (modally MNHN 1985.988, 2, Kampot River; MNHN 1985.996, 1, Stung Hong Kong. THE PHILIPPINES: CMK 9978, 2, Basey River, 9/9); longitudinal scale count 22–25* (mean 23); TRB 6–8 Samar. MALAYSIA: CAS 32976, 187, paratypes of Vaimosa (mean 8, 7 in holotype); predorsal scale count 6*–10 (mean perakensis, Chanderoh Dam, Perak; ex CAS 39485, 10, Kota Tinggi, 7); circumpeduncular scales 12–13 (13 only in one specimen). Johore; ex USNM 258941, 1, Sungei Sedili Besar, Johore; ZRC Gill rakers on outer face of fi rst arch 1+7 to 4+8 (modally 32013–26, 14, Penang; ZRC 35262–89, 28, Perak; CMK 20510, 3+6). Pterygiophore formula 3-12210 (in 19). Vertebrae 6, near Kota Tinggi; ZRC 27562–5 (in part), 1, Selangor; FMNH 10+15 (in two), 10+16 (in 17), 10+17 (in two). Neural 63039, 1, Sungei Saoh Sarawak; FMNH 68468, 1, Tawau, Sabah; spine of second and third vertebra narrow and pointed (in FMNH 68469, 3, Tawau, Sabah; FMNH 68470, 1, Tawau, Sabah; nine) or stout and pointed (in two). Two epurals (in 16). FMNH 68464, 5, Tawau, Sabah; ZSM/KEW 268, 1, on Kota Two (in nine) or three (in seven) anal pterygiophores before Kinabalu/Sandakan road, Sabah. BRUNEI: NTM S.14237-001, 1, Belalong; NTM S.14238-001, 1, east of Temburong. INDONESIA: haemal spine of fi rst caudal vertebra. Upper rear corner of CMK 9527, 1, Sungei Tulit, Kalimantan Timur, Borneo; BMNH scapula ossifi ed or not. Palatine not in contact with quadrate. 1975.8.15.387-389, 3, Ranu Klumdumgah E, Java; CMK 6098, 8, Pterygoid slender, may be equal to palatine in length. Lake Sawangan, Java; CMK 10816, 2, Rawa Pening, Java; CMK Quadrate forked. Metapterygoid small, with short dorsally 6104, 1, Maros, Sulawesi; RMNH 17118, 1, Sinaloa (possibly = oriented projection, no process reaching toward or contacting Sinalud, S of Singaraja, Bali; not Sinaloa, Mexico). NO LOCALITY: quadrate (illustrated in Larson, 2001: Fig. 64). RMNH 23266, 7; RMNH 23017, 1; RMNH 14050, 5. Body compressed posteriorly, rounded to somewhat depressed Diagnosis. – A moderately slender Pseudogobiopsis with anteriorly. Body depth at anal fi n origin 14.0–21.6% (mean second dorsal rays I,6–8 (modally I,6); anal rays I,5–7 18.4%) of SL. Head wider than deep, especially in mature (modally I,6); pectoral rays 16–19; longitudinal scales 22– males; HL 28.6–35.8% (mean 32.1%) of SL; head in 25; TRB 6-8; predorsal scales 6–10, large, reaching up to mature females appearing oval in cross-section, males with close behind eyes; head fl attened and jaws greatly enlarged head distinctly depressed. Depth at posterior preopercular in male; preopercular pores present, posterior portion of margin 43.4–63.0% (mean 53.1%) of HL. Width at posterior oculoscapular canal present, including TLCP pore; scales preopercular margin 60.2–84.3% (mean 69.0%) of HL; cheeks on body mostly ctenoid; fi rst spine of dorsal fi n longest and may be infl ated. Mouth terminal, slightly oblique, forming usually fi lamentous in males, second or third spine longest angle of 15–20° with body axis; jaws greatly enlarged in in females, but not elongate; upper half of body with scale mature males, reaching well back past eye and nearly reaching margins narrowly outlined with dark pigment, fi ve elongate angle of preopercle; immature males with jaws reaching to

Fig. 24. Pseudogobiopsis festivus, new species, female paratype, 29.5 mm SL, ZRC 40280, Kuching, Sarawak, Malaysia.

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below rear half of eye; mature females with jaws reaching girdle smooth with no ridge or fl ange; only two specimens to below front half of eye or just behind eye (in holotype observed with single small fl eshy knob. Gill rakers very short of Glossogobius oligactis, end of jaws protruding past and unspined, longest rakers near angle of arch; rakers on preopercular edge). Lips usually smooth, fl eshy fi mbriae may inner face of fi rst arch about twice height of outer rakers, as be present anteriorly on inner edge of upper lip (fi mbriae are outer and inner rakers on other arches. Tongue large, tip sometimes very close to outer edge); lower lip free at sides, deeply bilobed (rarely concave); occasionally tongue blunt fused across front, fl eshy symphysis slightly raised. Upper and reduced in size. Teeth small; teeth in outermost row of jaw 36.6–81.5% (mean 40.7% in females, 59.5% in males) upper jaw largest, curved and pointed, placed at front of of HL. Eyes dorsolateral, forming part of dorsal profi le, jaws; two rows of smaller pointed teeth behind this row; 20.4–30.2% (mean 27.4% in females, 24.4% in males) of one or two rows of small pointed teeth at side of jaw (very HL. Snout fl attened, rounded, 24.2–35.9% (mean 28.4%) little difference in size of teeth between males and females). of HL. Interorbital narrow, 11.1–19.1% (mean 13.8%) of Lower jaw with three or four rows of small pointed teeth HL. Caudal peduncle long, compressed, length 24.7–31.3% across front, teeth in inner rows tending to point posteriorly; (mean 28.5%) of SL. Caudal peduncle depth 10.3–14.1% teeth in innermost row on side of jaw largest and stoutest (mean 12.3%) of SL. (posteriormost few teeth largest); usually one or two rows of teeth at side of jaw in females; in males, only anterior half First dorsal fi n rounded, tips of spines free, fi rst spine nearly of lower jaw with teeth, posteriormost tooth largest. always longest in males (rarely second or third spine longest), spine elongate and fi lamentous, reaching back nearly to rear Predorsal scales moderately large, evenly sized, reaching of second dorsal fi n when depressed; females with no elongate to close behind eyes; anteriormost scale broader than those fi rst dorsal fi n spines, second or third spines longest; spines behind it. Operculum with large cycloid scales, lower quarter barely reaching second dorsal fi n origin when depressed. sometimes unscaled. Cheek always naked. Pectoral base First dorsal spine 15.1–15.3% (mean 15.2%) in females, covered with few large cycloid scales. Prepelvic area covered 14.1–41.0% (mean 25.4%) in males, of SL. Second dorsal with cycloid scales. Belly midline naked, remainder covered spine length 13.2–16.8% (mean 15.0%) of SL. Third dorsal with cycloid or ctenoid scales, or with patch of ctenoid scales spine length 11.9–15.0% (mean 13.9%) of SL. Second dorsal close to pelvic fi n base and rest of scales cycloid. Ctenoid and anal fi ns short-based, posteriormost rays longest, rays scales on side of body extending forward to pectoral base, falling well short of caudal fi n base when depressed. Pectoral sometimes extending dorsally over base. Anteriormost scales fi n slender, pointed, central rays longest, 20.4–27.9% (mean on body larger than those posteriorly. 25.0%) of SL; rays usually branched but for uppermost. Pelvic fi ns slender to oval, reaching to anus, 20.2–26.6% Genital papilla in male elongate and fl attened, narrowing (mean 23.2%) of SL. Caudal fi n relatively short, rounded toward tip, one or several tiny lobes at tip; in female, papilla posteriorly, 23.8–28.9% (mean 26.4%) of SL. short, rounded and bulbous, indented at tip.

No distinct mental fraenum, chin smooth, sometimes with Head pores present (Fig. 25). Pair of nasal pores, pair of fl eshy pad behind symphysis (usually discernible in freshly anterior interorbital pores, a median posterior interorbital preserved material). Anterior nostril placed just above pore, postorbital and infraorbital pore behind each eye, preorbital edge, in short tube oriented forward. Posterior oculoscapular canal and pore over preoperculum and three nostril rounded to oval, placed very close to front centre preopercular pores. One specimen from Cambodia (MNHN margin of eye. Gill opening wide, extending forward to 1985.996) with extra pore on each oculoscapular canal, close under opercle or just under preopercle. Inner edge of pectoral to infraorbital pore.

Fig. 25. Pseudogobiopsis oligactis, papillae pattern. 36 mm SL male, CMK 5385, Ranong, Thailand. Scale bar = 1 mm.

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Sensory papillae pattern longitudinal, as in Fig. 25. Cheek of body scale margins outlined by series of small blackish rows a, c and cp composed of large, widely spaced papillae, dots. No dark midlateral blotches visible, but posterior half and rows b and d of very small, closely spaced papillae. of body with about fi ve dusky internal bars. Head (facing Two to three s rows on snout, of one papilla each; middle away from viewer) with brownish bar extending from eye row papilla absent or reduced in size. Mandibular f row of toward snout tip, some brownish marks directly below eye two papillae, one on each side of symphysis. and diagonal brown bar extending from rear of the eye across cheek. Opercle, pectoral base and pectoral fi n yellowish. Colouration of fresh material. – Gerhard Ott made available Both dorsal fi ns translucent, with rows of elongate black photographs of captive specimens from the Chong-fa and light yellow spots (two rows on fi rst dorsal fi n; four or Waterfall on the Andaman coast of Thailand (Plate 2B). The fi ve on second); greatly elongate fi rst dorsal fi n spine broad head and body are translucent pearly grey, slightly yellowish (membranous?) and yellowish white. Caudal fi n translucent to gold-speckled dorsally; belly whitish; fi ve or six dusky yellowish with 10 rows of purplish brown small spots internal blocks of pigment just visible through body wall and coalescing posteriorly into narrow lines. several indistinct dusky saddles on dorsum (most distinct on posterior half of body); scales with dark margins, on lower Larson & Lim (2005: 143) also show a live specimen. half of body, scale margins outlined with black speckles and Freshly dead specimens in available slides are similar to dense black small round spots. Head pinkish-grey on lower preserved specimens, being whitish or whitish yellow with half, dorsal half slightly yellowish to gold-speckled. First brown and/or greyish-brown body markings and blackish dorsal fi n transparent, with alternating pale creamy yellow spots on the dorsal fi ns. and faint diffuse brown bands and small black oval spot near base of fi n posteriorly, elongate fi rst dorsal spine creamy Colouration of preserved material. – Head and body yellow, fi n margin translucent. Second dorsal fi n and caudal yellowish-white, light greyish-brown to pale brownish fi n with closely spaced irregular alternating bands of pinkish (depending upon preservation), paler on ventral half, with brown and creamy yellow to pinkish, fi n margins brownish. fi ve elongate, rectangular, brown blotches along midside Anal fi n whitish. Pectoral and pelvic fi ns translucent blueish of body; scale margins on upper half of body outlined white to nearly transparent. narrowly with brown; scales on lower half of body becoming less clearly outlined, with series of tiny black or brown A photograph of a living captive specimen, identifi ed as spots (conspicuous against light background) (Fig. 26). Boleophthalmus sp., appears in Wheeler (1979: Pl. 431). Five diffuse brown saddles usually visible, crossing back Fish light yellowish-grey, with whitish breast and abdominal and reaching down to midside of body, sometimes joining region, scale margins narrowly outlined with light greyish midlateral blotches. Posteriormost midlateral blotch usually brown (darker on back) on dorsal half of body; on ventral half wedge-shaped, extending onto caudal fi n base. Nape and

Fig. 26. Pseudogobiopsis oligactis, CMK 10862, 28.8 and 34.9 mm SL, locality unknown. From colour slide by Maurice Kottelat.

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upper half of body often with indistinct brown spotting Herre (1940), in his description of Vaimosa perakensis, and mottling. Ventral midline of caudal peduncle with thin described how the species rapidly increased in abundance black line, usually with three to four evenly spaced narrow along the edges of Chanderoh Dam (Malaysia) after it was black blotches present. built.

Head with three distinct brown markings: fi rst, short broad Schneider (2003) described captive spawning of specimens bar from front of eye to upper lip but not extending onto originally collected from Santubong Peninsula, Sarawak; the lip; second, curved broad to narrow streak from lower edge habitat was described as a small stream with pebble and rock of eye to jaw, bending sharply rearward and ending on substrate. In captivity, the male constructed burrows under middle of cheek; and third, broad diagonal bar from rear stones and became golden-yellow in colour with darker of eye extending backward to end of middle of cheek just scale margins (dark saddles and lines on the body vanish), a above second streak; third bar usually darkest and broadest. violet hue on the head and bright red fi ns. A ready to spawn Opercle with few diffuse brown spots, usually dark brown female had a pale body with black pelvic fi ns. About 1,000 irregular marking present near upper rear corner. Lips and eggs were deposited on the roof of the spawning cave and chin plain dusky grey to brown. Pectoral base with blackish larvae were planktonic. to brown spot or blotch across bases of uppermost few fi n rays. Breast and belly pale to dusky. Horsthemke (in litt.) observed that captive female specimens often moved up into the water column and rested on top of First dorsal fi n translucent to whitish with two to three rows large stones or on the leaves of aquatic plants, while a male of elongate black spots on spines and membranes (spots often was strictly benthic (and cryptic in behaviour). They ate live joining to form irregular rows); fi lamentous portion of fi rst food, but nothing larger than chironomid larvae. spine plain whitish. Second dorsal fi n similar, with four to seven rows of blackish spots coalescing on posterior half of Remarks. – Stigmatogobius neglectus Koumans is based fi n at least, forming wavy rows. Anal fi n plain dusky grey. on a Bleeker manuscript name, Pseudogobius neglectus, Caudal fi n translucent to dusky with 5–10 rows of small and listed as name only in Koumans (1931) and Bleeker elongate dusky spots or fi ne irregular dusky bars. Pectoral fi n (1983). Bleeker (1983: Pl. 438, Fig. 13) indicated that the translucent to dusky. Pelvic fi ns translucent dusky, darkest black spot drawn on the fi rst dorsal fi n on the fi gure of toward fi n bases. Pseudogobius neglectus was erroneous. Kottelat et al. (1993: pl. 70) reproduced this fi gure without comment. Comparisons. – This species was found to be frequently been confused with Pseudogobiopsis siamensis in museum Koumans (1940: 126) fi rst recognised that Glossogobius collections, as the two species can occur syntopically and campbellianus was a synonym of P. oligactis. However, he small female specimens look superficially similar. The also thought that Vaimosa siamensis was also a synonym headpore arrangement (preopercular pores and rear portion (Koumans 1940: 135), which it is not. of oculoscapular canal and pore present in P. oligactis, but absent in P. siamensis) and caudal fi n colour pattern (vertically Kottelat et al. (1993) combined P. oligactis and P. siamensis oriented rows of brown spots in P. oligactis vs. similarly under their account for Pseudogobiopsis campbellianus. The oriented grey to brown zigzag lines in P. siamensis) are specimens in their Fig. 70 are P. siamensis; and the batch of the simplest means of distinguishing the two. Additionally, specimens from which they came (CMK 7864) included both P. oligactis tends to be longer bodied with a somewhat species. They also listed this species (as Pseudogobiopsis fl attened head (especially in adult males), while P. siamensis neglectus) as a “... species to watch ...” due to its apparent is stockier, with the head depth at the preopercle greater rarity and lack of biological information about it (Kottelat than the width. et al., 1993: xxxvi).

Distribution. – Specimens are known from Thailand through Roberts (1993: 44, Fig. 51) published one of Kuhl & van the Indo-Malayan Archipelago to Indonesia. Two specimens Hasselt’s illustrations, of a fi sh from Java (artist unknown) are known from the Philippines (but see Remarks). Ng & which could be Pseudogobiopsis oligactis. It is relatively Tan (1999) reported the species for the fi rst time from the slender, has an elongate fi rst dorsal fi n spine, elongate black Endau River system in southern Peninsular Malaysia. blotches along the midside of the body, small brown spots on the whitish lower half of the body and fi nely barred This species appears to have become extinct in Singapore, caudal fi n. However, the mouth is not enlarged and there is with the only known specimens collected in 1959. Despite a black blotch on the pectoral base, features which resemble recent surveys of most habitats on the island by resident that of specimens of a probably new species not included biologists, P. oligactis has not been found since (Larson et within this paper. al., 2008). There appear to be some dwarf (i.e. sexually mature at a small Ecology. – Freshwater, usually from fast-flowing hill size) populations of this species. Thirteen specimens from streams, out of the main current. Syntopic with P. siamensis Borapet Swamp, Nakornsawan Province, Thailand (NTM in Malaysia. S.14241-001), include 23 mm SL males with enlarged mouths and a 19 mm SL female with nearly ripe eggs. Additionally,

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the only specimens from the Philippines which are of this 14, Sungai Piyam, Nanga Empanang, Kalimantan Barat; CMK species are two very small mature fi sh (21 mm SL male, 11781, 1, Sungai Barito, Kalimantan Tengah; MHNG 2537.36, 2, 17.5 mm SL female) from the Basey River, Samar (CMK N of Sempit, Kalimantan Tengah; CMK 9674, 1, Sungai Serdang, 9978). The male has an enormous mouth reaching to the Sumatra, Riau Province. rear of the preopercle and the female is fi lled with ripening ova. Both fi sh are very dark in colour (chocolate-brown, Diagnosis. – Slender Pseudogobiopsis, somewhat rounded with whitish scale centres) and show some indistinct vertical anteriorly, with second dorsal rays I,7–10 (modally I,8); anal brown bars on the side of the body. It is possible that these rays I,6–7 (modally I,7); pectoral rays 13–16; longitudinal two specimens belong to a separate species (although basic scales 23–27; TRB 7–9; predorsal scales 5–9, small, counts agree with P. oligactis), but additional material is reaching to above preopercular margin; headpores absent; required to confi rm this. mouth enlarged in males; scales on body mostly ctenoid; caudal fi n pointed; head and body yellow to whitish with scattered brown spots and fi ve brown bars crossing back Pseudogobiopsis paludosus (Herre, 1940) and sides, caudal and dorsal fi ns with rows of fi ne black (Figs 27–30; Tables 13–16, 19) spots; single very broad epural; known from fresh waters of Indo-Malaysian Archipelago. Ctenogobius paludosus Herre, 1940: 23, Pl. 18 (north of Kota Tinggi, Johore, Malay Peninsula). Description. – Based on 23 specimens, 16–30.5 mm SL. Calamiana sp. undet. – Roberts 1989: 168. An asterisk indicates counts of holotype of Ctenogobius Rhinogobius paludosus – Kottelat 1989: 19. paludosus (Fig. 27). Eugnathogobius paludosus – Larson 2001: 70. First dorsal V (in one), VI* (in 21), VII (in one); second dorsal Material examined. – MALAYSIA: Holotype of Ctenogobius I,7–10 (mean I,8*); anal I,6–7 (mean I,7*), pectoral rays paludosus , CAS 32998, 1(30.5), 5 miles (8 km) north of Kota Tinggi, 13*–16 (mean 15), segmented caudal rays 15–17*; caudal Johore, A. W. Herre, May 1937. ZRC 8411, 1(30), Sungai Sedili, Johore, 22 Feb.1968; CMK 7384, 8(12.5–29.0), Sungai Mupor, ray pattern 6/6 to 8/8 (modally 7/6*); branched caudal rays Johore, M. Kottelat, P. K. L. Ng & K. Lim, 22 Jan.1991; NTM 12–16 (mean 14, 13 in holotype); unsegmented (procurrent) S.16416-001, 2(18–25), C&S, same data as previous. INDONESIA: caudal rays 5/5 to 6/5; longitudinal scale count 23-27 (mean USNM 230332, 3(18–23), West Kalimantan, Sungai Gentu, near 25, 24 in holotype); TRB 7*–9 (mean 8); predorsal scale confl uence with Kapuas River, T. Roberts, 16 Aug.1976; CAS count 5–9 (mean 8*); circumpeduncular scales always 12*. 49462, 58(7.5–28.0), Sungai Mandai Kecil, near confl uence with Gill rakers on outer face of fi rst arch 1+6 to 3+6 (modally Kapuas River, 18 km west-south-west of Putissibau, T. Roberts and 2+6). Pterygiophore formula 3-12210 (in eight). Vertebrae S. Woerjoatmodjo, 1 Aug.1976; CMK 9009, 5(13–21), Sumatra, 10+15 (in three), 10+16 (in nine), 10+17 (in one). Neural Riau district, Sungai Kalesa, north of Seberida, A. J. Whitten, 22 spines of fi rst few vertebrae slender, pointed (in 10). One Feb.1992. (in 10) very broad epural; two in one specimen. Two (in Other material examined (but not used in description). 11) anal pterygiophores before haemal spine of fi rst caudal MALAYSIA: ZRC 8405, 1, by Gunung Panti, Johore; ZRC vertebra. Palatine not in contact with quadrate. Pterygoid 17095, 1, by Gunung Panti, Johore; ZRC 14011, 1, Kota Tinggi. long, equal to or longer than palatine. Metapterygoid low INDONESIA: USNM 230333, 7, Kapuas River tributary, Borneo; to moderate, well separate from quadrate in female, slightly USNM 230331, 7, Sungei Seriang, Sungei Palin tributary; CMK overlapping quadrate in adult male (suspensorium shifted 6933, 1, Sungai Sibau, Kalimantan Barat; CMK 6761, 3, Kapuas back and shortened by enlarged jaws) (Fig. 28). Quadrate River tributary, Kalimantan Barat; CMK 11598, 1, Sungai shallowly forked. Dentary short, narrow. Pectoral radials Letang, Kapuas River tributary, Kalimantan Barat; CMK 10141, poorly ossifi ed. Scapula unossifi ed.

Fig. 27. Pseudogobiopsis paludosus. Holotype of Ctenogobius paludosus Herre, CAS 32998, 30.5 mm SL male, Kota Tinggi, Johore, Malaysia.

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Body slender, compressed, more rounded anteriorly. Body First dorsal fi n triangular, pointed, third to fi fth spines longest depth at anal fi n origin 15.4–19.5% (mean 17.1%) of SL. or subequal, sometimes elongate, especially in large males Head rounded to slightly depressed, wider than deep, but (fi rst dorsal spine longest in one female), second or third not greatly so, HL 30.5–33.7% (mean 31.9%) of SL; cheeks spine longest in females, third modally longest in males; may be considerably infl ated in males. Depth at posterior spines slightly longer in males than females; spines usually preopercular margin 47.8–56.9% (mean 51.7%) of HL. Width reaching second dorsal fi n origin when depressed. Second at posterior preopercular margin 56.9–71.6% (mean 65.3%) dorsal spine longest in some females, length 15.3–17.3% of HL. Mouth terminal, lower jaw tip anteriormost, oblique, (mean 16.5%) of SL. Third dorsal spine length 14.0–15.7% forming angle of about 25–30° with body axis; jaws reaching (mean 15.1%) of SL. Fourth dorsal spine length in males past rear of eye in males and to below front half of eye in 15.0–27.3% (mean 22.8%) of SL. Fifth dorsal spine length females (well past rear of eye in holotype). Lips smooth, in males 19.2-35.0% (mean 25.8%) of SL. Second dorsal and usually narrow in males; without fl eshy fi mbriae along edge; anal fi ns short-based, pointed posteriorly, rays falling well lower lip free at sides, broadly fused across front. Upper short of caudal fi n base when depressed. Pectoral fi n pointed, jaw 32.1–62.5% (mean 34.3% in females, 52.4% in males) slender, central rays longest, 23.3–31.3% (mean 26.5%) of of HL. Eyes dorsolateral, close to snout tip, high on head SL; rays usually all branched. Pelvic fi ns long, oval, usually forming part of dorsal profi le, 22.1–35.2% (mean 29.3%) reaching to anus, 20.0–25.9% (mean 23.8%) of SL. Caudal of HL. Snout short, fl attened, 17.7–27.4% (mean 23.7%) of fi n slender, pointed, 25.2–34.8% (mean 29.7%) of SL. HL. Interorbital fl at, moderate to narrow, 8.5–22.7% (mean 18.8% in males, 14.4% in females) of HL. Caudal peduncle No mental fraenum, chin blunt, smooth, slightly protuberant. long, compressed, length 22.1–30.1% (mean 26.6%) of SL. Anterior nostril in very slender tube, placed just at preorbital Caudal peduncle depth 10.0–12.9% (mean 11.7%) of SL. edge, tube oriented forward and slightly downward, preorbital

Fig. 28. Jaws and suspensorium of Pseudogobiopsis paludosus. A, male, ex CAS 7384, Sungei Mupor, Johore, Malaysia; B, female, CMK 6735, Kalimantan Barat, Borneo (specimens from this lot possibly represent a separate species). Scale bar = 1 mm.

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occasionally produced forward to accommodate nostril. Female genital papilla quite large, swollen and rounded, with Posterior nostril small, oval, placed about midway between two lobes at tip (size probably due to large size of ripe ova). front centre margin of eye and anterior nostril. Gill opening Male genital papilla small, fl at and pointed. extending forward to just under opercle. Inner edge of pectoral girdle with smooth narrow edge (in 14) or with smooth bony Head pores absent. fl ange (in nine) which may be angled outward, rarely with low fl at lobes or fl aps present. Gill rakers on outer face of Sensory papillae pattern longitudinal, as in Fig. 29. Papillae fi rst arch usually short knobs, occasionally slender or reduced, in row p few, widely spaced. Papillae in cheek rows a, c shortest rakers on upper limb; rakers on inner face of fi rst and cp enlarged, widely separated; other papillae very small arch of similar size but stouter; outer and inner rakers on and fi ne. Cheek row b broken into two short sections, with other arches similar to those of fi rst arch. Tongue long, tip wide gap. Three s rows of papillae on snout, of one papilla blunt to rounded, occasionally slightly pointed. each. Mandibular f row of one papilla on each side. Papillae in row i along dentary few, widely separated. Upper jaw teeth in male: teeth in outermost row, across front of jaw to midside of jaw, large, sharp and curved, largest teeth Colouration of fresh material. – Colour slide by Maurice grouped at midside of jaw, rear third or less of jaw without Kottelat, of freshly dead specimens, showing colour very teeth; inner three or four rows consisting of small, curved, like that of preserved material: pale yellowish-white with sharp teeth. Upper jaw teeth in female: about three rows of blackish to dark brownish grey markings on body and small curved sharp teeth across front; one row of teeth at side blackish markings on fi ns. of jaw. In males, lower jaw with band of fi ve to seven rows of curved sharp teeth across front, band broader toward side Colouration of preserved material. – Head and body of front of jaw and one or two large curved outer teeth at yellowish-white to pale yellowish-brown, covered with small each side. Lower jaw teeth arrangement similar in females, dark brown round spots and fi ve oblique to nearly vertical but with four or fi ve rows of curved sharp teeth across dark brown bars crossing back and sides, evenly spaced front of jaw; one or two slightly enlarged teeth outermost, from just before fi rst dorsal fi n origin to just before upper toward each side of lower jaw. Tips of teeth in both sexes procurrent rays of caudal fi n (Fig. 30). Oblique bars usually sometimes tinted translucent orange or brown. ending on lower side of body, often intensifi ed on midside of body forming row of square dark brown blotches, slightly Predorsal scales cycloid, moderately sized, reaching forward offset from oblique bar. Dark brown spots on body of variable to at least preopercular margin or to between preopercular intensity, sometimes very fi ne and partly outlining scale margin and rear of eyes. Operculum naked or with one to few margins; on side of head, spots usually large and rounded. cycloid scales. Cheek always naked. Pectoral base naked or At midbase of caudal fi n, triangular to oblique elongate dark with one or two cycloid scales. Prepelvic area usually with brown blotch present; if oblique, blotch extending down and few cycloid scales, occasionally naked. Belly with cycloid back onto lower segmented caudal rays. Underside of head scales, occasionally naked. Side of body with ctenoid scales and belly plain pale whitish to pale brownish; small patch extending up to behind pectoral base. Anteriormost scales of dark brown spots on breast just anterior to pelvic fi ns in on side of body larger than those on caudal peduncle. males. Ventral midline of caudal peduncle with thin black line and three (occasionally four) narrow black blotches, Gut short, S-bend shape.

Fig. 29. Pseudogobiopsis paludosus, papillae pattern. 29 mm SL male, CMK 7384, Johor, Malaysia. Scale bar = 1 mm.

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evenly spaced, from midbase of anal fi n to just before lower Comparisons. – This species is most similar to P. festivus, procurrent rays of caudal fi n. new species from Sarawak; see Comparisons under that species. There may be another undescribed species related Side of head with two short oblique dark brown bars from to this species-group, see Remarks below. rear of eye extending down and backward, ending on middle of cheek; and one bar extending from lower front margin of Distribution. – Specimens are known from Peninsular eye to middle of jaws, crossing jaws to meet its counterpart Malaysia, western Borneo and the Riau Archipelago, on chin; bars blotchy and often nearly indistinguishable Sumatra. from small dark brown spots covering head. Opercle with dark brown spots often coalescing into dark blotches. Lips Ecology. – Known from freshwater habitats, including nearly uniform brown or with small dark brown spots. Nape clear, swift “blackwater” (brown to reddish tinted) streams and interorbital with small brown blotches and small spots. (Roberts, 1989). Pectoral base pale with scattered dark brown round spots. Roberts (1989) gives information on reproductive condition First dorsal fi n translucent to whitish, with broad dark grey of the specimens obtained during the Kapuas River survey, to black band across centre, intense black spot within band e.g. a 19.5 mm female specimen (out of CAS 49462) had toward rear of fi n usually present, and short narrow oblique 30 eggs, of 1.5 mm diameter. blackish streak from base of fi rst dorsal fi n crossing bases of fi rst three spines; fi rst dorsal spine always pale with two Remarks. – This species was described from “the type and small dark brown to black spots (regardless of intensity of only specimen, 30 mm. long” (Herre, 1940a). His illustration blackish bands on rest of fi n). Second dorsal fi n whitish to of the type, despite its simplicity, gives a good impression translucent, with three to fi ve rows of small dark brown to of the species, and it is apparently the only published black round to oval spots; unsegmented spine always pale illustration of the species. A specimen presently labelled as with two black spots. Anal fi n plain dusky to brownish, paratype (CAS 32988) is a species of Pseudogobius and is darker toward base. Pectoral fi ns translucent dusky to pale not a type specimen. brownish, rays darkest at bases. Pelvic fi ns plain dusky grey to brownish. Caudal fi n translucent whitish to pale dusky Twenty small specimens from Sanggau and Sintan on the with fi ve to seven rows of vertically oriented, dark brown to Kapuas River drainage in western Kalimantan (CMK 6735, black small spots or short lines; spots or short lines closest CMK 6799 and CAS 49461) were initially referred to this to fi n base sometimes joining to form one or two wavering species. However, they have some headpores, but are lacking vertical dark lines. preopercular canals and the posterior portion of oculoscapular canal. Three very small specimens in CMK 6735 have no Sensory papillae often orange-tipped. Peritoneum pale with pores or exhibit open canals only. All these specimens are broad brown dorsal saddle and scattered brown spots (as in similar in that the colour pattern shows narrow brown scale colour pattern of surrounding skin). margins over the pale back and sides, forming an irregular

Fig. 30. Pseudogobiopsis paludosus, CMK 7384, 20 and 29.5 mm SL, Johor, Malaysia. From colour slide by Maurice Kottelat.

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network pattern, and the fi ve brown saddles and lateral bars 2) anal pterygiophores before haemal spine of fi rst caudal are rather indistinct. Additional material, including mature vertebra (in 9). males, is required to determine the status of these specimens, which probably represent an undescribed species. These Body slender, compressed. Body depth at anal fi n origin specimens are not dealt with further in this paper. 17.1–18.6% (mean 17.9%) of SL. Head slightly depressed, wider than deep, but not greatly so, HL 25.8–31.6% (mean 28.7%) of SL. Depth at posterior preopercular margin 51.5– Pseudogobiopsis tigrellus (Nichols, 1951) 61.3% (mean 58.4%) of HL. Width at posterior preopercular (Fig. 31; Plate 2C; Tables 13–16, 20) margin 58.8–75.8% (mean 69.3%) of HL. Mouth subterminal, very slightly oblique, forming an angle of about 12° with Gobius tigrellus Nichols, 1951: 3, Fig. 2 (Bernhard Camp, Idenburg body axis; jaws reach to below just past middle of eye in River, West New Guinea). – Larson 2001: 71–72, Fig. 68 (as males and to below front half of eye in females. Lips fl eshy, incertae sedis). smooth. Upper jaw 34.3–47.2% (mean 35.0% in females, Ctenogobius tigrellus – Allen 1996: 19. 44.2% in males) of HL. Eyes dorsolateral, forming part of dorsal profi le, 25.0–29.9% (mean 27.7%) of HL. Snout short, Material Examined. Holotype, AMNH 18574, 1(21.5), 75 m altitude, Bernhard Camp, Idenburg River [tributary of Mamberamo rounded to slightly pointed, 25.4–30.8% (mean 27.7%) of River], West New Guinea, W. B. Richardson, Apr.1939. Paratypes, HL. Interorbital narrow, 8.3–13.3% (mean 10.2%) of HL. AMNH 15096, 9(16–22), same data as holotype. WAM P.31752- Caudal peduncle slender, compressed, length 25.6–30.8% 001, 6(21.0–25.5), Furu Creek, Mamberamo River, West Papua, (mean 28.5%) of SL. Caudal peduncle depth 11.4–13.5% 0.1–1.0 m, G.R. Allen, 4 Sep.2000; WAM P.31754-003, 1(24.5), (mean 12.0%) of SL. Dabra, Tiri Creek, Mamberamo River, West Papua, 0.1–1.0 m, G. R. Allen, 7 Sep.2000. First dorsal fi n triangular in males, triangular to slightly rounded in females, tips of spines free; spines slightly longer Description. – Based on eight specimens, 21.0–25.5 mm SL; in males than females; fi n barely reaches second dorsal osteological information from x-rays of type specimens. An origin when depressed. First dorsal spine always longest, asterisk indicates the counts of the holotype. spine length 12.2–16.7% (mean 14.2% in females, 15.7% in males) of SL. In females, second dorsal fi n short and First dorsal VI*; second dorsal I,7*; anal I,6–7* (modally triangular, anterior rays longest, posteriormost rays fall well I,6), pectoral rays 14*–16 (modally 15), segmented caudal short of caudal fi n base when depressed; in males second rays 17*; caudal ray pattern 9/8; branched caudal rays 14–15, dorsal fi n triangular, anterior rays tall but posteriormost rays in 7/7 pattern (broken in holotype); longitudinal scale count longer (about twice height of anteriormost ray), elongate and 23–25 (24 in holotype); TRB 7–8*; predorsal scale count reaching to caudal fi n base in two largest male specimens. 2–9 (1 in holotype); circumpeduncular scales 12*. Gill rakers Anal fi n rounded, posteriormost rays longest, especially in on outer face of fi rst arch 2+6 (in 1), 2+7 (in 1). Vertebrae males, but always falling short of reaching caudal fi n base. 10+16 (in 9), 10+15 (1). Neural spine of second vertebra Pectoral fi n slender, pointed, central rays longest, 20.0–29.4% slightly expanded at tip (in 2), or all pointed (in 1). Two (mean 26.2%) of SL; rays all branched. Pelvic fi ns oval, epurals (in 7); nearly fused in two. One to three (modally reaching to anus, 22.8–28.6% (mean 25.9%) of SL; frenum

Fig. 31. Pseudogobiopsis tigrellus, 25.2 mm SL male, WAM P.31752-001, Mamberamo River, West Papua, showing headpore and papillae pattern. Scale bar = 1 mm.

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Plate 2. A. Pseudogobiopsis festivus, new species, captive specimen from Bako National Park, Sarawak. Photograph by Peter Ng. B. Pseudogobiopsis oligactis, captive specimen from Andaman coast, Thailand. Photograph by Gerhard Ott. C. Pseudogobiopsis tigrellus, captive male specimen from Furu Creek, Mamberamo River, West Papua (one of WAM P.31752-001). Photograph by Gerry Allen.

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fl eshy, especially around pelvic spines. Caudal fi n oval to vermiculate and broken posteriorly, fi n margin narrowly slightly pointed, 12.2–16.7% (mean 14.7%) of SL. dense black, post-margin region plain yellowish white. In female, fi rst dorsal fi n similar but with four blackish irregular No mental fraenum, chin smooth to slightly raised. Anterior lines and broad blackish margin and dense black double spot nostril tubular, placed just behind upper lip, tube short, at rear of fi n. Second dorsal fi n pattern in males similar to oriented upward. Posterior nostril oval, placed close to that of fi rst dorsal fi n but fi n margin more diffusely blackish anterior margin of eye, above level of centre of eye. Gill and irregular lines more oblique and broken, especially on opening extends forward to under opercle. Inner edge of posterior half of fi n; in females, second dorsal fi n similar shoulder girdle smooth. Gill rakers on outer face of fi rst to fi rst dorsal fi n in that fewer dark lines present. Caudal arch very short and stubby, longest rakers near angle of arch. fi n transparent, rays yellowish, rear margin of fi n slightly Tongue tip blunt to slightly concave. Teeth in upper jaw in dusky, 8–10 vertical blackish lines crossing fi n, basalmost three or four rows, outer row teeth largest and stoutest (in line broadest and most intense, forming narrow blotch (within males especially); all teeth small, sharp and conical; largest line) at mid-base of fi n; ventralmost quarter of fi n unmarked. tooth at mid-side of jaw. Lower jaw small and sharp, in Anal fi n plain whitish yellow. Pectoral fi n transparent. Pelvic broad band of three to fi ve rows across front of jaw; side fi ns transparent, dusted lightly with melanophores. of jaw with two rows of slightly more upright sharp teeth; outermost row of teeth across front of jaw largest, stoutest Colouration of preserved material. – Preserved colour is and more curved. basically the same as live colour, with background colour yellowish white and brown to dark brown body markings, Predorsal scales small, cycloid; arrangement variable (one scale margins on upper half of body thinly outlined with fi ne fi sh with scales right up to behind eyes, others have scales dark speckling, dorsal fi n markings black to dark brown, up to above rear margin of preopercle at least on the side, caudal fi n barring dark brown. with only a few scales on midline). Operculum with one to several cycloid scales, usually partly embedded. Cheek, Comparisons. – The species resembles Pseudogobiopsis pectoral base and prepelvic area naked. Belly midline broadly oligactis in possessing the posterior portion of the naked. Side of body covered with ctenoid scales; scales on oculoscapular canal above the preopercle; the two species midbody slightly larger than others. are more similar to each other than other species of the genus. Its strongly barred colour pattern is distinctive, and Head pores as for Pseudogobiopsis oligactis, with three is unlike that of any other related species. preopercular pores (Fig. 31). Distribution. – Known only from the Mamberamo River Sensory papillae pattern with cheek rows a, c and cp system of West Papua. composed of few large, widely spaced papillae; rows b and d of small, closely spaced papillae (Fig. 31). Two s rows Ecology. – The species has been found only in “… small, on snout, of one papilla each, just behind upper lip, other clear, relatively fast-fl owing rainforest (mostly closed-canopy papillae close together by posterior nostril. Mandibular forest) creeks. The fi sh were invariably found away from papillae widely spaced, with mental f row consisting of one the main fl ow, around the edge of small pools to about 10 papilla on each side of symphysis. cm depth. They were seen solitary or in loose groups of 2–4 individuals, generally resting on sand or rock substrate” Colouration of fresh material. – Based on scans of colour (Jerry Allen, in litt.) slides of captive male and female by Gerry Allen (Plate 2C). Head and body translucent pearly grey, with 12–14 black to Remarks. – The holotype and nine paratypes have been blackish narrow vertical bars along side of body and posterior dehydrated at some stage and are not in the best condition, half of head, dorsum with short blackish lines and blotches so that at fi rst it was diffi cult to be sure as to what genus partly following scale margins, predorsal region with irregular they actually belonged (Larson, 2001). The capture of fresh black blotches and lines; side of body dusted with orange specimens by Jerry Allen provided the key to the identity and pink-gold speckles, most noticeable in between vertical of this species. black bars. Side of head with two oblique black lines crossing cheek from ventral edge of eye; opercle with vertical black bar posteriorly (continuation of anteriormost vertical bar on ACKNOWLEDGEMENTS nape) and two black blotches anteriorly (ventralmost is a blotch in male and oblique line in female, a continuation of My many thanks to curators and staff at the institutions who dorsalmost oblique line from eye). Interorbital and snout with loaned specimens, provided photographs and information: irregular blackish lines and spots and intensifi cation of the Jerry Allen, Conservation International (formerly of WAM); orange and pink-gold speckles. Iris silvery, becoming pale Eric Anderson, SAIAB (formerly RUSI); Dave Catania, CAS; golden dorsally, and speckled with brown. Tips of lips dull I-Shiung Chen, National Taiwan Ocean University, Keelung; yellowish with dusky blotches extending from snout tip. Barry Chernoff, formerly of FMNH; G. Duhamel, MNHN; Bill Eschmeyer, CAS; Phil Heemstra, SAIAB (formerly Dorsal fi ns translucent yellowish-white with black markings. RUSI); Masayoshi Hayashi, YCM; Doug Hoese, AMS; In male, fi rst dorsal fi n with 5–6 irregular lines, becoming Hans Horsthemke, Germany (especial thanks for German

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translations and information); J.-C. Hureau, MNHN; Zeehan Fowler, H. W., 1938. A list of the fi shes known from Malaya. Jaafar, ZRC; Susan Jewett, formerly of USNM; Jeff Johnson, Fisheries Bulletin, 1: 1–268. QM; Maurice Kottelat, Cornol, Switzerland; Friedhelm Herre, A. W. C. T., 1936. Eleven new fishes from the Malay Krupp, SMF; Kelvin Lim, ZRC; Mark McGrouther, AMS; Peninsula. Bulletin of the Raffles Museum, Singapore, 12: Nigel Merrett, formerly of BMNH; Gary Nelson, formerly 5–16. of AMNH; Peter Ng, ZRC; Sue Morrison, WAM; Martien Herre, A. W. C. T., 1940. New species of fi shes from the Malay van Oijen, RMNH; Gerhard Ott, Flensburg, Germany; H.-J. Peninsula and Borneo. Bulletin of the Raffles Museum, Paepke, formerly of ZMB; Gianluca Polgar, Università degli Singapore, 16: 5–26. Studi di Roma “La Sapienza”; Sandra Raredon, USNM; Herre, A. W. C. T., 1945. Two new genera and four new gobies Ukkrit Satapoomin, Biological Research Centre, Phuket; from the Philippines and India. Copeia, 1945 (1): 1–6. Bill Smith-Vaniz, formerly of ANSP; Heok Hui Tan, ZRC; Herre, A. W. C. T., 1950. Two new gobies from the Philippines Chavalit Vidthayanon, formerly NIFI; H. Wilkens, ZMH; Rex with notes on a third rare goby. Proceedings of the Biological Williams, NTM; Rick Winterbottom, ROM; Kai-Erik Witte, Society of Washington, 63: 73–76. University of Konstanz; Prachit Wongrat, KUMF; Thosaporn Herre, A. W., 1953. Check list of Philippine fi shes. United States Wongratana, Chulalongkorn University, Bangkok; Dr Wu Fish and Wildlife Service Research report, 20: 1–977. Han-Ling, Shanghai University; Tetsuo Yoshino, URM. Herre, A. W. C. T. & G. S. Myers, 1937. A contribution to the ichthyology of the Malay Peninsula. Bulletin of the Raffl es Museum, Singapore, 13: 5–75. LITERATURE CITED Hora, S. L., 1923. Fauna of the Chilka Lake. Fish (Part V). Memoirs of the Indian Museum, 5: 737–769. Alfred, E. R., 1966. The fresh-water fi shes of Singapore. Zoologische Verhandelingen, 78: 1–68. Hubbs, C. L. & K. F. Lagler, 1970. Fishes of the Great Lakes Region. University of Michigan Press: Ann Arbor. Allen, G. R., 1996. In: Kitchener, D. J. & Suyanto, A. (eds.) Proceedings of the fi rst international conference on eastern Jordan, D. S. & A. Seale, 1908. List of fi shes collected in the river Indonesian-Australian vertebrate fauna, Manado, Indonesia, at Buytenzorg, Java, by Dr. Douglas Houghton Campbell. November 22–26, 1994. Pp. 15–21. Proceedings of the U.S. National Museum, 33(1575): 535– 543. Aurich, H. J., 1938. Mitteilung XXVIII der Wallacea-Expedition Woltereck. Die Gobiiden. (Ordnung: Gobioidea). Internationale Kottelat, M., 1989. Zoogeography of the fi shes from Indochinese Revue der gesamten Hydrobiologie und Hydrographie, 38(1/2): inland waters with an annotated check-list. Bulletin Zoölogisch 125–183. Museum, 12(1): 1–56. Birdsong, R. S., E. O. Murdy & F. L. Pezold, 1988. A study of the Kottelat, M. & K. K. P. Lim, 1995. Freshwater fi shes of Sarawak vertebral column and median fi n osteology in gobioid fi shes and Brunei Darussalam: a preliminary annotated checklist. The with comments on gobioid relationships. Bulletin of Marine Sarawak Museum Journal, 48(69): 227–256. Science, 42(2): 174–214. Kottelat, M., A. J. Whitten, with S. N. Kartikasari & S. Wirjoatmodjo, Bleeker, P., 1875. Gobioideorum species insulindicae novae. 1993. Freshwater fi shes of Western Indonesia and Sulawesi. Archives Néerlandaises des Sciences exactes et naturelles, Periplus Editions Ltd: Indonesia. 10: 113–134. Koumans, F. P., 1931. A preliminary revision of the genera of the Bleeker, P., 1983. Atlas Ichthyologique des Indes Orientales gobioid fi shes with united ventral fi ns. Proefschrift Drukkerij Néêrlandaises, par M.- P. Bleeker. (Plates originally planned Imperator N.V. Lisse. Pp. 1–174. for planned tomes XI-XIV published here for the fi rst time). Koumans, F. P., 1932. Notes on gobioid fi shes (1–5). Zoologische Smithsonian Institution Press: Washington. Mededeelingen, 15: 1–16. Böhlke, E. B., 1984. Catalog of type specimens in the ichthyological Koumans, F. P., 1935. Notes on gobioid fi shes. 6. On the synonymy collection of the Academy of Natural Sciences, Philadelphia. of some species from the Indo-Australian Archipelago. The Academy of Natural Sciences of Philadelphia Special Zoologische Mededeelingen, 18: 121–150. Publication, 14: 1–246. Koumans, F. P., 1940. Results of a reexamination of types and Chatterjee, T. K., 1980. Record of Stigmatogobius hoevenii (Bleeker) specimens of gobioid fi shes, with notes on the fi shfauna of from the Gangetic Delta, west Bengal, with a key to the Indian the surroundings of Batavia. Zoologische Mededeelingen, 22: species of Stigmatogobius. Bulletin of the Zoological Survey 121–210. of India, 2(2&3): 229–231. Koumans, F. P., 1953. X. Gobioidea. In: Weber, M. & Beaufort, Eschmeyer, W. N. & R. M. Bailey, 1990. Genera of recent fi shes. L.F. de. (eds.). The Fishes of the Indo-Australian Archipelago. In: Eschmeyer, W.N. Catalog of the genera of recent fi shes. E. J. Brill: Leiden. 423 pp. Pp. 7–433. California Academy of Natural Sciences: San Larson, H. K., 1995. A review of the Australian endemic gobiid fi sh Francisco. genus Chlamydogobius, with description of fi ve new species. Fowler, H. W., 1934. Zoological results of the third de Schauensee The Beagle, Records of the Museums and Art Galleries of the Siamese Expedition, Part 1 – Fishes. Proceedings of the Academy Northern Territory, 12: 19–51. of Natural Sciences of Philadelphia, 86: 67–163. Larson, H. K., 1999. Allocation to Calamiana and redescription of Fowler, H. W., 1937. Zoological results of the third de Schauensee the fi sh species Apocryptes variegatus and Vaimosa mindora Siamese Expedition. Part VIII, – Fishes obtained in 1936. (Gobioidei: Gobiidae: Gobionellinae), with description of a new Proceedings of the Academy of Natural Sciences of Philadelphia, species. Raffl es Bulletin of Zoology, 47(1): 257–281. 89: 125–264.

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Larson, H. K., 1999a. Report to Parks Australia North, on estuarine Pezold, F., 1993. Evidence for a monophyletic Gobiinae. Copeia, fi sh monitoring of Kakadu National Park, Northern Territory, 3: 634–643. Australia. Unpublished report. 35 pp. Randall, J. E. & K. K. P. Lim, 2000. A checklist of the fi shes of Larson, H. K., 1999b. Report to Parks Australia North, on the the South China Sea. Raffl es Bulletin of Zoology, Supplement estuarine fi sh inventory of Kakadu National Park, Northern No. 8: 569–667. Territory, Australia. Museum and Art Gallery of the Northern Roberts, T. R., 1989. The freshwater fi shes of Western Borneo Territory Research Report 5: 1–50. (Kalimantan Barat, Indonesia). Memoirs of the California Larson, H. K., 2000. Report to Parks Australia North, on estuarine Academy of Sciences, 14: 1–210. fi sh monitoring of Kakadu National Park, Northern Territory, Roberts, T. R., 1993. The freshwater fi shes of Java, as observed by Australia. Unpublished report. 26 pp. Kuhl and van Hasselt in 1820–23. Zoologische Verhandelingen, Larson, H. K., 2001. A revision of the gobiid genus Mugilogobius 285: 1–94. (Pisces: Teleostei), and its systematic placement. Records of the Sanzo, L., 1911. Distribuzione delle papille cutanee (organi Western Australian Museum, Supplement No. 62: 1–233. ciatiformi) e suo valore sistematico nei Gobi. Mitteilungen aus Larson, H. K., 2002. Report to Parks Australia North, on estuarine der Zoologischen Station zu Neapel, 20: 249–328. fi sh monitoring of Kakadu National Park, Northern Territory, Schneider, M. von., 2003. Ein grosmaul aus Sarawak Eugnathogobius Australia. Unpublished report. 42 pp oligactis. Aquaristik-Fachmagazin & Aquarium heute 35(4): Larson, H. K. & K. P. Lim, 2005. A guide to gobies of Singapore. 66–68. National Science Centre, Singapore. Smith, H.M., 1931. Descriptions of new genera and species of Larson, H. K., Jaafar, Z. & K. P. Lim, 2008. An annotated checklist Siamese fi shes. Proceedings of the U.S. National Museum, of gobioid fi shes of Singapore. Raffl es Bulletin of Zoology, 79(2873): 1–48. 56(1): 135–155. Smith, H.M., 1945. The fresh-water fi shes of Siam, or Thailand. Lim, K. K. P. & H. K. Larson, 1994. A preliminary checklist of Bulletin of the United States National Museum, 188: 1–622. the gobiid fi shes of Singapore. In: Sudara, S., Wilkinson, C.R., Suvatti, C., 1981. Fishes of Thailand. Royal Institute of Thailand: and Chou, L.M. (eds.) Proceedings, Third ASEAN-Australian Bangkok [in Thai]. Symposium on Living Coastal Resources, Vol. 2: Research Papers. Chulalongkorn University: Bangkok. Swofford, D., 2002. PAUP: Phylogenetic analysis using parsimony. Version 4.0b10. Champaign: Illinois. Miller, P. J., 1987. Affinities, origin and adaptive features of the Australian desert goby Chlamydogobius eremius (Zietz, Tan, S. H. & H. H. Tan, 1994. The freshwater fi shes of Pulau 1896) (Teleostei: Gobiidae). Journal of Natural History, 21: Bintan, Riau Archipelago, Sumatera, Indonesia. Tropical 687–705. Biodiversity, 2(3): 351–367. Miller, P. J., 1989. The classification of bumble-bee gobies Tweedie, M. W. F., 1940. Additions to the collection of fi shes in (Brachygobius and associated genera) (Teleostei: Gobiidae). the Raffl es Museum. Bulletin of the Raffl es Museum, Singapore, Cybium, 13(4): 375–383. 16: 68–82. Mohsin, A. K. M. & M. A. Ambak, 1983. Freshwater fi shes of Watson, R. E. & H. Horsthemke, 1995. Revision of Euctenogobius, peninsula Malaysia. Penerbit Universiti Pertanian Malaysia: a monotypic subgenus of Awaous, with discussion of its natural Kuala Lumpur. history. Revue Française Aquariologie, 22(3–4): 83–92. Ng, H.-H. & H.-H. Tan, 1999. The fi shes of the Endau Drainage, Werner, U. 1981. Meergrundeln, skurrile Pfl eglinge im Süsswasser- Peninsular Malaysia with descriptions of two new species of Aquarium. Das Aquarium, 15(140): 74–79. catfi shes (Teleostei: Akysidae, Bagridae). Zoological Studies, Wheeler, A., 1979. Fishes of the world. An illustrated dictionary. 38(3): 350–366. Ferndale Editions, London. Nichols, J. T., 1951. Four new gobies from New Guinea. American Wu, H.-L. & Y. Ni, 1985. On two new species of Mugilogobius Museum Novitates 1539: 1–8. Smitt (Perciformes: Gobiidae) from China. Zoological Research, Peters, W. C. H., 1869. Über die von Hrn. Dr. F. Jagor in 6(4): 93–98 [in Chinese]. dem ostindischen Archipel. gesammelten und dem Königl. Zhong, J.-S. & I-S. Chen, 1997. A new species of the genus, zoologischen Museum übergebenen Fische. Monatsberichte Pseudogobiopsis (Pisces, Gobiidae) from China. Journal of der Königllich Preussischen Akademie Wissenschaften Berlin, the Taiwan Museum 50(2): 77–84. 1868: 254–281.

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13_Larson_Pg 127-181.indd 171 2/19/09 12:03:48 PM Larson: Review of Eugnathogobius and Pseudogobiopsis Characters 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 ? 0 0 0 0 0 4 2 3 3 4 3 3 0 0 2 1 2 1 1 1 1 0 1 1 1 0 1 0 4 5 6 7 8 9 0 0 8 9 0 1 2 3 0 6 7 2 3 5 8 9 0 5 4 0 4 0 3 0 2 1 0 9 8 7 6 5 1 2 3 4 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 2 1 0 0 0 1 1 0 1 0 0 0 0 1 1 0 0 0 1 0 1 0 1 0 0 1 1 0 0 1 2 1 0 0 1 1 1 0 1 1 1 0 1 0 0 1 0 1 0 2 1 0 1 0 1 1 1 2 0 1 1 1 1 0 1 1 0 1 0 1 0 2 1 0 0 0 1 1 1 2 0 1 1 1 0 1 1 0 0 1 0 0 1 1 1 1 1 0 0 1 2 0 1 0 0 0 2 0 1 0 1 0 2 1 0 1 0 1 0 1 2 2 1 0 1 0 0 0 1 1 0 1 1 1 ? 1 0 ? 0 1 0 0 1 0 0 1 0 0 1 1 1 1 1 1 0 0 1 1 0 1 1 1 0 1 0 1 0 0 0 1 2 0 0 0 0 0 1 2 0 0 0 0 1 1 1 1 2 1 0 0 0 0 1 0 1 1 1 1 1 0 1 0 0 0 1 0 0 0 1 0 0 1 1 0 1 1 1 1 1 2 0 1 0 1 0 0 1 0 0 1 1 1 1 1 1 ? 1 0 0 0 1 0 1 1 2 0 1 1 0 1 1 ? 0 0 1 1 1 0 1 0 ? 1 0 1 2 0 ? 1 1 1 0 0 2 1 1 0 1 0 1 1 1 ? 1 0 0 0 1 1 1 ? 1 0 1 2 0 1 0 1 0 1 2 0 1 1 0 0 ? 0 0 0 0 1 0 1 0 1 0 ? 1 1 0 1 1 0 2 1 1 1 1 ? 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 1 1 0 1 1 1 1 0 0 0 0 0 2 0 1 0 0 0 0 0 1 1 0 1 1 0 1 0 1 1 1 0 1 0 1 0 0 1 0 1 0 1 0 0 1 1 0 1 0 1 1 0 1 0 1 1 0 1 ? 1 0 1 0 1 0 1 0 ? 0 0 1 1 1 0 1 1 0 0 ? 1 1 1 1 0 0 1 1 0 1 1 1 1 0 0 1 0 2 ? 1 1 0 1 1 1 0 1 0 1 1 0 1 0 1 1 0 0 1 1 1 0 1 0 1 ? 1 1 0 0 0 0 1 1 1 0 0 2 ? 0 0 0 1 1 1 1 0 0 0 0 ? 0 0 0 0 0 ? 1 0 0 1 1 ? 1 0 1 0 1 ? 0 0 0 0 1 ? 0 0 1 ? 0 0 1 0 ? 1 0 0 ? 0 1 1 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 1 0 1 1 0 0 0 0 0 1 1 1 1 1 1 0 0 0 1 1 ? 0 1 1 0 0 1 1 0 0 borealis Micropercops Sineleotris chalmersi Calamiana illota Calamiana kabilia Calamiana mindora Calamiana polylepis species new Calamiana stictos , Calamiana variegata 1 1 1 0 Eugnathogobius microps species 0 1 2 0 1 0 1 1 1 new 1 1 0 Pseudogobiopsis festivus , 1 0 1 0 0 1 1 0 Pseudogobiopsis oligactis Pseudogobiopsis paludosus Pseudogobiopsis siamensis Pseudogobiopsis tigrellus Pseudogobius javanicus Pseudogobius melanostictus Pseudogobius olorum Redigobius balteatus Redigobius dispar Redigobius macrostoma Table 1. Character matrix of the 20 taxa used for phylogenetic analyses; characters described in full Larson (2001: 15–31), with the exception of character 41 (see text). Names used are those and Sineleotris are outgroups. originally assigned to each species. Missing data is indicated by a question mark. The odontobutids Micropercops Species

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Table 2. Nominal species of Calamiana, Eugnathogobius and Pseudogobiopsis and their status.

Nominal species Taxonomic allocation Gnathogobius aliceae Smith, 1945 Eugnathogobius kabilia (Herre, 1940) Gnathogobius aliceae Smith, 1945 Eugnathogobius kabilia (Herre, 1940) Glossogobius campbellianus Jordan & Seale, 1908 Pseudogobiopsis oligactis (Bleeker, 1875) Calamiana illota Larson, 1999 Eugnathogobius illotus (Larson, 1999) Vaimosa jurongensis Herre, 1940a Eugnathogobius siamensis (Fowler, 1934) Vaimosa kabilia Herre, 1940a Eugnathogobius kabilia (Herre, 1940) Calamiana magnoris Herre, 1945d Eugnathogobius kabilia (Herre, 1940) Glossogobius mas Hora, 1923 Eugnathogobius or Pseudogobiopsis Vaimosa mawaia Herre, 1936 Eugnathogobius siamensis (Fowler, 1934) Vaimosa mindora Herre, 1945 Eugnathogobius mindora (Herre, 1945) Eugnathogobius microps Smith, 1931 Eugnathogobius microps Smith, 1931 Stigmatogobius neglectus Koumans, 1932 Pseudogobiopsis oligactis (Bleeker, 1876) Gobiopsis oligactis Bleeker, 1875 Pseudogobiopsis oligactis (Bleeker, 1875) Vaimosa oratai Herre, 1940a Eugnathogobius siamensis (Fowler, 1934) Ctenogobius paludosus Herre 1940a Pseudogobiopsis paludosus (Herre, 1940c) Vaimosa perakensis Herre, 1940a Pseudogobiopsis oligactis (Bleeker, 1875) Mugilogobius polylepis Wu & Ni, 1985 Eugnathogobius polylepis (Wu & Ni, 1985) Vaimosa rivalis Herre, 1927 Redigobius or Pseudogobiopsis? Vaimosa siamensis Fowler, 1934 Eugnathogobius siamensis (Fowler, 1934) Tamanka tagala Herre, 1927 Pseudogobiopsis? Mugilogobius? Gobius tigrellus Nichols, 1951 Pseudogobiopsis tigrellus (Nichols, 1951) Tamanka umbra Herre, 1927 Mugilogobius or Pseudogobiopsis? Apocryptes variegatus Peters, 1868 Eugnathogobius variegatus (Peters, 1868) Pseudogobiopsis wuhanlini Zhong & Chen, 1997 Eugnathogobius siamensis (Fowler, 1934)

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Table 3. Frequency distribution of dorsal and anal fi n soft ray counts in Eugnathogobius species.

Second dorsal fi n rays Anal fi n rays Species 5 6 7 8 9 5 6 7 8 9 E. indicus, new species – – 8 3 – – 1 10 1 – E. illotus – 1 22 1 – – – 22 2 – E. kabilia – 2 17 3 – 1 4 17 1 – E. microps 1 12 – – – – 13 – – – E. mindora – 1 39 3 – – 1 42 – – E. polylepis – 1 6 37 – – – 3 40 1 E. siamensis – 36 – – – – 36 – – – E. stictos, new species – – – 13 1 – – 5 8 – E. variegatus – – 1 34 1 – 2 34 –

Table 4. Frequency distribution of pectoral fi n ray counts (right fi n) in Eugnathogobius species.

Species 14 15 16 17 18 19 20 E. indicus, new species – 3 8 – – – – E. illotus 1 3 15 5 – – – E. kabilia – 2 5 12 3 – – E. microps – – – – 7 5 1 E. mindora 10 20 10 3 – – – E. polylepis – – 8 32 3 – – E. siamensis – – – 2 13 18 2 E. stictos, new species – 7 5 1 1 – – E. variegatus – 10 23 2 – –

Table 5. Frequency distribution of transverse backward scale counts in Eugnathogobius species.

Species 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 E. indicus, new species – – – 5 3 1 2 – – – – – – – – – E. illotus – – – – 1 11 8 3 1 – – – – – – – E. kabilia – 3 4 6 5 4 – – – – – – – – – – E. microps – 4 9 – – – – – – – – – – – – – E. mindora – – 11 17 6 5 3 1 – – – – – – – – E. polylepis – – – – – 1 – 3 3 11 7 7 2 3 2 2 E. siamensis 6 28 2 – – – – – – – – – – – – – E. stictos, new species – – – – – – – – 1 2 2 5 – 3 – 1 E. variegatus – – – 1 4 16 8 7 – – – – – – – –

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13_Larson_Pg 127-181.indd 174 2/19/09 12:03:48 PM THE RAFFLES BULLETIN OF ZOOLOGY 2009 Eugnathogobius species. – – – – – – – – – – – – – 1 2 – 1 1 3 2 – – 1 8 1 4 1 6 3 4 – 2 – – 4 – 6 1 – 1 – – – – – – – – – – – 3 – 10 9 5 – – 4 2 1 – 4 – 1 3 5 – – 26 – 4 – – – – – – – 2 – 1 – – – – – – – – – – – – – 8 – 10 6 – 8 1 3 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – 1 – – – – – – 2 – 5 – – – 4 – 1 – – 1 – 3 – 2 1 – – – 6 – 2 1 6 – 3 – – – – – – – – – – – – – – – – – – – 22 23 24 25 26 27 28 29 30 39 42 31 37 40 41 32 33 34 35 36 44 47 48 49 50 51 52 53 54 62 55 61 63 56 57 58 59 60 Species Table 6. Frequency distribution of lateral scale counts in , new species new – E. indicus , – – – – – E. illotus – – – 1 – – 1 4 2 – 1 3 E. kabilia E. microps species E. mindora E. polylepis new E. siamensis – E. stictos , – – – E. variegatus – – – – – – – – Species , new species new – E. indicus , – – – – – E. illotus – – – – – – E. kabilia E. microps species E. mindora E. polylepis new E. siamensis – E. stictos , – 1 – E. variegatus 3 – 1 1 1 – – 1 1 2 –

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Table 7. Measurements (mm) of Eugnathogobius indicus, new species.

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 8.4 4.2 10.0 6.5 6.2 8.4 7.1 Head depth 4.6 2.6 5.8 3.7 3.4 4.6 4.0 Head width 6.1 3.4 8.0 4.9 4.4 6.1 5.2 Body depth 4.6 2.7 5.1 3.6 3.6 4.6 4.0 Body width 2.5 1.6 3.2 2.2 2.1 2.9 2.5 Caud. ped. length 5.9 3.8 9.0 5.9 5.9 7.7 6.6 Caud. ped. depth 3.3 1.9 3.8 2.6 2.6 3.3 3.0 Snout 2.0 1.1 3.0 1.7 1.5 2.1 1.8 Eye 2.1 1.2 2.2 1.6 1.5 2.1 1.8 Jaw 3.1 1.5 4.7 2.5 2.1 3.1 2.5 Interorbit 1.7 1.0 2.5 1.5 0.9 1.7 1.3 Pectoral 6.0 3.1 6.7 4.8 5.2 6.0 5.6 Pelvic 6.0 3.0 7.1 4.5 4.6 6.0 5.1 Caudal 7.3 4.0 10.0 6.1 6.4 7.3 6.8 Longest D1 spine 3.6 1.9 6.5 3.8 3.0 3.6 3.2 Caud. ped., Caudal peduncle

Table 8. Measurements (mm) of Eugnathogobius kabilia (Herre, 1940).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 10.5 6.4 15.2 9.6 4.4 10.5 8.0 Head depth 6.2 3.8 8.8 5.4 2.2 6.2 4.8 Head width 8.5 4.8 11.3 7.1 2.9 8.5 5.9 Body depth 7.0 3.6 9.5 5.7 4.7 7.0 5.9 Body width 4.3 2.1 6.7 3.7 3.6 4.5 4.2 Caud. ped. length 9.6 5.5 11.7 8.0 4.5 9.6 7.7 Caud. ped. depth 4.7 2.3 6.4 4.0 3.2 4.7 3.9 Snout 3.1 1.9 4.6 3.0 1.1 3.1 2.0 Eye 2.5 1.5 3.6 2.1 1.4 2.5 2.1 Jaw 4.3 3.4 12.5 7.3 1.4 4.3 3.2 Interorbit 2.5 1.6 6.0 2.7 1.0 2.5 1.9 Pectoral 7.1 5.0 11.0 7.0 5.2 8.0 6.8 Pelvic 6.3 4.3 8.0 5.7 3.0 6.3 5.2 Caudal 9.0 6.1 13.1 9.0 4.1 10.0 7.8 Longest D1 spine 5.1 3.1 5.4 4.8 3.2 5.1 4.2 Caud. ped., Caudal peduncle.

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Table 9. Measurements (in mm) of Eugnathogobius microps (Smith, 1931).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 9.2 5.8 9.2 7.5 6.2 8.7 7.6 Head depth 4.9 3.4 5.4 4.4 3.7 5.1 4.5 Head width 6.0 4.3 7.5 5.7 4.6 7.0 5.8 Body depth 5.2 3.6 5.9 4.6 3.9 5.7 5.0 Body width 3.1 2.3 4.1 3.1 2.8 3.5 3.3 Caud. ped. length 6.1 4.8 6.8 5.7 4.9 6.4 5.8 Caud. ped. depth 3.8 2.2 3.8 3.0 2.5 3.7 3.2 Snout 2.4 1.5 2.4 1.9 1.5 2.1 1.9 Eye 1.1 1.0 1.8 1.3 0.9 1.6 1.3 Jaw 6.4 3.1 6.4 4.8 2.8 4.7 3.7 Interorbit 2.2 1.6 2.5 2.1 1.7 2.4 2.1 Pectoral 6.6 4.2 6.6 5.4 4.8 5.6 5.1 Pelvic 6.5 4.5 6.6 5.6 4.9 6.2 5.6 Caudal 6.5 5.0 6.6 5.7 4.7 5.8 5.4 Longest D1 spine – 2.5 3.5 3.0 2.5 2.7 2.6 Caud. ped., Caudal peduncle.

Table 10. Measurements (mm) of Eugnathogobius polylepis (Wu & Ni, 1985).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 8.0 4.6 8.0 5.9 4.7 6.4 5.5 Head depth 4.2 2.5 4.2 3.2 2.4 3.5 3.0 Head width 5.4 3.2 5.6 4.3 3.1 4.6 3.9 Body depth 4.4 2.3 4.4 3.2 2.5 4.2 3.1 Body width – 1.1 3.0 2.1 1.5 2.8 2.2 Caud. ped. length 8.0 4.3 8.0 5.6 4.5 6.5 5.5 Caud. ped. depth 3.4 1.9 3.4 2.6 2.0 3.1 2.6 Snout 2.0 1.1 2.0 1.6 1.1 1.7 1.4 Eye 1.9 1.2 1.9 1.5 1.2 1.8 1.4 Jaw 3.4 1.7 3.4 2.3 1.6 2.3 2.0 Interorbit 1.4 0.9 2.1 1.4 0.8 1.8 1.3 Pectoral 6.0 3.4 6.0 4.3 3.4 4.8 4.1 Pelvic 4.2 2.5 4.2 3.2 2.6 3.5 3.0 Caudal – 3.7 5.8 4.8 3.8 5.1 4.6 Longest D1 spine – 1.5 2.6 2.1 1.3 2.2 1.8 Caud. ped., Caudal peduncle.

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Table 11. Measurements (mm) of Eugnathogobius siamensis (Fowler, 1934).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 10.4 5.7 11.6 9.6 5.5 9.4 7.4 Head depth 6.3 3.4 7.9 6.0 3.1 5.5 4.4 Head width 7.3 4.2 9.8 6.9 3.7 6.2 5.2 Body depth 6.0 3.9 7.9 6.3 3.3 5.5 4.6 Body width – 2.3 5.2 4.2 1.8 3.8 2.8 Caud. ped. length 9.6 5.1 10.5 8.5 5.2 8.3 6.6 Caud. ped. depth 4.2 2.3 4.7 3.9 2.0 3.4 2.8 Snout 2.7 1.6 3.9 2.7 1.3 2.8 1.9 Eye 2.7 1.8 3.0 2.5 1.8 2.6 2.2 Jaw 6.7 2.2 7.4 5.3 2.0 5.2 2.9 Interorbit 1.4 0.9 2.6 1.7 0.6 1.9 1.0 Pectoral 6.4 4.2 9.2 7.4 4.7 6.8 6.0 Pelvic 6.8 4.0 9.4 7.4 4.6 6.8 6.0 Caudal – 6.0 10.7 8.9 5.2 7.5 6.5 Longest D1 spine 4.4 4.6 5.6 5.1 3.3 4.4 3.7 Caud. ped., Caudal peduncle.

Table 12. Measurements (mm) of Eugnathogobius stictos, new species

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 6.0 4.5 6.0 5.1 4.9 6.0 5.3 Head depth 3.8 2.4 3.8 3.2 2.6 3.5 3.1 Head width 4.6 2.9 4.6 3.8 3.2 4.3 3.8 Body depth 3.6 2.6 3.7 3.1 3.2 3.7 3.4 Body width 2.2 1.6 2.3 1.9 1.6 2.3 1.9 Caud. ped. length 5.5 4.0 5.5 4.7 3.8 5.3 4.7 Caud. ped. depth 2.8 2.0 2.8 2.3 2.2 2.7 2.4 Snout 1.5 1.1 1.6 1.4 1.2 1.5 1.4 Eye 1.5 1.1 1.6 1.3 1.2 1.6 1.4 Jaw 2.8 1.7 2.8 2.2 1.7 2.1 1.9 Interorbit 1.0 0.6 1.2 0.9 1.0 1.3 1.2 Pectoral 4.4 3.1 4.4 3.7 3.9 4.4 4.1 Pelvic 3.5 2.4 3.5 2.9 2.7 3.0 2.9 Caudal 6.4 4.1 6.4 5.1 5.0 5.4 5.2 Longest D1 spine 2.5 1.3 2.5 1.9 1.4 1.4 1.4 Caud. ped., Caudal peduncle.

Table 13. Frequency distribution of dorsal and anal fi n soft ray counts in Pseudogobiopsis species.

Second dorsal fi n rays Anal fi n rays Species 6 7 8 9 10 5 6 7 8 P. festivus, new species – 32 – – – – – 31 1 P. oligactis 51 5 – – – 2 51 2 – P. paludosus – 6 11 5 1 – 10 13 – P. tigrellus – – 8 – – – 7 1 –

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Table 14. Frequency distribution of pectoral fi n ray counts in Pseudogobiopsis species.

Species 14 15 16 17 18 19 20 P. festivus, new species – – – – 5 24 3 P. oligactis – – – 4 23 25 4 P. paludosus 1 3 13 6 – – – P. tigrellus 1 5 2 – – – –

Table 15. Frequency distribution of transverse backward scale counts in Pseudogobiopsis species.

Species 6 7 8 9 P. festivus, new species – – 32 1 P. oligactis 1 11 44 – P. paludosus – 3 19 1 P. siamensis – 6 28 2 P. tigrellus – 1 7 –

Table 16. Frequency distribution of lateral scale counts in Pseudogobiopsis species.

Species 22 23 24 25 26 27 P. festivus, new species - - 5 26 1 - P. oligactis 7 41 6 2 - - P. paludosus - 1 9 9 3 1 P. tigrellus - 4 3 1 - -

Table 17. Measurements (mm) of Pseudogobiopsis festivus, new species.

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 10.2 4.4 10.2 6.7 6.1 8.1 7.3 Head depth 6.2 2.3 6.2 3.7 3.1 4.6 4.1 Head width 7.7 2.5 7.7 4.4 3.7 5.6 5.0 Body depth 6.0 2.4 6.0 3.8 3.6 5.1 4.6 Body width 4.3 1.6 4.4 2.8 2.7 4.3 3.6 Caud. ped. length 10.2 4.9 10.2 6.9 6.3 9.0 8.0 Caud. ped. depth 4.5 1.7 4.5 2.9 2.6 3.7 3.3 Snout 2.7 0.9 2.7 1.6 1.2 2.0 1.7 Eye 2.2 1.3 2.7 1.8 1.7 2.4 2.1 Jaw 6.7 1.4 6.7 3.3 2.1 3.2 2.6 Interorbit 3.0 0.5 3.0 1.2 0.9 1.5 1.2 Pectoral 10.0 3.7 10.0 6.1 5.3 7.7 6.6 Pelvic 8.2 3.4 8.2 5.3 4.9 6.8 5.9 Caudal 9.7 4.6 9.7 7.0 6.1 8.6 7.6 Longest D1 spine 7.0 3.0 8.2 5.0 3.6 5.7 4.7 Caud. ped., Caudal peduncle.

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Table 18. Measurements (in mm) of Pseudogobiopsis oligactis (Bleeker, 1875).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 8.1 5.7 15.2 10.1 7.1 12.3 9.2 Head depth 4.8 3.2 7.9 5.2 3.8 6.8 5.0 Head width 5.6 3.7 11.0 7.0 4.5 8.8 6.4 Body depth 5.5 3.8 9.3 5.6 3.7 8.1 5.3 Body width - 2.9 7.8 4.3 2.9 6.9 4.2 Caud. ped. leng. 6.6 5.7 12.6 8.5 6.8 11.2 8.6 Caud. ped. depth 2.8 2.6 6.2 3.8 2.7 5.2 3.6 Snout 2.0 1.5 4.7 2.9 1.9 3.4 2.6 Eye 1.8 1.7 3.2 2.4 2.1 3.1 2.5 Jaw 6.6 2.1 11.3 5.9 2.6 5.7 3.8 Interorbit 0.9 0.9 2.5 1.5 1.0 1.7 1.2 Pectoral 5.7 5.1 11.3 7.5 6.5 9.3 7.5 Pelvic 5.4 5.2 10.1 7.0 4.8 8.6 6.8 Caudal - 5.8 12.0 8.1 6.0 10.4 7.8 Longest D1 spine - 2.9 16.0 8.1 3.5 6.9 5.3

Table 19. Measurements (mm) of Pseudogobiopsis paludosus (Herre, 1940).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 9.5 5.4 9.6 8.0 5.4 6.4 5.9 Head depth 4.9 2.9 5.0 4.1 2.9 3.3 3.1 Head width 6.8 3.7 6.8 5.3 3.3 4.0 3.8 Body depth 5.2 3.0 5.2 4.1 3.0 3.7 3.3 Body width 4.4 2.2 4.4 3.0 2.0 2.9 2.3 Caud. ped. length 7.6 4.8 8.6 6.5 4.3 5.9 5.2 Caud. ped. depth 3.6 2.2 3.6 2.9 1.9 2.7 2.2 Snout 2.6 1.4 2.6 2.0 1.1 1.5 1.3 Eye 2.1 1.6 2.4 2.1 1.8 2.0 1.9 Jaw 5.7 2.2 5.7 4.3 1.8 2.3 2.0 Interorbit 1.8 1.1 2.0 1.5 0.5 1.1 0.9 Pectoral 7.3 4.3 7.5 6.6 4.7 5.4 5.0 Pelvic 6.9 4.1 6.9 5.8 3.8 5.2 4.6 Caudal 7.7 5.1 8.4 7.4 4.8 6.1 5.5 Longest D1 spine 4.7 3.9 7.7 6.2 2.6 3.3 3.1 Caud. ped., Caudal peduncle.

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Table 20. Measurements (mm) of Pseudogobiopsis tigrellus (Nichols, 1951).

Males Males Males Females Females Females Character Holotype Minimum Maximum Mean Minimum Maximum Mean Head length 6.8 5.9 7.5 6.9 6.0 6.7 6.4 Head depth 3.5 3.4 4.4 3.9 3.5 4.1 3.8 Head width 4.0 4.0 5.4 4.7 4.2 4.7 4.5 Body depth 4.0 3.6 4.7 4.2 3.6 4.3 4.0 Body width - 2.5 3.4 3.0 2.5 2.9 2.8 Caud. ped. leng. 6.0 6.0 7.4 6.8 5.5 7.4 6.4 Caud. ped. depth 2.9 2.6 3.1 2.9 2.4 2.9 2.7 Snout 1.8 1.5 2.1 1.9 1.7 2.0 1.8 Eye 1.7 1.7 2.0 1.8 1.7 2.0 1.8 Jaw 3.2 2.2 3.4 3.1 2.1 2.3 2.2 Interorbit 0.7 0.6 1.0 0.8 0.5 0.7 0.6 Pectoral 4.3 4.3 7.5 6.2 5.2 6.6 5.9 Pelvic 4.9 4.9 7.3 6.1 5.4 6.2 5.9 Caudal - 6.1 8.4 7.5 5.8 7.0 6.3 Longest D1 spine - 3.2 4.1 3.7 2.8 4.0 3.2 Caud. ped., Caudal peduncle.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 183–187 Date of Publication: 28 Feb.2009 © National University of Singapore

GEOSESARMA AEDITUENS, A NEW TERRESTRIAL CRAB (CRUSTACEA: DECAPODA: BRACHYURA: SESARMIDAE) FROM BALI, INDONESIA

Tohru Naruse Raffl es Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Republic of Singapore; Present address: Transdisciplinary Research Organization for Subtropical and Island Studies, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan Email: [email protected]

Zeehan Jaafar Raffl es Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Republic of Singapore

ABSTRACT. – A new species of Geosesarma is here described from the island of Bali, Indonesia. This is the fi rst species of this genus to be reported from the Island. Geosesarma aedituens, new species, is compared with its congeners, especially with the morphologically similar G. aurantium from Sabah, Borneo, G. serenei from Peninsular Malaysia and another four species occurring in the neighbouring island of Java. This species is found to be different based on the following diagnostic characters: extent of external orbital angles, movable fi nger on the chela, and G1 structure.

KEY WORDS. – Geosesarma aedituens, new species, taxonomy, Sesarmidae, Bali, Indonesia.

INTRODUCTION TAXONOMY

Terrestrial crabs of the genus Geosesarma De Man, 1892, Geosesarma De Man, 1892 presently number about 50 species are distributed from Southeast Asia to the islands of the Western Pacifi c. Although Geosesarma aedituens, new species new species are continually being described, (Manuel-Santos (Figs. 1–3) & Yeo, 2007; Yeo & Freitag, in prep), taxonomy at the genus level is still in a state of fl ux (Ng, 1988; Ng et al., Material examined. – Holotype: male, 7.6 × 8.9 mm, MZB 2004). Specimens of an undescribed species were recently Cru 1655, Bedugul, Tabanan, Bali, Indonesia, coll. Z. Jaafar, 5 collected from tropical montane forests of Bedugul, on the Sep.2005. island of Bali, Indonesia. There are no congeners of this species in Bali and this present study aims to describe this new species.

Specimens examined are deposited in the Muzium Zoologicum Bogoriense, Bogor, Indonesia (MZB) and the Zoological Reference Collection, Raffl es Museum of Biodiversity Research, National University of Singapore (ZRC). Measurements provided are of the carapace length (CL) by the carapace width (CW) in millimeters. The abbreviations G1 and G2 are used for the male fi rst and second gonopod respectively.

Fig. 1. Geosesarma aedituens, new species. Holotype, male, MZB Cru 1655.

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Paratypes: 6 males, 4.0 × 4.5 – 7.4 × 8.3 mm, 2 ovigerous females, southwest of Lahad Datu, eastern Sabah, ca. 4°58'S 118°10'E, East 7.2 × 8.2 mm & 7.2 × 8.6 mm, MZB Cru 1656, data as holotype; Malaysia, Borneo, coll. M. Manjaji, J. Payne & B. Perumal, 10 2 males, 6.9 × 7.9 mm & 7.3 × 8.5 mm, ZRC 2007.0527, data Jul.1994. Geosesarma bicolor Ng & Davie, 1995: holotype male, as holotype; 3 males, 5.0 × 5.7 – 6.3 × 7.3 mm, 4 females, 3.8 × 10.5 × 11.0 mm, ZRC 1995.0279, Citerjun, in creeks, under stones, 4.2 – 6.2 × 7.1 mm, 1 ovigerous female, 7.6 × 9.0 mm, MZB Cru Ujung Kulon, West Java, Indonesia, coll. C. Stewart, Jul.1993. 1657, Pura Kayu, Sugih, Bedugul, Tabanan, Bali, Indonesia, coll. Geosesarma serenei Ng, 1986: holotype male, 8.0 × 8.2 mm, ZRC Z. Jaafar, 4 Sep.2005; 2 males, 5.7 × 6.5 mm & 6.2 by 7.0 mm, 1964.9.8.1, Larut Hill, Maxwell Hills, Perak, Malaysia, coll. M. 1 female, 6.4 × 7.6 mm, 2 ovigerous females, 6.6 × 7.6 mm & W. F. Tweedie, 1950. 8.1 × 9.6 mm, ZRC 2007.0528, data as MZB Cru 1657; 1 male, 7.5 × 8.2 mm, ZRC 2006.0071, east of Bedugul, Mt. Bratan, Bali, Description. – Carapace (Figs. 1, 2a) almost squarish to Indonesia (8°16.861'S 115°10.829'E, 1240 m, within a log), coll. slightly trapezoidal, widest at bases of second ambulatory D. S. Sikes, 14 May 2005. legs, CW 1.09–1.20 CL (n = 25, mean = 1.15); dorsal surface weakly convex longitudinally and transversely, Comparative material. – Geosesarma aurantium Ng, 1995: holotype male, 10.6 × 11.7 mm, ZRC 1995.0283, in montane forest on rough, partially granulated, metagastric region laterally ground, Bukit Silam, ca. 760 m above sea level, ca. 20 km west- demarcated by shallow grooves, three pairs of relatively

Fig. 2 Geosesarma aedituens, new species. Holotype, male, MZB Cru 1655: a, carapace; b, right third maxilliped; c, left chela; d, movable fi nger of left chela, dorsal view; e, left third ambulatory leg. Scale bars = 1 mm.

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large granules present on anterior part of metagastric region Antennule with swollen basal article, each antennular and outer-posterior part of H-shaped gastric groove. Front fossa occupying one-third width of front; antennal basal wide, frontal width 0.44–0.51 fronto-orbital width (n = 24, article swollen laterally, basal article and endopod fi lling mean = 0.49), strongly defl exed at level of anterior margin gap between lateral margin of front and inner margin of of ocular peduncle, frontal margin bilobed, median region inner orbital tooth; eyes moderately developed, diverging concave, shallow, wide, lobe convex on inner third; two posteriorly when set in orbits. Third maxillipeds (Fig. 2b) pairs of epigastric cristae on defl exed part of front, median leaving rhomboidal gap when closed, merus as long as pair wider than lateral ones. Supraorbital margin diverging ischium; exopod short, reaching proximal third of merus, posteriorly, curve shallow. External orbital angle (= fi rst without fl agellum. anterolateral tooth) blunt, directed laterally, but with tip slightly curved anteriorly and upturned dorsally; second Chelipeds (Fig. 1) equal; male chelipeds more robust than anterolateral tooth separated from fi rst tooth by V-shaped those of female, merus with triangular cross-section, ventral cleft, upturned, blunt, smaller than fi rst tooth, sometimes outer margin rimmed, slightly corneous, lined with granules followed posteriorly by granules. Lateral margins slightly in small specimens, eroded in large specimens, ventral inner divergent posteriorly, almost subparallel, posterolateral margin granulated, dorsal margin incurving, outer surface surface sloping outwards, posterior margin as wide as front. sparsely granulated, inner surface with two longitudinal Epistome with posterior margin cristate, granulated, trilobed, lines of stiff setae, ventral one denser; carpus slightly rather median lobe longest. granulated, inner angle broad, not projected, proximal inner

Fig. 3 Geosesarma aedituens, new species. Holotype, male, MZB Cru 1655: a, abdomen and telson; b, right G1, ventral view. Scale bars = 1 mm.

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margin lined with stiff setae. Chela (Fig. 2c) massive, inner Etymology. – The specifi c epithet “aedituens” is Latin for and outer surfaces of palm swollen, outer surface sparsely “keeper of a temple” as many specimens were collected from granulated, except for bases of fi ngers, inner surface with a banks around an abandoned temple. The name is used as a crest from dorsal part of base of movable fi nger to base of noun in apposition. immovable fi nger, granules lined along crest; movable fi nger as long as palm, slender, slightly curving downwards, dorsal Remarks. – Geosesarma aedituens, new species, has a surface (Fig. 2d) with two cristae, outer without granules, relatively wide carapace and front and lacks a fl agellum on inner lined with numerous tiny granules, cutting edge with the exopod of the third maxilliped, characters that are shared four prominent white teeth, two on proximal fi fth, one on with G. aurantium Ng, 1995, from Sabah, Borneo and G. middle, one on distal fi fth, distal tip corneous; immovable serenei Ng, 1986, from Peninsular Malaysia. Geosesarma fi nger with cutting edge slightly raised on proximal third, aedituens can be easily differentiated from G. aurantium by distal two-thirds concave, forming narrow oblong gape. the distinct bilobed frontal margin of its carapace (vs frontal Ambulatory legs slender, third ambulatory leg (Fig. 2e) margin not distinctly bilobed); laterally-directed external longest, distal end of merus slightly exceeding external orbital angle (vs external orbital angle anteriorly-directed); orbital angle when folded anteriorly; tufts of setae present dorsal surface of the movable fi nger of the male chela with between coxae of cheliped and ambulatory legs, denser at an outer low ridge and an inner row of densely-set granules between fi rst and second ambulatory legs and second and third (ca. 41) (vs. dorsal surface of movable finger with 4–5 ambulatory legs; meri with anterior and posterior margins teeth near proximal end); and narrow terminal process of slightly raised at edges, subparallel, anterior margin rough, the G1 being less strongly bent (ca. 30°) [vs. broad, spade- subdistal tooth almost right angle; propodi with outer and like terminal process of the G1 being more strongly bent inner margins subparallel, dactyli terminated in sharp tooth, outwards (ca. 90°)] (present study; Ng, 1995: Figs. 1A, C, propodi to dactyli sparsely lined with black stiff setae. 3). Geosesarma aedituens can be distinguished from G. serenei by the convex dorsal surface of the carapace (vs. Male abdomen (Fig. 3a) seven segmented including telson; dorsal surface of the carapace not convex); slightly divergent base of first segment placed below posterior margin of lateral margins of the carapace (vs. lateral margins of the carapace; second segment narrower than fi rst, short; third carapace parallel); laterally-directed external orbital angle segment widest, third segment to telson bell-shaped; telson (vs. external orbital angle produced anterolaterally); dorsal slightly longer than sixth segment. G1 (Fig. 3b) slender, surface of the movable fi nger of the male chela with an outer gently curving outwards, chitinous distal process long, about low ridge and an inner row of densely-set granules (ca. 41) one-quarter of G1 length, compressed laterally, straight, bent (vs. dorsal surface of the movable fi nger with a few teeth outwards at ca. 30°. (ca. 5) near proximal end) and narrow terminal process of the G1 (vs. terminal process of G1 dorso-ventraly fl attened Eggs large, subcircular in shape, long axis 1.42–1.62 mm and broader) (present study; Ng, 1986: Fig. 2; Ng, 1988: (mean 1.51 mm, n = 10). One female (MZB Cru 1657, 7.6 Fig. 55A, D, F). by 9.0 mm) brooded only 17 eggs. Geosesarma aedituens is the fi rst species of this genus to Colouration. – Live colouration of dorsal carapace reddish- be reported from the island of Bali. Four of its congeners brown to deep purple. Chelipeds and ventral region orange- are recorded from the neighbouring island of Java viz.; G. red. Legs with dark transverse bands. Eggs deep red. bicolor Ng & Davie, 1995; G. confertum (Ortmann, 1894); G. noduliferum (De Man, 1892) and G. rouxi (Serène, 1968). Habitat and Ecology. – This species is found in montane Geosesarma aedituens, is distinguished from G. confertum, forests in the interior region of Bali, Indonesia. There are no G. noduliferum, and G. rouxi by its blunt external orbital known streams in the area, although Lake Bratan is within angles (vs. acute in G. confertum, G. noduliferum, and G. 3 km distance within all sites from which this species were rouxi [present study; De Man, 1892: 342, Pl. 20, Fig. 16; collected. Individuals were collected from burrows excavated Serène, 1968: Pl 2(4)]. Geosesarma aedituens can also be in raised soil banks near an abandoned temple and along distinguished from G. bicolor by the absence of fl agellum of footpaths and mounds within vegetated and forested areas. the exopod of the third maxillped in the former (vs. present Banks and mounds were heavily-covered with bryophytes in the latter) (Ng & Davie, 1995: Fig. 2C). and the substrate was often moist. Burrows appeared to be excavated by the crabs and were found at about one metre above the ground level upwards. Juveniles tended to reside ACKNOWLEDGEMENTS within the lower strata and mature males and females were most often found in burrows higher up on the bank. Burrow The authors wish to thank Professor Peter Ng and the two entrances were clean and well-maintained, sometimes covered anonymous reviewers for their comments on this manuscript. or sheltered with mats of bryophytes. When live, these We are grateful to Miss Debby Ng for help rendered in animals appeared cautious and shy. No individuals were the fi eld; and to Dr. Derek S. Sikes (University of Alaska found outside of their burrows. They can be seen near the Museum) for providing us with a specimen of the new burrow entrances quickly retreating at any perceived signs species. This study is supported by research grant number of threat (Z. Jaafar, pers. obs.). ARF R-154-000-334-112 from the National University of Singapore to Professor Peter Ng.

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LITERATURE CITED Ng, P. K. L. & P. J. F. Davie, 1995. The terrestrial sesarmine crabs of the genera Metasesarma and Geosesarma (Crustacea: Man, J. G., de, 1892. Decapoden des Indischen Archipels, in Weber, Decapoda: Brachyura: Grapsidae) of Ujung Kulon, West Jawa, M. (ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Indonesia. Tropical Biodiversity, 3(1): 29–43. Ost-Indien, 2: 265–527, Pls. 15–29. Ng, P. K. L., H.-C. Liu & C. D. Schubart, 2004. Geosesarma Manuel-Santos, M. R. & D. C. J. Yeo, 2007. A new species of hendon, a new species of terrestrial crab (Crustacea: Decapoda: Geosesarma from Palawan, Philippines (Crustacea: Decapoda: Brachyura: Sesarmidae) from Taiwan and the Philippines. Brachyura: Sesarmidae). Zootaxa, 1607: 63–67. Raffl es Bulletin of Zoology, 52(1): 239–249. Ng, P. K. L., 1986. Preliminary descriptions of 17 new freshwater Ortmann, A. E., 1894. Crustaceen, in Von Semon, R. (ed.), crabs of the genera Geosesarma, Parathelphusa, Johora and Zoologische Forschungsreisen in Australien und dem Malayischen Stoliczia (Crustacea, Decapoda, Brachyura) from South East Archipel. Systematik, Tiergeographie, Anatomie wirbelloser Asia. Journal of the Singapore National Academy of Science, Tiere. Denkschriften der medicinisch-naturwissenschaftlichen 15: 36–44. Gesellschaft zu Jena 8(1): 1–80, Pls. 1–3. Ng, P. K. L., 1988. The Freshwater Crabs of Peninsular Malaysia Serène, R., 1968. Note préliminaire sur de nouvelles espèces de and Singapore. Shinglee Press, Singapore. viii, 4 pls, 156 p. Sesarma (Decapoda Brachyura). Bulletin du Muséum national d’Histoire naturelle, 39(6)[1967]: 1084–1095. Ng, P. K. L., 1995. Geosesarma aurantium, a new sesarmine land crab (Crustacea: Decapoda: Brachyura: Grapsidae) from Sabah. Malayan Nature Journal, 49(2): 65–70.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 189–198 Date of Publication: 28 Feb.2009 © National University of Singapore

AN ASSESSMENT OF FOOD OVERLAP BETWEEN GIBBONS AND

Chuti-on Kanwatanakid-Savini and Pilai Poonswad Department of Biology, Faculty of Science, Mahidol University, Rama 6 Rd. Phayathai, Bangkok 10400 ,Thailand Thailand Project, Department of Microbiology, Faculty of Science, Mahidol University, Rama 6Rd., Phayathai, Bangkok 10400, Thailand Emails:[email protected] (CKS); [email protected] (PP)

Tommaso Savini Conservation Ecology Program, School of Bioresources & Technology, King Mongkut’s University of Technology Thonburi, 83 Moo. 8 Thakham, Bangkhuntien, Bangkok 10150, Thailand Email: [email protected] (Corresponding author)

ABSTRACT. – Hornbills and gibbons are considered to be potential competitors for food. They show considerable overlap in their frugivorous diet, foraging site (outer part of tree crowns), and reproductive period. However, the extent and nature of such overlaps have never been measured. Here we characterize the dietary overlap of the Great Hornbill, Wreathed Hornbill, Oriental Pied Hornbill, White-throated Brown Hornbill and White-handed Gibbon based on a long-term data on their feeding ecology at Khao Yai National Park, Thailand. Results from a Correspondence Analysis showed a slight overlap between the diets of the hornbills and gibbon, which may be attributed primarily on the dietary consumptions of Ficus spp. and, to a lesser extent, Polyalthia viridis. Due to the large crown size of Ficus and the overall low nutritional quality of its fruits, we hypothesized that a dietary overlap between the hornbills and gibbon for Ficus is energetically inconsequential for either groups. In contrast, P. viridis has comparatively small crowns, is present in the site in low densities, and produces fruits high in protein. Therefore, the overlap in diet between hornbills and gibbon for P. viridis may create the basis for competition, which in any event will be limited to pseudo-interference.

KEY WORDS. – Food overlap; Resources exploitation; Khao Yai National Park; Thailand.

INTRODUCTION coexisting species, face increase in energy requirement due to reproduction. In mammals, the physiology of reproduction, The coexistence of two different animal species using regular cycling, gestation, and nursing, place females under overlapping niches, defi ned by the full range of ecological substantial energetic pressure, (Lee & Bowman, 1995), and conditions and resources under which they can survive, is thus access to resources plays an important limitation in linked to their capacity for sharing such niches and their regulating overall reproductive success. Similarly, in birds, resources, in a way that allows both of them to survive and reproductive success has been related directly to the level of reproduce optimally (Edgerly et al., 1998; Park et al., 1998; dietary protein available during the nesting period (Beckerton Cross & Benke, 2002; Elmberg et al., 2003). & Middleton, 1982; Wikelski et al., 2000). Those general effects have been so far documented for food overlaps Overlap in the use of niches and their resources may cause between closely related coexisting species (Luiselli et al., competition that result in separation of the two species into 1999; Neale & Sacks, 2001; Mitchell & Bank, 2005), and specifi c parts of their original niche (Terborgh & Diamond, not closely related coexisting species (Morin, 1999; Brown 1970). However, competition will not arise when two et al., 1997). coexisting species that share resources are limited to different resources (Tilman, 1977) or to one unlimited resource Two separate long-term ecological research projects have (Camillo & Garofalo, 1989; Geoffrey, 1998; Krasnov et been conducted for more than two decades at Khao Yai al., 2005). National Park, Thailand, on the White-handed Gibbon (Hylobates lar) (Brockelman et al., 1998) and four sympatric Feeding overlap may acquire ulterior importance if species of hornbill, bicornis (Great Hornbill), Aceros accentuated during parts of the year when one, or more of the undulatus (Wreathed Hornbill), Anthracoceros albirostris

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(Oriental Pied Hornbill) and Anorrhinus austeni (White- from the moment when the female in the nest began to take throated Brown Hornbill) (Poonswad et al., 1998a). Both food from the male until all food items had been passed to animal groups have been observed to feed on the outer part of the female. We identifi ed and characterized every plant and the forest canopy and share the same niche (Carpenter, 1967; animal food item that was consumed. Kemp, 1995), where they select primarily ripe, sugar-rich, juicy fruits (Leighton, 1998; Poonswad et al., 1998a) with We defi ned gibbon diet using direct observations of fi ve bright colors (Suryadi et al., 1994; Kanwatanakid, 2000). well-habituated gibbon groups. For a period of two and a half Moreover, a recent investigation of gibbon reproduction has years (May 2001 to Sep.2003), two trained fi eld assistants also highlighted signifi cant overlap with hornbills during observed each group for a minimum of 5 days per month. that period of the year when they have a strong dependence Data on feeding activities were collected by continuous on resources (Poonswad, 1993; Savini et al., 2008). The observations during the time between leaving and re-entering objective of this study was to characterize the dietary overlap a night-tree (cf. Martin & Bateson, 1993), alternating every between the White-handed Gibbon and the four sympatric hour between males and females. Food sources were known hornbill species, as suggested by Kemp (1995), specifi cally by the observers or where identifi ed later from collected plant during the fi rst half of the year when both animal groups face samples (for details see Savini et al., 2008). strong feeding pressure due to their reproductive energetic demands (Poonswad, 1993; Savini et al., 2008). Resource abundance – The spatial distribution of plant resources was measured in order to quantify their abundance in the area by using 13 north–south transects (total length MATERIALS AND METHODS 19.4 km, range 620–2,100 m) located within the gibbon study area of 39.8 ha (0.398 km2). Along each transect, trees larger Study site and species – The study was conducted between than 10 cm diameter at breast height (DBH) were marked, May 2001 and Dec.2005 at Khao Yai National Park, Thailand measured, and identifi ed to species, so that a total of 19,524 (2,168 km2; 14°26'N 101°22'E), in slightly hilly terrain at individual trees were included in our analyses. We calculated about 600–890 m above sea level. The study area (Fig. 1) the amount of resources as the percentage of the total tree covered approximately 60 km2 and was inhabited by a large sample, based on both stem number and basal area. population of four sympatric hornbill species that have been studied in detail for the past twenty years (Poonswad, Statistical analysis – Correspondence Analysis (CA) using 1993; 1995; 1998a). The area was also inhabited by a large Minitab 15 (Minitab Inc.) was employed to compare feeding population of White-handed Gibbon, which has been studied assemblages between White-handed Gibbons and the four since the late 1970s (see Raemaekers & Raemaekers, 1985; hornbill species, following Ter Braak (1986) whereby Reichard, 2003; Bartlett, 2003; Brockelman et al., 1998). data points are projected on a two-dimensional space that Khao Yai supports mainly a seasonally wet, evergreen forest maximizes the correspondence between categories on the (Kitamura et al., 2005), which experiences a distinct dry rows and columns (Poulsen et al., 2002). CA also helps to season (November–April) and wet season (May–October). The mean annual rainfall from 1993–2002 was 2,360 mm (Kitamura et al., 2005). Average daily temperature varied annually between 18.7 to 28.3°C and mean humidity ranged from 64.6% to 77.1% during the dry and wet season. Although fi gs (Ficus spp.) produce syconia through out the year (Poonswad et al., 1998b), the diversity and abundance of non-fi g fruiting species are relatively high only in the rainy season and beginning of the dry season (Bartlett, 2003; Savini et al., 2008).

Diet composition – During the breeding season, the female hornbill imprisons herself in the nest site, a cavity in a tree trunk, and is fed by the male during the entire period of her egg incubation and chick development. The male moves constantly between the nest and the selected feeding sources (Kemp & Poonswad, 1993). We studied characteristics of food items consumed by hornbills in the fi eld by using direct observations of feeding behaviour at nests by continuous observation of the visiting male (Altman, 1974). At least six nests of each hornbill species were monitored each breeding season, for a total of 24 nests over a period of two breeding seasons (2004 and 2005). The fi rst author, with the help of research assistants, made feeding observations at each nest during four days per month for 10 hours per day (between 0500–1700 hrs). We scored each feeding visit by the male Fig. 1. Hornbill and gibbon study site.

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fi nd the best simultaneous representation of the diet records observation where some plant species were not fruiting each (Gorchov et al., 1995), with frugivorous species as column year. More in detail were the two components that accounted and fruit species as rows. Species showing similar diets for 35% of the variance (variance for Component 1 was will appear positioned close to each other. To determine 21% and Component 2 was 14%) when all plant species which food items generated overlaps, we removed targeted were considered (Fig. 4a). After excluding Ficus sp. and P. plant species and re-ran the CA analysis. We included in viridis from the analysis (Fig. 4b), we observed that both the CA all plant species and animals consumed per hour by components accounted for 33% of the variance (variance for gibbon and hornbills based on direct observation. We used Component 1 was 19% and Component 2 was 14%). When the Shannon-Wiener diversity index to defi ne the diversity only P. viridis was removed from the analysis (Fig. 4c), both of each species’ diet. components accounted for 35% of the variance (variance for Component 1 was 21% and Component 2 was 14%). After removing only the genus Ficus from the analysis, our result RESULTS was similar to removing both Ficus sp. and P. viridis from the analysis, where the grouping along the two components Diet composition – The general composition of the diet accounted for 33% of the variance (variance for Component during the breeding and/or nesting season in the fi rst half 1 was 19% and Component 2 was 14%; Fig. 4d). of the year (January to June), for both hornbill and gibbon species had a higher component of fi g and animal matter Resource abundance - Sample percentages of individual in the diet of hornbills, while the gibbon diet had a higher species of diet trees that overlapped did not vary much proportion of non-fi g fruits (Fig. 2; see Table 1 for species between the fi ve species (Table 2). However, the proportion details). Diet diversity, measured using the Shannon-Wiener of trees included in the non-overlapping portion of the diet diversity index, was: Gibbon (GB) = 2.1, Great Hornbill showed a distinct difference between the gibbon and the (GH) = 1.2, Wreathed Hornbill (WH) = 1.6, Oriental Pied four hornbill species, with the gibbons having a much higher Hornbill (PH) = 1.7 and White-throated Brown Hornbill (BH) proportion of their diet non-overlapping. This suggests that = 1.8; where higher numbers show a higher diet diversity. gibbons feed on a much larger set of resources that they do The mean estimate for hornbill Shannon-Wiener diversity not share with hornbills. index was 1.575 with a 95% confi dence interval of ±0.515 with the Gibbon value falling just above the interval. A comparison of the density and basal area between Ficus spp. and P. viridis, the two species composing most of the Food overlap – Fig. 3a shows the results for CA calculated diet overlap, indicated there was a larger amount of fi gs by including all plant species (n = 58) consumed by gibbons available, mainly as a low number of individuals but of a and hornbills. The two components account for 84% of large size. Conversely, P. viridis was present in the area the variance (variance for Component 1 was 66% and as a large number of individual trees but of much smaller Component 2 was 18%), separating BH and PH from WH size (Table 1). and GH, among hornbills, and from GB, which are isolated from the hornbills. After excluding from the analysis Ficus sp. and P. viridis (Fig. 3b), we observed a clear separation DISCUSSION between GB and the four hornbill species, which appeared grouped, with the two components accounting for 88% From a general dietary perspective, hornbills were less of the variance (variance for Component 1 was 69% and frugivorous, fed on a smaller number of plant species, Component 2 was 19%). To isolate the cause of this overlap, and fed more on insects, small mammals and birds than we analyzed the importance of the genus Ficus and of P. gibbons. Based on the correspondence analyses, overall we viridis separately. When removing only P. viridis from the detected only a moderate degree of overlap between gibbons analysis, we saw no obvious change from what was observed and the two larger species of hornbill (Great hornbill and in Fig. 3a, with the separation of the fi ve species along the Wreathed hornbill), while the two smaller hornbill species two component accounting for 85% of the variance (variance (Oriental pied hornbill and White-throated Brown hornbill) for Component 1 was 67% and Component 2 was 18%; Fig. appear more clearly separated. Even this moderate degree 3c). After removing only the genus Ficus from the analysis, of overlap fell away once trees of the genus Ficus and, in we obtained a result similar to as when Ficus sp. and P. a lower proportion, of the species P. viridis, were removed viridis was excluded, where the grouping along the two from the calculation. As a consequence of the relatively low components accounted for 87% of the variance (variance nutritional quality (O’Brien et al., 1998) and the abundant for Component 1 was 68% and Component 2 was 19%; distribution of the genus Ficus, we assumed a low importance Fig. 3d). These results suggest that the main component in for any such overlap to both animal groups. the overlap was represented by species of the genus Ficus, with a smaller proportion represented by P. viridis. Overall, both hornbills and gibbons persist in habitats of similar forest structure. Hornbills, as well as other large A similar pattern was observed when CA considered all the birds are able to observe the forest during fl ight and so detect individual nests observed for hornbill and each group observed fruits in the upper strata of the canopy (Kemp, 1995). This for gibbons (Fig. 4). Note that the two clusters observed in differs from most primate species, which are more likely all hornbill species are the consequence of two years nest to detect fruits in the middle strata of the canopy while

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traveling through the forest (Poulsen et al., 2002). However, (Poonswad, 1993; Savini et al., 2008). For the gibbon, this gibbons show the peculiar capacity to forage mainly on the period coincides with the pre-conception period when females outer part of tree canopy, a consequence of their specialized must acquire a certain amount of body reserves in order to morphology and physiology (Carpenter, 1967). start cycling regularly and therefore conceive successfully (Savini et al., 2008). For the hornbills, as for other bird We observed an overlap in the reproductive season between species, the nesting period is considered the period of the hornbills and gibbons at the study site during the fi rst half year where high costs are faced by both the reproductive of the year, when dependence on resources was higher male and female (Poonswad et al., 2004).

Fig. 2. Diet composition variation, during breeding season, between hornbill species [Great Hornbill (GH), Wreathed Hornbill (WH), Oriental Pied Hornbill (PH), and White-throated Brown Hornbill (BH)] and White-handed Gibbons (GB).

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Table 1. Species list in the diet of the four hornbill species hornbill species [Great Hornbill (GH), Wreathed Hornbill (WH), Oriental Pied Hornbill (PH), and White-throated Brown Hornbill (BH)] and White-handed Gibbons (GB).

Plant Density Chemical Composition Percentage in the Diet Basal area / hectare Protein Fat Calory Ca+ GH WH BH PH GB g% g% kal/g g%

Animal - - - - - 3.57 4.42 16.97 8.18 0.30 Acacia megaladena ------2.99 Adinadra integerrima 20.9 ------0.37 Aglaia elaeagnoidea 37.8 7.72 - 4259.77 0.46 - - - - 0.30 120.9 10.32 48.47 6493.3 0.59 0.64 0.48 0.67 0.76 - Aglaia spectabilis - 3.67 6.39 2200.77 0.31 0.93 2.46 0.26 0.20 - Alphonsea boniana 36.3 ------0.18 Anthocephalus chinensis ------0.08 Antiaris toxicaria 7.2 12.18 11.84 4670.6 0.4 0.03 - - - - Aphanamixis polystachya 66.8 8.01 - 6489.79 0.44 - 0.97 0.04 - - Aquilaria crassna 218 ------7.26 Arthocarpus lakoocha - 5.27 9.83 4113.76 0.4 0.10 - - - - Baccaurea ramiflora 125.1 ------1.33 Balakata baccata 586.2 4.04 1.63 3877.47 - - - - - 3.51 Beilschmiedia balansae - 4.65 6.93 2675.51 0.45 0.09 0.43 0.58 0.11 0.10 Beilschmiedia glabra 69.9 8.34 32.84 5969.97 0.17 0.00 0.07 0.29 0.02 - Beilschmiedia maingayi 70.02 9.92 - 4842.06 0.68 0.31 0.83 1.97 0.80 0.14 Beilschmiedia sp. - - - - - 0.06 0.01 0.04 0.03 - Bhesa robusta 12.2 12.75 - 6195.61 0.78 2.40 2.28 7.06 2.06 - Canthium glabrum - - - - - 0.71 - 0.32 0.73 - Carallia brachiata 68.8 ------0.62 - Cinnamomum subavenium 395.3 5.71 31.92 6390.68 0.3 19.12 12.84 37.06 15.57 2.89 Cleistocalyx opercolatus 768.9 5.83 ------4.41 Combretum acuminatum - - - - - 0.29 - - 4.57 - Cryptocarya impressa - 12.73 26.69 6512.13 0.52 0.16 2.81 1.50 1.20 - Desmos chinensis climber 5.38 1.87 4465.84 0.63 0.90 0.05 0.43 0.11 4.58 Dipterocarpus gracilis 873.7 ------0.07 Duabanga grandiflora ------2.51 Dysoxylum cyrtobotryum 143.3 7.4 16.33 4653.66 0.52 0.38 0.10 5.08 3.18 - Dysoxylum densiflorum - 11.84 - 6355.67 0.63 1.11 2.09 3.27 1.57 - Elaeagnus latifolia climber 4.81 0.12 1500.27 0.29 0.17 0.35 0.04 0.07 2.44 Ericybe elliptilimba climber ------4.90 Ficus sp. 108.2 4.84 5.07 3093.59 1.57 57.56 44.15 9.84 38.12 33.59 Gironniera nervosa 348.8 ------1.71 Gnetum montanum climber 13.2 2.33 4655.83 0.45 - - - 0.46 1.35 Horsfieldia glabra 6.6 3.56 41.61 6939.2 0.32 1.27 1.36 3.15 3.33 - Knema elegans 132.5 5.93 18.2 4955.44 0.68 0.00 0.59 0.12 0.23 0.74 Livistona speciosa - 4.16 30.61 5472.85 0.26 1.99 3.40 - - - Mangifera sp 21.3 ------0.07 Nauclea orientalis 205.5 ------0.01 Nephelium melliferum 250.4 5.8 11.61 3996.05 0.34 - - - - 8.02 PeninsularPeninsular_sp sp. ------0.79 Platea latifolia 63.3 3.24 - 2062.74 0.66 - - 0.11 0.28 0.04 Polyalthia jucunda - 11.59 6.36 4723.72 0.57 1.27 3.13 1.17 0.95 - Polyalthia viridis 56.1 9.9 12.78 4939.84 0.6 5.03 14.73 2.08 5.10 5.36 Prunus arborea 12.4 - - - - - 0.08 - - - Prunus javanicus 22.5 - - - - 0.25 1.06 0.58 0.66 1.35 Ratan sp. ground vegetation ------1.25 Sabia limoniacea ------0.73 Salacia chinensis climber ------0.31 Sarcosperma arboreum 206.9 - - - - - 0.05 1.28 - - Symplocos cochinchinensis 764.5 ------3.22 Sterculia balanghas ------0.05 - - - Sterculia sp. - - - - - 0.14 1.11 0.11 0.20 - Syzygium sp. - 2.03 - 4453.14 0.05 0.71 - 5.58 10.86 0.01 Ternstroemia wallichiana 8.7 - - - - 0.80 0.13 0.39 - - Walsura robusta 79.1 4.6 4.74 3654.17 0.42 - - - - 2.15 Ziziphus attopoensis ------0.04 0.94

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For several reasons, this small overlap in resources might suggested that the structure of fruits might infl uence their affect hornbills more than gibbons. The gibbon’s diet selection by gibbons, which prefer ripe fruits with juicy appears to be more diverse than that of hornbills. Overall, pulp in bright colors (red, orange and yellow) and with Asian primates are known among all primate species for one large generally well-protected seed. White-handed their high diversity in diet (Kappeler & Heymann, 1996). gibbons are also generalized frugivores, consuming various However, the higher diversity of primate diet versus hornbill morphological types of fruits because of the advantage of diet has also been observed in lowland rainforest of south- hands for manipulation and long digestive tracts that allow central Cameroon (Poulsen et al., 2002). In more detail, consumption of fruit species with hard covers and fl esh the diversity of the gibbon’s diet is a consequence of both attached to seeds (e.g. Choerospondias axillaris, Sandoricum lower selectivity by gibbons, in terms of fruit morphology, koetjape and Garcinia xanthochymus) (Kanwatanakid, 2000). and their travel mode, which allows them to move through Conversely, hornbills do not consume these fruit species the canopy by brachiating at relatively low energetic cost because their feeding adaptations are restricted by their bill (Usherwood & Bertram, 2003). From the point of view of morphology, which signifi cantly reduces their capability to fruit morphology, Kanwatanakid & Brockelman (2005) manipulate and extract food. Hornbills tend to select the

Fig. 3. Correspondence Analysis per animal species: a, with all plant species included (n = 58); b, with the genus Ficus and P. viridis excluded; c, with only P. viridis excluded; d, with only the genus Ficus excluded. Hornbill species [Great Hornbill (GH), Wreathed Hornbill (WH), Oriental Pied Hornbill (PH), and White-throated Brown Hornbill (BH)] and White-handed Gibbons (GB).

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Table 2. Amount of individual trees belonging to overlapping and non-overlapping diet species (data expressed in % over total tree population measured in the 40 hectare botanical plot). Hornbill species [Great Hornbill (GH), Wreathed Hornbill (WH), Oriental Pied Hornbill (PH), and White-throated Brown Hornbill (BH)] and White-handed Gibbons (GB).

GB GH WH OPHPH BH

Overlapping sp. 5.2 5.2 4.4 5.2 1.8 non-overlapping sp. 22.5 4.0 4.1 4.4 5.0

Fig. 4. Correspondence Analysis per hornbill nest and gibbon group: a, with all plant species included (n = 58); b, with the genus Ficus and P. viridis excluded; c, with only P. viridis excluded; d, with only the genus Ficus excluded. Hornbill species [Great Hornbill ◆, Wreathed Hornbill ■, Oriental Pied Hornbill ▲, and White-throated Brown Hornbill ■] and White-handed Gibbons ▲.

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ripest fruits that show split (dehiscent) husks when ripening more resources (Milinski & Parker, 1991), also referred to and high nutritional value (Poonswad et al., 1998a). During as pseudo-interference (Free et al., 1977). We do not exclude digestion, seeds do not pass through the digestive track and the presence of aggressive behaviour, but we expect the allow complete detachment of the nutritious fl esh from the limit of such events to be sporadic. From our study, we can seed. In birds in general, regurgitated seeds spend less time classify gibbons as “depleting competitors”, the species that in the digestive track than seeds defecated by other animals is more effi cient in reaching and depleting a food resource, (Johnson et al., 1985), to limit body weight and reduce fl ight and hornbills as “unsuccessful competitors” that may suffer costs. An alternative explanation for the low diet diversity from the exploitation effects (Milinski & Parker, 1991). in hornbills may be due to their larger home ranges (from 5.9 km2 for BH to 30.8 km2 for WH, Poonswad & Tsuji, 1994), compared to gibbon [on average 25 hectares (0.25 ACKNOWLEDGMENTS km2), Savini et al., 2008], which allows hornbills to select only high quality resources that can be found more easily We are grateful to Alan and Meg Kemp for facilitating the by covering a larger area. participation of CK-S and TS at the 4th International Hornbill Conference held on 6–10 November 2005 in South Africa, Due to the high cost of fl ying (Gill, 1989), the selection of where an early stage of this work was presented. G. A. Gale, foraging sites for birds is affected by the distance from the A. Kemp, A. Koenig, R. Corlett, L. P. Koh and S. Kitamura nesting site and quality of the resources (Drent & Daan, have provided useful comments in the various stages of the 1980). Hornbills will fl y preferentially to larger resources development of this manuscript. S. F. Yap and G. A. Gale that will provide them with suffi cient nutritional rewards. provided guidance in the statistical analysis. We would like On the other hand, the low cost of brachiation for gibbons to thank P. Chuylua, P. Desgnam, M. Kinglai, U. Martmoon, (Usherwood & Bertram, 2003) allows them greater mobility, C. Mungpoonklang, T. Ong-in, K. Plongmai, W. Soonpracon, resulting in multiple visits to a larger variety of sources due S. Sornchaipoon, U. Sunarat, A. Sungwal, P. Suwanlikid, to the relatively low cost of traveling. M. Tofani who assisted us during data collection in the fi eld (both for the gibbon and hornbill sections). We thank the Hornbill diet showed a relatively higher proportion of animal Department of National Park, Wildlife and Plant Conservation food items, which were absent in the gibbon’s diet, with the (CK-S and TS) and the National Research Council of complete absence of vertebrates reported in the Khao Yai Thailand (TS) for granting permission to conduct research population and insect consumption recorded at less than at Khao Yai National Park, and the park superintendent P. 1% over the study period (T. Savini, unpublished data). Vohandee, and his staff, for their assistance in the park. We may assume that the vertebrate component of hornbill’s This research was supported fi nancially by the Thailand diets may reduce the direct impact of resource overlap with Research Fund (the Royal Golden Jubilee PhD Program), gibbons for protein-rich fruit (e.g. P. viridis), and on which Mahidol University (Research Assistantships), and Thailand gibbons may be more successful foragers. The evolution of Hornbill Project (CK-S), and by the Max-Planck Institute the hornbill’s diet to include a higher quantity of vertebrates for Evolutionary Anthropology (Graduate Fellowship), and as a source of protein may also have arisen as a result of the Christian Vogel Fond (TS). competition with other frugivorous animals that feed on protein-rich plant parts. Our interpretation was supported partly by observations of Great Hornbill diets in southern LITERATURE CITED of Thailand (Budo Sungai Padi National Park), where the Great Hornbill consumed less than 5% of animal matter Altman, J., 1974. Observational study of behaviour: sampling during the late nesting period (Chaisuriyanun, 2005). This methods. Behaviour, 49: 227–267. amount is lower, although not signifi cantly lower, than the Bartlett, T. Q., 2003. Intragroup and intergroup social interactions 8% observed for the same species during the same nesting in white-handed gibbons. International Journal of Primatology, period in Khao Yai National Park. One explanation could be 24: 239–259. the lower gibbon density in the southern study area (WCS Beckerton, P. R., & A. L. A. Middleton, 1982. Effects of dietary Thailand, 2006) reduces competition for plant species with protein levels on ruffed grouse reproduction. Journal of Wildlife high nutritional value for proteins. Management, 46: 569–579. Brockelman, W. Y., U. Reichards, U. Treesucon & J. J. Raemaekers, Overall, our results show limited resource overlap between 1998. Dispersal, pair formation and social structure in gibbons gibbons and hornbills, mainly based on fi gs that are of low (Hylobates lar). Behavioural Ecology and Sociobiology, 42: quality generally, except for their calcium content (O’Brien 329–339. et al., 1998), but were relatively abundant at the study site. Brown, J. S., B. T. Kotler & W. A. Mitchell, 1997. Competition The recorded overlap might assume importance for the second between birds and mammals: A comparison of giving up species involved, P. viridis, because it shows a high value densities between crested lark and gerbils. Evolutionary Ecology in protein, which appears important during the breeding 11: 757–771. of both hornbills and gibbons, and was represented at the Camillo, E. & C. A. Garofalo, 1989. Analysis of niche of two study site by small crops. If the overlap of such a species sympatric species of Bombus (Hymenoptera Apidae) in South can generate competition between the two animal groups, eastern Brazil. Journal of Tropical Ecology, 5: 81–92. then we can consider it as a simple exploitation of one or

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Carpenter, C. R., 1967. A fi eld study in Siam of the behavior and Krasnov, B. R., N. V. Burdelova, I. S. Khokhlova, G. I. Shenbrot social relations of the gibbon (Hylobates lar). New York: The & A. Degen, 2005. Larval interspecifi c competition in two John Hopkins Press. fl ea species parasitics on the same rodent host. Ecological Chaisuriyanun, S., 2005. A Comparative study of diets of Great Enthomology, 30: 146–155. hornbill (Buceros bicornis) and Rhinoceros hornbill (Buceros Lee, P. C. & J. E. Bowman, 1995. Infl uence of ecology and energetic rhinoceros) during the breeding season in Budo Su-Ngai Padi on primate mothers and infants. In: Pryce, C. R., R. D. Martin & National Park Southern Thailand. [M.Sc. thesis in the Natural D. Skuse (eds.). Motherhood in human and nonhuman primates, Resource Management]. Bangkok. School of Bioresources Karger Press, Basel, Switserland..Pp. 47–58. and Technology. King Mongkut’s University of Technology Leighton, D. R., 1998. 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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 199Ð206 Date of Publication: 28 Feb.2009 © National University of Singapore

WHALE STRANDINGS IN INDONESIA, INCLUDING THE FIRST RECORD OF A HUMPBACK WHALE (MEGAPTERA NOVAEANGLIAE) IN THE ARCHIPELAGO

Putu Liza Kusuma Mustika School of Earth and Environmental Sciences, James Cook University, Townsville 4811, Queensland, Australia. Email: [email protected] (Corresponding author)

Pariama Hutasoit Reef Check Indonesia Jl. Tukad Balian 113 Renon, Denpasar 80234, Bali, Indonesia. Email: [email protected]

Cecilia Cicik Madusari Sekala Sustainable Development Consultancy, Denpasar – Bali, Indonesia. Jl. Pulau Kawe 39 Denpasar 80114, Bali, Indonesia Email: [email protected]

Februanty S. Purnomo Community Empowerment Offi cer, WWF Indonesia Solor Alor Project, Jl. Trans Lembata RT 15A / 07, Nubatukan, Lembata 86282 – East Nusa Tenggara Indonesia. Email: [email protected]

Adityo Setiawan Kaimana Program Monitoring Offi cer, Conservation International Indonesia, Jl. Batu Putih RT 4 Krooy, Kaimana 98654, West Papua, Indonesia Email: [email protected]

Katherina Tjandra Monitoring Specialist, Conservation International Indonesia, Jl. Arfak No. 45, Sorong 98413, West Papua, Indonesia. Email: [email protected]

Walesa Edho Prabowo Matalabiogama (Biology Nature Club), Faculty of Biology, Gadjah Mada University, North Sekip, Yogyakarta 55281, Indonesia. Email: [email protected]

ABSTRACT. – The paper presents whale stranding records in Indonesia from 1987 to 2007. Most identifi ed stranding species were sperm whales (Physeter macrocephalus), followed by short-fi nned pilot whales (Globicephala macrorhynchus). In total, almost half of all stranding events involved unidentifi ed cetaceans, indicating the need of more training on proper procedures of managing stranded whales and dolphins. Yet, despite an insuffi cient stranding network in the country, there was a well-recorded stranding of a young humpback whale (Megaptera novaeangliae) on 2 and 9 October 2007 in Bali, Indonesia. Colouration of the upper side of the pectoral fi n, calf length and time of stranding suggest that the calf belonged to a Southern Hemisphere population, possibly Australia’s. The humpback whale stranding in Bali was the one of the fi rst recorded incidences of the species’ presence in the Archipelago, hence signifying the importance of a good stranding network in Indonesia.

KEYWORDS. – Whale stranding, Humpback whale, Megaptera novaeangliae, Bali, Indonesia.

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INTRODUCTION times, including Bryde’s whale (Balaenoptera edeni), an TO WHALE STRANDINGS IN INDONESIA unidentifi ed baleen whale and a humpback whale. Rescuers of half of the stranding event failed to measure the animal’s Indonesian waters, with their deep trenches and narrow straits, length (Table 1), indicating that training on proper procedures support at least 12 species of dolphins and porpoises and 18 to manage a marine mammal stranding event is needed. species of whales, including the blue whale (Balaenoptera musculus), sperm whale (Physeter macrocephalus) and Most cetacean stranding events were recorded in Central humpback whale (Megaptera novaeangliae) (Rudolph et al., and Eastern Indonesia, particularly in Bali, Maluku Islands 1997; Kahn, 2002). Up to the early 20th century, information and West Papua (Fig. 1, Table 1). Only one stranding was on cetacean diversity in the country was mostly obtained recorded in Sumatra (unidentifi ed cetacean in Bengkulu). from stranding events recorded by Dutch scientists (Rudolph Four stranding events were recorded in Java (melon-headed et al., 1997), until more regular research was conducted to whale (Peponocephala electra) in Jakarta, sperm whale update the species list (e.g. Barnes, 1996; Kahn, 2002, 2004, (Physeter macrocephalus) in Yogyakarta, short-fi nned pilot 2005; Kreb, 2007). Stranding events, however, still hold an whales (Globicephala macrorhynchus) in Banyuwangi and important role in our understanding of cetacean populations unidentifi ed cetaceans in Surabaya). Since 1987, at least in Indonesia, notably for species such as the humpback nine stranding events were reported for Bali, mostly on the whale of which presence data was still obscure until the southern shores of the island. The fi rst well-recorded stranding late 20th century (see Rudolph et al., 1997). Although this of humpback whale also took place in Bali; however, this paper describes records of cetacean stranding events in the does not imply that there were no other humpback whale Archipelago, the focus is on whales and large delphinids, sightings in the Archipelago (see following subsections). This for there is no available information on the strandings of high number for Bali might be due to better communications small delphinids. and networks for stranding reports, and does not necessarily reflect a lack of stranding events in more remote areas The stranding data from 1987 to 2007 were compiled of Indonesia. The number of strandings recorded in Bali by Marine Mammals Indonesia, a mailing list group of emphasised the need for a good marine mammal stranding several individuals that are interested in the conservation network on the island, and an effort was initiated by WWF and research of marine mammals in the Archipelago. The Indonesia and APEX Environmental in mid-2002. Detailed educational materials for such training have been developed sources of this information include: electronic news, TV for Indonesia for on-going cetacean projects (Kahn, news, printed media, and personal observations. Table 1 2005). Because communication can be difficult at most lists 35 stranding events from 1987 to 2007, but only three of the stranding locations, local-level networks should be pieces of information were available from the 1980s and encouraged to minimise delays in rescue and data collection 1990s, i.e. the 1987 stranding in Jimbaran (Bali), the 1996 efforts. In spite of inadequate stranding data in Indonesia, stranding in Manado (North Sulawesi) and the 1997 stranding it was fortunate that the humpback whale stranding in Bali in East Kalimantan. This trend demonstrates that tracing (described later in this paper) received enough coverage to data from periods before the internet and electronic news result in suffi cient data for further analysis. archives were widely available is diffi cult. Data from 2000 onwards are more substantial, we feel this is part due to Pre-21st century sightings of humpback whales in Indonesia greater communication and access to records and part because are illustrated in Fig. 2. The first historical record of a of the greater attention given to cetacean stranding events humpback whale in Indonesia was Gervais’ record of a in Indonesia, as opposed to strandings in the Archipelago stranding in Pekalongan on the northern coast of Java Sea, having increased. There is a peak of recorded stranding on 12 Apr.1863. Records of Dutch whaling ships in the events in 2007 (a total of 10 strandings), yet we are not Indian Ocean from 1953Ð1958 also revealed humpback whale able to ascertain whether it simply an artifact of improved sightings westward of Aceh in Sumatra (in February, March, news coverage. July, and November) and in the Java Sea (in December). In the Pacifi c part of the Archipelago, humpback whales were Almost half of the stranding events in Indonesia involved seen in these same years around the Arafura Sea in March unidentified cetaceans (15 cases or 42.86%). Three of and the Flores Sea in September and December (Slijper et these were unidentifi able due to decay. Ten unidentifi ed al., 1964). However, photographic confi rmation of humpback strandings took place in remote places, but the remaining whale’s presence in Indonesia was not available until mid- fi ve happened in major towns or provinces (one in Surabaya 2005. The fi rst recorded sighting of humpback whales in in East Java and four in Bali). This trend signifi es that, in Indonesia in the 21st century was in ‘Batu Moncong’, north addition to geographical obstacles, insuffi cient attention to of Komodo Island inside the Komodo National Park. The stranding events may also contribute to the high number of humpback whale was sighted by a local guide (CN Dive) unidentifi ed species. There is no information whether or not in 1999 without photographic evidence. Afterwards, no the unidentifi ed strandings involved small delphinids. other sighting was recorded in Komodo and adjacent waters despite a long-term cetacean program in this area (Kahn, Most of the identifi ed cetaceans were sperm whales (Physeter 2002; Kahn, 2004), nor have humpbacks been sighted during macrocephalus, 10 cases or 28.57%), followed by short- other extensive cetacean survey work in East Indonesia that fi nned pilot whales (Globicephala macrorhynchus, 3 cases included the waters off Lesser Sundas, East Kalimantan, or 8.57%). Baleen whale strandings were only recorded three Sulawesi and West Papua (Kahn, pers. comm.).

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Another humpback whale was found on 19 Jun.2005 near km away from the original stranding spot. The chronologies Lamalera, and was harpooned by the local hunters. The of the stranding events are based on the accounts of the fi rst whale was approximately 8 m long and was heading west- two authors of this paper that witnessed both strandings and east when it was sighted by the hunters of “jonson” (a local participated in the rescue of the fi rst stranding event. dinghy with outboard engine) Ronnie. Before the whale was caught, the fi shers thought that the whale was a “seguni” Upon learning of the stranding event, the Tanah Lot (killer whale, Orcinus orca) due to its whitish ventral area. Authority immediately contacted the coast guard from As “Lama fa” (harpooner) Matheus Ebang harpooned the several adjacent beaches and local villagers. Considering whale, they learned that it was a “kelaru” (baleen whale) the whale’s resistance and the heavy surges in Tanah Lot, instead. The whale was hauled back to the village. At the the rescue team decided to haul the whale with a boat to beach, the villagers realised that the “kelaru” had distinctive the neighbouring Kedungu Beach (approximately 1 km features unlike they had seen before (“knobby snout as in away westward) and cut the nets there. After a few hours crocodile’s”). A photograph taken during the event revealed of rescue efforts, the whale was fi nally set free and swam that it was a humpback whale. Because interviews with the back out to sea around 1400 hrs. harpooner and villagers of Lamalera revealed no previous historical record of humpback whale sightings in Lamalera, There was another stranding a week later on 9 Oct.2007, the occurrence of this whale in the Savu Sea is particularly this time at Kelating Beach in Tabanan, 7 km northwest of interesting. Photographic evidence of the harpooned Kedungu Beach. Apparently, the whale had stranded the night humpback whale in Lamalera was provided by Benjamin before at approximately 0200 hrs and was already dead and Kahn, courtesy of Mrs. Udis Keraf. This photograph is the decaying by the time it was found by the locals. To prevent fi rst physical evidence of humpback whale occurrence in the foul smell and to avoid bad luck (for the whale stranded Indonesia. when the locals were having a religious ceremony), the Regent of Tabanan had ordered that the whale be burned on the spot with a simple religious ceremony (Fig. 4d). However, HUMPBACK WHALE STRANDING IN BALI some skin samples, measurements and pictures were taken before the carcass was all burned. Sex determination was The fi rst recorded sighting of humpback whale in Bali (and impossible, for it was already decaying (the whale must the second in Indonesia) was made on 2 Oct.2007 ca. 1000 have died offshore, perhaps 1Ð2 days after returning to the hrs, when a 6.1 m individual got entangled in a fi shing sea) and half-covered in tarpaulin, but the fl uke and the net approximately 350 m off Tanah Lot Beach, District of morphological characteristics left little doubt that it was Tabanan. It was found dead a week later on 9 Oct.2007, 7 truly a humpback whale. The whale’s size and timing of the

Fig. 1. Map of cetacean stranding events in Indonesia 1997Ð2007.

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16_Mustika_Pg 199-206.indd 201 2/19/09 12:06:25 PM Mustika et al.: Whale strandings, including a humpback whale, in Indonesia Fisheries/MMAF) Marine Affairs and Seputar Indonesia, Ria Fauziah (Ministry of 5 Sep.2003 Liputan 6 SCTV Putu Liza (WWF Indonesia) Putu Liza (WWF Indonesia) Budiono, RASI Foundation Bali Post Liputan 6 SCTV Tifa Siwalima Magazine, Budiono, RASI Budiono, RASI Foundation WWF Indonesia Ð Andreas H Muljadi (The Nature Conservancy) Ð RASI Foundation Ð Ð

Ð 1 Ð Ð Ð Ð Ð Ð Dead Dead Alive later died by villagers by by villagers 1 animal, dead 1 dead, burried Dead, then cut up 1 dead animal, 2 ton Alive when stranded,

samudra Ancol, died after 1 day.

Ð Ð 8 1 dead whale, then cut up Media Indonesia Ð Ð Ð 8 Ð Ð Ð Ð Ð 10 11 19 24 nned various 49 alive back to sea, nned Ð Ð n.a Ð Sperm whale Sperm whale Sperm whale Sperm whale Sperm whale Sperm whale

Short-fi Melon-headed whale 2.28 Ð 1, alive. Kept in Gelanggang Jodie Raditya

Ð

Risso’s dolphin Ð Ð Ð Short-fi pilot whale Sperm whale Ð Ð Ð sp. Baleen whale

Physeter macrocephalus

Globicephala Grampus grisseus Physeter Physeter macrocephalus Physeter macrocephalus Peponocephala electra Ð macrorhynchus pilot whale macrocephalus Ð

macrocephalus macrocephalus Balaenoptera Grampus grisseus Physeter Physeter macrocephalus Ð Globicephala macrorhynchus Physeter macrocephalus Ð

East Java, Indonesia East Kalimantan Central Maluku P. Rinca, TN Komodo

21 28 Aug.2005 Sorong, Papua

20 25 Aug.2005 Pulau Baai, Bengkulu 18 2 Mar.2005 19 5 Aug.2005 Skow Mabo Beach, Tanjung Priuk, 13 14 3 Sep.2003 22 May 2004 15 Larangan Beach, 16 Banyuwangi, 25 May 2004 Surabaya Kenjeran, Serangan, Bali 17 2004 24 Feb 2005 Berau, Maratua, Nusa Penida, Bali Jayapura Jakarta DKI

12 Sep.2003 Ngongurung, Dsn Kalimantan East 8 9 Apr.2001 Oct./Nov.2001 Bali Manokwari 10 Harbour, Papua, Benoa 11 7 Jul.2002 2002 Ð Kampung Kerora - Berau, Maratua, 7 2001 Bali Canggu, 4 2000 Bali Benoa, Tanjung Ð

Table 1. Data on cetacean stranding events in Indonesia. No. 1 2 Date 1987 Aug.1996 Manado Ria, Bali Tasik Jimbaran, Location Kalimantan Ð East Species Ð English name Length (m) Ð Condition Ð Source Ð 3 1997 5 Kaltim Mar./Apr.2001 6 Maratua, Sangkulirang, P. 2001 Ð Bali Perancak,

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16_Mustika_Pg 199-206.indd 202 2/19/09 12:06:26 PM THE RAFFLES BULLETIN OF ZOOLOGY 2009 (The Nature Conservancy) Andreas H Muljadi Kupang Post, 21 Oct.2007 Ichwan Susanto, Kompas Program) Indonesia-Raja Ampat Indonesia-Raja International (Conservation (The Nature Conservancy) Andreas H Muljadi Program) Indonesia-Kaimana International Anton Wijonarno (Conservation Conservation International already decaying (The Nature Conservancy) Walesa Edho Prabowo (16 Jan.2007), The Jakarta Post (16 Antara, 23 Aug.2006 Kompas, TNC report WWF Indonesia, Reef Check, Kompas, etc

Dead Dead Dead Dead Dead 8 stranded, 1 animal, dead, mostly skeleton mostly all released back Dead, decayed and decayed for one month 1 species dead, already Adityo Setiawan Dead, already decaying 1 dead, already decaying Anton Wijonarno

Ð Ð Ð already decayed, estimated 1 months M. Erdi Lazuardi Ð Ð 8 20 6.5 2Ð3 10.4 1.92, 2.1 nned various 93 stranded, 30 dead SCTV, Media Indonesia,

Ð Ð Ð

Humpback whale Sperm whale 6 1 alive. Found dead 7 days later. Reef Check Indonesia Ð

Bryde’s whale 12

Ð Sperm whale Sperm whale Ð

Pygmy killer various, Ð whale Short-fi

Ð Ð

macrocephalus Megaptera novaeangliae Physeter Ð

Balaenoptera edeni

Ð Physeter macrocephalus Physeter Ð macrocephalus

Feresa attenuata Feresa Ð

macrorhynchus pilot whale Globicephala

East Nusa Tenggara waiwo, Raja Ampat

Bitsyari Bay,

South Kalimantan

35 28 Nov.2007 Batanta Island 34 18 Oct.2007 Pangga Jarang Beach, Papua Manokwari, Ð 33 18 Oct.2007 Doreri Bay,

32 5 Oct.2007 Napier Beach,

31 3 Oct.2007 Boo Kecil Island 30 2 Oct.2007 Tanah Lot, Bali

29 29 Sep.2007 Namatota Island, Wakatobi Wangi-wangi, Papua Kaimana, 28 17 Mar.2007 Wandoka Beach,

25 Nov.2006 26 Wakatobi 15 Jan.2007 27 Trisik, Kulon Progo, 11 Mar.2007 Tomia, Wakatobi Yogyakarta

23 Apr.2006 19 Bali 24 Padanggalak, 20 Aug.2006 Teluk Tamiang, Sulawesi Central Table 1. Continued. No. 22 Date 16 Mar.2006 Donggala, Location Species name English (m) Length Condition Source

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Fig. 2. Historical records of humpback whales in Indonesian Archipelago and adjacent waters ca. 1953Ð1958. Estimated coordinates compiled from Slijper et al. (1964)

Fig. 3. Locations of humpback whale strandings in Bali

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stranding lead to the strong suggestion that it was the same the total length from snout to the end of the fl uke (6.1 m) whale that became entangled the previous week. and fl uke width (1.67 m). As humpback whale newborns are 4Ð5m length (Carwardine, 2002), we suggest that the whale Humpback whales are renowned for the distinctive long was a calf. As mentioned, we initially hoped that the ventral pectoral fi ns and the photographs taken during the stranding fl uke photograph would help in individual identifi cation. event on 2 Oct.2007 in Tabanan Bali indicate that the stranded However, young humpback whales are known to dramatically whale was indeed a humpback (Megaptera novaeangliae). change ventral fl uke colouration over time, hence dorsal fi n Fig. 4a, which was taken on 2 Oct.2007, shows knobs on and caudal peduncle are considered the more appropriate the snout of the whale, another characteristic of humpback identifi cation method for younger animals (Blackmer et al., whales. The upper side of the pectoral fi ns were black, 2000). Unfortunately, these details were not recorded. In indicating that the whale might originate from an Australian addition, as the calf was possibly on its fi rst journey with the population. There is no accurate photograph that describes mother, it is unlikely that any researchers had photographed the ventral colouration, but Fig. 4b partially shows the black it elsewhere along its migration route before it stranded. and white pigmentation on the abdomen. Fig. 4c, which was taken on the same day, shows clear colouration of the An interview with Wayan Sujana, Head of Tanah Lot fl uke underside that is potentially very useful for individual Authority revealed that the calf had been sighted for two identifi cation. Fig. 4c was sent to several humpback whale weeks before it was entangled in fi shing net on 2 Oct.2007. researchers in the region, including Western Australia, Hawaii Approximately two hours before the rescue, the locals saw and the Philippines to see whether the fl uke matched any another ‘big fi sh’ roaming the waters next to the entangled in their catalogues. calf. Even though the locals were unsure whether the other ‘fi sh’ was a whale or a shark, the fact that it did not disturb The second humpback whale stranding in Bali (9 Oct.2007) the entangled calf lead to the assumption that the ‘big provided opportunity to take some measurements, including fi sh’ might be the calf’s mother. There is also anecdotal

Fig. 4. A humpback whale (Megaptera novaeangliae) stranding in Bali, Indonesia: a, knobs and black upper fl ipper of the humpback whale (Photo by Pariama Hutasoit, Reef Check Indonesia); b, black and white colouration of the ventral (Photo courtesy of I Wayan Sujana, Tanah Lot Authority Bali); c, the white underside of humpback whale’s fl uke (Photo courtesy of I. Wayan Sujana, Tanah Lot Authority Bali); d, Cremation process of the dead humpback whale in Tabanan, Bali (Photo by Putu Liza Mustika, James Cook University, Australia).

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information of regular sightings of whales with similar Indonesia. Interviews with local inhabitants of southern morphological characteristics off Tanah Lot (Sujana, pers. Bali, Lamalera, Komodo and other islands should also be comm.). The ‘Jro Gde’ (or whales in Balinese) occasionally conducted to retrieve more information on cetacean sightings displayed flukes and sang during the night time. The in the Archipelago. whales were always seen annually, around September and October, roaming the southern waters heading eastward. Their sightings usually lasted for more than a week before ACKNOWLEDGEMENTS they disappeared towards the direction of Uluwatu cliffs, in southern Bali. Residents of the Bukit Peninsula in southern The authors wish to thank Marine Mammals Indonesia, Mr. Badung Regency also witnessed whales breaching close to I Wayan Sujana (Tanah Lot Authority), Mr. Wira Sanjaya the shore in late Aug.2007; the whales might be several (Bahtera Nusantara), Mr. Bagaskara (Sekala Foundation) humpbacks travelling the southern waters of Bali (Kahn, and Mr. Chris R. Shepherd (TRAFFIC Southeast Asia) unpublished data). for technical input and discussions. Special thanks to Dr. Danielle Kreb (RASI Foundation), Mr. Benjamin Kahn July is usually the mating season for Southern Hemisphere (APEX Environmental), Mr. Jo Marie Acebes. The input humpback whales, with births occurring in June of the of two reviewers greatly improved this paper. subsequent year. A calf is generally strong enough to migrate with its mother at three months old (Gill & Gibson, 1997), i.e. around September of the year it was born. Hence, the time LITERATURE CITED of Bali stranding and the blackish upper side of the fl ippers (Fig. 4a) (Gill & Gibson, 1997; Shirihai & Jarret, 2006) Barnes, R. H., 1996. Sea Hunters of Indonesia: Fishers and Weavers suggest that the stranded calf originated from a Southern of Lamalera. Oxford University Press, New York. Hemisphere population. However, since 2006, fi shers have Blackmer, A. L., S. K. Anderson, & M. T. Weinrich, M. T., observed humpback whales off Bontang, northeast of the 2000. Temporal Variability in Features Used to Photo-Identify Mahakam Delta in East Kalimantan, Indonesia (Kreb, pers. Humpback Whales (Megaptera novaeangliae). Marine Mammal comm.). Acknowledging previous sighting and stranding Science, 16(2): 338Ð354. records of humpback whales in the country, it is also Carwardine, M., 2002. Whales, dolphins and porpoises. Dorling possible that the Bali calf belonged to a population unique Kindersley Ltd., New York. 288 pp. to the Archipelago. Gill, P. & L. Gibson, 1997. Whales, Dolphins and Porpoises. Reader’s Digest Association, Inc., New York & Montreal. 160 pp. CONCLUSION Kahn, B., 2002. Visual and Acoustic Cetacean Surveys and Evaluation of Traditional Whaling Practices, Fisheries With regard to the cetacean check list for Indonesia, Rudolph Interactions and Nature-based Tourism Potential. Alor and et al. (1997: 8) once stated that “Pending irrefutable evidence, Solor, WWF Indonesia Wallacea Program. 36 pp. we regard the occurrence of three species as still unconfi rmed: Kahn, B., 2004. Indonesia Oceanic Cetacean Program Activity Stenella coeruleoalba, Balaenoptera acutorostrata and Report: October–December 2003. Apex Environmental, TNC Megaptera novaeangliae”. Stenella coeruleoalba was sighted Indonesia. 29 pp. in the waters off Berau District in East Kalimantan in 2007 Kahn, B., 2005. Indonesia Oceanic Cetacean Program Activity (Kreb, 2007), while Balaenoptera acutorostrata in Indonesia Report: April–June 2005. Apex Environmental, Cairns. 31 was confi rmed by Kahn (2002). Irrefutable evidence of the pp. humpback whale as the last species unconfi rmed by Rudolph Kreb, D., 2007. Conservation and diversity of marine cetaceans in et al. (1997) is now available through the data presented in the Berau Archipelago, East Kalimantan, Indonesia. Technical this paper. Progress Report, fi eldwork period October 2007. Samarinda, RASI Conservation Foundation. 16 pp. Increased interest and coverage of cetacean stranding Rudolph, P., C. Smeenk, & S. Leatherwood, 1997. Preliminary events in Indonesia resulted in the recording of the stranded checklist of cetacea in the Indonesian Archipelago and adjacent humpback calf in Bali. It is important to strengthen the waters. Zoologische Verhandelingen, 312: 3Ð48. country’s stranding network by conducting meetings that Shirihai, H. & B. Jarrett, 2006. Whales, Dolphins and Seals. A&C include training on the proper methods of handling stranded Black London. 384 pp. cetaceans and procedures to collect data during the rescue. Slijper, E. J., W. L. v. Utrecht, & C. Naaktgeboren, 1964. Stranding events also provide excellent opportunities to Remarks on the Distribution and Migration of Whales, Based further study rare species such as humpback whales in on Observations from Netherlands Ships. Bijbragen Tot De Djerkunde, 34: 3Ð93.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 207–211 Date of Publication: 28 Feb.2009 © National University of Singapore

OBSERVATIONS ON BREEDING AND MOULTING OF THE GREY-EYED BULBUL, IOLE PROPINQUA, IN THAILAND

Andrew J. Pierce Conservation Ecology Program, School of Bioresources & Technology, King Mongkut’s University of Technology Thonburi, 83 Moo. 8 Thakham, Bangkhunthien, Bangkok. 10150. Thailand. Email: [email protected]

ABSTRACT. – I describe for the fi rst time observations on the nest, breeding and moult of the Grey-eyed Bulbul, Iola propinqua, a common species in evergreen forests of central Thailand. In 2007, two active nests were found and monitored at Khao Yai National Park. Only females built the nest and incubated eggs while the male assisted the female to provision nestlings and defend the nest. The overall nesting season lasted from mid-January to at least early May. Adults underwent a typical complete post-breeding moult with primary moult taking ca. 126 days from the end of May until early October. Juveniles underwent a complete moult of similar duration shortly after leaving the nest following which they were indistinguishable in the fi eld from adults. There was little or no overlap in the breeding and moulting cycles.

KEYWORDS. – Breeding, grey-eyed bulbul, Iole propinqua, moult, nest.

INTRODUCTION intensive nest searches for all bird species during the same period, no nests were seen until two active nests were found The life history of many tropical birds is poorly known in 2007. Here I describe these nests and their contents and and this is particularly true within South-east Asia (Sodhi, present details of the moult cycle of GBU none of which 2006). With the continuing declines in forest cover and forest have been previously documented. quality (FAO, 2007) it is essential that the basic ecology of species is understood for long-term management of bird communities. To provide more details of the ecology of METHODS forest birds in Thailand a long-term study has been conducted in and around the 30 ha. (0.3 km2) Mo-singto permanent Nests of all species in and around the study area were forest plot, Khao Yai National Park (KYNP), Nakhon Nayok actively searched for and monitored following the guidelines Province, (14°26'N 101°22'E) since 2002. The plot is situated of Martin & Guepel (1993) with precautions to reduce the in mature seasonally-wet evergreen forest at 723–817 m possibility of leading predators to the nest as recommended elevation (Brockelman, 1998). The average annual rainfall by Robinson et al. (2000). The GBU were caught using is 2,326 mm, most of which falls between May and October adapted mist nets; short 2–4 m four shelf nets or 6 m two (Kitamura et al. 2002). shelf nets were positioned along streams during the afternoon when birds came to drink and bathe, often in amongst dense The Grey-eyed Bulbul, Iole propinqua, (hereafter GBU) tangles of overhanging vegetation and woody climbers. ranges from South-west China through Myanmar, much of The GBU were occasionally susceptible to wing strain and Indochina and Thailand. At KYNP, it is a common resident apparent haemorrhaging so handling time was minimized found mainly in the mid- to upper story of forest and forest and the birds released immediately upon showing signs of edge. Elsewhere in its range, it may also inhabit secondary stress. Birds were fi tted with one numbered aluminium ring growth, up to an elevation of 1,525 m (Robson, 2000). They and two plastic colour rings in unique combinations to allow are highly frugivorous and were the most frequently observed individual recognition in the fi eld. A copy of all banding species at fruiting trees during a frugivory study on the Mo- and biometric data collected was deposited at the Wildlife singto plot (Sankamethawee, in prep.). Intensive surveys of Research Division, Department of National Parks, Wildlife, the study area using distance sampling during 2002–2006 and Plant Conservation, Bangkok. found GBU to be one of the most abundant species, with densities of approximately one individual per hectare (Gale I adopted the frequently used method of measuring primary et al., in litt.). Despite its abundance, and implementation of moult visually (Humphrey & Parkes, 1959). Each feather

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was assigned a score of 0 (old feathers) through to 5 (fully a thick layer of large apical leaves. The nest measured 65 grown new feathers). Thus, a bulbul with 10 primaries yet × 63 mm across with a total depth of 43 mm (the internal to start moulting has a moult score of 0 while one that has cup was too disturbed to measure accurately). The outside fi nished has a score of 50 (only right wing measurements of the nest was constructed of leaves held together with were used but both wings were checked to make sure the spiders’ web and horse hair-like fungal hyphae. A distinct moult was normal rather than accidental). As most passerines nest lining was composed of fi ne aerial rootlets of woody have a moult score that increases linearly with time (Ginn & climbers (Figs. 1, 2). Melville, 1983) this method gives a good indication of the progress of moult although it tends to slightly over estimate The eggs, viewed with binoculars from the ground with the total duration (Oschadleus & Underhill, 2006). In this the aid of a mirror, were off-white (pale fawn or creamish) analysis, birds with a moult score of 0 or 50 were only with small sparsely scattered dark fl ecks that were more included if they had started body moult or were still growing concentrated around the widest part of the egg forming a their innermost secondary, indicating that primary moult was slight ring effect. about to begin or had recently fi nished (Fogden, 1972). The same scoring system was used for the 6 secondaries, three Although these were the only nests found during this study tertials and all twelve rectrices giving total scores of 30, 15 several other observations of breeding activity were recorded. and 60 respectively. Birds carrying nesting material were recorded in all months from mid-January to late April. All records of GBU carrying nest material were of birds fl ying upwards and disappearing RESULTS into the sub-canopy or above suggesting that the two nests found were at the lower end of their height range, which is Two GBU nests found during 2007 were the fi rst to be probably why no other nests were found despite the fi nding of described for this species. Both were in Dipterocarpus over 300 nests of Puff-throated Bulbul, Alophoixus pallidus, gracilis Blume (Dipterocarpaceae) trees. The fi rst nest was found nearly complete on 15 Mar. when an adult was seen to make three visits within twenty minutes in order to add nest lining. On 19 Mar. the adult was seen incubating and continued to do so until at least 31 Mar. but by 2 Apr. the nest had been destroyed, probably depredated. The nest was ca. 10 m above the ground, built at the fork of a thin branch about 0.5 m from the trunk of the tree. It was shielded from above by leaves of a woody climber which encompassed the main tree. The nest tree was situated at the edge of a gap in the forest, about 20 m in diameter, caused by a tree fall.

The second nest was found in a 9.5 m tree on 25 Apr. by following an adult bird back to its nest. The nest tree was at the edge of a small gap less than 10 m across. Examination of the nest contents on 27 Apr., using a combination of ladder, pole and mirror, showed it to contain three eggs. The female was sitting on the nest on 29 Apr., 3 and 5 May. On 8 May both adults were seen bringing small invertebrate prey items to the nest and the heads of at least two nestlings were visible. During these feeding observations an adult Moustached Barbet Megalaima incognita landed on the nest tree and both bulbuls, alarm-calling, rapidly fl ew at it causing it to fl y off. Although the female was seen brooding on 10 May, during one hour of observation on 12 May no adults were seen in the area and the nest was considered to have failed. On 17 May, the nest was collected. It contained one decomposed nestling which had been fed on by Diptera larvae which had pupated within the nest structure. Some primary feathers of the nestling had just emerged from their sheaths suggesting it was 7–8 days old when it died.

The collected nest was a neat cup attached with spiders’ web between two thin (7 mm diameter), almost parallel, branches that came out from the main stem such that the rim of the nest was level with these branches. It was 6 cm from the stem and ca. 50 cm from the top of the tree below Figs. 1, 2. The nest of Grey-eyed Bulbul Iola propinqua.

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during the study period. A few recently fl edged birds (short in March 2005. Males caught during the breeding season tails, limited fl ight capabilities and begging behaviour) were had distinct cloacal protuberances but no brood patch. This observed, with the latest being 8 May 2006. This would indicates that only females incubate which is consistent with indicate that the complete breeding season was from mid- other bulbuls including Puff-throated Bulbul in which only January to early May making it one of the earlier nesting the female builds the nest and incubates eggs (Pierce at al., species in the study area (Pierce et al., in prep). 2007). Further proof of this came from the fi rst GBU nest where the nest-builder was accompanied by a second bird No direct evidence was obtained for the incubation and that neither carried nest material nor perched at the nest and nestling period. However, similar-sized birds with open cup was likely therefore to be a mate guarding male. nests on the Mo-singto plot, including Black-crested Bulbul, Pycnonotus melanicterus, and Puff-throated Bulbul, have a Moult. – Adults had a complete moult at the end of the total nest period from fi rst egg to fl edging of 25–28 days. breeding season. Regression of moult score against date for (Pierce et al., 2004, Pierce et al., in prep.). 44 adult birds measured in 2004 showed an average moult period of 126 days (18 weeks) from late May until early The bird building the second nest was known to be a female October (Fig. 3). The progression of moult was typical of as it had an obvious brood patch when fi rst caught and ringed

Fig. 3. Primary moult scores by date for adult Grey-eyed Bulbuls at Khao Yai National Park, Thailand in 2004.

Fig. 4. Primary moult score of Grey-eyed Bulbul plotted against secondary moult score.

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most passerines (Ginn & Melville, 1983) with a few head and body feathers being replaced shortly before the onset Rectrices were moulted from the centre outwards. They of regular-descendant primary (P) moult starting with the were replaced more quickly than the primaries (Fig. 6) innermost primary (P1). Between 1–3 primaries were in active starting when the primary score was > 10 and fi nishing moult at any time simultaneously (n = 46, median = 2) and by a score ca. 40 with 2–6 feathers on each side moulting there was no discernable variation the numbers of primaries simultaneously. in moult throughout the whole period of replacement. Juvenile GBU also had a complete moult. Ten juveniles in Secondary moult started after primary moult had progressed active moult were examined in 2004 and although this was to a score of > 10 beginning with the outermost secondary too small a sample to show precise moult periods the data and continued ascendantly (Fig. 4). The innermost secondary suggested that it was of a similar timing, duration and pattern was the last remex to fi nish growing, shortly after the outer to that of adults. Undergoing a complete post-juvenile moult most primary became full grown. Moult of the tertials is consistent with many tropical passerines (Wells, 2007; started with the centremost shortly after the fi rst primary Round, 2008), but unlike most temperate passerines which was dropped followed by the outer and fi nally the inner have distinct post-juvenile plumages following a partial tertial. All three tertials were fully grown by the time P8 moult (Svensson, 1992). Juveniles could be aged by their was dropped (Fig. 5). richer brown colouration but became indistinguishable from

Fig. 5. Primary moult score of Grey-eyed Bulbul plotted against tertial moult score.

Fig. 6. Primary moult score of Grey-eyed Bulbul plotted against rectrice moult score.

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adults after their fi nal old remex (innermost secondary) was study of animal diets in Khao Yai National Park. In Poonswad, dropped. The adult remiges, before replacement, were worn P. (ed.), The Asian hornbills: ecology and conservation. Thai at the tips unlike juveniles’ which, although sometimes Studies in Biodiversity, No. 2. Pp. 307–310. castellated (chipped) (Sutherland, 2000), had unworn fringes. Fogden, M. P. L., 1972. The Seasonality and Population Dynamics of Bare part colouration warrants further study as there is likely Equatorial Forest Birds in Sarawak. Ibis, 114(3): 307–343. variation in iris or bill colour as noted with other bulbuls Food and Agriculture Organization of the United Nations, 2007. (Round, 2008). This may allow aging after the completion State of the World’s Forests 2007, FAO Rome. (accessed of post-juvenile moult. There was no evidence of suspended Jul.2008). or partial moult of remiges as I have observed in other Ginn, H. B. & G. R. Melville, 1983. Moult in Birds, BTO Guide bulbuls (Black-crested Bulbul, Pycnonotus melanicterus, 19. BTO, Tring. and Streak-eared Bulbul, Pynonotus blanfordi). Humphrey, P. S., & K. C. Parkes, 1959. An approach to the study of molts and plumages. Auk, 76(1): 1–31. The data indicate that in GBU there is little if any overlap Kitamura, S., T. Yumoto, P. Poonswad, P. Chuailua, K. Plongmai, in their breeding and moulting cycles as has been found for T. Maruhashi & N. Noma, 2002. Interactions between fl eshy many other passerines due presumably to the high energetic fruits and frugivores in a seasonal forest in Thailand. Oecologia, costs of both activities. Fogden (1972), working in Sarawak, 133: 559–572. noted that moult rarely overlapped with breeding and this Marini, M. Â. & R. Durães, 2001. Annual patterns of molt and included observations of individuals of more than 20 species reproductive activity of passerines in South-central Brazil. that were known to have begun moulting after their young had Condor, 103(4): 767–775. fl edged. Marini & Durães (2001) found that moult of remiges Martin, T. E. & G. R. Geupel, 1993. Nest-monitoring Plots: Methods and breeding overlapped in less than 2% of passerines in a for locating nests and monitoring success. Journal of Field south-central Brazil. A few individual passerines at Khao Yai Ornithology, 64(4): 507–519. have been observed moulting while still feeding nestlings Oschadleus, H. D. & L. G. Underhill, 2006. Breeding seasonality [Puff-throated Bulbul and Black-naped Monarch, Hypothymis and primary moult in weavers in eastern South Africa. Durban azurea (pers. obs.)] but their nestlings would have fl edged by Museum Novitates, 31: 24–31. the time the majority of feather tracts started to moult. Fogden Pierce, A. J., K. Tokue, K. Pobprasert & P. D Round, 2004. (1972) suggests that low food availability in November (the Observations on the breeding of the Puff-Throated Bulbul lean season) in Sarawak may be the reason for moult being Alophoixus pallidus in north east Thailand. Folktail, 20: completed before this time. Ongoing investigations of fruit 101–102. phenology and availability at Khao Yai, where the Pierce, A. J., K. Tokue, K. Pobprasert & W. Sankamethawee, climate is more seasonal than Sarawak, may show there is 2007. Cooperative breeding in the Puff-throated Bulbul an even more pronounced lean season which in turn may Alophoixus pallidus in Thailand. Raffl es Bulletin of Zoology, affect breeding and moult seasonality. 55(1): 199–201. Robinson, W. D., T. R. Robinson, S. K. Robinson & J. D. Brawn, 2000. Nesting success of understory forest birds in central ACKNOWLEDGEMENTS Panama. Journal of Avian Biology, 31(1): 51–164. Robson, C., 2000. Field guide to the birds of south-east Asia, New I thank W. Sankamethawee for assisting with the ringing Holland Publishers Ltd., United Kingdom. of birds, P. Wonkson for fi nding the second nest and all Round, P. D., 2008. Birds of the Bangkok area, White Lotus Co., members of the Khao Yai Avian research team for searching Ltd. Bangkok. for and monitoring nests. I am grateful to W. Brockleman Southeast Asian Biodiversity for facilitating access to the Mo-singto plot. P. Round, Sodhi, N. S. & B. W. Brook, 2006. in Crisis, Cambridge University Press, Cambridge, UK. G. Gale and one anonymous reviewer provided valuable comments on earlier drafts of this manuscript. This research Sutherland, W. J., 2000. The Conservation Handbook: Research was supported by grant BRT 346004 from the Biodiversity, Management and Policy, Blackwell Science, U.K. Research & Training Program, Thailand and logistical Svensson, L., 1992. Identifi cation Guide to European Passerines, support was provided by King Mongkut’s University of 4th edition. Svensson, Stockholm. Technology, Thonburi, Bangkok. I am grateful to the Wells, D., 2007. The Birds of the Thai-Malay Peninsula, Volume Khao Yai Training Center, Training Division, Department 2. Christopher Helm. of National Parks, Wildlife and Plants Conservation for providing accommodation and The Wetland Trust (U.K.) for supplying rings.

LITERATURE CITED

Brockelman, W. Y., 1998. Long term ecological research plot for the

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 213–221 Date of Publication: 28 Feb.2009 © National University of Singapore

BAT DIVERSITY IN THE VEGETATION MOSAIC AROUND A LOWLAND DIPTEROCARP FOREST OF BORNEO

Daisuke Fukuda Center for Ecological Research, Kyoto University, Hirano 509-3, Otsu, 520-2113, Japan

Oswald Braken Tisen Sarawak Forestry Corporation, Level 12, Offi ce Tower, Hock Lee Center, Jalan Datuk Abang, Abdul Rahim, 93450 Kuching, Sarawak, Malaysia

Kuniyasu Momose Faculty of Agriculture, Ehime University, 3-5-7, Tarumi, Matsuyama, 790-8566, Japan

Shoko Sakai Center for Ecological Research, Kyoto University, Hirano 509-3, Otsu, 520-2113, Japan Email: [email protected] (Corresponding author)

ABSTRACT. – Tropical rainforests in Southeast Asia are decreasing rapidly because of conversion to agricultural lands, which in turn leads to an increase in mosaic landscapes. Little is known about the effects of these anthropogenic changes on bat fauna or feeding behaviour. To better explore anthropogenic effects on bats (Chiroptera), we investigated the diversity and feeding habits of bats in an intact lowland dipterocarp forest and surrounding areas of varying vegetation types. The total sampling effort using mist nets (32,795 m2h) resulted in the capture of 495 bats, representing 28 species. Simpson’s index of diversity was relatively high in primary forests (7.86) compared to secondary forests (3.38), orchards (3.65), and oil palm plantations (1.24). The capture rate of two frugivorous bats clearly differed among the four vegetation types, and these species were never caught in orchards or oil palm plantations. In addition, the capture rate of microchiropterans was notably lower in oil palm plantations than in the other three vegetation types. These results indicate that many bat species rarely use agricultural land for feeding, and these plant communities are not suitable for maintaining bat diversity. However, the data on the feeding habits of three megachiropterans that were frequently captured in orchards and oil palm plantations indicate that agricultural plants are their primary food sources. Orchards and oil palm plantations may be inadequate habitats for most frugivorous and insectivorous bats, but these plant communities may provide important food sources for certain species of megachiropterans.

KEY WORDS. – Lambir Hills National Park, megachiropterans, oil palm plantation, orchard, secondary forest, Sarawak.

INTRODUCTION The effects of these anthropogenic changes on mammals have been studied across several taxonomic groups (e.g., Ickes, Tropical forest landscapes are changing rapidly because of 2001; Cox et al., 2003; Kinnaird et al., 2003; Naughton- human activities. Approximately half of the potential tropical Treves et al., 2003). Among these groups, bats (Chiroptera) closed-canopy forest has already been removed and converted are regarded as one of the most vulnerable taxa. There are to other uses (Wright, 2005). Deforestation rates were 3.9 × about 1,000 known species of bats, nearly 25% of which are 106 ha/yr in tropical America, and 2.3 × 106 ha/yr in tropical globally threatened (Mickleburgh et al., 2002). According Asia during the 1990s (Hansen & DeFries, 2004). Wright to the IUCN Red List (IUCN, 2007), nine bat species (2005) predicted that in the near future, tropical landscapes are confi rmed extinct and 247 species are threatened (i.e. will only remain as islands of secondary and protected old- categorized as “Critically Endangered”, “Endangered” or growth forests isolated within a sea of land converted for “Vulnerable”). Major threats to bat populations include the human use and agriculture. loss or reduction in quality of foraging habitat and habitat fragmentation (Racey & Entwistle, 2002).

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The order Chiroptera is divided into two suborders: the In Malaysia, as of 2003, oil palm plantations covered 3.5 Megachiroptera with a single family, Pteropodidae (ca. 163 × 106 ha, and this number continues to increase. These species) and the Microchiroptera with 17 families (ca. 814 plantations constitute 11% of the total land area of the entire species in total; Corbet & Hill, 1992). Megachiropterans are country and 62% of all cultivated agricultural land (Brown known as Old World fruit bats and are distributed across & Jacobson, 2005). Africa, tropical Asia, India, Australia and their surrounding oceanic islands. Megachiropterans are relatively large (20– Such anthropogenic changes can create mosaics of fragmented 1,500 g) and predominantly feed on plants (fruits, nectar, vegetation, thereby greatly affecting the diversity, abundance, pollen, fl owers, and leaves). In contrast, microchiropterans and feeding behaviour of bats. Megachiropterans that inhabit are found on every continent except Antarctica, are relatively these mosaic landscapes are expected to feed on crops, small (1.5–150 g), and exhibit more diverse feeding because some agricultural plants may serve as food sources, habits (e.g., insectivorous, frugivorous, nectarivorous, and the bats can fl y long distances from mosaic to mosaic. ichthyophagous, and sanguivorous; Altringham, 1996). However, little is known about the effects and extent of the impacts of these anthropogenic changes. Although several Recent studies have indicated that many bat species studies have investigated bat fauna in Southeast Asia (e.g., play important roles in tropical rainforests. For example, Zubaid, 1993; Francis, 1994; Abdullah et al., 1997; Hall et pollination by bats is a phenomenon restricted to the tropics al., 2004; Heaney et al., 1991, 1999; Rickart et al., 1993), and subtropics. Megachiropterans visit at least 141 plant few study have compared bat fauna among plant communities species, including a number of commercially important plants under different land-uses including agricultural fi elds within (e.g. Durio, Ceiba and Parkia) for nectar or pollen (Fujita & a single landscape. Tuttle, 1991; Marshall, 1985). Bats may be more energetically costly to attract than other plant pollinators, but these costs We investigated the bat community in mosaic landscapes are offset by the extensive distribution of pollen provided by formed by the expansion of agricultural lands. Specifi cally, widely foraging bats (Start & Marshall, 1976). Some species we explored differences in the density and diversity of of bats travel long distances in one night (e.g., Eonycteris megachiropterans between primary forests and agricultural spelaea travels at least 38 km, Rousettus amplexicaudatus lands within a single landscape. These bat species are travels 25 km; Marshall, 1983). Megachiropterans feed upon important pollinators and seed dispersers of many plant 145 genera of fruits and presumably disperse the seeds of species and are critical to the of forests. We the majority of the fruits consumed (Marshall, 1985). Small also discuss the impact of increasing mosaic landscapes on seeds ingested by megachiropterans are transported over bat-plant interactions, focusing on nectarivorous bats. substantial distances, and larger seeds are frequently carried with fruits to feeding roosts. In Palaeotropical regions, fl ying foxes (family Pteropodidae) likely play a unique role in METHODS the longer-distance dispersal of the large-seeded fruits of dominant canopy trees on oceanic islands (Rainey et al., Study site. – We conducted our research in and around the 1995). In the Neotropics, where megachiropterans do not Lambir Hills National Park (LHNP), Sarawak, Malaysia (Fig. occur, nectarivorous or frugivorous microchiropteran species 1; 4°2'N,113°50'E, ca. 150 m in altitude). One characteristic belonging to the family Phyllostomidae act as pollinators or of the site was the high heterogeneity of vegetation. We seed dispersers. In addition, approximately 70% of extant selected four types of vegetation for bat censuses: primary bat species are insectivorous and prey on a diverse range of forests, secondary forests, orchards, and oil palm plantations. insects (e.g., Lepidoptera, Diptera, Coleoptera and Hemiptera; The primary forests were intact lowland mixed dipterocarp Jones & Rydell 2002). forests within the LHNP. The park covers an area of 6,949 ha (69.49 km2) and the height of emergent trees sometimes Southeast Asia is a hotspot of bat diversity. Like many exceeds 70 m. Shanahan & Debski (2002) recorded 10 species groups of organisms, the number of bat species per unit area of bats (fi ve megachiropterans and fi ve microchiropterans) is large in tropical zones, especially in the Amazon basin in the park. The secondary forests were young forests that and Southeast Asia (Altringham, 1996). In Borneo, bat is the had developed after slash-and-burn agriculture conducted most diverse order of mammals in terms of the number of by Iban villagers. Census points were established in three species, accounting for 42% of all mammal species (Payne forests of varying ages: a 7-year-old forest dominated & Francis, 1998). by Vitex pinnata, a ca. 30-year-old forest dominated by Artocarpus elasticus, and a > 60-year-old forest dominated by Southeast Asia also has the highest deforestation rate of any Artocarpus elasticus (Nakagawa et al., 2006). Forest height major tropical region, and currently, more than 50% of the varied among census points (2–25 m). These forests were land area in Asia is under agriculture (Zhao et al., 2006). surrounded by ponds, paddy fi elds, isolated intact forests, The conversion of forests to cash-crop plantations (e.g., oil and rubber (Hevea brasiliensis) forests. The orchards were palm, rubber, and cocoa) is thought to be one of the major small [< 5 ha (< 0.05 km2)] areas with many cultivated causes of the current high deforestation rates in the region plants (e.g., Durio kutejensis, Nephelium lappaceum, (Primack & Corlett, 2005). Since the 1970s, areas planted Carica papaya, Cocos sp., Musa sp., Parkia sp., Artocarpus with African oil palm (Elaeis guineensis), which is native to integer, Lansium domesticum, Piper sp., and Saccharum sp.) West Africa, have expanded in both Malaysia and Indonesia. established by Iban villagers. Tree height ranged from 1 to

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10 m. The orchards were located near villagers’ houses and m2h in the orchards (three census points), and 3,382 m2h in were surrounded by ponds, paddy fi elds, rubber forests, and the oil palm plantations (fi ve census points). bamboo groves. The oil palm plantations were large-scale [ca. 4,000 ha (40.0 km2)], continuous plantations of mature We identifi ed bat species according to Payne & Francis African oil palm managed by a corporation or Iban villagers. (1998). Because some studies have suggested the presence of Vegetation consisted of a complete monoculture, and the two cryptic species within Cynopterus brachyotis that differ heights of oil palms were 10–20 m. The plantations shared signifi cantly in genetic and morphological characteristics borders with the primary and secondary forests. as well as habitat preferences (Francis, 1990; Abdullah et al., 2000; Abdullah, 2003), we divided the species into Field methods. – Censuses were conducted four times between two categories (C. brachyotis I and C. brachyotis II). We Apr.2005 and Aug.2006 in all vegetation types except oil classifi ed individuals of C. brachyotis based on their forearm palm plantation. In the oil palm plantation we conducted length: in adult C. brachyotis I (larger form), the forearm censuses only for twice according to the availability of length ranged from 60 to 66 mm, and for adult C. brachyotis permission. We set mist nets on the ground along trails in II (smaller form), the forearm length was approximately the forests or forest edges. In the primary forests, we also 55 mm, but < 60 mm (M.T. Abdullah, pers. comm.). If set nets on canopy walkways (at a height of 15–35 m) for individuals of the species were juveniles and their forearm fi ve nights. There were no signifi cant differences between lengths were less than 60 mm, they were recorded as ‘C. data from the ground and the walkways, so the data were brachyotis (unidentifi able)’. We also used mark-recapture combined. We used two to four nets (24 to 36-mm mesh; 6–12 methods to assess bat movements and fi delity to a vegetation m long; 6.0 m high; eight shelves) per night and occasionally type. Prior to releasing bats, we attached aluminum bands harp traps (The Austbat Harptrap, Faunatech and Austbat, with running numbers (Lambournes Ltd., Leominster, UK) Australia; 4.2 m2 in area; four nights in primary forests to the forearm. and two nights in orchards). Traps were set at sunset and checked at 15-min intervals. Sampling continued until 2300 Feeding habits of megachiropterans. – We surveyed hrs unless it rained. We recorded age, sex, morphological the feeding habits of megachiropterans to determine measurements, and reproductive state of the captured bats. the availability of cultivated plants as food sources. Age class was determined by the degree of fusion of the Pollen samples were collected from the faces of captured epiphyseal plates on the phalanges (Kunz, 1988). Bats with megachiropterans using transparent sticky tape (18 × 57 unfused epiphyseal plates were regarded as juveniles. The mm) immediately after the removal of bats from the traps. number of census points and sampling effort varied among We also collected pollen from 21 species of plants growing vegetation type: 14,179 m2h [area of traps (m2) × sampling in the vicinity of the census points as reference samples. time (h)] in the primary forests (six census points), 8,707 Pollen from the megachiropterans were observed under a m2h in the secondary forests (three census points), 6,526 light microscope and identifi ed to genus by comparison to reference samples. Morphological features of the pollen were recorded following Erdtman (1966). Prior to identifi cation, we washed the pollen sample with 10% KOH at 55°C for 10 min, mounted it on glass slides with glycerin jelly, and sealed the cover glass with clear nail polish. We recorded a pollen genus only when > 10 grains of that type were present.

In the oil palm plantations, we tested if C. brachyotis I consumed fruits of the oil palm. An individual of the species and the fruit were placed into a cloth bag. After one hour, we checked whether the fruit was bitten or not. This test was repeated for three individuals.

Data analysis. – A Chi-square test (Sokal & Rohlf, 1973) was used to compare the number of captures of megachiropterans, microchiropterans, and each individual species among vegetation types. In the Chi-square tests, the observed and expected numbers of captures were compared. The expected number of captures was calculated assuming the capture rate was equal for each vegetation type. We did not distinguish seasons and census points, since we did not detect any signifi cant differences.

Species accumulation curves were depicted to examine Fig. 1. Study area and location of census points in four vegetation saturation of species numbers. Simpson’s index of diversity types in and around Lambir Hills National Park. Inset: the location of Lambir Hills National Park, Borneo, indicated by an arrow. (Simpson, 1949) was calculated for each vegetation type.

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Table 1. Capture rate of bats (individuals per 10,000m2h). Data for species with small sample sizes (< 20 individuals) are not shown. The rightmost column presents results of the Chi-square test: * = P < 0.05; ** = P < 0.01; *** = P < 0.001; - = test not carried out to due insuffi ciently large sample size.

Primary forests Secondary forests Orchards Oil palm plantations Signifi cance Megachiropterans 55 86 208 272 *** Microchiropterans 40 26 49 6 ** Each species Megachiroptera Balionycteris maculata 16 2 0 0 – Cynopterus brachyotis I 4 49 106 201 *** C. brachyotis II 8 11 6 9 – Eonycteris spelaea 4 1 44 12 *** Macroglossus minimus 5 3 31 0 *** Penthetor lucasii 13 3 0 0 – Microchiroptera Hipposideros cervinus 23 13 11 3 * Glischropus tylopus 4 1 31 0 –

Unidentifi able individuals of C. brachyotis (see above) were types (Table 1). The capture rates of Balionycteris maculata, assigned to C. brachyotis I and C. brachyotis II based on Penthetor lucasii, and Hipposideros cervinus were the highest the proportion of the two forms in the community. in primary forests. Capture rates were lower in secondary forests than in primary forests for all species except C. brachyotis I and II. In the orchards, the capture rates of RESULTS Eonycteris spelaea, Macroglossus minimus and Glischropus tylopus were very high compared to the other three vegetation Capture rate of bats. – The total sampling effort (32,795 types. In contrast, B. maculata and P. lucasii were not m2h) resulted in the capture of 495 bats representing 28 observed in the orchards. In the oil palm plantations, the species in five families (Appendix 1). The capture rate capture rate of C. brachyotis I was notably higher than in the of megachiropterans differed significantly (P < 0.001) other vegetation types. However, the capture rates of other among the four vegetation types and was particularly high bat species were low or zero in the plantations. in the oil palm plantations and orchards compared to the primary and secondary forests (Table 1). The capture rate Diversity of the bat community. – The species accumulation of microchiropterans also differed signifi cantly (P < 0.001) curves did not reach asymptotes but demonstrated clear among vegetation types and was lower in the oil palm differences among the four vegetation types (Fig. 2). plantations compared to the other three plant communities Simpson’s index of diversity also varied among plant (Table 1). communities (Table 2). Simpson’s index for megachiropterans was highest in primary forests, lower in secondary forests For the eight bat species with relatively large sample sizes and orchards, and lowest in the oil palm plantations. (> 20 individuals), capture rate varied among vegetation Cynopterus brachyotis I was the most common and abundant species in all vegetation types, except primary forests. This species accounted for 44% of all bats in the secondary forests, 41% in the orchards, and 72% in the oil palm plantations. In the primary forests, the dominant species was Hipposideros cervinus (24% of all bats), and C. brachyotis I accounted for only 4% of bats.

Recaptures. – We recaptured 25 bats of eight species (Table 3). Nineteen recaptures of seven species occurred at the same census point, and 16 of these were recaptures after 31 days or longer. This result indicate that some individuals of captured species have strong site fi delity. Recaptures at different census points in a short time (< 5 days) occurred four times. We also observed bat movement from orchards to the secondary forest twice. Even though the data were scarce, long-distance recaptures (> 1 km) demonstrated Fig. 2. Species accumulation curves indicating the cumulative that C. brachyotis I and M. minimus are capable of long number of species encountered relative to the total number of distance travel. individuals captured in each vegetation type.

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Table 2. Simpson’s index of diversity in the four vegetation types. (N: number of individuals, S: number of species, 1/D: Simpson’s index of diversity). Note that 1/D was not calculated for microchiropterans in oil palm plantations due to the insuffi ciently large sample size.

Primary forests Secondary forests Orchards Oil palm plantations Megachiropterans N 78 75 136 92 S 8 10 6 3 1/D 5.00 2.09 2.54 1.19 Microchiropterans N 57 23 32 2 S 11 8 5 2 1/D 2.95 3.50 2.24 All bats N 135 98 168 94 S 19 18 11 5 1/D 7.86 3.38 3.65 1.24

Table 3. Recaptures of bats. Numbers indicate the distance (m) between the original and recapture census points. * = transfer from the orchard to the secondary forest.

Days from fi rst capture to recapture Species 1–5 6–30 31–100 101–300 > 300 Megachiroptera Balionycteris maculata 0 0, 0 0 Cynopterus brachyotis I 0, 170, 1,600* 0, 0 0 0, 0 0, 0 C. brachyotis II 210 430* C. brachyotis (unidentifi able) 210 0 Macroglossus minimus 1,320 0 0, 0 Microchiroptera Glischropus tylopus 0 Rhinolophus trifoliatus 0 Megaderma spasma 0

Table 4. Proportion of individuals with pollen (> 10 grains), total number of pollen types, and number of pollen types per individual (mean ± SD and ranges in parenthesis) for each species. Data for species with small sample sizes (< 10 individuals) are not shown.

No. of sampled- Individuals with Total no. of pollen No. of pollen Species individuals pollen (%) types types/individual Balionycteris maculata 25 0 Cymopterus brachyotis I 146 5 4 1 C. brachyotis II 24 0 C. brachyotis (unidentifi able) 37 8 3 1 Eonycteris spelaea 38 58 5 1.7 ± 0.72 (1–3) Macroglossus minimus 29 76 5 1.7 ± 0.84 (1–3) Penthetor lucasii 19 0

Feeding habits of megachiropterans. – Pollen samples on E. spelaea (36 total adhesions). In M. minimus, fi ve types were collected from 85.3% of megachiropteran individuals. of pollen were observed, of which two were identifi ed as At times, pollen adhesion to the rostrums of nectarivorous Musa and Parkia. In total, these two plants accounted for bats (E. spelaea and M. minimus) was apparent upon visual 32% of the total pollen adhesions in M. minimus (37 total examination. The high rates of pollen adhesion to samples adhesions). Both E. spelaea and M. minimus used Musa and from E. spelaea (58%) and M. minimus (76%) indicated that Parkia, while four types of pollen (Durio sp., Unidentifi ed these species frequently visited fl owers (Table 4). In contrast, 2, 3, and 4) were unique to each species. pollen rarely adhered to the other bat species (0–8%). Five types of pollen were found on E. spelaea, of which three Fruits of the oil palm were bitten by C. brachyotis I each were identifi ed as Durio, Musa and Parkia. In total, these time they were enclosed with the bats. The exocarp and three plants accounted for 64% of the total pollen adhesions mesocarp of the fruit were heavily gnawed.

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DISCUSSION nectarivorous bats may be the abundance of fl owers in the orchards. The pollen examination indicated that three species The diversity of the bat community in LHNP did not differ of cultivated plants (Durio sp., Musa sp., and Parkia sp.) greatly from bat diversity in other old growth lowland were the primary food sources of E. spelaea and M. minimus, dipterocarp forests, although comparable studies are limited. although we did not distinguish between wild and domestic When accounting for differences in sample size, the number species within the genera. The particularly high capture rate of megachiropteran species in LHNP (eight species; 78 of C. brachyotis I in the oil palm plantations may indicate that individuals) was roughly similar to other study sites in they feed on oil palm fruits, as indicated by the preliminary Malaysia: Krau Wildlife Reserve (8; 352; Hodgkison et al., feeding experiment, although we do not know to what extent 2004a), Sepilok Forest Reserve (6; 75; Francis, 1994), and the bats depend on oil palm fruits. As Kinjo et al. (2003) Pasoh Forest Reserve (4; 23; Francis, 1994). In addition, the noted, agricultural lands can be important feeding locations number of microchiropteran species in LHNP (11 species; for certain species of megachiropterans. 57 individuals) did not differ greatly from the diversity in Sepilok (14; 53) or Pasoh (13; 71). The capture rate The capture rate of megachiropterans differed signifi cantly of megachiropterans in LHNP (55 individuals per 10,000 among vegetation types, with high rates in the oil palm m2h) was nearly equal to that in Krau (49) but higher plantations and orchards and low rates in the secondary and than in Sepilok (32) and Pasoh (14). The capture rate of primary forests. This trend was nearly opposite to the pattern microchiropterans in LHNP (40) was nearly equal to that of of bat diversity. The observed high capture rate and low Pasoh (44) and higher than in Sepilok (23). Because there species diversity of megachiropterans in agricultural lands were no large differences in bat diversity or capture rates were similar to the pattern reported by Hall et al. (2004) between LHNP and other sites, our sampling method appears although their sampling sites were scattered across Southeast valid. Thus, the bat community in LHNP can be regarded as Asia, and the crop species studied differed from those in our representative of old growth lowland dipterocarp forests. study. In the Philippines, Heaney et al. (1989) also measured high capture rates and low diversity of megachiropterans in To our knowledge, ours is the fi rst study to demonstrate clear agricultural lands relative to lowland forests. Thus, these differences in the diversity and abundance of bats among patterns may be common throughout Southeast Asia. Similar forests and agricultural lands within a single landscape in patterns have also been found in the Neotropics. For example, Southeast Asia. Relative to primary and secondary forests, Medellin et al. (2000) observed both low species richness the number of megachiropteran species was somewhat low of phyllostomid bats and high dominance of particular bat in orchards and notably low in the oil palm plantations. species in agricultural lands (shaded cacao plantations and In addition, the capture rates of two frugivorous bats (B. cornfi elds with other crops) compared to adjacent rainforests. maculata and P. lucasii) clearly differed among the four Phyllostomid bat diversity was also low in coffee plantations vegetation types, and these two species were not recorded in relative to small fragmented forests (Numa et al., 2005). the orchards or the oil palm plantations. These results indicate that these megachiropteran species rarely use agricultural The capture rate of microchiropterans was lower in the oil lands for feeding; thus, the vegetation is not suitable for palm plantations but did not differ among the other three maintaining a diversity of megachiropterans. Moreover, there vegetation types. One possible cause of the strikingly low may be no species unique to agricultural lands, because those capture rate of microchiropterans in the oil palm plantations species for which more than two individuals were captured may have been the low abundance of insects. Although studies were also recorded at least once in the primary or secondary concerning the abundance of prey insects in this vegetation forests (Appendix 1). Megachiropteran diversity in natural type are rare, Chung et al. (2000) reported low beetle forests is thought to be maintained by various factors, abundance and diversity in oil palm plantations compared including the diversity of food sources (Hall et al., 2004; to logged and primary forests in Sabah, Malaysia. Hodgkison et al., 2004a), the availability of roosts for tree- roosting bats (Zubaid, 1993), and the heterogeneity of forest The drastic increase of C. brachyotis I in the oil palm structure (Hall et al., 2004; Hodgkison et al., 2004b). The oil plantations was a particularly intriguing result. The population palm plantations clearly lacked these characteristics. size of C. brachyotis I in the oil palm plantations may indeed be large, considering that its capture rate was extraordinarily Orchards and oil palm plantations may be inadequate habitats high and the plantations cover a vast area (ca. 388,500 ha in for most frugivorous and insectivorous bats; nevertheless, Sarawak as of 2004; Brown & Jacobson, 2005). High capture the capture rates of some species were high in these plant rates and dominance of C. brachyotis in agricultural lands communities. The high capture rate of G. tylopus in the have been reported repeatedly in other regions (Abdullah orchards may have been due to an abundance of bamboo et al., 1997; Hall et al., 2004). For example, Abdullah et al. groves in the vicinity of the census points. This bat roosts (1997) measured a very high capture rate of C. brachyotis in the hollow stems of dead or damaged bamboo (Payne (570 individuals per 10,000 m2h) in Cocos and Musa & Francis, 1998; Yasuma & Andau, 2000). Two species plantations in Indonesia, where C. brachyotis accounted for of nectarivorous bats (E. spelaea and M. minimus) were 93% of total captures. Our recapture data indicated that C. captured at signifi cantly higher rates in orchards than in brachyotis was capable of movement between vegetation the other vegetation types, and E. major was only captured types. In addition, a fecal analysis suggested movement in an orchard. One cause of these high capture rates of between the oil palm plantations and adjacent forests. Ficus

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seeds were found in approximately half of the feces samples (eds.), Proceedings of the International Conference on In-Situ of C. brachyotis I captured in oil palm plantations (n = 11), and Ex-Situ Biodiversity Conservation in the New Millennium. but no Ficus trees were present in this vegetation type (D. Yayasan Sabah, Kota Kinabalu, Malaysia. Pp. 403–408. Fukuda, pers. obs.). Since the census points of the plantations Abdullah, M. T., H. Siswanto, A. Widiyanto, A. Setiabudi & are located within the maximum distance of recapture of the Firmansyah, 1997. Abundance, diversity and distributional species (1600 m, Table 3) from the edge, the bats might have records of bats in disturbed habitats in Kalimantan Barat, eaten Ficus fruits outside of the plantation. We can not deny Indonesia. Sarawak Museum Journal, 11: 75–84. the possibility that the original composition of the forest bat Altringham, J. D., 1996. Bats: Biology and Behaviour. Oxford community would be altered by expanding bat populations University Press. 272 pp. in surrounding oil palm plantations. Such situation has not Appanah, S., 1985. General fl owering in the climax rain forests of been reported for bats in Southeast Asia, but Ickes (2001) Southeast Asia. Journal of Tropical Ecology, 1: 225–240. noted hyper-abundance of wild pigs (Sus scrofa) at a lowland Brown, E. & M. F. Jacobson, 2005. Cruel Oil: How Palm Oil dipterocrp forest that was probably caused by an abundant Harms Health, Rainforest and Wildlife. Center for Science in year-round food supply of oil palm fruits from plantations the Public Interest. Washington, D.C. bordering the forest. Chung, A. Y. C., P. Eggleton, M. R. Speight, P. M. Hammond & V. K. Chey, 2000. The diversity of beetle assemblages in different Cultivated plants in orchards may be abundant and consistent habitat types in Sabah, Malaysia. Bulletin of Entomological food sources, contributing to the maintenance of nectarivorous Research, 90: 475–496. bats populations. This may be especially true in the lowlands Corbet, G. B. & J. E. Hill, 1992. Order Chiroptera. In: Corbet, of Southeast Asia, because lowland dipterocarp forests in G. B. & J. E. Hill (eds.), The Mammals of the Indomalayan the region are known for the supra-annual fl owering and Region. Oxford University Press, Oxford. Pp. 54–161. fruiting phenologies of many tree species (Appanah, 1985; Cox, M. P., C. R. Dickman & J. Hunter, 2003. Effects of rainforest Sakai et al., 1999; Sakai, 2002). Megachiropterans may fragmentation on non-fl ying mammals of the Eastern Dorrigo rely on cultivated plants in agricultural lands during inter- Plateau, Australia. Biological Conservation, 115: 175–189. fl owering periods. Hodgkison et al. (2004a) reported seasonal Erdtman, G., 1966. Pollen and spore morphology. In: Erdtman, fl uctuations in the capture rates of certain megachiropteran G. (ed.), Pollen Morphology and Plant Taxonomy. Hafner species that consumed ‘big bang’ phenology food items in Publishing Company, New York and London. Pp. 11–24. an old-growth lowland mixed dipterocarp forest. However, Francis, C. M., 1990. Trophic structure of bat communities in if the majority of a bat population is drawn to cultivated the understory of lowland dipterocarp rain forest in Malaysia. plants, this food supply in orchards may actually reduce both Journal of Tropical Ecology, 6: 421–431. bat density and ecological services such as pollination and Francis, C. M., 1994. Vertical stratification of fruit bats seed dispersion in surrounding areas. This study suggests (Pteropodidae) in lowland dipterocarp rainforest in Malaysia. signifi cant changes in megachiropteran population densities Journal of Tropical Ecology, 10: 523–530. and feeding habits among natural and cultivated vegetation Fujita, M. S. & M. D. Tuttle, 1991. Flying foxes (Chiroptera: types. Therefore, the effects of human activities on plant Pteropodidae): Threatened animals of key ecological and reproduction in natural forests as a result of changes in economic importance. Conservation Biology, 5: 454–463. pollination and seed dispersion by megachiropterans are Hall, L. S., G. G. Grigg, C. Moritz, B. Ketol, I. Sait, W. Marni & important subjects for future research. M. T. Abdullah, 2004. Biogeography of fruit bats in Southeast Asia. Sarawak Museum Journal, 80: 191–284. Hansen, M. C. & R. S. DeFries, 2004. Detecting long-term global ACKNOWLEDGMENTS forest change using continuous fi elds of tree-cover maps from 8-km Advanced Very High Resolution Radiometer (AVHRR) We thank Kanehiro Kitayama, Hidetoshi Nagamasu and data for the years 1982–99. Ecosystems, 7: 695–716. Mohd Tajuddin Abdullah for reviewing earlier drafts of Heaney, L. R., P. D. Heideman, E. A. Rickart, R. B. 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Hodgkison, R., S. T. Balding, A. Zubaid & T. H. Kunz, 2004b. Racey, P. A. & A. C. Entwistle, 2002. Conservation ecology of Habitat structure, wing morphology, and the vertical stratifi cation bats. In: T. H. Kunz & M. B. Fenton (eds.), Bat Ecology. The of Malaysian fruit bats (Megachiroptera: Pteropodidae). Journal University of Chicago Press, U.S.A. Pp. 680–743 of Tropical Ecology, 20: 667–673. Rainey, W. E., E. D. Pierson, T. Elmqvist & P. A. Cox, 1995. The Ickes, K., 2001. Hyper-abundance of native wild pigs (Sus scrofa) role of fl ying foxes (Pteropodidae) in oceanic island ecosystems in a lowland dipterocarp rain forest of peninsular Malaysia. of the Pacifi c. In: P. A. Racey & S. M. Swift (eds.), Ecology, Biotropica, 33: 682–690. Evolution and Behavior of Bats: the Proceedings of a Symposium IUCN, 2007. IUCN Red List Categories. IUCN Species Survival Held by the Zoological Society of London and the Mammal Commission. http://www.iucnredlist.org/ Society: London, 26Ð27 November 1993. Oxford University Press, Oxford. Pp. 47–62. Jones, G. & J. Rydell, 2002. Attack and defense: interactions between echolocating bats and their insect prey. In: T. H. Rickart, E. A., L. R. Heaney, P. D. Heideman & R. C. B. Utzurrum. Kunz & M. B. Fenton (eds.), Bat Ecology. The University of 1993. The distribution and ecology of mammals on Leyte, Chicago Press. Pp. 301–345. Biliran, and Maripipi islands, Philippines. Fieldiana: Zoology, new series, 72: 1–62. Kinjo, K., A. Nakamoto, M. Izawa, M. Baba, T. Doi & Boeadi, 2003. Preliminary report on the bat fauna in an agricultural fi eld Sakai, S., 2002. General fl owering in lowland mixed dipterocarp in West Java, Indonesia. Bulletin of the Kitakyushu Museum forests of South-east Asia. Biological Journal of Linnean of Natural History and Human History. Series A, Natural Society, 75: 233–247. History, 1: 51–55. Sakai, S., K. Momose, T. Yumoto, T. Nagamitsu & H. Nagamasu, Kinnaird, M. F., E. W. Sanderson, T. G. O’brien, H. T. Wibisono & 1999. Plant reproductive phenology over four years including G. Woolmer, 2003. Deforestation trends in a tropical landscape an episode of general flowering in a lowland dipterocarp and implications for endangered large mammals. Conservation forest, Sarawak, Malaysia. American Journal of Botany, 86: Biology, 17: 245–257. 1414–1436. Kunz, T. H., 1988. Age determination in bats. In: T. H. Kunz (ed.), Shanahan, M. & I. Debski, 2002. Vertebrates of Lambir Hills Ecological and Behavioral Methods for the Study of Bats. National Park, Sarawak, Malaysia. Malaysian Nature Journal, Smithsonian Institution Press, Washington, D.C. Pp. 47–58. 56: 103–118. Marshall, A. G., 1983. Bats, flowers and fruit: evolutionary Simpson, E. H., 1949. Measurement of diversity. Nature 163: relationships in the Old World. Zoological Journal of Linnean 688. Society, 20: 115–135. Sokal, R. R. & F. J. Rohlf, 1973. Introduction to Biostatistics. W. H. Marshall, A. G., 1985. Old World phytophagous bats (Megachiroptera) Freeman and Company, San Francisco and London. 368 pp. and their food plants: a survey. Zoological Journal of Linnean Start, A. N. & A. G. Marshall, 1976. Nectarivorous bats as pollinators Society, 83: 351–369. of trees in western Malaysia. In: J. Burley & B. T. Styles (eds.), Medellin, R. A., M. Equihua & M. A. Amin, 2000. Bat diversity Tropical Trees: Variation, Breeding and Conservation, Linnean and abundance as indicators of disturbance in neotropical Society Symposium Series 2. Linnean Society of London and rainforest. Conservation Biology, 14: 1666–1675. Academic Press, London. Pp. 141–150. Mickleburgh, S. P., A. M. Huston & P. A. Racey, 2002. A review Wright, S. J., 2005. Tropical forests in a changing environment. of the global of bats. Oryx, 36: 18–34. Trends in Ecology and Evolution, 20: 553–560. Nakagawa, M., H. Miguchi & T. Nakashizuka, 2006. The effects of Yasuma, S. & M. Andau, 2000. Order Chiroptera. In: Japan various forest uses on small mammal communities in Sarawak, International Cooperation Agency and Sabah Wildlife Malaysia. Forest Ecology and Management, 231: 55–62. Department (ed.), Mammals of Sabah, Part 2, habitat and ecology. Japan International Cooperation Agency and Sabah Naughton-Treves, L., J. L. Mena, A. Treves, N. Alvarez & V. C. Wildlife Department. Pp. 62–156. Radeloff, 2003. Wildlife survival beyond park boundaries: the impact of slash-and-burn agriculture and hunting on mammals Zhao S., C. Peng, C., H. Jiang, D. Tian, X. Lei & X. Zhou, 2006. in Tambopata, Peru. Conservation Biology, 17: 1106–1117. Land use change in Asia and the ecological consequences. Ecological Research, 21: 890–896. Numa, C., J. R. Verdú & P. Sánchez-Palomino, 2005. Phyllostomid bat diversity in a variegated coffee landscape. Biological Zubaid, A. 1993. A comparison of the bat fauna between a primary Conservation, 122: 151–158. and fragmented secondary forest in Peninsular Malaysia. Mammalia, 57: 201–206. Payne, J. & C. M. Francis, 1998. A Field Guide to The Mammals of Borneo. The Sabah Society, Kota Kinabalu, Sabah, Malaysia. 332 pp. Primack, R. & R. Corlett, 2005. Tropical Rain Forests: An Ecological and Biogeographical Comparison. Blackwell Publishing, Oxford. 336 pp.

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18_Fukuda_Pg 213-221.indd 220 2/19/09 12:07:40 PM THE RAFFLES BULLETIN OF ZOOLOGY 2009 1 = c 186 c 0 0 2 4 0 0 30 21 4 39 0 2 3 28 0 2 0 0 25 2 0 0 2 0 3 26 0 0 2 2 0 0 0 4 0 4 8 4 17 41 C. brachyotis ; 68 c 0 3 0 9 2 4 0 0 0 0 0 0 1 2 0 2 0 20 29 20 captured No. 69 c 1 1 3 3 1 9 0 0 2 1 0 3 1 0 1 0 0 4 0 10 43 11 7 1 51 11 7 1 c 1 0 7 5 6 0 2 1 2 0 5 6 2 0 2 4 2 18 23 32 135 98 168 94 495 Primary Secondary Orchards Oil palm Total palm Oil Primary Secondary Orchards

b

Fruit Fruit Insects Insects Insects Insects Insects Insects Insects Insects Insects Insects Fruit, nectar Nectar, pollen Nectar, pollen Nectar, pollen Fruit, nectar, pollen Fruit, nectar, pollen Diet

a

NA NA 9 ± 1 6 ± 0 9 ± 1 4 ± 0 6 ± 0 4 ± 0 8 ± 1 9 ± 0 89 ± 8.8 15 ± 1.5 35 ± 5.7 48 ± 7.4 76 ± 1.0 27 ± 3.8 Fruit, nectar, pollen 11 38 ± 5.0 14 ± 2.2 19 ± 0.5 Fruit, nectar, pollen 33 ± 4.0 Fruit, nectar, pollen 14 ± 2.0 19 ± 3.6 Insects, spiders, small vertebrates 4 = based on Yasuma & Andau (2000), although the literature does not distinguish between two forms of b (mean, ± SD) a

NA 76 ± 0 86.16 ± 2.67 47.45 ± 1.16 39.98 ± 1.45 59.63 ± 3.80 63.61 ± 2.58 84.18 ± 0.13 57.80 ± 1.37 41.63 ± 1.75 49.19 ± 1.01 28.89 ± 0.90 29.91 ± 0.09 42.35 ± 0.34 49.98 ± 1.51 54.32 ± 0.27 (mean, ± SD) Forearm length (mm) Body weight (g) able) NA = data of juveniles were excluded; a c name Scientifi Rousettus spinalatus Microhiroptera HIPPOSIDERIDAE cervinus Hipposideros Macroglossus minimus Macroglossus Penthetor lucasii Rousettus amplexicaudatus Eonycteris spelaea Dyacopterus spadiceus 77.5 1 Eonycteris major 74 0 0 Fruit 1 0 C. brachyotis (unidentifi Chironax melanocephala Chironax 45.50 ± 1.20 dominant species in the vegetation type. Hipposideros diadema Hipposideros C. brachyotis II dyacorum Hipposideros 39.55 ± 0.05 Appendix 1. Species list of bats captured in and around Lambir Hills National Park during this study.

Megachiroptera PTEROPODIDAE Balionycteris maculata Cynopterus brachyotis I 62.36 ± 1.87 forests forests plantations 48.15 NA Insects 0 1 0 0 1 0 0 galeritus Hipposideros 1 ridleyi 48.15 Hipposideros NA Insects 0 VESPERTILIONIDAE tylopus Glischropus blanfordi Hesperoptenus 25.05 ± 0.05 NA = data not available; 31.09 4 Insects 0 1 0 0 0 1 1 0 0 Insects 0 4 Kerivoula pellucida 0 1 1 Murina suila 31.09 Philetor brachypterus 32.70 1 0 0 tenuis 29.20 Pipistrellus Insects 5 0 11 pachypus NA 0 Tylonycteris 0 1 Insects 1 1 26.80 robustula Tylonycteris NA Insects 0 RHINOLOPHIDAE 1 5 Rhinolophus borneensis 0 0 Insects 0 42.12 ± 0.20 Rhinolophus sedulus Rhinolophus trifoliatus MEGADERMATIDAE Megaderma spasma Total

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BOOK REVIEW

Private Lives: An The book is divided into chapters, but these are by no means Exposé of Singapore’s systematic, although there is a certain order but this is more Mangroves. Peter K. L. ecological; the chapters have unusual titles that make the Ng, Wang Luan Keng, reader curious to discover what is behind, and he will not Kelvin K. P.Lim, 2008. be disappointed. The Raffl es Museum of Biodiversity Research, The book is not written by one author. Many contributors, Singapore, 249 pages. each with his own speciality, share their knowledge with the ISBN: 978-981-05-8717- reader, and that makes the book still more valuable. 8. It starts with the mangrove plants and all aspects of their This book is the second biology, follow chapters on the mangrove lobster community volume in a new series and the very rich life in the canopy of the mangrove forest. called “Private Lives”, Another chapter is on the intertidal zone in front of the edited by the Raffles mangrove trees, and a special chapter treats the mud-dwelling Museum of Biodiversity animals, mostly molluscs and crustaceans, and shows that Research. In this series mud is not at all a desert but a rich environment. The next the different biotopes of the coastline of Singapore Island chapter contains the fi sh fauna, and the migrating birds and adjacent regions are treated simply and explicatively, which make their stop-overs in the mangrove biotopes to destined for the grand public but also for scientists. The take profi t from the rich offer of food are not forgotten. subject of the first volume were the sandy and rocky Do you know what a “top dog” is (by the way we are also seashores, and this volume is about the mangroves, a much one!)? This is also the title of another chapter in which more specialized biotope. The books of this series do not the feeding strategies of the big predators in the mangrove give scholarly information on fauna and fl ora in form of biotopes are explained. The infl uence of men to the mangrove “dry” descriptions with the corresponding pictures but take biotopes in Singapore is illustrated in the last part of the all kind of interesting facts “from the life” (as the title book, among others explaining what people take from the promises) and tell stories. And about mangroves there are mangroves for food, health etc., the last news of scientifi c many stories to tell. However, the mangrove biotopes are research on the mangrove biotopes with interesting new very diffi cult in access; the bottom is often soft mud, the species recently described, and a vivid plea for protection roots block the way and serve as barricades, the air is very of the last remaining mangrove biotopes, which is most hot, damp and humid, and the mosquitoes are stinging. So, important because 95% of the original mangrove covering the life out there cannot be observed “like that” and is really of Singapore Island are now gone. “private”. The merit of this book is to bring that mangrove life to light and to tell us about it in an easily readable and There are many strange or curious things in the mangroves, understandable text, with numerous excellent photos of and the book is full of small histories and surprising facts. plants, live animals and biotope situations, often close-up Can you imagine a frog that lives in salty sea water? Or a fi sh photos never seen before in that manner. that walks on land and watches at you with twinkling eyes? Or a bivalve that climbs on trees up to the leaves? Or that Singapore is situated within the “golden triangle of there are lobsters that even build some sort of “skyscraper”- biodiversity” in the centre of the Indo-Pacifi c realm, together like condominiums? And a fi sh that uses its mouth as a water with Indonesia, Malaysia and the Philippines, that means pistol to catch prey? All that exists in the mangroves, and the species diversity is extremely high, and the mangroves it is nicely explained in the book, accompanied by superb make no exception, they are among the most species rich photos. There are many other strange facts accumulated, mangrove communities on the planet, in number of mangrove e.g. on the propagation of the mangrove trees, on the birds trees, associated plants and animal species. All aspects of living around and in them, on the molluscs living on and mangrove life are covered in the book, from tiny algae to underneath them, on the worms, the reptiles, the fi shes and the huge mangrove trees and from small invertebrates to especially the crustaceans which are a speciality of many birds and mammals at the end of the food chain. of my colleagues in the Raffl es Museum.

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The book is said to have a “Singapore-centric approach” as I read it, and I recommend it warmly. Go into a bookshop is expressed in the Prologue. But that is an understatement, and have a look. And enjoy reading! the book has no restriction at all and with the many general explanations, it is extremely interesting and useful for all naturalists, especially those interested in tropical biotopes, animals and plants. So the book should be much more widely diffused as initially thought and fi nd readers not only in Rudo von Cosel Singapore and the neighbouring countries. Muséum national d’Histoire naturelle Département de Systématique et Evolution This is one of the best popular books on mangroves I have Taxonomie et Collections ever seen. I will not say more now. I was fascinated when Case Postale 51, F-75231 Paris cedex, France.

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BOOK REVIEW

The Ecology of a It covers a detailed history of the origins of the site for Tropical Rainforest KMNP. The main fi eld station is located near Long Alango in Kayan Mentarang village at Lalut Birai. There are chapters devoted to the National Park in the physical environment and forest ecology. Some interesting Heart of Borneo. excerpts are included and interspersed within the volume, e.g. Ekologi Hutan Hujan ancient sarcophaguses built by the native people, Bornean Tropis Taman Nasional Red Carnivore. Several interesting groups of plants are Kayan Mentarang di discussed, e.g. Zingiberaceae, Raffl esiaceae, Amorphallus, Jantung Kalimantan. bamboos, palms and orchids. The mammals and birds sections By Wulffraat, S., P. are covered in greater detail, with almost every major group Tatengkeng & A. Salo, or species illustrated. The other faunal groups are covered 2006. WWF – Indonesia, in less detail. xxii+250 pages. ISBN: 979-99919-2-7. The appendices consisted of checklists of mammals, birds, amphibians, reptiles, fi shes and crustaceans. The taxonomy This is an A4-sized used is not the latest, but nonetheless illustrates the rich bilingual (English/Bahasa Indonesia) volume covering the biodiversity that abounds in KMNP. ecological aspects of one of the largest protected areas in Borneo. It is the Kayan Mentarang National Park (KMNP), This publication defi nitely provides interested naturalists with 1,360,500 hectares, situated in the province of East and scientists an interesting read into the biodiversity of Kalimantan, along the borders of Malinau District and the Borneo. Malaysian State of Sarawak.

This book consists of 19 Chapters and is richly illustrated: Introduction; The research area; Landscape of the upper Enggeng; Land use history; Climate; Phenology; Growth, Tan Heok Hui recruitment and mortality; Regeneration; Forest productivity; Department of Biological Sciences Leaf litter decomposition; Reforestation; Specific plant National University of Singapore groups; Mammals; Birds; Amphibians and reptiles; Fishes and Kent Ridge 119260 crustaceans; Notes on insects; References; and Appendix. Republic of Singapore

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BOOK REVIEW

A guide to the Specimens listed and photographed in this guide book were Amphibians and Reptiles obtained through several methods – by visual sightings, hand of Tasek Merimbun catching, pitfall and sticky traps. Specimens obtained were Heritage Park Brunei photographed and specimenised. These voucher specimens Darussalam. Editors: were deposited in the Brunei Museum. I. Das, Samhan bin Nyawa and J. K. Charles, A total of 23 amphibian and 31 reptile (2 crocodilians, 3 2008. Brunei Museums turtles, 15 lizards and 11 snakes) species were recorded. Department, xvii+92 Each species is illustrated with a colour photograph of pages. ISBN: 99917-30- its habitus, material examined and several diagnostic 32-X. characters. It ends off with a checklist of the herpetofauna, dated 1 December 2007. Two lists of relevant literature and This is a compact 23 x 16.5 websites pertaining to Brunei Herpetology are also provided. cm volume covering the A section on snake-bite and its management is appended. amphibians and reptiles Pictures of the activities of the workshop are appended in of the Tasek Merimbun the last few pages. Heritage Park (TMHP) situated in Tutong District of Brunei Darussalam. This Although this volume is richly illustrated, the colour publication stems directly from a workshop conducted at separation was not ideal, and this mars this otherwise TMHP in December 2006, organized by the Brunei Museums compact guide book. Department – “Inventory and Training Workshop on Herpetological Research and Management Techniques”. This is definitely a must-have for nature lovers and herpetologists. The purpose of this guide book is for visitors to appreciate the amphibian and reptile fauna of TMHP and it is printed Tan Heok Hui in a user friendly format. Although, the basis of this Department of Biological Sciences publication is from the brief workshop, additional records National University of Singapore should be forwarded to the editors for future incorporation Kent Ridge 119260 into the checklist. Republic of Singapore

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BOOK REVIEW

covering a region with multi-authorship). It basically covers the country reports on the current state of their coral reefs. It had been a monumental effort for such a publication to come into fruition.

From this report, the world has effectively lost 19% of the original area of coral reefs; with a further 15% of projected loss in the next 10–20 years. However, 46% of the world’s reefs are regarded as healthy and not under immediate threat of destruction. These projected fi gures are not without caveats, as explained in the text.

From the various regional reports, the Southeast Asia region is here highlighted as it contains the largest area of coral reefs (34% of the world’s total) and is within the scope of Status of Coral Reefs of the World: 2008. Edited by the Raffl es Bulletin of Zoology. Clive Wilkinson. Global Coral Reef Monitoring Network and Reef and Rainforest Research Centre, Townsville, Amongst the noteworthy is that the conditions of coral Australia, vi + 296 pages (www.gcrmn.org). ISSN 1447-6185. reefs had improved in Thailand, Philippines, Vietnam and (Accompanying CD with much more extended material) Singapore, but declined in Indonesia and Malaysia. Timor- Leste is included for the fi rst time as the tenth Southeast Asian This is a compact soft-cover volume (18 × 25 cm) with a country. The 2004 Indian Ocean tsunami caused localized colourful front and back cover. The front cover depicts three coral reef damage in Indonesia, Thailand and Malaysia. The young boys in a village market posing with a large specimen region’s fi sh stock had been overfi shed, and more than 50% of Napolean Wrasse (Cheilinus undulatus), a marine coral of the region’s mangroves have been lost. reef fi sh listed as endangered on Appendix II of CITES. The back cover depicts a healthy coral reef scene, but includes The accompanying CD contains more material in PDF in the top left corner a submerged polar bear. The polar format pertaining to the coral reef status. Additional material bear had been added in to illustrate the deleterious impacts on Middle East, Indian Ocean, Asia, Pacifi c, Caribbean; of global warming (more details on page 13). Both covers and including a 13 page list of suggested reading material are also displayed with the logos of 20 agencies directly encompassing numerous web resources. supporting this joint publication effort. All in all, it makes both gloomy and heartening reading, This current volume is the fifth global report since the and it is good to know that there are many volunteers and inception of the Global Coral Reef Monitoring Network agencies out there monitoring and documenting the reefs and (GCRMN) in 1996; and it was recently launched in February making the effort to make a change. A defi nite must-have (2009) in Singapore. All the previous reports (in 1998, 2000, for marine biologists and conservationists. 2002 and 2004) had aimed to present the current status of the world’s coral reefs, the threats to the reefs, and the initiatives undertaken to arrest the reef decline. The current report does the same and had called upon the voluntary contributions of 372 experts from 96 countries (which is a vast increment of Tan Heok Hui contributions from previous reports). Department of Biological Sciences National University of Singapore The report covers 95 countries, states and territories; which Kent Ridge 119260 had been divided into the various 20 chapters (each chapter Republic of Singapore

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BOOK REVIEW

Mangrove the mangroves on Bohol is covered with more detail. Next, Environments and information of the Arbatan River, where the main basis of Molluscs: Abatan this study is formed, is well presented to the reader with neat River, Bohol and illustrations and photographs of various vegetation types. Panglao Islands, Defi nitions of the mangrove environment and the zonation Central Philippines. of vegetation are also clearly explained. Pierre Lozouet & Jean- Claude Plaziat. 2008. Part II – Molluscs in the Mangal, is possibly the main part of 160 pp. Conchbooks, the text that will interest most potential readers of this book. Hackenheim, Germany. The general occurrence of mollusc distribution is discussed ISBN 978-3-939767-16- in detail, and includes useful information of molluscs 9. in the mangal environment. The occurrence of different species relative to the mangrove ecosystem is discussed in The Indo-West Pacific detail, and includes defi nitions as to what constitutes real region is by far the richest mangal malacofauna in the strict sense, something often in terms of mollusc species misinterpreted by various authors. Aided with the information diversity, but despite more gleaned from Part I, the reader can easily relate to most than 200 years of malacology, the identifi cation of mangal of the discussions such as, for example, the zonation of molluscs, and knowledge of their habits in the mangrove molluscs. Examples of mollusc species are also given under ecosystem, remains poor. Containing a small part of the all subsections allowing the reader to easily form an idea of results of the Panglao Marine Biodiversity Project, a large the kind of species to expect and their respective distribution scale comprehensive survey undertaken by a team of research in the mangrove habitat. Accompanying photographs serve scientists from many ASEAN countries, Europe and the well to break the monotony of the text for the casual reader United States, this book covers the mangrove environments in with an attention defi cit, and those of molluscs in situ and relation to the molluscs. Molluscs inhabiting the mangroves their living environs are particularly of interest, allowing have received considerably less attention compared to their a closer look at the fascinating animals for the armchair counterparts inhabiting the coral reefs, and the present book naturalist. This section is concluded with some notes on fi lls an important niche that is seldom visited in detail by the importance of the mangrove ecosystem in the economic other authors. Thumbing through the plates, it becomes sense, and includes, besides the molluscs, the exploitation of apparent that the main focus of the book is on showcasing other material, and other generally interesting information the mangal malacofauna diversity. Although the contents for ecologists and the like. of the present book are based on material and studies of only a small area in the Central Philippines, the information A list of the included mollusc species arranged in systematic within is nevertheless relevant from a regional perspective order is provided for in Part III – Taxonomic Inventory as similarities in biotope, and species composition are often of Mangal Molluscs and other Mangrove-Associated shared throughout the region. The present book satisfi es the Environments, and is clearly meant complement the colour most basic need of species identifi cation, which is often a plates in Part V, and for the serious student, references are frustrating affair faced by students with an interest in the provided in Part IV. All species photographed are represented mangrove ecosystem, as well as containing useful information by beautiful colour photographs, and short notes such as on the mangrove ecosystem. the locations and habitat where each species were collected are supplied as accompanying text. However, diagnostic The book is organised into several parts, starting with an descriptions for the photographed species are not provided introduction of the characteristics of the Indo-West Pacifi c and the means of species identifi cation provided for by this biogeographical region, followed by a mention of the book lies solely in the plates. Depending on how keen the methods employed for the survey. Part I – The Mangal individual reader’s eyes are, differences between similar Ecosystem, contains interesting general information on the looking species may not be easily discerned. This is possibly mangrove ecosystem. There is cursory mention of the state the only blemish in an otherwise fl awless book. Nevertheless, of the mangroves in the Philippines, but the situation of as most malacological students are accustomed to studying

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plates, this does not in any way detract from the usefulness that molluscs are not well represented in the mangroves. My of this book, and the importance of the role that it fulfi ls to attention is also unwittingly drawn to the fact that a number showcase the mangal malacofauna. of species rarely or never before depicted in popular books are included. These inclusions make the book even more In recent years, there is an increasing shift in attitudes attractive for the professional and amateur malacologist alike, towards the mangrove ecosystem. The importance of the and this beautifully illustrated book is a defi nite must-have role of mangrove ecosystem in the scheme of things is for the library of any self respecting malacologist. increasingly being appreciated beyond the ugly reputation as dirty muddy mosquito infested swamps. The authors are commended for having the foresight to demonstrate Tan Siong Kiat the actual diversity of the malacofauna in a specialised but Raffl es Museum of Biodiversity Research often ill-defi ned ecosystem, and students of the mangrove Department of Biological Sciences ecosystem will appreciate this book as a useful reference National University of Singapore and a means of identifying their fi nds. The impressive array Block S6, #03-01, Science Drive 2 of species illustrated, apart from pleasantly stimulating the Singapore 117546 eye and mind, will no doubt change many a reader’s mind Republic of Singapore

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OPINION

BOTULA KLEEMANNI VALENTICH-SCOTT, 2008, ACTUALLY A MEMBER OF LITHOPHAGA (LEIOSOLENUS)

Karl Kleemann Department of Palaeontology, University of Vienna, Centre for Earth Sciences, Althanstrasse 14, A1090 Vienna, Austria Email: [email protected]

INTRODUCTION the shell rim does not apply to Botula but to Lithophaga (Leiosolenus). Botula kleemanni Valentich-Scott, 2008 (Valentich-Scott & Tongkerd, 2008) does not show the generic characters of Regarding the original description, the following is stressed: Botula, particularly a kidney-shaped and smooth shell with Anterior is semicircular not “truncate”. “Beaks very infl ated”. prominent terminal beaks, as given by Soot-Ryen (1955, Those of the holotype may be more elevated from the 1969) and Kleemann (2007). ellipsoidal outline than in most other Leiosolenus species. But they need a terminal position and somewhat coiled form to support a determination as Botula. Both these features HISTORICAL BACKGROUND are lacking in the holotype of kleemanni (Valentich-Scott & Tongkerd, 2008: 193, Fig. 2G–H). The beaks are clearly sub- Mörch (1853: 55) introduced Botula as subgenus of terminal, the ventral margin is straight to slightly convex and Lithophaga (for details see Kleemann, 2007: 4). Although not concave as in Botula. The posterior shell end is somewhat Jousseaume (1888: 216) had placed Botula in generic rank, attenuated and distinctly less high than the anterior, while Dall (1898) treated Botula as a subgenus of Modiolus in Botula both ends have about the same height. and designated Modiola cinnamomea Lamarck, 1819, as the type species of Botula (Kleemann, 2007: 4). Several “Holotype (specimen described above)” (Valentich-Scott & authors, e.g. Soot-Ryen (1969), Savazzi (1982), Arnaud & Tongkerd, 2008: 194) must read fi gured above. The legend Thomassin (1990) and Darga (1992), considered Botula a cannot be regarded as a description. It reads “G–H, holotype, member of Modiolinae instead of Lithophaginae, to which it SBMNH 83123, length 12.35 mm: G, external left valve; H, belongs (Wilson & Tait, 1984; Coan et al., 2000; Kleemann, internal right valve”. In the description, the shell length is 2007). given as “12.21 mm, plus 1.22 mm of periostracum extending past shell”. Thus, there is a difference of 1.08 mm between the initially given (total) length in the legend and the sum TAXONOMY of shell length plus extension, amounting 13.43 mm.

Lithophaga (Leiosolenus) kleemanni The general similarity of the holotype of kleemanni with (Valentich-Scott, 2008) other Leiosolenus species is obvious because of the shell form and its postero-dorsal cover wich is usually mainly Botula kleemanni Valentich-Scott, 2008: 194, Fig. 2G–H, ? Fig. calcareous and of a species-specifi c pattern (Kleemann, 1977, 2F, non Fig. 2I. 1980, 1982, 1984, 1986, 1990, 1994, 2002, 2008; Kleemann & Hoeksema, 2002). “Shell ellipsoid-elongate; beaks subterminal; periostracum usually dark brown, with heavy fi brous mat posterodorsally, The similarity of kleemanni with the Caribbean Lithophaga extending past the posterior margin (Fig. 2G)” (Valentich- (Leiosolenus) appendiculata (Philippi, 1846; synonyms Scott & Tongkerd, 2008: 194). This shell confi guration, the Lithodomus bisulcatus Orbigny, 1853 and bi-excavatus Reeve, position of the beaks, the shell cover and its extension beyond 1857) is particularly striking (Kleemann, submitted).

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“F, in situ in living coral” (Valentich-Scott & Tongkerd Darga, R., 1992. Geologie, Paläontologie und Palökologie der 2008: Fig. 2), does not show a live coral. No remains of SO-bayerischen unterpriabonen (O-Eozän) Riffkalkvorkommen coral tissue can be recognized on the skeleton surface, having des Eisenrichtersteins bei Hallthurn (N Kalkalpen) und the same grey colour as the skeleton. A freshly split-open des Kirchbergs bei Neubeuern (Helvetikum). Münchner Geowissenschaftliche Abhandlungen, A23: 1–66. live coral would reveal coral tissue on the coral surface, distinctly darker in colour compared with the whitish skeletal Gmelin, J. F., 1791. Vermes. In: Caroli a Linné Systema naturae interior below the uppermost live part. The sample appears per regna tria naturae, Ed. 13, Lipsiae, 1(6): 3021–3910. dry when the picture was taken and the coral was likely Jousseaume, F., 1888. Description des Mollusques recueillis par M. dead when collected. Covered by detritus (and a fi brous Le Dr. Faurot dans la mer Rouge et le golfe d’Aden. Mémoires mat?), the exposed bivalve is posteriorly fl ush with the de la Société Zoologique de France, 1: 165–223. coral surface. In contrast, Botula generally penetrates deep Kleemann, K. H., 1977. A new species of Lithophaga (Bivalvia) into the substrate, as can be seen in the adjacent Fig. 2E from the Great Barrier Reef, Australia. Veliger, 20: 151–154. (Valentich-Scott & Tongkerd, 2008). Kleemann, K. H., 1980. Boring bivalves and their host corals from the Great Barrier Reef. Journal of Molluscan Studies, Odd records of Botula in live coral, e.g. Savazzi (1982), 46:13–54. have to be considered as occurrences in dead parts of live Kleemann, K. H., 1982. Ätzmuscheln im Ghetto? Lithophaga coral (Kleemann, 1996). Valentich-Scott & Tongkerd (2008) (Bivalvia) aus dem Leithakalk (Mittel-Miozän: Badenian) claim that both, Botula cinnamomea and kleemanni, inhabit von Müllendorf im Wiener Becken, Österreich. Beiträge zur live coral, the latter exclusively. Paläontologie von Österreich, 9: 211–231. Kleemann, K. H., 1984. Lithophaga (Bivalvia) from dead coral “Fig. 2I, facultative of living animal” (Valentich-Scott from the Great Barrier Reef, Australia. Journal of Molluscan & Tongkerd, 2008), clearly depicts the extended inhalant Studies, 50: 192-230. siphon and the shell ends of a specimen lacking any posterior Kleemann, K., 1986. Lithophagines (Bivalvia) from the Caribbean shell extensions or a fi brous cover as in Fig. 2G. Only some and the Eastern Pacifi c. Proceedings of the 8th International detrital sediment adheres on the shells. Therefore, I do not Malacological Congress, Budapest 1983, 1986: 113–118. consider it as a member of Lithophaga kleemanni. Kleemann, K.,1990. Boring and growth of chemically boring bivalves from the Caribbean, Eastern Pacifi c and Australia’s Valentich-Scott & Tongkerd (2008: 194) “Comparisons. Great Barrier Reef. Senckenbergiana maritima, 21:101–154. – Easily separated from Botula cinnamomea and other Kleemann, K., 1996. Biocorrosion by bivalves. P.S.Z.N. I: Marine members of the genus, by the heavy periostracal mat found Ecology, 17: 145–158. along the posterodorsal margin of the shell.” How true, Kleemann, K., 2007. Catalogue and Bibliography of Recent and as kleemanni is a member of Lithophaga (Leiosolenus), it fossil Botula (Bivalvia: Mytilidae). Zootaxa, 1508: 1–48. should be compared with those species. Kleemann, K., 2008. Lithophaga (Leiosolenus) purpurea (Bivalvia: Mytilidae): one species becomes three. Club Conchylia Informationen, 39(3/4): 32–45 ACKNOWLEDGEMENTS Kleemann, K., (in press). Application to conserve Modiola cinnamomea Lamarck, 1819 as the type species of mytilid bivalve For the honour and undeserved respect I received from Paul Botula Mörch, 1853. Bulletin of Zoological Nomenclature. Valentich-Scott I feel sorry to correct him. Kleemann, K., (submitted). Lithodomus bisulcata Orbigny, 1853, a junior synonym of Modiola appendiculata Philippi, 1846. Club Conchylia Informationen, 40(1/2). LITERATURE CITED Lamarck, J. B. P. A. de M. de, 1819. Histoire naturelle des animaux sans vertebres, Verdière, Paris, 6: 343 pp. Arnaud, P. M. & B. A. Thomassin, 1990. Habits and morphological adaptations of mytilids (Mollusca: Bivalvia) from coastal Mörch, O. A. L., 1852–53. Catalogus Conchyliorum quae reliquit and reefal environments in southwest Malagasia (Indian D. Alphonso D’Aguirra et Gadea Comes de Yoldi, Copenhagen, Ocean). In: Morton, B. (ed.) The Bivalvia – Proceedings of Fasc. 2: iv + 74 pp. a Memorial Symposium in Honour of Sir Charles Maurice Orbigny, A. d’, 1841-53. Mollusques. In: Sagra, Histoire physique, Yonge, Edinburgh, 1986, Hong Kong University Press, Hong politique et naturelle de l’île de Cuba, 2: 1–380, Atlas. 28 + Kong, 333–344 pp. 1 Colour Pls. Coan, E. V., Scott, P. V. & F. R. Bernard, 2000. Bivalve Seashells Philippi, R. A., 1846-47. Modiola. In: Abbildungen und of Western North America Marine Bivalve Mollusks from Beschreibungen neuer oder wenig gekannter Conchylien, 2(5): Arctic Alaska to Baja California. Santa Barbara Museum of 1/147–4/150, Pl. 1 (1846); 3(1): 5/18–8/22, Pl. 2 (1847). Natural History Monographs Number 2, Studies in Biodiversity Reeve, L. A., 1857–58. Monograph of the genus Lithodomus. Number 2: viii + 764. In: Conchologia Iconica, 10: 5 Pls. (+ descriptions). London, Dall, W. H., 1889. A preliminary catalogue of the shell-bearing Reeve Brothers. [Pl. 1, Mar. 1858, Pl. 2–4 Oct.1857, Pl. 5, marine mollusks and brachiopods of the south-eastern coast Jan.1858]. of the United States, with illustrations of many of the species. Savazzi, E., 1982. Commensalism between a boring mytilid bivalve Bulletin of the United States National Museum, 37: 7–221, and a soft bottom coral in the Upper Eocene of northern Italy. 74 Pls. Paläontologische Zeitschrift, 56(3/4): 165–175.

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Soot-Ryen, T., 1955. A report on the family Mytilidae (Pelecypoda). Valentich-Scott, P. & P. Tongkerd, 2008. Coral-boring bivalve Allan Hancock Pacifi c Expeditions, 20: 1–175. molluscs of southeastern Thailand, with the description of a Soot-Ryen, T. 1969. Superfamily Mytilacea Rafi nesquae, 1815. new species. Raffl es Bulletin of Zoology, Supplement No. 18: In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology. 191–216. Part N1, Mollusca 6, Bivalvia, The Geological Society of Wilson, B. R. & R. Tait, 1984. Systematics, anatomy and boring America and University of Kansas Press, Lawrence, Kansas, mechanisms of the rock-boring mytilid bivalve Botula. N271–N280. Proceedings of the Royal Society of Victoria, 96: 113–125.

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THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 237 Date of Publication: 28 Feb.2009 © National University of Singapore

OPINION

RESPONSE TO “BOTULA KLEEMANNI VALENTICH-SCOTT, 2008, ACTUALLY A MEMBER OF LITHOPHAGA (LEIOSOLENUS)”

Paul Valentich-Scott Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105, USA Email: [email protected]

I appreciate having an opportunity to respond to Dr. In addition, Dr. Kleemann suggests that Fig. 2I in our Kleemann’s hypothesis that Botula kleemanni Valentich- publication (Valentich-Scott & Tongkerd, 2008) is not an Scott, 2008, might be better included in another genus. While image of Botula kleemanni. This image is of a paratype his suggestions have merit, I fi nd many of his conclusions specimen of this species (SBMNH 80178). It has very little to be lacking, and will outline the discrepancies below. of the periostracal matt, but is defi nitely a B. kleemanni.

Dr. Kleemann based his fi ndings on two photographs of the So, in fact, this species represents characters of both Botula holotype. These images include an exterior lateral view and and Leiosolenus. It was well outside the scope of our original an internal lateral view. Sadly, he has not examined any publication to erect a new genus, but perhaps this will be of the type material of the species in question. Additional accomplished by subsequent authors. paratypes, showing variation within the species, also were not examined. Had Dr. Kleeman examined these specimens he would have noticed the following characters. LITERATURE CITED

1. There is signifi cant variation in the placement of the Kleemann, K., 2009. Botula kleemanni Valentich-Scott, 2008, beaks, from completely terminal as in Botula in the strict actually a member of Lithophaga (Leiosolenus). Raffl es Bulletin sense, to subterminal (e.g. the holotype specimen). of Zoology, 57(1): 221–223. 2. The beaks are exceedingly infl ated in many specimens, Valentich-Scott, P. & P. Tongkerd, 2008. Coral-boring bivalve as in Botula. molluscs of southeastern Thailand, with the description of a 3. The ventral margin is incurved in some specimens, and new species. Raffl es Bulletin of Zoology, Supplement No. 18: straight in others. 191–216. 4. The periostracal matt on the posterior end is not like the typical calcifi ed concretions of Leiosolenus. Botula kleemanni has a softer, more fi brous matt, frequently with sand inclusions.

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