THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 183-187 Date of Publication: 31 Dec.2005 © National University of Singapore
FIRST RECORD OF THE GENUS AMPHIENTULUS TUXEN, 1981 (PROTURA: BERBERENTULIDAE) FROM CHINA, WITH DESCRIPTION OF A NEW SPECIES
Yan Xiong School of Life Science, East China Normal University, Shanghai 200062, China Department of Biology, Hainan Normal University, Haikou 571158, China
Rongdong Xie and Wenying Yin Institute of Entomology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Shanghai 200032, China Email: [email protected] (WY)
ABSTRACT. Ð Amphientulus sinensis, a new species from Guangdong and Hainan, south China is described. Comparison between the new species and Amphientulus ruseki (Nosek, 1978) from Madagascar is provided.
KEY WORDS. Ð Protura , Amphientulus, first record, new species, China.
INTRODUCTION Paratypes Ð 6 females, 3 males, 2 maturi juniores (mounted) (SIE) (P-GD-021209) and (P-GD-030225); 1 female, 1 male (ZRC), The genus Amphientulus was established by Tuxen (1981) same locality as holotype, coll. Mr. Qiang Liu, 25 Feb.2003, 9 for some species removed from Berberentulus. It belongs to Dec.2002; 1 female (SIE) (P-HN-021217), Mt. Jianfengling the genera of the family Berberentulidae with reduced striate (18°23'N 108°36'E), 340 m alt., Hainan Province, China, coll. Mr. band on abdominal segment VIII, with the subapical and the Qiang Liu & Mrs. Yan Xiong, 17 Dec.2002. delicate median apical seta on abdominal legs II-III, and with simple canal of maxillary gland. All species in genus Description. Ð Adult: Amphientulus are characterized by the presence of foretarsal sensillia b’ (missing in Berberentulus). They often have the Body length 883.4µm (830-908µm, n=10), width 165.2µm hook-shaped design on the “striate band” on abdominal (152-176µm, n=10). segment VIII and the lateral apical seta instead of the median one on abdominal legs II-III. There are six Australian species, Head Ð Elliptic, length 94.5µm (88-98µm, n=10), width one Korean and one Madagascarian species in genus 66.9µm (59-72µm n=10). Additional setae absent, Amphientulus. The present paper reports a new species from postpseudocular seta present. Dorsal side of head with 2+2 China. The specimens have been collected in Guangdong and short and thin sensory setae, ventral side with no differentiated Hainan through the soil faunistic surveys in China from 2001 setae. Sensilla of maxillary palpus thin, subequal. Labial to 2003. Type specimens are deposited in the collection of palpus reduced to three setae and a claviform sensillum. Institute of Entomology, Shanghai Institutes for Biological Pseudoculus almost circular, slightly broader than long, 7-8 Sciences (SIE), Chinese Academy of Sciences, and the X 8-9 m. PR=11.8. Canal of maxillary gland straight and Zoological Reference Collection (ZRC) of the Raffles µ Museum of Biodiversity Research, National University of smooth, with heart-shaped calyx, short terminal filament and Singapore. slightly developed terminal widening. Length 15.6µm (14- 17µm, n=10), as long as half of the branch of fulcrum (Figs. 1, 2, 5, 7, 8). TAXONOMY Thorax Ð Chaetotaxy on thoracic tergites II-III with two Amphientulus sinensis, new species anterior setae (A2 and A4), seta P1a, P2a and P5 very short, (Figs. 1-17) less than one-tenth of P1 in length. P2a nearer to P3 than to P2. Seta P4a on mesonotum, P3a on metanotum.Thoracic Material examined. Ð Holotype-female (mounted) (SIE) (P-GD- sterna without pores (Figs. 3, 4). 020921), from Mt. Dinghu (23°9'N 112°30'E), 290 m alt., Guangdong Province, China, coll. Mr. Qiang Liu, 21 Sep.2002. Foretarsus length 84.0µm (80-87µm, n=10), claw 26.3µm
183 Xiong et al.: A new species of Amphientulu from China
Figs. 1-10. Amphientulus sinensis, new species. 1, labial palpus; 2, pseudoculus; 3, mesonotum; 4, metanotum; 5, anterior part of head, dorsal view; 6, urotergite VI; 7, filamento di sostegno; 8, head chaetotaxy; 9, urosternite VI; 10, abdominal legs II, III. Scale bar: Figs. 1-2 = 5 µm; Figs. 3-10 = 10µm.
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Figs. 11-17. Amphientulus sinensis, new species. 11, foretarsus in interior view; 12, urosternite VII; 13, urotergite VII; 14, foretarsus in exterior view; 15, furotergite VIII-XII; 16, urosternite VIII-XII; 17, female squama genitalis. Scale bar: Figs. 11-17 = 10µm.
185 Xiong et al.: A new species of Amphientulu from China
Table 1: Chaetotaxy of Amphientulus sinensis, new species.
Dorsal Ventral Formula Composition of setae Formula Composition of setae Thorax I 4 Ð 4 A1, 2, M1, 2 4A 2, 4 6P1, 2, 3 II 6 A2, 4, M 7 Ð 2 Ac, 2, 3, 4, M 16 P1, 1a, 2, 2a, 3, 4, 4a, 5 4 P1, 2 III 6 A2, 4, M 7 Ð 2 Ac, 2, 3, 4, M 16 P1, 1a, 2, 2a, 3, 3a, 4, 5 4 P1, 2 Abdomen I 4 A1, 2 3 P1, 2 10 P1, 1a, 2, 2a, 3 4 P1, 2 II-III 6 A1, 2, 5 3 Ac, 2 16 P1, 1a, 2, 2a, 3, 4, 4a, 5 5 Pc, 1, 2 IV-VI 6 A1, 2, 53Ac, 2 16 P1, 1a, 2, 2a, 3, 4, 4a, 5 8 P1, 1a, 2, 3 VII 4 A4, 5 3 Ac, 2 16 P1, 1a, 2, 2a, 3, 4, 4a, 5 8 P1, 1a, 2, 3 VIII 6 A1, 3, 5 4 1, 2 15 Mc, P1, 1a, 2, 2a, 3, 3a, 5 0 IX 12 1, 1a, 2, 2a, 3, 4 4 X121, 1a, 2, 2a, 3, 4, 4 XI 4 2, 4 4 XII 9 6
Table 2. Comparison of the new species with Amphientulus ruseki (Nosek, 1978).
A. sinensis, new species A. ruseki Nosek, 1978 Foretarsal sensillium b distally to level of c nearly on level of c The ratio of exterior sensilla b>a>e>f>c>d>ga>b>d>e>c>f>g Foretarsal sensillium c nearly on level of d distally to level of d Foretarsal sensillium e nearer to f half way of f and g Foretarsal sensillium g reaching base of empodium not reaching base of empodium Urotergite I setae A5 absent A5 present Urotergite VII setae A2 absent A2 present Urotergite VIII setae Mc present Mc absent Urosternite XI 2+2 setae 3+3 setae TR= 3.2 2.8 PR= 11.8 19
(24-27µm, n=10), slender, no tooth. TR=3.2; Empodium without P3a; accessory seta as thin, hair-like, less than short, EU=0.15. S-shaped seta subequal to the claw. Dorsal one-fifth of P1 in length (Figs 6, 9). Tergite VII with A4 and sensillum t-1 claviform, t-2 long and thin, t-3 small, jar- 5, P3a absent; tergite VIII with the central seta Mc, p3a shaped, BS=0.4-0.5. Interior sensillum a« proximal to level present; tergite IX-X with six pairs of setae, seta 1a shorter of t-1, broad, sword-like; b« thin and long, its apex surpassing than seta 1; XI with 2+2 setae, both short. Pore al dorsally base of δ5; c« also thin and long, subequal to b«, its apex to A5 on tergite II-V, on VI and VII ventrally to it. Sternite surpassing base of δ6. Seta δ4 proximally to base of c«, VII with no Pc, sternite VIII without posterior setae, sternite subequal to β1. Exterior sensillum a slender and long, IX-XI with 2+2 setae. Pore on sternite VII situated asymmetrically near base of P1a (Figs. 12, 13, 15, 16). reaching almost level of γ3; b extremely long, reaching base of γ4; c nearly on level of d, longer than d. Seta e situated Abdominal legs II-III with two setae each, no terminal vesicle, nearly level with f, longer, reaching base of γ5, g shorter, but apical seta shorter than subapical one. On abdominal leg II, its apex reaching base of empodium. Length ratio of exterior the apical seta less than half of the subapical in length, the sensilla as: b>a>e>f>c>d>g (Figs. 11, 14). apical seta less than one-third of the subapical seta on abdominal leg III (Fig. 10). Striate band on abdomen VIII Abdomen Ð Chaetotaxy as shown in Table 1. Abdominal reduced, with undulant faint striae along the proximal margin, tergite I with two pairs of anterior setae (A1, and A2). Tergites comb consisting of 6-8 small, irregular teeth (Fig. 15). Female II-VI each with three pairs of anterior setae, A1, 2, and 5, squama genitalis with pointed acrostyli (Fig. 17).
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Etymology. Ð Named sinensis alluding to China, the place help in our taxonomic work. We thank Mr. Guoliang Xu for where the species was found. supplying some specimens to us. This work is supported by the National Natural Sciences Foundation of China (NO: Remarks. Ð The new species seems to be similar to 30130040). Amphientulus durumagi, described by Imadaté (1973) from Korea, but the latter is distinct in having double rows of setae on urosternite VIII and seta A4 on urotergite on VI from the LITERATURE CITED new species. The new species is closely similar in some respects to Amphientulus ruseki from Madagascar (Nosek, Imadaté, G., 1973. Contribution to the Proturan Fauna of Korea. 1978). However, they can be distinguished from each other Annales Historico-naturalis Musei Nationalis Hungarici, 65: by the relative length and the position of foretarsal sensilla 151-155. as well as by the chaetotaxy on abdomen. The differences Nosek, J., 1978. Madagascarian Protura I. Taxonomy. Bulletindes among them are listed in Table 2. Muséum national d'Histoire Naturelle. Paris. Écologie générale, 41: 3-28. Distribution. Ð Guangdong and Hainan Province, southern Tuxen, S. L., 1967. Australian Protura, their phylogeny and China. zoogeography Zeitschrift für zoologischen Systematik und Evolutionsforschung, 5 (1/2): 1-53. Tuxen, S. L., 1981. The systematic importance of “the striate band” ACKNOWLEDGMENTS and the abdominal legs in Acerentomidae (Insecta: Protura). With a tentative key to acerentomid genera. Entomologica We owe our very cordial thanks to Mr. Qiang Liu for scandinavica, 15: 125-140. collecting the specimens in our fieldwork. Our thanks are Yin, W. Y., 1999. Fauna sinica, Arthropoda: Protura, Beijing, given to Mr. Yiming Yang and Mrs. Yunxia Luan for their Science Press. Pp. 1-510.
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 189-209 Date of Publication: 31 Dec.2005 © National University of Singapore
A NEW SUBGENUS AND SIX NEW SPECIES OF NEPOMORPHA (INSECTA: HETEROPTERA) FROM YUNNAN, CHINA
Nico Nieser Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands.
Ping-ping Chen c/o Dept. of Entomology National Natural History Museum (Naturalis), Postbus 9517, 2300 RA Leiden, The Netherlands.
Chang Man Yang c/o Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore, 119260, Singapore.
ABSTRACT. Ð Five new species of Micronecta Kirkaldy, 1897b (Micronectidae), from Xishuangbanna, the extreme southwest of Yunnan Province, China are described: M. erythra, M. janssoni, M. lobata, M. melanochroa and M. ornitheia. Further, Anisops pseudostali, new species (Notonectidae) is described from the vicinity of Kunming. Micronecta khasiensis Hutchinson, 1940, and M. waltoniana Hutchinson, 1940, are removed from subgenus Mesonecta Poisson, 1938, and together with M. matsumurai Miyamoto, 1965, M. polhemusi Nieser, 2000, and M. melanochroa, new species are placed in the newly described subgenus Unguinecta (type species M. polhemusi). In addition, eight species: Micronecta drepani Nieser, 2000, M. guttatostriata Lundblad, 1933, M. jaczewskii Wróblewski, 1962, M. lemnae Nieser, 2000 (Micronectidae), Cercotmetus asiaticus Amyot & Serville, 1843 (Nepidae), Anisops breddini Kirkaldy, 1901, A. tahitiensis Lundblad, 1934, and Enithares stridulata Brooks, 1948 (Notonectidae) are recorded for the first time from China. Finally, some records of species already known from China are given.
KEY WORDS. Ð China, Yunnan, Micronectidae, Nepidae, Notonectidae, new subgenus, new species, new records.
INTRODUCTION species have a more limited distribution, and are usually restricted to more or less stagnant waters associated with This study is part of the results of a survey on the aquatic streams; such species may be predominantly brachypterous. insects of Xishuangbanna, in the southwest of Yunnan Apparently, the Micronecta specimens studied in this paper province, China. The survey was a joint research project mostly belong to the latter category. between the Raffles Museum of Biodiversity Research (National University of Singapore) and the Xishuangbanna The genus Micronecta in China has been poorly studied. Tropical Botanic Garden (Yunnan, China). This publication According to the checklist of Jansson (1995) fifteen species reports on the Micronectidae, Nepidae and Notonectidae occur in China, of which seven are known from Taiwan. collected from the area. The most interesting part of the However, Jansson overlooked the papers by Yang (1966, samples is the nine species of Micronecta Kirkaldy. 1967) in which three more species were described from Taiwan. This brings the total of Chinese species to eighteen, Species of Micronecta are small Nepomorphan bugs with a of which ten occur in Taiwan. The basic work for the study length from less than 1 to 4 mm, but in Asia only three species of Chinese Micronecta is by Chen (1960), who incorporated actually reach a length of over 3 mm. They live in stagnant thirteen of the species occurring in China in a key, and also or nearly stagnant waters. Some species are very widespread. gave short descriptions and redescriptions of four Chinese In the tropics, these are macropterous or with a significant species. Subsequently, Jansson (1995) and Wróblewski fraction of macropterous specimens. They occur in habitats (1968) cited numerous nomenclatural changes. In addition, in agricultural fields as well as in marshes or stagnant ponds one should consult Wróblewski (1960) who gives excellent in stream beds, e. g. M. quadristrigata Breddin, 1905. Other redescriptions of two species widespread in eastern China.
189 Nieser et al.: Nepomorpha from Yunnan
Further, Miyamoto (1965) described M. matsumurai from M. (Micronecta) janssoni SW China (Yunnan) Taiwan, and Lundblad (1934) described M. hummeli from new species Sichuan, which is the only reference useful for its M. (? Micronecta) lemnae SW China, Thailand, identification. Yang (1966, 1967) described three new species Nieser, 2000 Malay Peninsula M. (Micronecta) lenticularis SE China (Taiwan) from Taiwan, and presented a key to Taiwanese species in Chen, 1960 his 1967 publication. Finally, Hua (2000) mentions in his list M. (Micronecta) lobata SW China (Yunnan) of Chinese insects six species not incorporated in the checklist new species below: M. quadristriata is a misspelling for M. quadristrigata. M. (Micronecta) obtusa SE China (Taiwan) M. dione Distant, 1911 and M. proba Distant, 1911 (both not Yang, 1966 1910) are synonyms of M. scutellaris; M. quadriseta M. (Micronecta) ornitheia SW China (Yunnan) Lundblad, 1933c is a synonym of M. sedula; M. striata new species (Fieber, 1844) is a synonym of M. siva and M. thyesta Distant, M. (Micronecta) tuberculata SE China (Taiwan) 1911 (not 1910) is a synonym of M. grisea (Fieber, 1844) Yang, 1966 M. (Micronecta) unguiculata SE China (Taiwan) (see Jansson, 1995). Yang, 1967 M. (Micronecta) wui wui N China, E Asia Of the nine species of Micronecta collected in Lundblad, 1933c Xishuangbanna, five are new to science. The other four M. (Sigmonecta) quadristrigata SE China, Iran, species previously described are all new records for China, Breddin, 1905 tropical Asia, N Australia bringing the total number of Micronecta recorded from China M. (Unguinecta) matsumurai SE China (Taiwan) up to twenty-seven ( see checklist below). These four species Miyamoto, 1965 newly recorded for China are known from Thailand, Vietnam, M. (Unguinecta) melanochroa SW China (Yunnan) the Malay Peninsula or Java (Indonesia). This illustrates that new species the fauna of this southern part of Yunnan has its main affinities to tropical Southeast Asia. MATERIALS AND METHODS In the checklist below, the province name between brackets indicates that a species has been recorded so far only from Measurements are in mm and based on five specimens if that province. available, of each sex from the series containing the holotype. They are presented as the mean (
For nomenclature above the species level see Stysˇ & Jansson Micronecta (Micronecta) erythra, new species (1988). (Figs. 1-11)
The following abbreviations have been used for depositories: Material examined. Ð Holotype (CAS) - brachypterous male, CAS Institute of Zoology, Academy of Sciences, Beijing, CHINA: Yunnan, Xishuangbanna, 119 km from Jinghong to China. Mengla, rocky stream, 1 Jun.2002, coll. C. M. Yang & P. Chew, NCTN Nieser Collection, Tiel, The Netherlands. YCM310. RMNH National Natural History Museum (Naturalis), Paratypes: same data as holotype 1 female (ZRC); Yunnan, Leiden, The Netherlands. Xishuangbanna, Mengla, 55th stream, 30 May.2002, coll. C. M. ZRC Zoological Reference Collection of the Raffles Yang & P. Chew, YCM303, 1 male (NCTN); Yunnan, Museum of Biodiversity Research, Singapore. Xishuangbanna, Mengla, mountain stream by waterfall, 18 May.2000, coll. L. Cheng, LC001, 1 male (ZRC); km 10 Menglun to Mengyang, 21 May.2000, coll. L. Cheng, LC014, 1 male (ZRC); TAXONOMY Yunnan, pool at waterfall 65 power station, 21 May.2000, coll. L. Cheng, LC016, 1 male, 4 females (CAS, ZRC), 1 male, 1 female FAMILY MICRONECTIDAE (NCTN). All brachypterous.
Micronecta Kirkaldy, 1897b Description. - Based on brachypterous specimens in alcohol. In dorsal view a light reddish brown, shiny, rather small ovate Micronecta (Ctenonecta) jaczewskii Wróblewski, species with its greatest width at the caudal apex of claval new record commissure (Fig. 1).
Micronecta (Ctenonecta) jaczewskii Wróblewski, 1962: 178-180. Dimensions. Length, male
Micronecta (Lundbladella) guttatostriata Lundblad, new record
Micronecta guttatostriata Lundblad, 1933a: 101-102. Micronecta (Lundbladella) guttatostriata - Wróblewski 1967: 240- 243.
Material examined. Ð CHINA: Yunnan, Xishuangbanna, km 10 Menglun to Mengyang, 21 May.2000, coll. L. Cheng, 1 male, 1 female (ZRC); Yunnan, Morhan, stream near grazing field, 3 km from Laotian border, 23 May.2000, coll. L. Cheng, 1 male (ZRC). All brachypterous. First record for China.
Distribution. Ð Indonesia (Java), Malay Peninsula, Thailand, Vietnam (Nieser, 2002b) and SW China.
Micronecta (Micronecta) drepani Nieser, new record
Micronecta drepani Nieser, 2000: 278-281.
Material examined. Ð CHINA: Yunnan, Menglun, pool at waterfall 65 power station, 21 May.2000, coll. L. Cheng, 2 males macropterous, 4 larvae (ZRC). First record for China.
Distribution. Ð Thailand, SW China. Fig. 1. Micronecta erythra, holotype, brachypterous male, dorsal view, body length 1.65 mm. Scale: 0.1 mm.
191 Nieser et al.: Nepomorpha from Yunnan
Colour. Dorsally reddish to light brown; head sordid yellow, VIII five short two very long hair-like and one or two long eyes greyish brown. Pronotum unicolorous medium brown bristles. Pala (Fig. 9) with three dorsal bristles; upper row of except for caudolateral angles and posterior margin yellow; palm with about 13 bristles the distal one larger; ventral row scutellum sordid yellow. Hemielytra light brown with a with about 18 bristles which are becoming thicker distally. reddish hue due to small red dots which are especially dense Leg measurements as in table 1. proximally on clavus, covering the hyaline mark and along costal margin; distal margin of right membrane grey, Male. Fore leg (Fig. 9), femur with two dorsoapical spines, continuing in a grey stripe along posterior margin to apex of one dorsally in apical third and two in basal half. Tibia with clavus; left membrane entirely grey; embolium yellowish with one spine in apical third. Palar claw clavate to urn-shaped an ill-defined darker stripe near inner margin; no brown (Fig. 10). Prestrigilar lobe (Fig. 6) well differentiated, apically patches at costal margins. Venter and legs yellowish. The acutely pointed. Strigil small (Fig. 5) subrectangular, with holotype has a pair of large blackish marks on corium just one comb bearing about 35 densely packed teeth; next to it distally of the apex of clavus, one at costal margin and one there is a dark patch so that the strigil appears larger at first at the opposite margin. In the paratypes these marks are sight (Fig. 4). Mediocaudal lobe of seventh abdominal sternite smaller and less distinct. (Fig. 8) well developed, with a blunt apex and four well Structural characteristics. Length 1.9 times width. Width of defined bristles on its surface. Free lobe of left part of segment head and pronotum subequal, synthlipsis twice (1.7-2.1) the eight (Fig. 7) elongate, apical margin evenly rounded; with posterior margin of an eye. Pronotum poorly developed, about 17 bristles in lateroapical part which is slightly not produced. four (3.8-4.3) times as wide as long, lateral margins short. Right paramere (Fig. 2) with its shaft evenly curved, tapering Hemielytra densely but shallowly pitted on clavus and corium, in basal half, parallel-sided in apical half, apex rounded, basal with small pale spinules in most of the pits of corium. Spines process with about 30 stridulatory ridges. Left paramere (Fig. laterally on abdominal segments: VI two short, one or two 3) with a broad shaft with a small projection halfway on the long (Fig. 4); VII three short, one long or two short two long; side, opposite to the basal lobe.
Figs. 2-8. Micronecta erythra, new species, paratype, male; 2) right paramere; 3) left paramere; 4) right part of tergite VI; 5) strigil, scale 0.05 mm; 6) prestrigilar lobe; 7) free lobe of left part of tergite VIII; 8) mediocaudal lobe of sternite VII. Scale: 0.1 mm.
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Figs. 9-11. Micronecta erythra, new species, paratypes; 9) male, foreleg; 10) apex of male pala; 11) female, receptaculum seminis.
193 Nieser et al.: Nepomorpha from Yunnan
Table 1. Leg measurements* of Nepomorpha in mm.
femur tibia tars1 tars2 claw Anisops pseudostali fore leg male 1.421.880.92 0.41 middle leg male 2.48 2.05 0.93 0.52 0.28 hind leg male 3.42 3.03 1.20 1.10 fore leg female 1.45 1.70 0.63 0.39 0.41 middle leg female 2.56 2.13 0.91 0.56 0.28 hind leg female 3.68 3.16 1.25 1.26 Micronecta (M.) erythra fore leg male 0.31 0.19 0.19 middle leg male 0.73 0.24 0.38 0.14 hind leg male 0.50 0.44 0.50 0.20 0.13 fore leg female 0.38 0.39 middle leg female 0.69 0.23 0.36 0.16 hind leg female 0.50 0.42 0.48 0.19 0.11 Micronecta (M.) janssoni fore leg male 0.30 0.13 0.14 middle leg male 0.65 0.23 0.37 0.20 hind leg male 0.47 0.38 0.42 0.16 0.13 fore leg female 0.25 0.23 middle leg female 0.64 0.23 0.35 0.20 hind leg female 0.46 0.37 0.37 0.14 0.12 Micronecta (M.) lobata fore leg male 0.31 0.16 0.18 middle leg male 0.69 0.22 0.35 0.19 hind leg male 0.47 0.37 0.42 0.16 0.14 Micronecta (M.) ornitheia fore leg male 0.24 0.12 0.13 middle leg male 0.59 0.20 0.27 0.20 hind leg male 0.37 0.31 0.32 0.13 0.10 fore leg female 0.21 0.20 middle leg female 0.56 0.20 0.28 0.18 hind leg female 0.39 0.31 0.31 0.13 0.12 Micronecta (Unguinecta) melanochroa fore leg male 0.32 0.19 0.19 middle leg male 0.73 0.24 0.38 0.14 hind leg male 0.50 0.44 0.50 0.20 0.13 fore leg female 0.39 0.39 middle leg female 0.69 0.23 0.36 0.16 hind leg female 0.50 0.42 0.48 0.17 0.11 femur tibia tars1 tars2 claw The measurement of the fore leg in female Micronecta refers to the joint tibia and tarsus.
Female. Receptaculum seminis urn-shaped (Fig. 11). paramere of the same type but without the small projection halfway the shaft. Moreover, M. drepani has a large trowel- Macropterous form unknown. shaped palar claw, and is somewhat larger (male length 1.9- 2.1). Etymology. Ð Erythros (Greek adjective meaning reddish or red) referring to the reddish hue on the hemielytra. Micronecta (Micronecta) janssoni, new species Comparative notes. Ð This new species is structurally similar (Figs. 12-21) to M. hummeli Lundblad (1934) from northeast Sichuan, which has a virtually identical male fore leg, a similar free Material examined. Ð Holotype (CAS) - macropterous male, lobe of left part of segment eight, and is of the same size. CHINA: Yunnan, Xishuangbanna, 119 km from Jinghong to However, the right paramere of M. hummeli is apically hooked Mengla, rocky stream, 1 Jun.2002, coll. C. M. Yang & P. Chew, and not evenly curved, and the left paramere is of the same YCM310. general shape but has a distinct wide incision below the small Paratypes, same data as holotype, 14 males, 19 females (CAS, ZRC), projection on the shaft, more or less as in M. ornitheia new 6 males, 6 females (NCTN); in addition, 31 larvae were collected. species (Fig. 33), see also under that species. The right Yunnan, Xishuangbanna, Mengla, 55th stream, 30 May.2002, coll. paramere of M. erythra has a somewhat similar shape as in C. M. Yang & P. Chew, YCM303 (ZRC); Yunnan, Xishuangbanna, M. drepani Nieser, 2000 from Thailand, which has a left waterfalls near Mengyuan, 1 Jun.2002, coll. C. M. Yang & P. Chew,
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YCM309, 1 male (ZRC); Yunnan, Xishuangbanna, Menglun, Man- tongue-like, obtusely rounded; strigil (Figs. 18, 19) oval, with Er stream, 11 Jul.2001, coll. H. M. Liu, ME3-004F, 1 female (ZRC); a distinct stalk and one comb with about 90 densely packed Mengla, river by Jing Meng Yuan village near Mengyuan, 24 teeth. Mediocaudal lobe of seventh abdominal sternite (Fig. May.2000, coll. L. Cheng, LC033, 1 male, 5 females (ZRC). All 15) short, with an acute apex and four rather weak bristles macropterous. on its surface. Free lobe of left part of segment eight (Fig. 20) elongate, gradually broadening towards apex, apical Description. Ð Macropterous form, based on specimens in margin sinuate; with about 17 bristles in latero-apical part. alcohol. In dorsal view, generally a medium reddish brown, Right paramere (Fig. 16) with a broad shaft and a hooked medium sized, broadly ovate species with its greatest width apex, beset with three tooth-like structures, basal process with at the level of the caudal apex of claval commissure. about 40 stridulatory ridges. Left paramere (Fig. 17) rather broad with a small notch halfway the shaft and about three Dimensions. Length, male
Male. Fore leg (Fig. 13): femur with two apical spines, one Material examined. Ð Holotype (ZRC) - brachypterous male, in apical third and two in basal half; tibia with three spines dissected, parts glued on card, CHINA: Yunnan, Xishuangbanna, in apical third; palar claw a simple clavate flap (Fig. 14). Menghai, Mengkuan River, Km 57 Dalou to Menghai road, 19 Prestrigilar lobe (Fig. 21) well differentiated, apical part May.2000, coll. L. Cheng (LC006).
195 Nieser et al.: Nepomorpha from Yunnan
Description. Ð In dorsal view a light brown, shiny, medium lateral half of left membrane smoky grey. Venter medium sized ovate species, with its greatest width about one third of greyish brown, legs yellowish. the length of claval commissure anteriorly of its caudal tip. Structural characteristics. Body twice as long as wide. Dimensions. Male (holotype only), length 2.10, width 1.04, Pronotum very slightly wider (1.05) than head; synthlipsis width of head male 0.76, synthlipsis 0.36, posterior width of 1.7 times the posterior margin of an eye. Pronotum poorly an eye 0.21, width of pronotum 0.80, ocular index 1.8. developed, four times as wide as long, lateral margins short. Hemielytra shiny, densely and distinctly pitted on clavus and corium, with very small spinules in most of the pits of corium. Colour. Dorsally light brown; head light brown, eyes Spines laterally on abdominal segments: VI left side three castaneous. Pronotum unicolorous light brown, yellowish short, one long; right side two short one long and one band along posterior margin poorly differentiated; scutellum additional thin and intermediate (Fig. 27); VII three short, light brown. Hemielytra light brown; hyaline stripe at base one long; VIII four short, two very long bristle-like, and an of clavus rather short; corium mid way with a broad but additional one short bristle-like. Pala (Fig. 25) with three indistinct dark transverse band; embolium yellowish with a dorsal bristles; upper row of palm with 12 bristles; ventral greyish stripe at base and a brownish stripe at level of the row with 23 bristles, distally only slightly thicker than dark transverse band; caudal margin of right membrane and proximally. Leg measurements as in table 1.
Figs. 12-15. Micronecta janssoni, new species, paratypes, 12) female, receptaculum seminis; 13) male, foreleg; 14) apex of male pala; 15) male, mediocaudal lobe of sternite VII. Scale: 0.1 mm.
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Male. Fore leg (Fig. 25): femur with two dorsoapical spines, but its apical lobe is broader. In addition, M. johorensis and one dorsally in apical third and two in basal half; tibia with M. dentifera are smaller (length 1.8 or less), and have the a bristle about half way of dorsal margin and one near right paramere either with a narrow parallel shaft (M. apicodorsal angle; palar claw elongate, nearly parallel-sided dentifera) or basally broader and tapering towards apex (M. (Fig. 26). Prestrigilar lobe (Fig. 29) well differentiated, medio- johorensis). This new species does not run well in the key to apical projection short with a blunt tip. Right side of sixth SE Asian species by Nieser (2000). In view of the apical lobe tergite with very few submarginal bristles (Fig. 27). Strigil of the left paramere one could end in couplet 14: M. johorensis (Fig. 28) small, subcircular, with one comb with about 60 (see above), or in couplet 18: M. ludibunda Breddin, 1905 densely packed teeth. Mediocaudal lobe of seventh abdominal and M. malayana Leong, 1966, which have the shafts of both sternite (Fig. 31) well developed, with an acute apex and five parameres narrowly parallel-sided and hemielytra with well defined bristles on its surface. Free lobe of left part of distinct, reticulate or longitudinally striped patterns. segment eight (Fig. 30) elongate, nearly parallel-sided, apical margin shallowly and broadly indented; without bristles in lateroapical part which is not produced. Aedeagus without Micronecta (Micronecta) ornitheia, new species specialized teeth. Right paramere with its shaft widened in (Figs. 32-41) apical half and constricted just before its apex, basal process strongly projecting laterally, with about 20 stridulatory ridges Material examined. - Holotype (CAS) - macropterous male, which are based on the medial side of the process (Figs. 22, CHINA: Yunnan, Xishuangbanna, Mengla, 55th stream, 30 23). Left paramere (Fig. 24) with a broad shaft and an irregular May.2002, coll. C. M. Yang & P. Chew (YCM303). shape. Paratypes (adults only) - Same data as holotype, 4 males, 13 females (CAS, ZRC), 3 males, 3 females (NCTN); Yunnan, Xishuangbanna, Female and macropterous form unknown. 119 km from Jinghong to Mengla, rocky stream, 1 May.2002, coll. C. M. Yang & P. Chew, 1 male (ZRC); Yunnan, Xishuangbanna, Etymology. Ð Lobatus (Latin adjective, meaning lobate) refers forest stream, 9 km from Mengla road to Menglun, 18 May.2000, to the strongly developed basal process on right paramere. coll. L. Cheng (LC003), 5 males, 13 females (ZRC), 2 males, 4 females (NCTN); Xishuangbanna, Menglun, stream near Manpao Comparative notes. Ð The left paramere is similar to that of village, km 57 Menglun to Jinhong, 21 May.2000, coll. L. Cheng, M. johorensis Fernando, 1964 and M. dentifera Nieser, 2002a, 4 males, 2 females, 1 larva (ZRC), 1 male (NCTN); km 10 Menglun
Figs. 16-21. Micronecta janssoni, new species, paratype, male; 16) right paramere; 17) left paramere; 18) right part of tergite VI; 19) strigil, scale: 0.05 mm; 20) free lobe of left part of tergite VIII; 21) prestrigilar lobe. Figs. 16-18, 20-21, scale: 0.1 mm.
197 Nieser et al.: Nepomorpha from Yunnan to Mengyang, 21 May.2000, coll. L. Cheng (LC014), 1 male (ZRC); Colour. Dorsally generally light brown; head yellowish with Menglun, small waterfall near 55 power station, 21 May.2000, coll. an elongate orange spot medially between eyes, eyes L. Cheng (LC015), 10 males, 9 females (ZRC), 4 males 4 females castaneous; pronotum unicolorous light brown except for (NCTN); pool at waterfall 65 power station, 21 May.2000, coll. L. caudolateral angles and posterior margin narrowly yellow. Cheng (LC016), 25 males, 14 females (CAS, ZRC), 8 males, 8 females (NCTN); Xishuangbanna, clear forest stream, km 13 Hemielytra light brown with irregular reddish markings on Menglun to Mengyang, 22 May.2000, coll. L. Cheng (LC018), 1 costal third of corium, most distinctly in apical half, in male, 4 females (ZRC); Yunnan, Morhan, ditch by rice field near addition, indistinct slightly darker reddish brown to brown army base, 2 km from Laotian border, 23 May.2000, coll. L. Cheng patches on corium and membrane, some of which form a (LC021), 3 males, 2 females (ZRC); Yunnan, Shangyong, river by broken transverse zigzag band halfway corium; embolium roadside, Dalongha village, 23 May.2000, coll. L. Cheng (LC025), with a longitudinal dark greyish stripe, two medium brown 1 male (ZRC); Mengla, river by Jing Meng Yuan village near patches at costal margins, one just caudally of nodal furrow Mengyuan, 24 May.2000, coll. L. Cheng (LC033), 7 males 4 females (ZRC), 3 males 2 females (NCTN). All macropterous. and one at base of membrane; hyaline mark at base of clavus short, about 0.1 long not V-shaped and marked with reddish. Description. Ð Macropterous form, based on specimens in Thorax and abdomen ventrally and legs yellowish, in male alcohol. In dorsal view, generally a light brown, small, most of abdomen medium brown and aedeagus blackish, in elongate ovate species with its greatest width at the level of female ovipositor with brown marks; dorsal scent glands the caudal apex of claval commissure. black.
Dimensions. Length, male
Figs. 22-24. Micronecta lobata, new species, holotype, male; 22) right paramere; 23) right paramere, medial view; 24) left paramere. Scale: 0.1 mm. 198 THE RAFFLES BULLETIN OF ZOOLOGY 2005 short two very long hair-like. Pala (Fig. 39) with three dorsal paramere (Fig. 32) with a widened apex, vaguely reminding bristles; upper row and ventral row of palm each with about a bird head, basal process with about 30 stridulatory ridges. 17 bristles; the apical two bristles of both upper and lower Left paramere broad with a distinct notch in basal part (Fig. row are somewhat thicker than the remaining bristles in their 33). row. Leg measurements as in table 1. Female. Fore leg with the same set of spines as male; palar Male. Fore leg (Fig. 39): femur with two apical spines, one claw bristle-like, distinctly more strongly developed than the in apical third and two in basal half; tibia with three spines bristles in the palmar rows. Receptaculum seminis club- in apical third. Palar claw a simple clavate flap (Fig. 40). shaped (Fig. 37). Prestrigilar lobe (Fig. 36) well differentiated; strigil (Figs. 34, 35) oval, with a distinct stalk and one comb with about Brachypterous form unknown. 55 densely packed teeth. Median lobe of seventh abdominal sternite (Fig. 41) short, with an acute apex and three to four Etymology. Ð Ornitheios (Greek adjective: from a bird, bird- rather weak bristles on its surface. Free lobe of left part of like) refers to the characteristic apical part of the right segment eight (Fig. 38) elongate, gradually broadening paramere which is vaguely resembling a bird head. towards apex; with about 18 bristles in lateroapical part. Right
Figs. 25-31. Micronecta lobata, new species, holotype, male; 25) foreleg; 26) apex of pala, scale: 0.05 mm; 27) right part of tergite VI; 28) strigil, scale: 0.05 mm; 29) prestrigilar lobe; 30) free lobe of left part of tergite VIII; 31) mediocaudal lobe of sternite VII. Figs. 25, 27, 29- 31, scale: 0.1 mm.
199 Nieser et al.: Nepomorpha from Yunnan
Comparative notes. Ð With the key to SE Asian species absent; antennae three segmented, with the last segment (Nieser, 2000) this species might run to M. pocsi Wróblewski, largest and swollen; pronotum posteriorly convex; scutellum 1967, from Vietnam, which is slightly smaller (length 1.55- exposed; hemielytra with sparse pale pubescence, embolar 1.72 for males and 1.61-1.78 for females), and its right groove without nodal furrow; fore tibia and pala in males paramere is with a different shape: with the shaft broader in separate, in females fused; in both sexes, apical bristle of its middle part, the apical part less widened, and with a notch ventral palar row strongly developed into a secondary claw at its base. The left parameres of M. ornitheia, M. pocsi, M. on the pala (Fig. 50); claw of hind leg placed apically on hummeli Lundblad, 1934, from China (Sichuan) and M. tarsal segment two; seventh abdominal sternite with two stout singhpruthii Lundblad, 1933b, from India (Punjab) are quite long hairs. Male: prestrigilar lobe with a broadly rounded similar, but the latter two have the apex of the right paramere medial apex (Fig. 46); strigil present; free lobe of left part of hooked and not widened. The only species with a similar right segment eight with a slightly produced laterocaudal angle and paramere is M. sanctaecatherinae Hutchinson, 1940, from left parameres with a long and narrow, apically tapering shaft India and Sri Lanka. However, this species is larger (length (Fig. 43). Usually dark coloured except for M. waltoniana. 2.2-2.4), and has reddish spots on pronotum and hemielytra, Males have a row of small spines along ventral (flexor) margin moreover, it has a totally different left paramere with a of the fore femur except for M. khasiensis. Moreover, the widened tip giving it a club-shaped appearance. right parameres have a generally similar shape with a slightly curved, over most of its length more or less parallel, shaft with an obtusely pointed apex. Micronecta (Unguinecta), new subgenus Species included. Ð Micronecta (Unguinecta) polhemusi Type species. Ð Micronecta polhemusi Nieser, 2000. Nieser, 2000, from Thailand and the Malay Peninsula (type species); M. (U.) khasiensis Hutchinson, 1940, from India Description. Ð Medium sized Micronecta. Length, males 2.1- (Assam) and Vietnam; M. (U.) matsumurai Miyamoto, 1965, 2.8, females 2.5-3.0; rostrum with transverse sulcations; ocelli from China (Taiwan); M. (U.) melanochroa, new species,
Figs. 32-38. Micronecta ornitheia, new species, paratypes; 32-36 male; 32) right paramere; 33) left paramere; 34) right part of tergite VI; 35) strigil, scale: 0.05mm; 36) prestrigilar lobe; 37) female, receptaculum seminis; 38) male, free lobe of left part of tergite VIII. Figs. 32- 34, 36-38. scale: 0.1 mm.
200 THE RAFFLES BULLETIN OF ZOOLOGY 2005 from China (Yunnan); and M. (U.) waltoniana Hutchinson, 1938 from Madagascar. Mesonecta was based on the 1940 from India (Assam). following characters: claw of male pala flanked with a secondary claw; free lobe of left part of segment eight evenly Etymology. Ð Unguinecta [unguis (Latin: nail or claw) + necta rounded apically; both parameres with sharp apical points. (Greek suffix: swimmer)], means “clawed swimmer”, Hutchinson (1940) added M. khasiensis and M. waltoniana referring to the distinctly developed secondary claw on the to the subgenus Mesonecta without indicating on which pala. characters the classification was based. However, in his key to subgenera he mentioned the characters of the secondary Discussion. Ð In Micronecta the truly reliable characters for palar claw, inner angle of free lobe of left part of tergite VIII specific identification and delimitation of subgenera are the obtusely rounded, and seventh sternite with two large bristles, male sexual characters, which will be emphasized below. adding that the status of the last character is unknown in M. pilosa. Wróblewski (1962) pointed out that the only character Poisson (1938) established the monotypic subgenus connecting these three species is the secondary palar claw Mesonecta to include Micronecta (Mesonecta) pilosa Poisson, but left them in the subgenus Mesonecta.
Figs. 39-41. Micronecta ornitheia, new species, paratype, male; 39) fore leg; 40) apex of pala; 41) mediocaudal lobe of sternite VII. Scale: 0.1 mm.
201 Nieser et al.: Nepomorpha from Yunnan
In our opinion, Mesonecta has to be rectricted to its single (0.1) yellow; scutellum brown. Hemielytra medium brown original species, M. (M.) pilosa which differs from the but, when closed, appearing dark brown due to the underlying subgenus Unguinecta in the following characters: The left dark grey abdominal dorsum, without darker patches; distal paramere of M. pilosa has a broad, slightly sinuate shaft which margin of right membrane yellowish; embolium yellowish has a wide truncate apex pointing to one side; the right to light brown with a thin blackish stripe near inner margin; paramere has a narrow, straight shaft with a sharply pointed without brown patches at costal margins; hyaline mark at base apex; the free lobe of left part of segment eight is evenly of clavus short, about 0.12 long, its V-shape obscured by rounded. Consequently in Mesonecta both parameres and the reddish marking. Thorax and abdomen ventrally dark greyish free lobe of left part of segment eight are of entirely different brown. Legs pale yellowish with some small dark brown types compared to those of Unguinecta. In addition, M. pilosa marks, notably in distal half of pala and at distal edge of has a light colour, a length of 4 mm, and its hemielytra are middle tarsus. quite densely beset with black bristles, whereas the species of Unguinecta measure 2-3 mm and have sparse pale bristles Structural characteristics. Length 2.1-2.2 times width. Head on the hemielytra, except in M. polhemusi where the bristles slightly narrower than pronotum, synthlipsis 1.7-2.0 times are pale but more dense. Therefore, we conclude that as wide as the posterior margin of an eye. Pronotum well Mesonecta is a different subgenus. developed, dorsally convex with well developed lateral margins, 2.3-2.4 times as wide as long. Hemielytra smooth Yang (1967) described M. unguiculata from Taiwan in which with only sparse and small pale spinules, mostly in distal half the male also has a secondary palar claw. Compared to the of corium. Spines laterally on abdominal segments: VI two species of Unguinecta, this species differs in being distinctly short, three long and a long bristle (Fig. 44); VII five short, smaller (length of male 1.6); having a broad left paramere; three long; VIII three short two very long hair-like on left a different type of prestrigilar lobe with an obtuse but distinct part, and five short two very long hair-like on right part. Pala process pointing medially; and four large bristles on seventh (Fig. 49) with five dorsal bristles; upper row of palm with abdominal sternite. In our opinion, in spite of the secondary about 18 bristles the distal one somewhat larger; ventral row palar claw, the four large bristles on seventh abdominal with about 32 very fine bristles, the distal one conspicuously sternite and the male sexual characters place this species in larger than the proximal ones, developed into a secondary subgenus Micronecta. Yang (1967) correctly placed M. claw. Leg measurements as in table 1. unguiculata in the subgenus Micronecta. We consider that M. unguicula is closely related to M. (M.) lenticularis Chen, Male. Fore leg (Fig. 49): femur with two apical spines, of 1960, which is commonly distributed in Taiwan, instead of which one very small, and one in apical third, three spines M. matsumurai as indicated by Yang (1967). in basal half; in addition, there are some very small spines along ventral margin, several of which were represented by scars only in the microscopic slide of the paratype; tibia with Micronecta (Unguinecta) melanochroa, new species three long spines in apical third; palar claw triangular (Fig. (Figs. 42-50) 50). Prestrigilar lobe (Fig. 46) poorly differentiated, apically broadly rounded. Strigil (Figs. 44, 45) oval, with a distinct Material examined. Ð Holotype (ZRC) - macropterous male, stalk and one comb with about 50 densely packed teeth. CHINA: Yunnan, Xishuangbanna, Lancang river, 24 km from Median lobe of seventh abdominal sternite (Fig. 47) well Jinghong, potholes on large boulder, 10 m above river, 6 Jun.2002, developed, with a rounded apex and two bristles on its surface. coll. C. M. Yang & D.Q. Li. Free lobe of left part of segment eight (Fig. 48) elongate, apical margin sinuate; with about 22 bristles in lateroapical Paratypes, same data as holotype, 1 male (teneral, NCTN), 1 female (ZRC). All macropterous. part which is slightly produced. Right paramere (Fig. 42) with its shaft somewhat swollen in the middle and with a blunt Description. Ð Macropterous form, based on specimens in apex, basal process with about 20 stridulatory ridges. Left alcohol. In dorsal view, a dark brown, medium sized, rather paramere (Fig. 43) with a narrow shaft. parallel-sided species, with its greatest width just anteriorly of the caudal apex of claval commissure. Female. Palar claw bristle-like, twice as thick as the apical bristle in ventral row. Dimensions (measurements of the holotype mentioned first). Length, male 2.31-2.38, female 2.32; width, male 1.09 Brachypterous form unknown. (paratype not measured), female 1.05; width of head, male 0.83-0.82, female 0.82; synthlipsis, male 0.42-0.40, female Etymology. Ð Melanochroos (Greek adjective, meaning of 0.43; posterior width of an eye, male 0.25-0.22, female 0.21; dark colour) refers to the dark colour of this species. width of pronotum, male 0.92-0.90, female 0.90; ocular index, male 2.05-1.90, female 2.21. Comparative notes. Ð This species is closely related with the other species of subgenus Unguinecta: M. khasiensis, M. Colour. Dorsally generally dark brown; head yellowish, eyes matsumurai, M. polhemusi, and M. waltoniana. In males the castaneous, rostrum dark brown to blackish; clypeal area light triangular palar claw of M. melanochroa is distinctive, in M. to medium brown. Pronotum unicolorous medium brown khasiensis and M. waltoniana the male palar claw is narrow, except for caudolateral angles and posterior margin broadly virtually parallel-sided (Hutchinson, 1940: Figs. 171, 180);
202 THE RAFFLES BULLETIN OF ZOOLOGY 2005 in M. matsumurai it is narrowly clavate (Miyamoto, 1965: Material examined. Ð CHINA: Yunnan, Xishuangbanna, Mekong ppl. 43 fig. c) and in M. polhemusi clavate (Nieser, 2000: basin, Mengkuan river, 19 May.2000, coll. H. H. Tan, 1 male (ZRC). Fig. 21); in addition, the last two species have only about 23 bristles in ventral palmar row. The right paramere of M. Distribution. Ð India, Sri Lanka, Thailand, Vietnam, West matsumarai is more slender, and the right paramere of M. Malaysia, Sumatra, China, Japan. polhemusi has a distinct bulge at the base of the shaft. M. khasiensis and M. waltoniana have the shaft of the right paramere parallel-sided, not slightly swollen in the middle. Laccotrephes pfeiferiae (Ferrari) With the key to SE Asian species (Nieser, 2000) M. melanochroa might run to M. khasiensis. Nepa pfeiferiae Ferrari, 1888: 187. Laccotrephes robustus - Lundblad 1933a: 26-27; Polhemus 1995: 15 (partim). Laccotrephes pfeiferiae - Polhemus & Keffer 1999: 2-3. FAMILY NEPIDAE Material examined. Ð CHINA: Yunnan, Simao, Mekong basin, 22 Laccotrephes Stål, 1866 May.2000, coll. H. H. Tan, 1 female (ZRC); CHINA: Yunnan, Baoshan, Irawaddy basin, Gaoligong mountain, 28 May.2000, coll. Laccotrephes grossus (Fabricius) H. H. Tan, 1 female (ZRC).
Nepa grossa Fabricius, 1787: 277. Distribution. Ð Burma, Thailand, West Malaysia, Sumatra, Nepa kohlii Ferrari, 1888: 180-181. Java, S China including Taiwan. Laccotrephes grossus - Lundblad 1933a: 21 (synonymizes L. kohlii with L. grossus); Polhemus 1995: 15 (checklist, distribution); Remarks. Ð Until recently L. pfeiferiae was considered to be Polhemus & Keffer 1999: 3. a synonym of L. robustus Stål, 1871. Polhemus & Keffer
Figs. 42-48. Micronecta melanochroa, new species, paratype, male; 42) right paramere; 43) left paramere; 44) right part of tergite VI; 45) strigil, scale: 0.05 mm; 46) prestrigilar lobe; 47) mediocaudal lobe of sternite VII; 48) free lobe of left part of tergite VIII. Figs. 42-44, 46- 48, scale: 0.1 mm.
203 Nieser et al.: Nepomorpha from Yunnan
(1999) separated these species again, presented differential 67 kmfrom Mengla, 1 Jun.2002, coll. C. M. Yang & P. Chew, 2 characteristics for the five species in the L. grossus-group females; Yunnan, Xishuangbanna, Mangle stream, by paddy field, occurring in the Malay Archipelago, and stated that L. 4 Jun.2002, coll. C. M. Yang & L. Gao, 3 larvae. First record for robustus is restricted to the Philippines, and other records of China. L. robustus actually refer to L. pfeiferiae. Distribution. Ð Sundaland, Thailand (Lansbury, 1973), SW China. Cercotmetus Amyot & Serville, 1843
Cercotmetus asiaticus Amyot & Serville, new record Ranatra Fabricius, 1790 Ranatra lansburyi Chen, Nieser & Ho Cercotmetus asiaticus Amyot & Serville, 1843: 441. Cercotmetus asiaticus - Lansbury 1973: 89-92 (redescription). Ranatra lansburyi Chen, Nieser & Ho, 2004: 89. Material examined. Ð CHINA: Yunnan, Xishuangbanna, Mengkuan River, 19 May.2000, coll. H. H. Tan, 1 male, 4 larvae (ZRC); Material examined. Ð CHINA: Yunnan, Xishuangbanna, Mengla Yunnan, Xishuangbanna, Man-Er stream, Mengla (21.4°N, (21°4'N 101°5'E), stream QD2, 11 Jul.2001, coll. H. M. Liu, 1 male, 101.5°E), 11 Jul.2001, coll. H. M. Liu, 1 female (ZRC); Yunnan, (paratype, ZRC). Manchang stream, 63 km from Jinghong to Menglun, 30 May.2002, coll. C. M. Yang & P. Chew, 1 female; Yunnan, Xishuangbanna, Distribution. Ð China (Yunnan), Thailand (Chiang Mai).
Figs. 49-50. Micronecta melanochroa, new species, paratype, male; 49) foreleg; 50) apex of pala. Scale: 0.05 mm.
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Ranatra sterea Chen, Nieser & Ho Borneo, The Philippines and S Japan (Miyamoto, 1964; Nieser & Chen, 1991; Nieser, 2004). Ranatra sterea Chen, Nieser & Ho, 2004: 98-99.
Material examined. Ð CHINA: Yunnan, Baoshan (25°1'N 99°1'E), Anisops pseudostali, new species Irawaddy basin, Gaoligong mountain, 28 Sep.2000, coll. H. H. Tan, (Figs. 51-53) 2 females (holotype, ZRC, paratype NCTN). Material examined. Ð Holotype (NCTN, to be transferred to RMNH) Distrinution. - China (Yunnan). - macropterous male, CHINA: Yunnan Prov., 10 km NE Kunming, Jindian water reservoir, ca. 200x750m, edges flooded grassland, 27 Aug.1989, coll. P. Chen & N. Nieser. Ranatra unicolor Scott Paratypes: 2 macropterous females, same data as holotype (NCTN); Ranatra unicolor Scott, 1874: 452. Yunnan Prov., Baoshan on road from Fengqing to Baoshan, 23 Ranatra unicolor - Lansbury 1972: 312-314 (redescription). May.2000, coll. Y. X. Cai & H. H. Tan, THH0085, 1 male (ZRC).
Material examined. Ð CHINA: Yunnan, Lijiang, Yangtze basin, Description. Ð Macropterous form, based on pinned Lashihai lake, 25 May.2000, coll. H. H. Tan, 1 male (det. I. specimens which have been stored in alcohol. A large, Lansbury); Yunnan, Dali, Mekong basin, Jianhu lake, 25 May.2000, whitish, rather parallel-sided species with its greatest width coll. H. H. Tan, 1 male (det. I. Lansbury). posteriorly of the caudal apex of scutellum.
Distribution. Ð Japan, China, southern Russia, Central Asia, Dimensions (the holotype is the larger male). Length, male Caucasus, Saudi Arabia, Iran, Iraq (Chen et al., 2004). 9.00-9.48, female 9.41-9.61; width of head, male 2.12-2.13, female 2.16-2.23; humeral width of pronotum male 2.53-2.60, female 2.57-2.70; anterior width of vertex, male 0.53-0.54, FAMILY NOTONECTIDAE female 0.59-0.60; synthlipsis, male 0.33-0.37, female 0.38- 0.40; ocular index based on vertex, male 0.67-0.68, female Anisops Spinola, 1837 0.74-0.75.
Anisops breddini Kirkaldy, new record Colour. Dorsally sordid white to pale yellowish, eyes greyish brown; apex of abdomen brown, shining through the hyaline Anisops breddini Kirkaldy, 1901a: 5-6. wings. Ventrally head pale yellow, fourth rostral segment Anisops breddeni Brooks, 1951: 439-441 (misspelling, redescription). black; thoracic venter yellowish with variable darker patches; Anisops breddini - Nieser, 2004: 85 (synonymy). abdomen blackish except for median keel and connexiva yellow. Legs pale yellow, anterior face of middle femur with Material examined. Ð CHINA: Yunnan, Xishuangbanna, Mengkuan a broad, longitudinal brown stripe. River, 19 May.2000, coll. H. H. Tan, 1 male (det. I. Lansbury); Menglun, Lily pond, botanical garden, 25 May.2000, coll. L. Cheng, Structural characteristics. In dorsal view the outline of the 3 larvae IV-V (ZRC); Xishuangbanna, pool near power station, 6 head is trapezoid with posterior margin V-shaped, lateral Jun.2002, coll. C. M. Yang, 1 female (ZRC). First record for China. margins virtually straight and converging anteriorly, anterior margin almost straight, and vertex not or only very slightly Distribution. Ð India, Sri Lanka, Burma, Thailand, Vietnam, protruding. Width of head 0.8 times the humeral width of West Malaysia, Indonesia (Java, Sulawesi, New Guinea), and pronotum and about four times the anterior width of vertex SW China. (3.7-4.0). Along median axis, length of head two thirds the median length of pronotum (0.59-0.63). Labrum elongate, slightly longer than its width at base (0.33/0.30), without Anisops kuroiwae Matsumura obvious pilosity, apex obtuse. Frontal surface of third rostral segment with long procumbent hairs, more strongly Anisops kuroiwae Matsumura, 1915: 109. Anisops batillifrons Lundblad, 1933c: 463-464. developed in males than in females. Median length of Anisops batillifrons - Brooks 1951: 420-423 (redescription). pronotum 1.3-1.5 times the length of an eye; humeral width Anisops kuroiwai - Miyamoto, 1964: 67-68 (misspelling, of pronotum slightly less than twice its median length (1.8- synonymizes A. batillifrons with A. kuroiwae). 1.9). Surface of pronotum smooth without impressions or carinae; its lateral margins diverging posteriorly, slightly over Material examined. Ð CHINA: Yunnan, Xishuangbanna, Mengkuan half as long as median length (0.54-0.59); posterior margin River, 19 May.2000, coll. H. H. Tan, 2 males, 2 females (ZRC); strongly sinuate. Length of leg segments as in table 1. Nanian River, 19 May.2000, coll. H. H. Tan, 1 male (ZRC); Male structural characteristics. Synthlipsis two thirds the Xishuangbanna, Lanchang River, 24 km from Jinghong, potholes anterior width of vertex (0.66). Tylus bare, with a semicircular on boulder, 10 m above river, 6 Jun.2002, coll. C. M. Yang & D. Q. Li, 2 males, 2 females (ZRC). transverse depression around a wart-like tubercle medially at base of labrum. Rostral prong (Fig. 51) about half as long Distribution. Ð From India and Sri Lanka through Indochina as third rostral segment, with base originating about halfway and the southern part of China to West Malaysia, North and apex pointed. Stridulatory comb on fore tibia (Figs. 52,
205 Nieser et al.: Nepomorpha from Yunnan
53) consisting of 16 teeth of roughly the same size. Fore tarsus surface of third rostral segment, which are present in females without small pegs on its inner surface. of A. pseudostali though distinctly less developed than in males. Anisops A. campbelli is smaller than A. pseudostali, Female structural characteristics. Synthlipsis two thirds the length of males 7.3-8.1, length of females 8.1-8.4. In addition, anterior width of vertex (0.63-0.68). Except for sexual the basal teeth of the stridulatory comb in A. campbelli are much smaller than the apical teeth, whereas in A. pseudostali characters females are identical with males. the difference between basal and apical teeth is only slight (Fig. 53). Etymology. Ð The name pseudostali refers to its similarity to A. stali Kirkaldy, 1904. Anisops tahitiensis Lundblad, new record Comparative notes. Ð Anisops pseudostali is very similar to A. campbelli Brooks, 1951 (India and Burma) and A. stali, Anisops tahitiensis Lundblad, 1934: 121-123. a widespread species occurring from Australia through Anisops tahitiensis - Brooks 1951: 376-378 (redescription); Lansbury Indonesia and the Philippines to S. Japan (Okinawa) and S. 1964: 217-218 (distribution, discussion of variability). China (Guangdong, Taiwan) (Nieser & Chen, 1991; Liu & Material examined. Ð CHINA: Yunnan Prov., Xishuangbanna Zheng, 1991). Males of A. stali have proximally on the middle County, Km 10 from Menglun to Mengyang, 21 May.2000, coll. L. tibiae a projection beset with thickly-set stout setae, this Cheng, 1 male (ZRC). First record for China. projection is lacking in males of A. pseudostali. In addition, the vertex of A. stali males has a short cephalic projection Distribution. Ð From Andaman Islands, Vietnam, SW China anteriorly on vertex, which is lacking in A. pseudostali. and Peninsular Malaysia through Malesia to Australia, Tahiti, Females of A. stali lack the procumbent hairs on frontal Guadalcanal and Okinawa (Lansbury, 1964; Nieser, 2004).
Figs. 51-53. Anisops pseudostali, new species, holotype, male; 51, head in lateral view; 52) foreleg; 53) tibial comb; 51 and 52, Scale: 1 mm.
206 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Enithares Spinola, 1837 Distribution. Ð Thailand, Laos, Vietnam, SW China.
Enithares ciliata (Fabricius) Nychia Stål, 1859 Notonecta ciliata Fabricius, 1798: 524-524. Enithares ciliata - Lansbury 1968: 413-416 (redescription, Nychia sappho Kirkaldy, new record synonymy). Nychia marshalli var. sappho Kirkaldy, 1901b: 809-810. Material examined. Ð CHINA: Yunnan, Menglun, Botanic garden, Nychia sappho - Lansbury 1985: 4-9 (redescription, synonymy). lily pond, 20 May.2000, coll. L. Cheng, 1 female, 1 larva (ZRC); same data, pool, 24 May.2000, coll. L. Cheng, 4 males, 1 female, Material examined. Ð CHINA: Yunnan, Menglun, Xishuangbanna 1 larva (CAS, ZRC) (all det. I. Lansbury). First record for Yunnan. Tropical Botanic garden, Lily pond, botanical garden, 25 May.2000, coll. L. Cheng, 2 males, 2 females, brachypterous, 5 larvae (CAS, Distribution. Ð Mauritius, Bhutan, India, Sri Lanka, SE Asia, ZRC). First record for China. SE and SW China (Lansbury, 1968; Liu & Zheng, 1991). Distribution. Ð N Australia, Papua New Guinea, Indonesia, Philippines, Thailand, Burma, SW China. Enithares sinica (Stål) Remarks. Ð Nychia limpida Stål, 1859, was described from Notonecta sinica Stål, 1854: 241. Whampoa island, Guangdong, China. It is known only by Enithares sinica - Lansbury 1968: 378-380 (redescription). the holotype which is such a poor condition that, according to Lansbury (1985: 1-2) “It is inpractical to propose N. limpida CHINA: Yunnan, Xishuangbanna, Mekong basin, Dalou, Nanian as a prior name for species described post 1859. The River at border between China and Myanmar, 19 May.2000, coll. relationship of limpida to other species of Nychia must remain L. Cheng, 1 male (ZRC); same, coll. Y. X. Chai & H. H. Tan, 4 speculative until additional material is available from larvae; Mekong basin, Mengkuan River, 19 May.2000, coll. Y. X. mainland China.” Cai & H. H. Tan, 1 males, 4 females (CAS, ZRC); Yunnan, Xishuangbanna, 13 km road from menglun to Mengyan, 22 May.2000, coll. L. Cheng, 3 males, 1 female, 1 larva (CAS, ZRC); Yunnan, Baoshan (25.1°N 99.1°E) Irawaddy basin, Gaoligong ACKNOWLEDGEMENTS mountain, 28 Nov.2000, coll. H. H. Tan, 1 male, 1 females (ZRC) (all det. I. Lansbury). Thanks are due to Dr. I. Lansbury, Oxford, England for identifying part of the material and to H. Zettel, Vienna, Distribution. Ð China except for the NE and NW, Japan, Austria and an anonymous reviewer who made valuable Vietnam, Laos (in Natural History Museum at Vienna, comments on the manuscript. Yang would like to thank H. Austria, H. Zettel personal communication). M. Liu, director of the Xishuangbanna Tropical Botanic Gardens (Yunnan) and his staff for their cooperation and support during our field work in Xishuangbanna. She is Enithares stridulata Brooks, new record grateful to D. Li (NUS) for logistic arrangements in Yunnan, to Lanna Cheng, Pony Chew, D. Li, Matthew Lim and H. H. Enithares stridulata Brooks, 1948: 37-38. Tan for collecting material and to P. K. L. Ng for his interest Enithares stridulata - Lansbury 1968: 366-368 (redescription). and support on this project.
Material examined. Ð CHINA: Yunnan, Xishuangbanna, 29 km from Menghai to Dalou, stream by rice field, 19 May.2000, coll. L. LITERATURE CITED Chang, 1 male (ZRC); Yunnan, Menglun, clear flowing creek, 21 May.2000, coll. L. Cheng, 1 female (ZRC); Yunnan, Menglun, pool Amyot, C. J. B. & A. Serville, 1843. Histoire Naturelle des Insectes at waterfall, 21 May.2000, coll. L. Cheng, 3 females, 21 larvae (CAS, Hémiptères, Paris: i-lxxvi, 1-675. ZRC); Yunnan, Xishuangbanna, 13 km from menglun to mengyang, clear forest stream, 22 May.2000, coll. L. Cheng, 1 male, 2 females, Breddin, G., 1905. Übersicht der javanischen Micronecta-Arten 5 larvae (ZRC); Yunnan, mekong basin, Simao, ca. 2 km from (Rhynchota). Societas Entomologica, 20: 57. Ning’er to Yuanjiang stream, 21 May.2000, coll. Y.X. Cai & H.H. Brooks, G. T., 1948. New species of Enithares (Hemiptera, Tan, 1 male, 2 larvae (ZRC)Yunnan, Xishuangbanna, Mengmoe Notonectidae). Journal of the Kansas Entomological Society, stream, 30 May.2002, coll. C.M. Yang & P. Chew, 1 male, 2 females, 21: 37-54. 1 larva (ZRC); Yunnan, Xishuangbanna, Mengka, 55th stream, 30 Brooks, G. T., 1951. A revision of the genus Anisops (Notonectidae, May.2002, coll. C.M. Yang & P. Chew, 1 male (ZRC), 1 male Hemiptera). The University of Kansas Science Bulletin, 34: 301- (NCTN); Yunnan, Xishuangbanna, Mengka, Power station, canal 519. from stream, 31 May.2002, coll. C.M. Yang & P. Chew, 1 male, 2 females, 1 larva (ZRC); Yunnan, Xishuangbanna, Mengka, Chen, L. C., 1960. A study of the genus Micronecta of India, Japan, Taiwan and adjacent regions. Journal of the Kansas waterfalls near Mengyuan, 1 Jun.2002, coll. C.M. Yang & P. Chew, Entomological Society, 33: 99-118. 3 males, 5 females, 2 larvae (CAS, ZRC), 1 male, 1 female (NCTN); Yunnan, Man-Zhang stream, 12 Apr.2001, coll. H. M. Liu, 2 males Chen, P.-p., N. Nieser, & J.-Z. Ho, 2004. Review of Chinese (ZRC); Yunnan, Man-Er stream, 11 Jul.2001, coll. H. M. Liu, 1 Ranatrinae (Hemiptera: Nepidae), with descriptions of four new male (ZRC). New record for China. species of Ranatra Fabricius. Tijdschrift voor Entomologie, 147: 81-102.
207 Nieser et al.: Nepomorpha from Yunnan
Distant, W. L., 1911. The fauna of British India including Ceylon Lindberg, H., 1922. Neue Micronecta-Arten (Hem. Het.). Notulae and Burma. Rhynchota 5. Taylor & Francis, London. Pp. i-xii, Entomologicae, 2: 114-117. 1-362. Liu, G.-Q. & L.-Y., Zheng, 1991. Checklist of Chinese Notonectidae Fabricius, J. C., 1787. Mantissa insectorum 2. Hafniae. Pp. 1-382. (Heteroptera). Occasional Papers of the Natural History Fabricius, J. C., 1798. Entomologia systematica. Supplementum 1. Museum of Tianjin, 1991 (8): 43-44 (in Chinese). Hafniae. Pp. 1-572. Lundblad, O., 1933a. Zur Kenntnis der aquatilen und semiaquatilen Fernando, C. H., 1964. A new species of Micronecta (Hemiptera: Hemipteren von Sumatra, Java und Bali. Archiv für Corixidae) from Malaya. Proceedings of the Royal Hydrobiologie, Suppl. 12, Tropische Binnengewässer 4: 1-195, Entomological Society of London (B), 33: 17-18. 263-498, pls. i-xxi. Ferrari, E. von, 1888. Die Hemipteren-Gattung Nepa (sens. natur.). Lundblad, O., 1933b. Eine neue Micronecta-Art aus Vorderindien. Annalen des Kaiserlich-Koeniglichen Naturhistorischen Entomologisk Tidskrift, 54: 217-219, pl.10. Hofmuseums, 3: 161-194. Lundblad, O., 1933c. Some new or little known Rhynchota from Fieber, F. X., 1844. Entomologische Monographien. Leipzig & China. Annals and Magazine of Natural History, (10) 12: 449- Prague. Pp. i, 1-138. 464. Horváth, G., 1905. Hémiptères nouveaux de Japon. Annales Lundblad, O., 1934. Schwedisch-chinesische wissenschaftliche Historico-Naturales Musei Nationalis Hungarici, 3: 413-423. Expedition nach den nordwestlichen Provinsen Chinas. Arkiv för Zoologi, 27A (14): 1-31, pls I-II. Hua, L.-Z., 2000. List of Chinese Insects 1. Guangzhou. Pp. 212- 216. Matsumura, S., 1905. Die Wasser-Hemipteren Japans. Journal of Hutchinson, G. E., 1940. A revision of the Corixidae of India and the Sapporo Agricultural College, 2: 53-66. adjacent regions. Transactions of the Connecticut Academy of Matsumura, S., 1915. Uebersicht der Wasser-Hemipteren von Japan Arts and Sciences, 33: 339-476, pls. i-xxxvi. und Formosa. Entomological Magazine, Kyoto, 1: 103-119. Jansson, A., 1989. Stridulation of Micronectinae (Heteroptera, Miyamoto, S., 1964. On the names of two species of Anisops from Corixidae). Annales Entomologici Fennici, 55: 161-175. the Ryukyus, with the designation of lectotype of A. kuroiwai Jansson, A., 1995. Family Corixidae Leach, 1815 Ð water boatmen. Matsumura. Kontyû, 32: 67-68. Aukema, B. & Rieger, C., 1995. Catalogue of the Heteroptera Miyamoto, S., 1965. Notes on Formosan Corixidae (Hemiptera). of the Palaearctic Region, 1: 26-56. Kontyû, 33: 483-492. Jakovlev, V. E., 1881. Contributions to the Hemipteran fauna of Motschulsky, V., 1863. Essai d’un catalogue des insectes de l’île Russia and adjacent regions. VIII. New species from Asiatic Ceylan (Suite). - Bulletin de la Société Impériale des Naturalistes Russia. Bulletin de la Société Impériale des Naturalistes de de Moscou, 36: 1-153. Moscou, 56: 206-214. (In Russian with German diagnoses of Nieser, N., 1975. The water bugs (Heteroptera: Nepomorpha) of new species). the Guyana region. Studies on the Fauna of Suriname and other Kirkaldy, G. W., 1897a. Notes on the genus Sigara, Fabr. Guyanas, 16: 1-308, pls. 1-24. (Rhynchota). The Entomologist, 30: 238-240. Nieser, N., 2000. Three new species of Micronecta from Thailand Kirkaldy, G. W., 1897b. Synonymic notes on aquatic Rhynchota. with a key to SE Asian species. (Heteroptera: Corixidae). The Entomologist, 30: 258-260. Journal of the New York Entomological Society, 107 (1999): Kirkaldy, G. W., 1901a. Miscellanea Rhynchotalia. The 277-288. Entomologist, 34: 5-6. Nieser, N., 2002a. Four new species of Micronecta (Insecta: Kirkaldy, G. W., 1901b. On some Rhynchota, principally from New Heteroptera: Micronectidae) from the Malay Peninsula. The Guinea, (Amphibicorisae and Notonectidae). Annali del Museo Raffles Bulletin of Zoology, 50(1): 9-16. Civico di Storia Naturale Giacomo Doria, 20: 804-810. Nieser, N., 2002b. Guide to Aquatic Heteroptera of Singapore and Kirkaldy, G. W., 1904. Über Notonectiden (Hemiptera). Wiener Peninsular Malaysia. IV Corixoidea. The Raffles Bulletin of Entomologische Zeitung, 6: 93-135. Zoology, 50(1): 263-274. Lansbury, I., 1968. The Enithares (Hemiptera-Heteroptera: Nieser, N., 2004. Guide to aquatic Heteroptera of Singapore and Notonectidae) of the Oriental region. Pacific Insects, 10: 353- Peninsular Malaysia. 3. Pleidae and Notonectidae. The Raffles 442. Bulletin of Zoology, 52(1): 79-96. Lansbury, I., 1972. A review of the oriental species of Ranatra Nieser, N. & P.-p. Chen, 1991. Naucoridae, Nepidae and Fabricius (Hemiptera-Heteroptera: Nepidae). Transactions of the Notonectidae, mainly from Sulawesi and Pulau Buton Royal Entomological Society of London, 124: 287-341. (Indonesia). Tijdschrift voor Entomologie, 134: 47-67. Lansbury, I., 1973. A review of the genus Cercotmetus Amyot & Poisson, R. A., 1938. Les Micronectes de la région Malgache. Revue Serville, 1843 (Hemiptera-Heteroptera: Nepidae). Tijdschrift Française d’Entomologie, 5: 135-144. voor Entomologie, 116: 83-106. Polhemus, J. T., 1995. Family Nepidae. B. Aukema & C. Rieger Lansbury, I., 1985. Notes on the identity of Nychia Stål (Hemiptera- (Eds.) Catalogue of the Heteroptera of the Palaearctic Region, Heteroptera: Notonectidae) in Australia. The Beagle, Occasional 1: 14-18. Papers of the Northern Territory Museum of Arts and Sciences, Polhemus, J. T. & S. L. Keffer, 1999. Notes on the genus 2: 1-9. Laccotrephes Stål (Heteroptera: Nepidae) the Malay Leong, C. Y., 1966. A taxonomic study of the malayan Corixidae Archipelago, with the description of two new species. Journal (with the description of Micronecta malayana sp. nov. Bulletin of the New York Entomological Society, 107: 1-13. of the National Museum of Singapore, 12: 83-90.
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Scott, J., 1874. On a collection of Hemiptera-Heteroptera from Japan. Stys, P. & A. Jansson, 1988. Check-list of recent family- group and Descriptions of various new genera and species. Annals and genus-group names of Nepomorpha (Heteroptera) of the World. Magazine of Natural History, (4) 14: 426-452. Acta Entomologica Fennica, 50: 1-44. Spinola, M. M., 1837. Essai sur les insectes Hémiptères L. ou Wróblewski, A., 1962. Notes on Micronectinae from Viêt-Nam Rhyngotes F. et à la section des Hétéroptères Duf.: 1-383. (Heteroptera, Corixidae). Bulletin de l’Académie Polonaise des Graviers, Geneva. Sciences Cl. II, 10: 175-180. Stål, C., 1854. Nya Hemiptera. Öfversigt af Kungliga Wróblewski, A., 1967. Further notes on Micronectinae from Viêt- Vetenskapsakademiens Förhandlingar, 8: 231-255. Nam (Heteroptera, Corixidae). Polskie Pismo Entomologiczne, Stål, C., 1859. Hemiptera. Species novas descripsit. Kongliga 37: 229-251. Svenska Fregattens Eugenies resa omkring Jorden under befál Wróblewski, A., 1968. Notes on oriental Micronectinae af C.A. Virgin ären 1851-1853. - Zoologi 1 Insecta: 219-298. (Heteroptera, Corixidae). - Polskie Pismo Entomologiczne, 38: Stål, C., 1866. Hemiptera Africana 3(1865): 1-200. Nordstetiana, 229-251. Holmiae. Yang, C. T., 1966. Two new Micronecta from Taiwan (Hemiptera, Stål, C., 1871. Hemiptera Insularum Philippinarum. Bidrag till Corixidae). Plant Protection Bulletin, 8: 213-216. Philippinska oarnes Hemipter-fauna. Öfversigt af Kungliga Yang, C. T., 1967. A new and an unrecorded species of Micronecta Vetenskapsakademiens Förhandlingar, 27 (1870): 607-776. with a key to Taiwanese species (Hemiptera, Corixidae). Bulletin of the Institute of Zoology Academia Sinica, 6: 43-46.
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 211-220 Date of Publication: 31 Dec.2005 © National University of Singapore
EMPIDINE DANCE FLIES FROM SINGAPORE AND SOUTHERN MALAYSIA (DIPTERA: EMPIDIDAE: EMPIDINAE)
Christophe Daugeron Muséum national d’Histoire naturelle, Département Systématique et Evolution, USM 601 & CNRS, UMR 5202, 45 rue Buffon, 75005 Paris, France Email: [email protected]
Patrick Grootaert Royal Belgian Institute of Natural Sciences, Department of Entomology Rue Vautier, 29, 1000 Brussels, Belgium Email: [email protected]
ABSTRACT. Ð Five new species belonging to the Empis (Coptophlebia) hyalea-group are described, namely E. (C.) neesoonensis, new species, E. (C.) producta, new species, E. (C.) tortuosa, new species, E. (C.) triseta, new species and E. (C.) verruca, new species. A key to all known species of the group is provided for Singapore and Southern Malaysia.
KEY WORDS. Ð Diptera, Empididae, Empidinae, Empis (Coptophlebia) hyalea-group, new species, Oriental region, Singapore, Malaysia, key to species.
INTRODUCTION (KBIN-IRScNB), the Raffles Museum of Biodiversity Research (ZRC), and the Muséum national d’Histoire The Empis (Coptophlebia) hyalea-group was recently naturelle, Paris (MNHN). recognised as a monophyletic group within the tribe Empidini of the subfamily Empidinae (Daugeron, 2002). Since this first The morpho-anatomical terms follow McAlpine (1981), publication on the group, forty-four species were described Sinclair et al. (1994), Cumming et al. (1995) and Daugeron from the Afrotropical (Central African Republic, Ivory Coast (1997). However, there is a need to refer to the detailed and Guinea), Oriental and Australasian (China, Indonesia, diagnosis recently given for the E. (C.) hyalea-group Japan, Laos, Malaysia, New Caledonia, Papua New Guinea, (Daugeron & Grootaert, 2005) as several characters, common Singapore, Thailand and Vietnam) regions (Daugeron & to most species, are not repeated in the descriptions proposed Grootaert, 2003a, b, 2005). Although the group is represented here. The abbreviations used in the figures are as follows: worldwide, it seems to be much more diversified in tropical cerc, cercus; ej ap, ejaculatory apodeme; epn, epandrium; hyp, and subtropical areas of the Oriental and Australasian regions hypandrium; ph, phallus. where it is the main representative of the tribe Empidini. A cladistic analysis recognised six main smaller taxonomic units within the group (Daugeron & Grootaert, 2005). In this paper TAXONOMY we continue our taxonomic inventory of the E. (C.) hyalea- group in Southeast Asia with the description of five new Empis (Coptophlebia) neesoonensis, new species species recently collected in Singapore and southern (Figs. 1A, 2) Malaysia. Material examined. Ð Holotype - male, Singapore, Nee Soon, primary rain forest, sample 22054, coll. P. Grootaert, 4 Dec.2002 MATERIALS AND METHODS (ZRC). Paratype Ð 1 female with same data (ZRC). This study is mainly based on material preserved in alcohol and collected by one of us (P.G.) during several field trips Other material Ð 1 male in bad condition (head missing), Singapore, in Malaysia and Singapore (2000, 2002, 2003). The material NUS campus, sample 23082, coll. P. Grootaert, 22 Nov.2003 (KBIN- is deposited in the Royal Belgian Institute of Natural Sciences IRScNB).
211 Daugeron & Grootaert: New empidine dance flies
Etymology. Ð The name of the species is derived from the Thorax brown, scutum subshiny. Acrostichals missing; type-locality. dorsocentrals uniserial ending with 1 strong, long bristle in prescutellar depression. Legs (Fig. 1A) brown. Left mid and Diagnosis. Ð Small blackish species with all bristles black, hind legs missing. All femora with rather short bristles. Fore wing clear with median veins almost not visible, fore tibia tibia deformed, somewhat S-shaped with about 6 spine-like deformed with about 6 spine-like anterodorsal bristles in basal anterodorsal bristles in basal half; first fore tarsomere with half not longer than tibia depth. 1 strong, long anterolateral bristle at base. Mid tibia with 3 strong, rather long dorsal bristles (at basal quarter, about at Description. Ð Male Ð Head. Occiput, antenna dark brown, middle, at tip); first mid tarsomere with 1 strong, long second and third flagellomeres aristiform. Face blackish. anterodorsal bristle at apical tip. Hind tibia swollen apically Palpus apparently brown. Labrum length about twice head with strong, long dorsal bristles, several distinct ventral height. bristles at apical half; hind tarsus swollen, first four tarsomeres
Fig. 1. Male fore leg in frontal view: A, tibia and first tarsomere of E. (C.) neesoonensis; B, tibia and tarsus of E. (C.) producta; C, tibia and first tarsomere of E. (C.) tortuosa; D, tibia and first tarsomere of E. (C.) verruca.
212 THE RAFFLES BULLETIN OF ZOOLOGY 2005 with long dorsal bristles. Wing (length = 2.2 mm) clear. Discal Empis (Coptophlebia) producta, new species cell and median veins almost not visible. A1 abbreviated. (Figs. 1B, 3, 4, 5) Halter brown. Abdomen brown, sternites with pair of distinct bristles posteriorly. Hypopygium (Fig. 2). Epandrium Material examined. Ð Holotype - male, Singapore, Pulau [= Island] triangular with strong, long bristles at tip. Hypandrium Ubin, Chek Jawa, sample 22059, coll. P. Grootaert, 11 Dec.2002 pointed. Phallus thin. (ZRC).
Paratypes Ð 3 males with same data, 1 male, Singapore, Pulau [= Female similar to male except for the following characters: Island] Ubin, 11 Dec.2002, mangrove, sample 23119, coll. P. palpus almost yellow. Dichoptic with all ommatidia of the Grootaert (ZRC); 35 males, Singapore, Pulau [= Island] Ubin, sample same size. Fore femora with distinct dorsal pennation at base; 23080, coll. P. Grootaert, 19 Nov.2003 (ZRC, KBIN-IRScNB, first fore tarsomere with short dorsal pennation. Mid and hind MNHN). femora with long dorsal and ventral pennation. Mid tibia with distinct dorsal pennation and some distinct ventral pennate Etymology. Ð From the Latin word productus in reference to bristles at base. Hind tibia with long dorsal and ventral the dorsal part of the epandrium that is considerably produced pennation. posteriorly.
Remarks. Ð Empis neesoonensis belongs to the E. jacobsoni Diagnosis. Ð Medium-sized blackish species with all bristles clade as defined recently (Daugeron & Grootaert, 2005). black, fore tibia deformed with row of anterodorsal spine- Within this clade E. neesoonensis seems to be closely related like bristles on basal third, first fore tarsomere elongated, to E. cameronensis (Malaysia) and differs from this species following three tarsomeres swollen with numerous long, fine by the following characters: smaller species; anterodorsal dorsal bristles, wing with visible veins, epandrium produced spine-like bristles of fore tibia shorter than tibia depth; first posterodorsally. mid tarsomere not whitish; median veins almost not visible; epandrium rather pointed at tip, not rounded, with ventral Description. Ð Male Ð Head. Occiput dark brown blackish, bristles less numerous basally. antenna black, first flagellomere somewhat conical, second
Fig. 2. Male hypopygium of E. (C.) neesoonensis, lateral view. Scale = 4.2 mm.
213 Daugeron & Grootaert: New empidine dance flies and third flagellomeres aristiform. Palpus brown, labrum Remarks. Ð This species belongs to the E. jacobsoni clade length 1.5x head height. Thorax blackish. Dorsocentrals as defined recently (Daugeron & Grootaert, 2005). Within uniserial ending with 2 strong, long bristles in prescutellar this clade it is closely related to E. padangensis (Indonesia: depression. Legs (Fig. 1B). Fore tibia deformed with row of Sumatra) with fore tarsus distinctly less rounded than in this anterodorsal spine-like bristles on basal third ending in 2-4 species. Both species occur in the same kind of coastal habitat, longer bristles (well visible in anterior view); first fore i.e. in mangroves (Fig. 5). tarsomere elongated, almost as long as the following four tarsomeres together, with 1 distinct anterolateral and posterodorsal bristles at base and at tip respectively; following Empis (Coptophlebia) tortuosa, new species three tarsomeres swollen with numerous long rather fine (Figs. 1C, 6A, B) dorsal bristles, 1 strong, long ventral bristle. Mid tibia with 3 strong, long ventral and dorsal bristles; first mid tarsomere Material examined. Ð Holotype - male, Singapore, Nee Soon, sample with 1 strong, long posterodorsal bristle at base and at tip. 20030, coll. P. Grootaert, 5 Oct.2000 (ZRC). Hind tibia and tarsus with several strong, long postero- and anterodorsal bristles, hind tarsus somewhat thick. Wing Paratypes Ð 8 males, Singapore, Nee Soon, sample 25070, coll. P. Grootaert, 19 Apr.2005 (ZRC, KBIN-IRScNB, MNHN). (length = 2.4 mm) feebly tinged with brown, all veins well visible except M1, A1 abbreviated. Halter dark brown. Etymology. Ð From the Latin word tortuosus in reference to Abdomen dark brown, paler, with distinct short bristles at the shape of bristles of the epandrium and male sternite 8. base. Hypopygium (Fig. 4). Epandrium rather truncate at tip, projected posterodorsally, with row of very strong, long Diagnosis. Ð Medium-sized brownish species, fore tibia bristles on the dorsal margin, several shorter, finer bristles strongly deformed at base, legs brownish with mid tarsus at tip. Hypandrium somewhat rounded at tip. Phallus rather distinctly paler, hind tibia with a characteristic cluster of 4 long, narrowed at tip. very long ventral bristles basally, sternite 8 with numerous long, tortuous ventral bristles directed anteriorly, ventral Female unknown. margin of epandrium with a characteristic indentation subapically.
Description. Ð Male Ð Head. Occiput dark brown, antenna brownish, second and third flagellomeres aristiform. Face blackish. Palpus clear yellowish, labrum length 2.3 x head height. Thorax. Acrostichals and dorsocentrals broken but apparently uniserial. Proepisternum with 1 fine, short bristle, prosternum with 1 long lateral bristle. Legs (Fig. 1C) brown, mid tarsus almost yellowish. Fore tibia strongly deformed at base with 2 distinct anterodorsal bristle (at middle, at apical third), 2 rather strong, long posterolateral bristles; first fore tarsomere with 1 strong long antero- and posterolateral bristles, 1 strong, long posteroventral bristle. Mid femur with 2 long ventral bristles at middle; mid tibia with 2 and 3 strong, long ventral and dorsal bristles respectively; first mid tarsomere with 2 strong, long anterodorsal bristles. Hind tibia with a cluster of 4 very long ventral bristles basally, rather strong at base, finer at tip, 4 strong, long dorsal bristles (1 basally, 1 at apical third, 2 apically); hind tarsus somewhat swollen, first hind tarsomere almost as long as the following four tarsomeres together, with 3 strong, long dorsal bristles, following tarsomeres with 2 long dorsal bristles. Wing (length = 2 mm) feebly tinged with brown, discal veins almost not visible, A1 abbreviated. Abdomen brownish. Sternites with distinct pair of bristles posteriorly. Sternite 8 (Fig. 6A) with numerous strong, long, tortuous ventral bristles directed anteriorly. Hypopygium (Fig. 6B). Epandrium subtriangular, long, with a ventral indentation subapically, strong, long, more or less tortuous ventral bristles, 1 very strong bristle at tip, several strong, long dorsal bristles. Hypandrium somewhat elongated, fine at tip. Phallus somewhat recurved at tip
Female unknown. Fig. 3. Habitus of E. (C.) producta, male.
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Remarks. Ð This species belongs to the E. hyalea clade. Etymology. Ð From the Latin word seta in reference to the Within this clade it probably forms a small monophyletic presence of three characteristic dorsal bristles on the male group with E. vitisalutatoris (Singapore) and E. thiasotes fore tibia. (Indonesia: Java) on the basis of the presence of a characteristic ventral indentation of the epandrium. Diagnosis. Ð Small-sized dark brown species with all bristles black, fore tibia with 3 distinct dorsal bristles, cercus with a characteristic ventral projection at tip. Empis (Coptophlebia) triseta, new species (Fig. 7) Description. Ð Male Ð Head. Occiput dark brown blackish, antenna blackish, second and third flagellomeres aristiform. Material examined. Ð Holotype - male, Malaysia, Kota Tinggi, Labrum length about 1.5 times head height. Thorax dark riverbed, sample 22041, coll. P. Grootaert, 11 Oct.2000 (KBIN- brown with most bristles missing, dorsocentrals apparently IRScNB). uniserial. Legs brown. Fore tibia with 3 distinct dorsal bristles (at middle, at apical third, at apical quarter), the first two
Fig. 4. Male hypopygium of E. (C.) producta, lateral view. Scale = 4.2 mm.
215 Daugeron & Grootaert: New empidine dance flies longer than tibia depth; first fore tarsomere with 1 strong, Empis (Coptophlebia) verruca, new species rather long anterolateral bristle at base, 1 strong anteroventral (Figs. 1D, 8) bristle at base. Mid femur with strong, long ventral bristles at basal half; mid tibia with 2 strong, long dorsal bristles (at Material examined. Ð Holotype - male, Singapore, Nee Soon, basal and apical quarter), 1 strong, long ventral bristle at sample 22057, coll. P. Grootaert, 5 Oct.2000 (ZRC). basal third; first mid tarsomere with 1 strong, long posterodorsal bristle apically. Hind tibia with row of strong, Paratypes Ð 1 male with the same data (ZRC); 6 males and 1 female, Singapore, Nee Soon, primary rain forest, sample 22054, coll. P. long dorsal bristles, several strong, rather long ventral bristles Grootaert, 4 Dec.2002 (ZRC, KBIN-IRScNB, MNHN); 1 male, at apical half; hind tarsus swollen; first hind tarsomere with Singapore, Nee Soon, sample 23102, coll. P. Grootaert, 3 Dec.2002 2 pairs of strong, long dorsal bristles, following tarsomeres (ZRC). with distinct pair of dorsal bristles apically. Wing (length = 2 mm) feebly tinged with brown, A1 abbreviated. Halter Etymology. Ð From the Latin word verruca in reference to brown. Abdomen brown. Sternites with distinct pair of the presence of a tubercle on the male fore tibia. bristles posteriorly. Hypopygium (Fig. 7). Cercus with a characteristic short ventral projection at tip. Epandrium Diagnosis. Ð Medium-sized brownish species with fore tibia subtriangular with some strong bristles ventrally and at tip. Hypandrium short, pointed at tip. Phallus rather long, thin. deformed and bearing an anterodorsal tubercle at basal quarter. Female unknown. Description. Ð Male Ð Head. Occiput dark brown blackish, Remarks. Ð The presence of characteristic dorsal bristles on antenna blackish, second and third flagellomeres aristiform. the fore tibia indicates that this species is a possible member Palpus brown, labrum length almost twice head height. of the E. hyalea clade although the fore tibia is not distinctly Thorax dark brown. Proepisternum with 1 fine, short bristle, deformed. Only the particular shape of cercus tip allows to proepisternum with 1 rather long bristle. Dorsocentrals be sure of the species identification. uniserial, strong, long. Legs (Fig. 1D) brown. Fore tibia deformed at base with a characteristic anterodorsal tubercle at basal quarter, 1 rather strong dorsal bristle at middle, longer than tibia depth; first fore tarsomere with 1 strong, long anterolateral bristle at base, 1 strong, long dorsal bristle apically. Mid tibia with 3 strong dorsal bristles (at basal quarter, near middle, at tip), the last two long, 1 strong ventral bristle at basal third; first mid tarsomere with 2 strong, long dorsal and ventral bristles basally, 1 strong, very long posterodorsal bristle apically, following three mid tarsomeres with 1 strong, long dorsal bristle apically. Hind tibia with row of strong, long dorsal bristles; first hind tarsomere with pair of strong, long dorsal bristles apically, following four tarsomeres with pair of distinct dorsal bristles apically. Wing (length = 2.6 mm) feebly tinged with brown, A1 abbreviated. Halter brown. Abdomen brown. Sternites with pair of distinct bristles posteriorly. Hypopygium (Fig. 8). Epandrium distinctly triangular with 3 strong, long bristles at tip. Hypandrium long, pointed at tip. Phallus somewhat narrowed apically.
Female similar to male except for the following characters: wing more distinctly brown. Fore femur distinctly paler than coxa and tibia; fore tibia with dorsal pennation as long as tibia depth; first fore tarsomere with short dorsal pennation except apically; mid femur, hind femur and tibia with long dorsal and ventral pennation; mid tibia with short dorsal pennation; first hind tarsomere with short dorsal pennation basally.
Remarks. Ð This species belongs to the E. hyalea clade. It can be distinguished from all other species by the presence Fig. 5. Photographs showing the typical coastal habitat of E. (C.) producta: mangroves of Singapore (Pulau Ubin). of a characteristic tubercle on fore tibia.
216 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Fig. 6. Male postabdomen of E. (C.) tortuosa, lateral view: A, sternite 8; B, hypopygium. Scale = 4.2 mm.
217 Daugeron & Grootaert: New empidine dance flies
Empis (Coptophlebia) vitisalutatoris ventrals, 2 strong, long dorsals, hypandrium long with bristly Daugeron & Grootaert, 2005 hairs at tip.
This species was described from nine males (Daugeron & Female unknown. Grootaert, 2005: p. 367) collected mainly by one of us (P.G.) during two successive field trips (2002 and 2003) and also found in the collection of ZRC. Key to species of the Empis (Coptophlebia) hyalea- group from Singapore and Southern Malaysia Material examined. Ð Holotype - male, Singapore, Pulau [= Island] (males only) Ubin, Chek Jawa, sample 22059, coll. P. Grootaert, 11 Dec.2002 (ZRC). 1. Male fore tibia with several anterodorsal spine-like bristles on basal half ...... 2 Paratypes Ð 3 males, with the same data; 1 male, Pulau [= Island] ÐMale fore tibia without anterodorsal spine-like bristles on basal Ubin, Chek Jawa, mangrove, sample 22061, coll. P. Grootaert, 13 half...... 3 Dec.2002 (KBIN-IRScNB); 1 male, Singapore, Pulau [= Island] 2. Male fore tibia with anterodorsal spine-like bristles not longer Ubin, Chek Jawa, sample 23080, coll. P. Grootaert, 19 Nov.2003 than tibia depth, first fore tarsomere not elongated (Fig. 1A), (KBIN-IRScNB); 1 male, Singapore, Pulau [= Island] Ubin, Chek wing pale with median veins almost not visible, epandrium not Jawa, malaise trap sample 23097, coll. P. Grootaert., 2 Nov.2003 projected posterodorsally, without a dorsal row of very strong, (MNHN); 1 male, Singapore, NUS campus, near woods King long bristles (Fig. 2) ...... Edward VII Hall, sample 23093, coll. P. Grootaert, 30 Nov.2003 ...... neesoonensis (Singapore: Nee Soon: swamp forest) (MNHN); 1 male, Singapore, Nee Soon Forest, on Vitis flower, coll. ÐMale fore tibia with row of spine-like anterodorsal bristles D. H. Murphy, 8 Nov.1979 (ZRC). ending with distinctly longer bristles, first fore tarsomere almost as long as the next four tarsomeres together (Fig. 1B, 3), median Diagnosis. Ð Blackish species of middle size with clear wing, veins visible, epandrium projected posterodorsally, with a dorsal labrum more than twice head height, fore tibia strongly row of very strong, long bristles (Fig. 4) ...... deformed at base, without anterodorsal spine-like bristles on ..... producta (Singapore: Pulau Ubin, Chek Jawa: mangrove) basal half, wing rather clear with all veins well visible, sternite 3. Male fore tibia without tubercle ...... 4 8 with several long posterior bristles, epandrium indented Ð Male fore tibia with an anterodorsal tubercle at basal quarter (Fig. 1D) ...... verruca (Singapore: Nee Soon: swamp forest) ventrally, with 1 very strong bristle at tip, numerous strong
Fig. 7. Male hypopygium of E. (C.) triseta, lateral view. Scale = 4.2 mm.
218 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Fig. 8. Male hypopygium of E. (C.) verruca, lateral view. Scale = 4.2 mm.
219 Daugeron & Grootaert: New empidine dance flies
4. Hind tibia without a cluster of 4 very long basoventral bristles, LITERATURE CITED sternite 8 and epandrium with normal or only a few tortuous bristles ...... 5 Cumming, J. M., B. J. Sinclair & D. M. Wood, 1995. Homology ÐHind tibia with a cluster of 4 very long basoventral bristles, and phylogenetic implications of male genitalia in Diptera- sternite 8 and epandrium with numerous very long, more or less Eremoneura. Entomologica Scandinavica, 26: 120-151. tortuous bristles (Figs. 6A, B) ...... tortuosa (Singapore: Nee Soon: swamp forest) Daugeron, C., 1997. Découverte du sous-genre Xanthempis Bezzi 5. Species of small size, fore tibia not distinctly deformed (in lateral en Afrique du Nord et description de trois espèces nouvelles view) with 3 distinct dorsal bristles (at middle, at apical third, (Diptera: Empididae). Annales de la Société entomologique de at apical quarter) ...... France (N.S.), 33: 155-164...... triseta (Malaysia: Kota Tinggi: rain forest) Daugeron, C., 2002. Monophyly of the Empis (Coptophlebia) Ð Species of larger size, fore tibia distinctly deformed at base (in hyalea-group with description of a new species from New lateral view) with 1 distinct dorsal bristle at middle...... Caledonia (Diptera, Empididae, Empidinae). In: Najt, J. & P. .. vitisalutatoris (Singapore: Nee Soon: swamp forest; Pulau Grandcolas (eds.), Zoologica Neocaledonica 5. Mémoires du Ubin, Chek Jawa: rain forest) Muséum national d’Histoire naturelle, 187: 33-38. Daugeron, C. & P. Grootaert, 2003a. The Empis (Coptophlebia) hyalea-group from Thailand, with a discussion on the world ACKNOWLEDGEMENTS distribution of this species-group (Diptera: Empididae: Empidinae). European Journal of Entomology, 100: 167-179. This research was partially supported through a European Daugeron, C. & P. Grootaert, 2003b. New species of the Empis Community Marie Curie Fellowship (grant HPMF-CT-2000- (Coptophlebia) hyalea-group (Diptera: Empididae: Empidinae) 00718) and a Research Project Action 1 (Belgian Federal from Guangdong province in China. Bulletin de l’Institut Royal Office for Scientific, Technical and Cultural Affairs) (C.D.). des Sciences Naturelles de Belgique, Entomologie, 73: 57-66. We thank M. Leclercq (KBIN-IRScNB) and G. Hodebert Daugeron, C. & P. Grootaert, 2005. Phylogenetic systematics of (MNHN) for drawing the habitus of E. producta and the male the Empis (Coptophlebia) hyalea-group (Insecta: Diptera: fore legs respectively. The second author thanks P. Ng, Darren Empididae). Zoological Journal of the Linnean Society, 145: Yeo and Yang Chang Man for their hearty hospitality at the 339-391. National University of Singapore. The research of the second McAlpine, J. F., 1981. Morphology and terminology - adults. author was possible thanks to a grant under the Raffles Chapter 2. In: McAlpine J. F. (ed.), Manual of Nearctic Diptera. Museum Fellowship Scheme in 2002 and 2003. Vol. 1. Agriculture Canada, monograph 27. Agriculture Canada, Ottawa. Pp. 9-63. Sinclair, B. J., J. M. Cumming & D. M. Wood, 1994. Homology and phylogenetic implications of male genitalia in Diptera - Lower Brachycera. Entomologica Scandinavica, 24: 407-432.
220 THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 221-229 Date of Publication: 31 Dec.2005 © National University of Singapore
FOUR NEW AND ONE NEWLY RECORDED SPECIES OF THE JUMPING SPIDERS (ARANEAE: SALTICIDAE: LYSSOMANINAE & SPARTAEINAE) FROM (SUB)TROPICAL CHINA
J. X. Zhang College of Life Sciences, Hebei University, Baoding, Hebei 071002, China Department of Zoology, 6270 University Boulevard, University of British Columbia, Vancouver, BC V6T 1Z4, Canada Email: [email protected]
Daiqin Li Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543 College of Life Sciences, Hubei University, Wuhan 430062, Hubei Province, People’s Republic of China Email: [email protected] (Corresponding author)
ABSTRACT. – Five species of jumping spiders belonging to four genera of subfamilies Lyssomaninae (Onomastus) and Spartaeinae (Phaeacius, Portia and Yaginumanis) from China were diagnosed, described and figured. These widely distributed genera range from subtropical China to tropical Southeast Asia. Four species were found to be new to science, Onomastus nigrimaculatus (female, male), Phaeacius yixin (male), Portia taiwanica (female, male) and Yaginumanis wanlessi (female); the species Phaeacius malayensis Wanless, 1981 (female, male), is newly recorded from China.
KEY WORDS. – Salticidae, Lyssomaninae, Spartaeinae, spiders, new species, Southeast Asia; China.
INTRODUCTION The Spartaeinae, defined by Wanless (1984a) who erected six new genera and revised most genera of this subfamily The family Salticidae, commonly called jumping spiders, is (Wanless, 1978, 1979, 1981a, b, 1984a, b, 1987), is now the largest and most diverse spider group, currently containing composed of 14 existing and five fossil genera. more than 5000 described species in 550 genera (Platnick, Approximately, 93 species of the existing genera are now 2005). Among them, about 370 species in 89 genera are recorded from the Old World, of which 18 species in eight reported from China. genera are known from Chinese fauna (Platnick, 2005). Members of this subfamily are diagnosed by the presence of The Lyssomaninae and Spartaeinae are usually considered the palpal tegular furrow in male. Being similar to as two singular subfamilies because they lack many derived lyssomanines, the spartaeines usually have large posterior features that mark the bulk of the Salticidae (Maddison, 1995). median eyes (PME), but they have been reduced in four The Lyssomaninae was even proposed as a separate family, genera of Lyssomaninae (Lyssomanes, Chinoscopus, the Lyssomanidae, by Blackwall (1877). After examining the Pandisus and Onomastus) and two genera of Spartaeinae morphological characters, Galiano (1976) redefined this group (Cyrba and Gelotia). and concluded that they merit no more than subfamilial rank within the Salticidae. The Lyssomaninae currently contains The present paper contains partial results of a taxonomic seven genera and 125 species, most of which have been study, which aims to investigate biodiversity of spiders in recently revised (Galiano, 1980, 1998; Wanless, 1980a, b, subtropical and tropical China and Southeast Asia. Here, we c). Five genera are known from the old world (Asemonea O. describe four new species and one new record of the P-Cambridge, Goleba Wanless, Macopaeus Simon, subfamilies Lyssomaninae and Spartaeinae. Onomastus Simon, and Pandisus Simon), of which only Asemonea has been reported from China. Members of this subfamily are usually adapted to an arboreal existence MATERIALS AND METHODS associated with green leaves. Although they are commonly distinguished from ordinary salticids by two diagnostic All measurements given in this paper are in millimetres. Palp characters: eyes in four rows and the tracheal system confined measurements are shown as: total length to the abdomen, Wanless (1980c) pointed out that the (femur+patella+tibia+tarsus). Leg measurements are shown subfamily is actually polyphyletic.
221 Zhang & Li: New species of jumping spiders from China as: total length (femur+patella+tibia+metatarsus+tarsus). Diagnosis. – The new species resembles Onomastus Dissected epigynes were digested in lactic acid for 10-30 min. complexipalpis Wanless, 1980 (Wanless, 1980b: 187, Fig. 4) or in 10% KOH for approximately 24 h at room temperature, in the shape of male palp, but can be distinguished from the rinsed in distilled water, stained in ethanol solution of latter in abdomen with pairs of dark spots dorsally (Fig. 1A); chlorazol black E under control and mounted in glycerin. median apophysis of male palp wide in basal part, with a Specimens including types are deposited in the Raffles middle curved flange, and spur of embolic guide shorter and Museum of Biodiversity Research (formerly Zoological wider (Fig. 1B-D). It is also similar to Onomastus nigricaudus Reference Collection, or ZRC), National University of Simon, 1900 (Wanless, 1980b: 181, fig. 1A-G) in male palp Singapore; Museum of Hebei University, Hebei, China with a femoral apophysis, but differs from the latter in (MHBU); and National Museum of Natural History, abdomen with pairs of dark spots dorsally (Fig. 1A); median Taichong, Taiwan (NMNH). apophysis of male palp not branched, with distal part thin and dagger shaped, spur of embolic guide wide (Fig. 1B-D); The abbreviations used are: ALE – anterior lateral eyes, AME female epigynum with a median septum which widened – anterior median eyes, PLE – posterior lateral eyes, PME – posteriorly, and spermatheca connected with an oval structure posterior median eyes. anteriorly, which is doubted to be the copulatory duct (Fig. 1E, F). It can also be distinguished from Onomastus kanoi Ono, 1995 (Ono, 1995: 157, figs. 16-21) by the long and thin TAXONOMY distal part of median apophysis, smaller embolic guide, larger spur of embolic guide and blunt femoral apophysis of male Onomastus Simon, 1900 palp (Fig. 1B-D).
Onomastus Simon, 1900: 29 (Type species: Onomastus nigricaudus Description. – Males. Total length 4.25-4.70. Length of Simon, 1900, by original designation); Wanless, 1980b: 179. holotype 4.60: cephalothorax 2.20 long, 1.45 wide, 1.05 high; abdomen 2.25 long, 0.90 wide. Carapace (Fig. 1A) pale Diagnosis. – The genus can be easily distinguished from other yellow, eye area grayish. Fovea indistinct. Eye sizes: AME genera of the subfamily Lyssomaninae by fovea indistinct, 0.50, ALE 0.30, PME 0.03, PLE 0.25. First eye row 1.00 posterior median eyes minute, male palpal organ with a wide, second row 1.45 wide, third eye row 1.10 wide, fourth subtegular apophysis (labelled as “y” by Wanless, 1980b), eye row 0.96 wide, eye area 1.09 long. Clypeus height 0.16. and female lacking epigynal ducts. Chelicerae, endites, labium and sternum pale yellow. Chelicerae with 5 teeth on promargin and 8-10 denticles on Remarks. – Members of this genus are mainly distributed in retromargin. Endites, labium and sternum yellowish. Coxae Japan, South and Southeast Asia. Up till now, six species and trochanters of legs yellowish, other segments deeper in have been recorded in the world (Platnick, 2005), not colour. Legs with some long spines and hairs. Spination of including the new species described in this paper. This genus leg I: metatarsi v 4-2-0, p 1-0-0, r 1-0-0; tibiae v 4-4-4, p 1- is first reported from China. 0-0, r 1-0-0, d 1-1-0; patellae v 0-1-0, p 0-1-0, r 0-1-0; femora p 0-0-1, r 0-0-1, d 1-1-1. Measurements of palp and legs: palp 2.50 (0.80+0.38+0.28+1.04); I 6.03 (1.65+2.45+1.30+0.63), II 5.27 (1.50+2.08 +1.16+0.53), III Onomastus nigrimaculatus, new species 5.55 (1.60+1.75+1.60+0.60), IV 6.90 (1.90+2.20+2.05+0.75). (Fig. 1A-F) Leg formula: 4132. Abdomen long oval and white yellow, with some short hairs; dorsum with two pairs of dark patches. ° Holotype. – Male (ZRC•ARA•494), Mingfeng Valley (18 44.635'N Subtegular apophysis (y) of palpal organ (Fig. 1B-D) long 108°50.667'E), Jianfengling National Park, Hainan Province, China, and finger shaped; median apophysis large with its distal part coll. D. Li, 15 Dec.2003. long and dagger-like; embolic guide (eg) terminating with a Paratypes. – 1 male (ZRC•ARA•495), same data as holotype; 1 large spur (s). male (ZRC•ARA•496)(OS 001/LD 2003), Mingfeng Valley (18°44.635'N 108°50.667'E), Jianfengling National Park, Hainan Females. Total length 4.60-4.72. A female total length 4.60: Province, China, 15 Dec.2003; 1 male (ZRC•ARA•497)(OS 022/ cephalothorax 2.05 long, 1.30 wide, 1.15 high; abdomen 2.65 LD 2003), Mingfeng Valley (18°44.635'N 108°50.667'E), long, 1.00 wide. Eye sizes: AME 0.50, ALE 0.30, PME 0.04, Jianfengling National Park, Hainan Province, China, 15 Dec.2003; PLE 0.22. First eye row 1.00 wide, second row 1.43 wide, 1 female (ZRC•ARA•498)(OS 008/LD 2003), Mingfeng Valley third eye row 1.04 wide, fourth eye row 0.95 wide, eye area ° ° (18 43.016'N 108 50.228'E), Jianfengling National Park, Hainan 1.14 long. Clypeus height 0.08. Retrolateral surface of distal Province, China, 14 Dec.2003; 1 female (ZRC•ARA•499)(OS 033/ femur I, prolateral and retrolateral surface of distal tibiae with LD 2003), Mingfeng Valley (18°44.635'N 108°50.667'E), a black patch respectively. Spination of leg I: metatarsi v 4- Jianfengling National Park, Hainan Province, China, 15 Dec.2003; 1 female (ZRC•ARA•500)(OS 009/LD 2003), Tian Chi 2-0, p 1-0-0, r 1-0-0; tibiae v 4-4-4, p 1-0-0, r 1-0-0, d 0-1- (18°43.982'N 108°52.186'E), Jianfengling National Park, Hainan 0; patellae v 0-1-0, r 0-1-0; femora p 0-0-1, r 0-0-1, d 1-1- Province, China, 13 Dec.2003; 2 females and 2 males 1. Measurements of legs: I 6.45 (1.85+2.75+1.30+0.55), II (ZRC•ARA•501)(ONO 002/LD 2004), Mengla Natural Reserve 5.45 (1.65+2.10+1.20+0.50), III 5.50 (1.55+1.80+1.60+0.55), (21°37.63'N 101°35.22'E), Pubang, Mengla County, Yunnan IV (1.85+2.20+2.05+0.60). Leg formula: 4132. Other Province, China, coll. D. Li, 7 May.2004. characters similar to those of male. Median septum of
222 THE RAFFLES BULLETIN OF ZOOLOGY 2005 epigynum (Fig. 1E, F) widened posteriorly, spermatheca Remarks. – A small salticid genus containing 11 species from small. South and Southeast Asia (Prószynski,´ 2003).
Etymology. – The specific name is from the Latin Phaeacius malayensis Wanless, 1981 “nigri+maculatus”, and refers to the dorsal dark patches on (Fig. 2A-E) abdomen. Phaeacius malayensis Wanless, 1981a: 205, Figs. 6A-E, 7A-C.
Phaeacius Simon, 1900 Materials examined. – 1 female and 1 male, Xishuangbanna Tropical Botanic Garden (ZRC•ARA•502)(21°55.85'N 101°15.25'E), Phaeacius Simon, 1900: 32 (Type species: Phaeacius fimbriatus Menglun, Mengla County, Yunnan, coll. D. Li, 7 May.2004. Simon, 1900, by original designation); Wanless, 1981a: 199; 1984a, 190; Wijesinghe, 1991: 249. Diagnosis. – This species is very similar to Phaeacius fimbriatus Simon, 1900 (Wanless, 1981a: 202, Figs. 4A-H, Diagnosis. – This genus can be easily distinguished from the 5B), but differs from the latter in male palp with a pronounced closely related genus Portia by the somewhat flattened body; apophysis lying between the ventral and retrolateral massive retrolateral tibial apophysis of male palp, filamentous apophyses (Fig. 2B, C), and median septum of female secondary conductor (sc, labeled as “M1” in Wanless, 1984a); epigynum narrower (Fig. 2D). and the shape of female epigynum.
Fig. 1. Onomastus nigrimaculatus, new species. A, male; B, left palp, prolateral view (the black arrow refers to the median apophysis, and the white arrow refers to the subtegular apophysis); C, same, retrolateral view; D, same, ventral view (the black arrow refers to the spur of embolic guide, and the white arrow refers to the embolic guide); E, epigynum, ventral view; F, same, dorsal view. Scales: A = 1.0 mm, B-D = 0.2 mm, E, F = 0.1 mm.
223 Zhang & Li: New species of jumping spiders from China
Description. – Male. Total length 9.00: cephalothorax 4.50 pronounced apophysis present between the ventral and long, 3.50 wide, 1.70 high; abdomen 4.50 long, 2.40 wide. retrolateral apophyses. Carapace (Fig. 2A) deep brown, red brown around fovea, with many fine hairs. Eye sizes: AME 0.56, ALE 0.31, PME Female. Total length 8.60: cephalothorax 4.10 long, 3.15 0.25, PLE 0.34. First eye row 2.03 wide, second row 1.88 wide, 1.60 high; abdomen 4.50 long, 2.20 wide. Eye sizes: wide, third eye row 2.03, eye area 2.00 long. Clypeus height AME 0.50, ALE 0.31, PME 0.22, PLE 0.31. First eye row 0.13. Chelicerae red brown, with 3 promarginal and 4 1.97 wide, second row 1.75 wide, third eye row 1.84 wide, retromarginal teeth. Endites and labium deep red brown. eye area 1.78 long. Clypeus height 0.09. Spination of leg Sternum yellow brown, with indistinct gray patches. Legs I: metatarsi v 2-0-0, p 1-1-1, r 1-1-0; tibiae v 2-3-2, p 1-1- yellow brown, with a few dark markings. Spination of leg 0, r 1-1-0, d 1-1-0; patellae p 0-1-0, r 0-1-0; femora p 0-1- I: metatarsi v 2-0-0, p 1-1-0, r 1-1-2, d 0-1-0; tibiae v 2-3- 1, d 0-1-4. Measurements of legs: I 9.39 2, p 1-1-0, r 1-1-0, d 1-1-1; patellae p 0-1-0, r 0-1-0; femora (2.69+3.63+2.13+0.94), II 9.69 (2.75+3.69+2.44+0.81), III p 0-1-1, r 0-0-1, d 0-2-2. Measurements of palp and legs: 9.38 (2.75+3.25+2.50+0.88), IV 10.69 palp 4.38 (1.41+0.53+0.47+1.97); I 10.60 (2.94+3.75+3.06+0.94). Leg formula: 4213. Other characters (2.97+4.13+2.44+1.06), II 10.76 (3.19+3.95+2.52+1.10), III similar to those of male. Median septum of epigynum (Fig. 10.72 (3.05+3.75+2.84+1.08), IV 12.16 2D, E) narrow in the middle and widened at the posterior (3.40+4.16+3.56+1.04). Leg formula: 4231. Abdomen oval, part, spermatheca almost oval, accompanied with a scattered with lots of short hairs; dorsum earthy yellow, black membranous sac. marginally, with a brownish patch at the centre; venter black brown. Embolus of palpal organ (Fig. 2B, C) long and robust, Distribution. – China (Yunnan; new record), Malaysia, secondary conductor (sc) translucent long and slender, a Singapore, Sumatra.
Fig. 2. Phaeacius malayensis Wanless, 1981. A, male; B, left palp, ventral view (the black arrow refers to the secondary conductor); C, same, lateral view (the black arrow refers to the pronounced apophysis between the ventral and retrolateral apophyses); D, epigynum, ventral view; E, same, dorsal view. Scales: A = 1.0 mm, B-E = 0.2 mm.
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Phaeacius yixin, new species retrolateral tibial apophysis of male palp and the presence of (Fig. 3A-E) pronounced apophysis between ventral and retrolateral apophyses (Fig. 3D, E). Materials examined. – Holotype: Male (ZRC•ARA•503), Institute of Tropical Forest (18°41.890'N 108°47.373'E), Jianfengling, Description. – Males. Length of holotype 8.64: Hainan, China, coll. Y. X. Li, 13 Dec.2003. cephalothorax 4.24 long, 3.20 wide, 1.76 high; abdomen 4.40 long, 2.48 wide. Carapace (Fig. 3A) deep brown, red brown Non-types: 2 female juveniles (ZRC•ARA•504), same data as behind eyes region and earthy yellow laterally, with many holotype; 1 male juvenile (ZRC•ARA•505), Institute of Tropical Forest (18°41.890'N 108°47.373'E), Jianfengling, Hainan, China, short fine hairs. Eye sizes: AME 0.59, ALE 0.31, PME 0.22, coll. D. Li, 12 Dec.2003. PLE 0.34. First eye row 2.12 wide, second row 1.88 wide, third eye row 2.04 wide, eye area 1.97 long. Clypeus height Diagnosis. – The new species resembles Phaeacius 0.09. Chelicerae (Fig. C) red brown, with 3 promarginal and malayensis Wanless, 1981, but can be easily distinguished 5 retromarginal teeth. Endites and labium red brown. from the latter by the pronounced apophysis between the Sternum and legs yellow. Legs with a few dorsal dark stripes ventral and retrolateral apophyses almost triangular, and the and many long spines. Spination of leg I: metatarsi v 2-0- different shape of retrolateral tibial apophysis (Fig. 3D, E). 0, p 1-1-0, r 1-1-2, d 0-1-0; tibiae v 2-3-2, p 1-0-1, r 1-0-1, It differs from Phaeacius yunnanensis Peng & Kim, 1998 d 1-1-1; patellae p 1-0-0, r 1-0-0; femora p 0-1-1, r 0-0-1, d (Peng & Kim, 1998: 411-413, Figs. 1-3) in the shape of 0-2-2. Measurements of palp and legs: palp 3.96
Fig. 3. Phaeacius yixin, new species. A, male; B, carapace, lateral view; C, left chelicera, retrolateral view; D, left palp, ventral view (the black arrow refers to the pronounced apophysis between the ventral and retrolateral apophyses); E, same, lateral view. Scales: A, B = 1.0 mm, C-E = 0.2 mm.
225 Zhang & Li: New species of jumping spiders from China
(1.28+0.56+0.40+1.72); I 9.57 (2.63+3.63+2.25+1.06), II Diagnosis. – Spiders of this genus can be easily distinguished 9.94 (2.88+3.75+2.31+1.00), III 9.81 (2.81+3.50+2.50+1.00), from others of the subfamily Spartaeinae by the presence of IV 11.20 (3.13+3.88+3.13+1.06). Leg formula: 4231. developed tufts on dorsal abdomen, fan-like fringes on legs, Abdomen oval, covered with many fine hairs; dorsum earthy and dorsal flange on male palpal cymbium. yellow, with lateral margins black, centre with a brownish foliform patch; venter gray brown, black in front of genital Remarks. – A total of 16 species have been reported (Platnick, groove, with two rows of muscle attachment spots from 2005), being distributed in the Oriental and Ethiopia regions. genital groove to spinnerets. Ventral tibial apophysis of male palp (Fig. 3D, E) robust and round, retrolateral apophysis long with a median process, pronounced apophysis between Portia taiwanica, new species ventral and retrolateral apophysis almost triangular. (Fig. 4A-G)
Etymology. – The specific name is a noun in apposition taken Materials examined. – Holotype: Male (ZRC•ARA•506), Kenting from the first name of the collector Y. X. Li, who was only National Park, Taiwan, coll. D. Li, 8 Jun.2001. 11 years old when he first collected the type specimen. Paratypes: 1 female (NMNH-Ar-95-00339), Taiwan Phoenix valley Bird Park, coll. J. D. Lee, 27 Sep.1995; 1 female (NMNH-Ar-95- 00214), Mt. Lanran, Taiwan, coll. J. D. Lee, 26 Jun.1995. Portia Karsch, 1878 Diagnosis. – The new species resembles Portia assamensis Portia Karsch, 1878: 774 (Type species: Portia schultzi Karsch, Wanless, 1978 (Wanless, 1978: 105, Figs. 10D-F, 11D-F) in 1878, by original designation and monotypy); Wanless, 1978: the shapes of male palp and female epigynum, but differs 84; 1984a: 191; Peng & Li, 2002b: 255. from the latter in the membrane at the base of embolus
Fig. 4. Portia taiwanica, new species. A, male; B, left palp, ventral view; C, same, retrolateral view; D, same, dorsal view; E, cheliceral teeth of female, retrolateral view; F, epigynum, ventral view; G, same, dorsal view. Scales: A = 1.0 mm, B-G = 0.2 mm.
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(labeled as “M1” in Wanless, 1984a) large, embolus shorter, Remarks. – Chang & Tso (2004) reported the species Portia retrolateral tibial apophysis finger shaped (Fig. 4B), and the fimbriata (Doleschall, 1859) from Taiwan. But judging from posterior depression of female epigynum with oblique lines the illustrations they provided, we believe that Chang & Tso’s and thickened border (Fig. 4F). It is also similar to Portia specimens are probably Portia taiwanica, new species, rather fimbriata (Doleschall, 1859) (Wanless, 1978: 99, Figs. 7A- than P. fimbriata. G, 8A-F) in the shape of male palp, but can be distinguished from the latter by female clypeus with white beards; embolus Etymology. – The specific name refers to the type locality. of male palp shorter, retrolateral tibial apophysis thin and finger-like (Fig. 4B). It also differs from Portia quei Zabka, 1985 (Zabka, 1985: 438, Figs. 497-501; Peng et al., 1993: Yaginumanis Wanless, 1984 188, Figs. 660-666) in the shorter embolus and retrolateral tibial apophysis of male palp (Fig. 4B), and the shape of Yaginumanis Wanless, 1984a: 152 (Type species: Boethus posterior depression of female epigynum (Fig. 4F). sexdentatus Yaginuma, 1967, by original designation and monotypy). Description. – Males. Length of holotype 9.00: cephalothorax 4.40 long, 3.44 wide, 2.72 high; abdomen 4.56 Diagnosis. – This genus is similar to Spartaeus in abdominal long, 2.08 wide. Carapace (Fig. 4A) deep red brown, with pattern, but can be distinguished from the latter in: legs robust grayish lateral and mesal bands composed of short hairs. and with less ventral spines on tibia I; distal modification of Clypeus lacking white beards. Eye sizes: AME 0.88, ALE the male palpal tegulum (labelled as “M3”by Wanless, 1984a) 0.40, PME 0.28, PLE 0.40. First eye row 2.60 wide, second with a pleated region, retrolateral apophysis of male palp with row 2.28 wide, third eye row 2.36 wide, eye area 1.94 long. a strong lobe. Clypeus height 0.50. Chelicerae red brown, with 3 promarginal and 4 retromarginal teeth. Endites and labium Remarks. – A small genus represented by only two species deep red brown, with distal margins in lighter colour. Sternum (Prószynski, 2003), distributed in China and Japan. Judging yellow brown, with darker margins, covered with numerous from the eyes pattern and the shape of eyigynum described white fine hairs and a few black long setae. Legs deep brown, by Peng & Li (2002a), Yaginumanis cheni Peng & Li, 2002 with many spines. Tibiae and patellae with ventral fringes, from Guangxi, China, might belong to a genus of the tibiae III and IV with short dorsal fringes. Measurements of subfamily Lyssomaninae. palp and legs: palp 4.04 (1.40+0.52+0.48+1.64); I 19.60 (4.64+6.80+6.08+2.08), II 12.80 (3.52+4.48+3.52+1.28), III 10.72 (3.20+3.52+2.96+1.04), IV 16.56 Yaginumanis wanlessi, new species (4.32+5.20+5.76+1.28). Leg formula: 1423. Abdomen oval; (Fig. 5A-E) dorsum yellow brown, with black lateral margins and some indistinct patches; venter deep brown, with a dark wide band Materials examined. – Holotype: Female (MHBU-Ar-46), Qiankun behind the epigynal groove. Retrolateral tibial apophysis of (28°37'N 106°16'E), Zihuai Town, Hejiang County, Luzhou City, male palp (Fig. 4B-D) finger shaped; cymbium with a Sichuan Province, coll. J. X. Zhang, 30 Jul.2003. prolateral flange; embolus long and slender. Paratype: 1 female (ZRC•ARA•507), Dagou Natural Reserve (32°367'N 105°13'E), Qingchuan County, Guangyuan City, Sichuan Females. Total length 9.50-9.75. A female total length 9.75: Province, coll. J. X. Zhang, 13 Jul.2003. cephalothorax 4.50 long, 3.75 wide, 2.38 high; abdomen 5.25 long, 2.75 wide. Carapace red brown, with a few indistinct Diagnosis. – This new species resembles Yaginumanis brown stripes. Clypeus with lots of white beards. Eye sizes: sexdentatus (Yaginuma, 1967) (Wanless, 1984a: 153, Fig. 6A- AME 0.88, ALE 0.41, PME 0.28, PLE 0.41. First eye row J), but can be distinguished from the latter by: posterior 2.69 wide, second row 2.50 wide, third eye row 2.63 wide, depression of the epigynum with dark surroundings, eye area 2.28 long. Clypeus height 0.53. Palps with many copulatory openings at the lateral sides of posterior white hairs. Chelicerae (Fig. 4E) red brown, promargin with depression, and posterior portion of the spermathecae larger 3 teeth, retromargin with 4 small teeth and 2 denticles. than that of the latter (Fig. 5D, E). Measurements of legs: I 14.19 (3.88+5.00+3.75+1.56), II 11.76 (3.38+4.00+3.00+1.38), III 10.77 Description. – Female. Total length 5.88-7.25. Holotype (3.13+3.63+2.88+1.13), IV 16.64 (4.38+5.38+5.50+1.38). total length 7.25: cephalothorax 3.06 long, 2.16 wide, 1.62 Leg formula: 4123. Dorsum of abdomen deep brown, with high; abdomen 4.32 long, 2.70 wide. Carapace (Fig. 5A) three yellow brown patches composed of long hairs, posterior black brown, eye area darker in color with many white fine part with a mesal rectangular grayish patch, with its centre hairs and several black long setae, the area behind PLSs earthy dark brown; venter yellow brown, with two pairs of deep yellow. Eye sizes: AME 0.57, ALE 0.30, PME 0.22, PLE brown patches behind the epigynal groove. Other characters 0.29. Anterior eye row 1.80 wide, median row 1.58 wide, similar to those of male. Epigynum (Fig. 4F-G) weakly posterior row 1.67 wide, eye area 1.43 long, clypeus height sclerotized, the posterior depression with dark margins; 0.13. Chelicerae (Fig. 5C) red brown, with a brown patch spermatheca large and round. in the front surface, promargin with 3 teeth and retromargin
227 Zhang & Li: New species of jumping spiders from China with 5 teeth. Endites and labium red brown, with distal inner generous assistance was provided by Xishuangbanna Tropical margins of endites and tip of labium white. Sternum yellow, Botanic Garden (XTBG), Chinese Academy of Sciences in with darker margins. Spination of leg I: metatarsi v 2-2-2; China. Special thanks are extended to Hongmao Liu, Yuping tibiae v 2-2-2, p 0-1-1; patellae p 0-1-0; femora d 0-2-4. Leg She, Liming Li, Jian Zhang, Jinghua Mao and Zhihua Li from measurements: I 5.95 (1.80+2.30+1.17+0.68), II 5.95 XTBG for generous assistance to the second author’s study (1.85+2.16 +1.22+0.72), III 6.30 (1.80+2.16+1.53+ 0.81), IV leave in XTBG without which this study would not have been 8.15 (2.25+2.75+2.25+0.90). Leg formula: 431=2. Dorsum possible. Special thanks also go Mingsheng Zhu and Maobin of abdomen with an indistinct brownish band in the middle Gu for assistance with the expedition to Hainan in December of anterior part, lateral sides brown, with some indistinct 2003 and also for assistance with field work in Hainan, China. stripes anteriorly; venter pale yellow, scattered with some Shyh-min Chao from National Museum of Natural Science brown patches. Epigynum (Fig. D, E) with an oval depression in Taiwan is grateful for giving access to the collection. We at the back part; copulatory duct short, spermatheca large and also wish to thank Shuqiang Li for assistance with export bean-shaped. permits from China. Import permits were provided by the CITES, Agri-food & Veterinary Authority of Singapore. Etymology. – The specific name is a patronym in honor of F. R. Wanless, who has made great contributions to the study of jumping spiders. LITERATURE CITED
Blackwall, J., 1877. A list of spiders captured in the Seychelle Islands ACKNOWLEDGEMENTS by Professor E. Perceval Wright, M. D., F. L. S.; with descriptions of species supposed to be new to arachnologists. This study was in part supported by grants from National Proceedings of the Royal Irish Academy, (2) 3: 1-22. University of Singapore ARC to D. Li (R-154-000-060-112 Chang, Y. H. & I. M. Tso, 2004. Six newly recorded spiders of the and R-000-072-112), NSFC to D. Li (30470229), and in part genera Araneus, Larinia, Eriophora, Thanatus, Portia and by NSFC to Mingsheng Zhu (30170118). Valuable and Dolichognatha (Araneae: Araneidae, Philodromidae, Salticidae
Fig. 5. Yaginumanis wanlessi, new species. A, female; B, carapace, lateral view; C, left chelicera, retrolateral view; D, epigynum, ventral view; E, same, dorsal view. Scales: A, B = 1.0 mm, C-E = 0.2 mm.
228 THE RAFFLES BULLETIN OF ZOOLOGY 2005
and Tetragnathidae) from Taiwan. Acta Arachnologica, Tokyo, Wanless, F. R., 1978. A revision of the spider genus Portia (Araneae: 53: 27-33. Salticidae). Bulletin of the British Museum (Natural History), Galiano, M. E., 1976. Comentarios sobre la categoria sistematica 34: 83-124. del taxon Lyssomanidae (Araneae). Revista del Museo Argention Wanless, F. R., 1979. A revision of the spider genus Brettus de Ciencias Naturales Bernardino Rivadavia (Ent.), 5: 59-70. (Araneae: Salticidae). Bulletin of the British Museum (Natural Galiano, M. E., 1980. Revision del genero Lyssomanes Hentz, 1845 History), 35: 183-190. (Araneae, Salticidae). Opera Lilloana, 30: 1-104. Wanless, F. R., 1980a. A revision of the spider genus Macopaeus Galiano, M. E., 1998. Revision of the genus Chinoscopus (Araneae, (Araneae, Salticidae). Bulletin of the British Museum (Natural Salticidae, Lyssomanidae). Bulletin of British Arachnological History), 38: 219-223. Society, 11: 1-9. Wanless, F. R., 1980b. A revision of the spider genus Onomastus Karsch, F., 1878. Exotisch-araneologisches. Zeitschrift für die (Araneae, Salticidae). Bulletin of the British Museum (Natural gesamten Naturwissenschaften, 51: 332-333, 771-826. History), 39: 179-188. Maddison, W., 1995. The Tree of Life Web Project: Salticidae. URL: Wanless, F. R., 1980c. A revision of the spider genera Asemonea http://tolweb.org/tree/phylogeny.html and Pandisus (Araneae: Salticidae). Bulletin of the British Museum (Natural History), 39: 213-257. Ono, H., 1995. Four East Asian spiders of the families Eresidae, Araneidae, Thomisidae and Salticidae (Arachnida, Araneae). Wanless, F. R., 1981a. A revision of the spider genus Phaecius Bulletin of the National Science Museum Tokyo (A), 21: 157- (Araneae: Salticidae). Bulletin of the British Museum (Natural 169. History), 41: 199-212. Peng, X. J. & S. Q. Li, 2002a. One new species of the genus Wanless, F. R., 1981b. A revision of the spider genus Cocalus Yaginumanis from Mt. Shiwandashan, Guangxi, China (Araneae: (Araneae: Salticidae). Bulletin of the British Museum (Natural Salticidae). Acta Zootaxonomica Sinica, 27: 238-240. History), 41: 253-261. Peng, X. J. & S. Q. Li, 2002b. Chinese species of the jumping Wanless, F. R., 1984a. A review of the spider subfamily Spartaeinae spider genus Portia Karsch (Araneae: Salticidae). Pan-Pacific nom. n. (Araneae: Salticidae) with descriptions of six new Entomologist, 78: 255-264. genera. Bulletin of the British Museum (Natural History), 46: 135-205. Peng, X. J. & J. P. Kim, 1998. Four species of jumping spiders (Araneae: Salticidae) from China. Korean Journal of Biological Wanless, F. R., 1984b. A revision of the spider genus Cyrba Sciences, 2: 411-414. (Araneae: Salticidae) with the description of a new presumptive pheromone dispersing organ. Bulletin of the British Museum Peng, X. J., L. P. Xie, X. Q. Xiao & C. M. Yin, 1993. Salticids in (Natural History), 47: 445-481. China (Arachnida: Araneae). Hunan Normal University Press, Changsha, 270 pp. Wanless, F. R., 1987. Notes on spiders of the family Salticidae. 1. The genera Spartaeus, Mintonia and Taraxella. Bulletin of the Platnick, N. I., 2005. The World Spider Catalog, version 5.5. British Museum (Natural History), 52: 107-137. American Museum of Natural History. URL: http:// research.amnh.org/entomology/spiders/catalog81-87/index.html Wijesinghe, D. P., 1991. New species of Phaeacius from Sri Lanka, Sumatra and the Philippines (Araneae: Salticidae). Bulletin of Prószynski,´ J., 2003. Salticidae of the World. URL: K “http:// British Arachnological Society, 8: 249-255. spiders.arizona.edu/salticid/main.htm” http:// spiders.arizona.edu/salticid/main.htm Yaginuma, T., 1967. Three new spiders (Argiope, Boethus and Cispius) from Japan. Acta Arachnologica, Tokyo, 20: 50-64. Simon, E., 1900. Etudes arachnologiques. 30e Mémoire. XLVII. Descriptions d’espèces nouvelles de la famille des Attidae. Annales de la Société Entomologique de France, 69: 27-61.
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 231-235 Date of Publication: 31 Dec.2005 © National University of Singapore
EXYRIAS AKIHITO, A NEW SPECIES OF CORAL-REEF GOBY (GOBIIDAE) FROM THE WESTERN PACIFIC
Gerald R. Allen Western Australian Museum, Francis Street, Perth, W.A. 6000, Australia
John E. Randall Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817-2704, USA
ASTRACT. Ð Exyrias akihito, is described from 16 specimens, 23.5-111.2 mm SL, collected at the Great Barrier Reef of Australia, Papua New Guinea, Indonesia, Philippines, Yaeyama Islands of southern Japan, and from a photographic record from Palau It is most similar to the wide-ranging Indo-Pacific E. belissimus, differing in having longer filamentous dorsal spines, usually 17 (vs. 18) pectoral rays, and in colour. Unlike the other three members of the genus, which frequent silt or mud bottoms of estuaries or turbid inshore reefs, the new species is found in relatively clear water in the vicinity of coral reefs to depths of at least 43 m.
KEY WORDS. Ð Fish taxonomy, Gobiidae, Exyrias, western Pacific.
INTRODUCTION of the caudal fin (posterior edge of hypural plate); head length is measured from the same anterior point to the posterior edge The gobiid genus Exyrias Jordan & Seale and its monotypic of the opercle flap; head depth is taken vertically at the level relative Macrodontogobius Herre were reviewed by Murdy of the preopercle margin and the head width is also taken at (1985). He recognized three species of Exyrias: E. belissimus this point; body depth is taken at two locations: between the (Smith) from the east coast of Africa (also known from the origin of the pelvic fins and the base of the first dorsal fin, Red Sea according to Dor, 1984) to the Samoa Islands; E. and between the origin of the anal fin and the base of the ferrarisi described as new from the Philippines, but now also second dorsal fin; snout length is measured from the anterior reported from Indonesia (Allen & Adrim, 2003), and E. end of the upper lip to the anterior margin of the eye; eye puntang (Bleeker) from the eastern Indian Ocean and western diameter is the horizontal fleshy diameter; interorbital width Pacific. These three gobies are relatively large, bottom- is the least fleshy width; upper-jaw length is taken from the dwelling species, frequently found in shallow, turbid waters anterior edge of the upper lip to the posterior end of the of estuaries and near-shore reefs. The present paper describes maxilla; lower-jaw length is taken from the anterior edge of a fourth species from the western Pacific that has sometimes the lower lip to the posteriormost extension of the lower been confused with E. belissimus. It was first collected by a jaw; caudal-peduncle depth is the least depth, and caudal- team from the Australian Museum on the northern Great peduncle length is the horizontal distance between verticals Barrier Reef of Australia in 1979. Subsequent collections have at the posterior base of the anal fin and the caudal-fin base; been made in recent years at Papua New Guinea, Indonesia, lengths of fin spines and rays are measured to their bases; Philippines, and the Yaeyama Islands of southern Japan. The caudal-fin length is the horizontal length from the posterior second author’s underwater photograph of the species in edge of the hypural plate to a vertical at the tip of the longest Palau, reproduced here, provides the record for this richest ray; pectoral-fin length is the length of the longest ray; pelvic- archipelago of Micronesia. The new species is typically found fin length is measured from the base of the outermost ray to in clearer water on sand or sand-rubble substrata in the vicinity the tip of the longest ray; counts of scales in longitudinal of sheltered coral reefs. series begin at the posterior upper attachment of the opercular membrane, continue posteroventrally to the fifth transverse scale row (starting from the mid-dorsal line), and then in a MATERIALS AND METHODS straight line midlaterally to the last scale, which is generally beyond the caudal-fin base (i.e. includes 2-3 scales on base Lengths of specimens are given as standard length (SL) of fin); transverse scale counts are taken from the origin of measured from the anterior end of the upper lip to the base the second dorsal fin ventroposteriorly to the anal fin base;
231 Allen & Randall: New species of Exyrias from Western Pacific also a second count is made from the anal-fin origin Exyrias sp. Ð Allen et al, 2003: 321, middle left fig. (Indonesia and dorsoposteriorly to the second dorsal-fin base; predorsal Philippines). scales are counted between the first dorsal-fin origin to where Exyrias sp. Ð Hayashi & Shiratori, 2003: 94 (Japan). they end just posterior to the interorbital space. Exyrias species Ð Kimura & Matsuura, 2003: 183, upper fig. (Bitung, Sulawesi). Exyrias sp. Ð Senou, Suzuki, Shibukawa & Yano, 2004: 240, upper Counts and proportions appearing in parentheses apply to the and lower figs. (Ishigaki, Iriomote Islands, Japan). range of values for paratypes. Selected proportional measurements are expressed as percentage of the standard Material examined. Ð Holotype - NSMT-P 71648, 67.1 mm SL, length (SL) in Table 1. Type specimens have been deposited Amitori Bay, Iriomote Island, Yaeyama Islands, Okinawa Prefecture, at the Australian Museum, Sydney (AMS), Biological Japan, 28 m depth, Y. Ikeda, K. Sugiyama & K. Hagiwara coll., 30 Laboratory of the Imperial Household, Tokyo (BLIH), Jun.2001. Bernice P. Bishop Museum, Honolulu (BPBM), National Science Museum, Tokyo (NSMIT), and Western Australian Paratypes Ð AMS I.20956-028, 100.3 mm SL, Tijou Reef (approximately 13 04'S 143 57'E), Great Barrier Reef, Queensland, Museum, Perth (WAM). ° ° 3-12 m, rotenone, AMS party coll., 23 Feb.1979; AMS I.22581- 011, 2 ex., 38.4-111.2 mm SL, Escape Reef (approximately 15°49'S 145°50'E), Great Barrier Reef, Queensland, 10-14 m, AMS party Exyrias akihito, new species coll., 29 Oct.1981; BLIH 19830335, 29.7 mm SL, El Nido, Palawan (Figs. 1-6; Table 1) Island, Philippine Islands, H. Masuda & A. Ono coll., 6-25 Mar.1983; BLIH 19960130, 38.4 mm SL, Amitori Bay, Iriomote Exyrias belissimus (in part) Ð Murdy, 1985: 4, Pl. 1B, Fig. B (Java). Island, Yaeyama Islands, Okinawa Prefecture, Japan, Y. Ikeda & Exyrias sp. 1 Ð Kuiter & Tonozuka, 2001: 669, Figs. A-D (Java, A. Iwata coll., 9 Jul.1996; BLIH 20010003, 34.9 mm SL, same Bali, and Sulawesi). locality as BLIH 1996130, 35 m, K. Yano coll., 18 Jun.2000; BLIH Exyrias sp. Ð Allen & Adrim, 2003: 58 (Bali and Manado, Sulawesi). 20010004, 25.2 mm SL, same locality as BLIH 19960130, 26 m, Y. Ikeda, K. Sugiyama & K. Hagiwara coll., 1 Jul.2001; BLIH 20010005, 38.2 mm SL, same locality and as BLIH 19960130, 27 m, Y. Ikeda, K. Sugiyama & K. Hagiwara coll., 1 Jul.2001; BLIH 20010006, 39.0 mm SL, same locality as BLIH 19960130, 27 m, Y. Ikeda, K. Sugiyama & K. Hagiwara coll., 28 Jun.2001; BLIH
Fig. 1. Holotype of Exyrias akihito, male, NSMT-P 71648, 67.1 mm SL, 28 m, Amitori Bay, Iriomote Island, Yaeyama Islands, Okinawa Prefecture, Japan (Y. Ikeda).
Fig. 3. Underwater photograph of Exyrias akihito, about 50 mm total length, 15 m, off Lawadi Village, Milne Bay Province, Papua New Guinea (G. Allen).
Fig. 2. Head of paratype of Exyrias akihito, 81.3 mm SL, showing cutaneous papilla system (solid dots) and pores (pale dots and Fig. 4. Underwater photograph of Exyrias akihito, about 60 mm arrows). total length, Seribu Islands, Java (J. Randall).
232 THE RAFFLES BULLETIN OF ZOOLOGY 2005
20010010, 23.5 mm SL, Funauki Bay, Iriomote Island, Yaeyama P29928-019, 7: 36-77 mm, Cocos-Keeling Islands; WAM P31650- Islands, Okinawa Prefecture, Japan, 43 m, K. Yano coll., 27 001, 94 mm, Cassini Island, Kimberleys, Western Australia; WAM May.2001; BPBM 28781, 37.2 mm SL, Pulau Pari, off Pulau Tikus, P30309-010, 69 mm, Cassini Island; WAM P27662-030, 74 mm, Seribu Islands, Java, Indonesia, outer reef in 23 m depth, quinaldine, Clerke Reef, Rowley Shoals, Western Australia; WAM P29054- J. E. Randall coll., 5 Nov.1982; WAM P.31214-001, 66.6 mm SL, 003, 77 mm, West Island, Ashmore Reef, Timor Sea; WAM P30412- Padoz Reef (approximately 5°10'S, 145°49'E) Madang Lagoon, 007, 2: 44-49 mm, Bohaydulong Island, Bodgaya Islands, Sabah; Papua New Guinea, 15-18 m, rotenone, G. R. Allen coll., 16 WAM P31546-004, 30 mm, Gam Island, Raja Ampat Islands, Irian Oct.1996; WAM P.32255-003, 2 ex. specimens, 27.2-33.9 mm SL, Jaya. Kawe Island (approximately 0°02'S, 130°08'E), Raja Ampat Islands, Papua Province, Indonesia: large bay on east side of island, 8-14 m, Diagnosis. Ð Pectoral rays 17; segmented caudal-fin rays quinaldine, G. R. Allen coll., 18 Nov.2002; WAM P.32615-001, 17; branched caudal-fin rays typically 13, rarely 12, 14, or 81.3 mm SL, male, Lawadi Village (10°15.81'S, 150°43.14'E), Milne 15; longitudinal scale series 28 (28-30); dorsal-fin spines Bay Province, Papua New Guinea, outer reef in 15 m, quinaldine elongate and filamentous, the second longest, 1.59-2.07 in and hand net, G. R. Allen coll., 20 Nov.2003. SL; generally light bluish grey in life with small yellow to Comparative material. Ð Exyrias bellismus: BPBM 29677, 4: 43-55 brownish orange spots on head and body, those on body mm, Pulau Pari, Seribu Islands, Java, Indonesia; BPBM 15767, 87 mostly conjoined to form longitudinal stripes along scale mm, Madang, Papua New Guinea; BPBM 32567, 3: 36-77 mm, rows; four pairs of dark brown spots on lower side of body, Nagada Harbour, Papua New Guinea; BPBM 1283, 116 mm, followed by a single spot at caudal-fin base; a few smaller Shortland Island, Solomon Islands; BPBM 15696, 2: 50-51 mm, dark brown spots on body, mostly in a row dorsally and one Yacht Harbor, Guadalcanal, Solomon Islands; BPBM 21855, 2: 57- ventrally; dorsal and caudal fins covered with numerous, 73 mm, Salu Islet, Singapore, 2: 57-73 mm; BPBM 29386, 85 mm, relatively large, orangish to yellowish brown spots. Low Isles off Port Douglas, Queensland; BPBM 9740, 2: 62.5-69 mm, Uru Island, Palau; BPBM 9754, 3: 19-58 mm, Uru Island, Palau; BPBM 31405, 2: 66.5-74.5 mm, Auluptagel, Palau; BPBM Description. Ð Dorsal-fin rays VI-I,10; anal-fin rays I,9; 9956, 3: 54-82 mm, Phonpei, Caroline Islands; BPBM 29384, 25 pectoral-finl rays 17; pelvic-fin rays I, 5; segmented caudal- mm, Phonpei, Caroline Islands, W side of Phonpei Passage; WAM fin rays 17; branched caudal-fin rays 14 (12-15); longitudinal scale series 28 (28-30); transverse-scale count from origin of second dorsal fin ventroposteriorly to anal-fin base 9; transverse-scale count from anal-fin origin dorsoanteriorly to first dorsal-fin base 9; predorsal-scale count 8 (7-8); vertebrae 10 + 16.
Mouth terminal; jaws extending to a vertical at anterior edge of pupil; jaw teeth small and conical, densely packed in several rows, those in outermost row the largest; tongue broad with straight anterior margin, broadly attached anteriorly to floor of mouth. Anterior extent of gill opening reaching to below middle of opercle. Arrangement of cutaneous papilla system and cephalic sensory pores as shown in Fig. 2.
Body moderately elongate and laterally compressed, more strongly compressed posteriorly; body depth at pelvic-fin base 4.05 (3.67-4.15) in SL; body depth at anal-fin origin 4.17 (3.68-4.82) in SL. Head laterally compressed, its width slightly less than its depth at level of preopercular margin. Fig. 5. Underwater photograph of Exyrias akihito, about 70 mm total length, Palau, 15 m (J. Randall). Head length 3.37 (3.24-3.44) in SL. Snout length 2.47 (2.17- 3.31), eye diameter 3.75 (3.38-4.23), and interorbital width 20.74 (15.69-30.94), all in head length. Distance between
Fig. 6. Underwater photograph of Exyrias belissimus, about 80 mm total length, 10 m, Milne Bay Province, Papua New Guinea (G. Fig. 7. Exyrias belissimus, 77 mm SL, Pohnpei, Caroline Islands Allen). (J. Randall).
233 Allen & Randall: New species of Exyrias from Western Pacific
Table 1. Proportional measurements of selected type specimens of Exyrias akihito expressed.
Holotype Paratype Paratype Paratype Paratype Paratype NSMT-P WAM AMS AMS BLIH BPBM 71648 P.32615 I.22581 I.20956 20010006 28781 Standard length (mm) 67.1 81.3 111.2 100.3 39.0 37.2 Head length 29.7 30.1 29.1 29.3 30.9 30.0 Head width 18.4 18.6 17.2 18.2 15.7 17.7 Head depth 22.1 21.4 31.2 23.2 20.7 20.8 Body depth at pelvic origin 24.7 26.2 27.2 27.3 24.1 25.4 Body depth at anal origin 24.0 24.5 26.1 27.1 20.7 24.4 Caudal-peduncle depth 15.4 14.5 15.6 16.1 14.5 15.5 Caudal-peduncle length 18.9 20.5 21.6 23.1 24.4 26.3 Snout length 12.0 12.9 13.3 13.5 9.3 10.2 Orbit diameter 7.9 7.2 6.9 6.9 9.1 8.9 Interorbital width 1.4 1.9 1.9 1.6 1.6 1.0 Upper jaw length 10.7 9.2 9.8 9.9 8.5 8.7 Lower jaw length 11.9 10.8 10.5 11.1 10.8 9.7 Snout to 1st dorsal origin 31.2 33.0 32.0 31.0 33.1 34.2 Snout to 2nd dorsal origin 53.7 53.4 55.6 52.4 54.6 54.4 Snout to anal fin origin 56.3 57.9 59.3 57.7 53.6 55.6 Snout to pelvic fin origin 32.4 31.5 31.5 30.9 32.3 31.0 Longest dorsal spine 62.8 56.5 50.6 48.4 55.0 49.7 Longest soft dorsal ray 28.9 33.2 30.1 29.8 22.8 26.7 Longest soft anal ray 30.0 30.9 28.7 30.1 24.1 23.9 Pectoral-fin length 34.8 33.6 31.3 31.0 32.2 33.7 Pelvic-fin length 28.5 25.2 28.2 25.0 27.0 27.0 Caudal-fin length 46.2 44.4 44.5 47.0 43.6 45.6
snout and origin of first dorsal fin 3.20 (2.92-3.23), between spots; a series of blackish blotches dorsally on body below snout and origin of second dorsal fin 1.86 (1.80-1.91), base of dorsal fins; an irregular row of smaller dark brown between snout and origin of anal fin 1.78 (1.69-1.87), and spots on lower side of body; a row of still smaller spots on between snout and origin of pelvic fins 3.09 (3.10-3.22), all upper side of body; a broad dark brown bar below eye, and in SL. a prominent dark brown spot on opercle; dorsal and caudal fins greyish brown with traces of numerous spots; anal and Membranes of first dorsal fin deeply incised (especially paired fins grey-brown. between first and second spines), the dorsal spines elongate and filamentous, second spine longest, 1.59 (1.77-2.07) in Color in life (from underwater photographs of Figs. 2-4): SL; longest (penultimate) ray of second dorsal fin 3.46 (3.01- orange-yellow to brownish yellow with pale bluish grey to 4.39), longest ray (penultimate) of anal fin 3.34 (3.24-4.19), pale blue spots, most conjoined to form longitudinal series pectoral-fin length 2.87 (2.96-3.23), pelvic-fin length 3.50 along scale rows; edges of scales narrowly dark brown, (3.55-4.01), caudal-fin length 2.16 (2.13-2.29), all in SL. especially dorsally on body; four pairs of small dark brown to black spots on scale row at level of upper edge of pectoral- Color of holotype in alcohol after three and a half years of fin base, these spots surrounded by a zone of orange-yellow preservation: generally yellowish tan with four pairs of dark or brownish yellow; a single dark brown to black spot on brown spots on scale row at level of upper edge of pectoral- caudal-fin base in line with double dark spots, followed by fin base; a single dark brown spot at caudalÐfin base in line one or two lesser dark spots on base of fin; a row of small with double dark spots; fins translucent tan. Most of the dark brown spots along back at base of dorsal fins, another paratypes are similar in colour except for the most recently on lower side, and a few still smaller dark brown spots along collected specimens (WAM P.32615-001), which differ as upper side, all surrounded by yellow or brownish orange; head follows: overall pale brown, the scale edges a little darker with yellow to brownish orange spots, mostly forming with characteristic row of paired dark brown spots (each pair irregular rows, some projecting obliquely downward and within a brown blotch) and single dark brown spot at caudal- forward from eye; dorsal and caudal fins translucent bluish fin base; the latter spot closely followed by two more dark grey with round to oval yellowish to orangish brown spots,
234 THE RAFFLES BULLETIN OF ZOOLOGY 2005 mostly with slightly darker edges, many as large or larger average count of branched caudal-fin rays for 16 specimens than pupil; anal fin pale translucent greenish or yellowish of E. akihito is 13.1. with similar but fewer and more elongate spots, mainly in basal half of fin; second dorsal and anal fins sometimes with Distribution and habitat. Ð Exyrias akihito is currently a narrow pale blue margin; paired fins pale greenish or known from the northern Great Barrier Reef, Papua New yellowish to nearly white. Senou et al. (2004: 240, upper fig.) Guinea, Indonesia (Bali, Java, north Sulawesi, and Raja illustrated a large adult in color. The pale blue spots are Ampat Islands), Philippines (Palawan), and the Yaeyama relatively larger, the edges of the scales more conspicuously Islands of southern Japan. It is typically found in clear water dark brown, the double dark spots much less distinct, and the in the vicinity of coral reefs, usually on sand or rubble dusky-edged orange-yellow spots in the median fins more bottoms, at depths between 10-43 m. By contrast, the three numerous and relatively smaller, most smaller than pupil. A other species of Exyrias are generally found in shallow water photograph in Hayashi & Shiratori (2003: 94, lower fig.) with soft silt or mud bottoms in estuaries or on turbid coastal shows a broad dusky bar below the eye (still containing reefs. An exception in depth is a specimen speared by the orange-yellow spots). second author in the lagoon off Madang, Papua New Guinea at a depth of 50 m. Etymology. Ð The new species is named akihito in honour of the Emperor of Japan, in recognition of his significant contribution to our knowledge of gobiid systematics. Many ACKNOWLEDGMENTS of the type specimens of E. akihito were supplied by the Biological Laboratory of the Imperial Household in Tokyo. We are very grateful to Yuji Ikeda, of BLIH, for sending us The name akihito is treated as a noun in apposition. most of the type specimens of E. akihito that were utilized in this study. He recognized the species as new and was Comparisons. Ð Exyrias akihito is easily differentiated from planning to describe it, but agreed to forward us the BLIH the other members of the genus by its extremely elongate specimens as well as loan material from other institutions dorsal-fin spines. It is also distinguished on the basis of its when he learned of our study. The first author is also grateful paler coloration, particularly the numerous yellow or to Rob Vanderloos, owner of the charter dive boat Chertan, brownish orange spots on the head and body. for his generosity in providing an opportunity to join the dive cruise at Milne Bay on which a paratype was collected. Roger This new goby is most similar to E. belissimus (Figs. 7-8), Steene of Cairns, Australia assisted with fish collections on with which it has been confused in the past. For example, this same cruise. Murdy (1985, Pl. l fig. B) illustrated a juvenile of the new species as E. belissimus. The latter species has the same double-row of dark brown spots on the side (though vertically LITERATURE CITED elongate and often obscured by dark brown bars, especially in adults), and the dorsal spines are filamentous. However, Allen, G. R. & M. Adrim., 2003. Coral reef fishes of Indonesia. they are not as long as those of E. akihito. The third or fourth Zoological Studies. 42(1): 1-72. dorsal spine is longest in E. belissimus. It measures 20.2- Allen, G. R., R. Steene, P. Humann & N. Deloach., 2003. Reef 36.9% SL in eleven Bishop Museum specimens from 44.7- Fish Identification - Tropical Pacific. New World Publications, 87.0 mm. The second dorsal spine is longest in E. akihito, Jacksonville, FL. 457 pp. 49.7-62.8% SL in the six type specimens of Table 1. Dor, M., 1984. Checklist of the fishes of the Red Sea. The Israel Acacemy of Sciences and Humanities, Jerusalem. 437 pp. There is also a clear modal difference in the number of Hayashi, M. & T. Shiratori, 2003. Gobies of Japanese Waters. TBS pectoral-fin rays. All type specimens of E. akihito possess Buritanica, Tokyo. 223 pp. (In Japanese). 17 rays, compared to the usual count of 18 for E. belissimus. Kimura, S. & K. Matsuura, 2003. Fishes of Bitung. Ocean Research Murdy (1985) reported an average count of 17.8 for E. Institute, University of Tokyo. vi + 244 pp. belissimus, however he included at least three specimens of the new species (AMS I.20956-028 and AMS I.22135-001) Kuiter, R. H. & T. Tonozuka, 2001. Photo Guide to Indonesian in the Material Examined for E. belissimus. Therefore, it is Reef Fishes. Zoonetics, Seaford, Australia. Part 3. iv + 623- 893 pp. possible that Murdy’s mean count for pectoral-fin rays may have included one or more specimens of E. akihito. We have Murdy, E. O., 1985. A review of the gobiid fish genera Exyrias counted 25 specimens of E. belissimus; 22 have 18 rays, and and Macrodontogobius, with description of a new species of Exyrias. Indo-Pacific Fishes, no. 10: 1-14. one each has 16, 17, and 19 rays. There is also an average difference in the number of branched caudal-fin rays. Murdy Senou, H., T. Suzuki, K. Shibukawa & K. Yano, 2004. A (1985) gave the mean as 14.2 for E. belissimus (n = 12), 14.5 Photographic Guide to the Gobioid Fishes of Japan. Heibonsha, Ltd., Tokyo. 534 pp. (In Japanese). for E. ferrarisi (n = 2), and 13.7 for E. puntang (n = 10). The
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 237-242 Date of Publication: 31 Dec.2005 © National University of Singapore
A NEW SPECIES OF FRESHWATER GOBY (TELEOSTEI: GOBIIDAE: STIPHODON) FROM PULAU TIOMAN, PAHANG, PENINSULAR MALAYSIA
I-Shiung Chen Institute of Marine Biology, National Taiwan Ocean University, Keelung, 202, Taiwan, Republic of China Email: [email protected]; [email protected] (Corresponding author)
Heok Hui Tan Department of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 119260, Republic of Singapore Email: [email protected]
ABSTRACT. Ð An amphidromous goby, Stiphodon aureorostrum, new species, is described from the freshwater streams of Pulau Tioman off the east coast of Peninsular Malaysia. It is characterised by a combination of the following features: I/9 second dorsal fin rays; I/10 anal fin rays, 15-16 pectoral fin rays, 35-37 longitudinal scale rows, 4-11 predorsal scales, first dorsal fin with prolonged filamentous rays; pectoral fin with 8-10 vertical rows of greyish black spots, body with distinct 9-10 narrow, dark grey bars in male; golden longitudinal band on anterior and lateral aspects of snout on both sexes in life.
KEY WORDS. Ð Gobiidae, Stiphodon, Peninsular Malaysia, new species.
INTRODUCTION primary freshwater fishes among the 48 recorded species of inland fishes (fide Ng et al., 1999; Lim & Ng, 1999) with There is a high diversity of freshwater gobioid fishes two endemic species, viz. Sundoreonectes tiomanensis inhabiting the basins of the west and tropical Pacific which (Balitoridae) and Clarias batu (Clariidae). Among them, comprise of amphidromous and landlocked species (Chen & gobioids are the largest group, with 13 genera and 17 species Fang, 1999). Members of the genus Stiphodon Weber (1895) recorded. Ng et al. (1999) reported the presence of Stiphodon are small amphidromous freshwater gobiid fish with short, atropurpureus (Herre, 1927) from Tioman, the first record transverse infraorbital papillae pattern and showing distinct of the genus for the area. After further surveys in 1999 and sexual dimorphism in colour pattern and dorsal fin shape; 2001, we found and confirmed another species of Stiphodon. the male always with iridescent or bright colour pattern and It is herein described as a new species - Stiphodon usually with prolonged fin rays in the first dorsal fin; the aureorostrum. female always with two longitudinal dark stripes on the body and no elongated rays in the first dorsal fin. Some studies on Stiphodon have been conducted in the Philippine islands MATERIAL AND METHODS (Herre, 1927; Watson & Kottelat, 1995); in Taiwan and the Ryukyu islands (Watson & Chen, 1998); and Hainan Island Specimens were collected with a cast net and also by utilising (Wu & Ni, 1985). Less is known of this taxon in Sundaic two hand-nets while snorkelling. All counts and Southeast Asia, with only one recent record from the island measurements follow Chen & Shao (1996), except for the of Tioman, off the east coast of Peninsular Malaysia (Ng et maximal extension of the predorsal scale series which is al., 1999). defined herein as the scale counts from the origin of the first dorsal fin base extending to the right side of the anterior tips Pulau Tioman is located about 45 km off the southern part of biforked squamation (PreD-max). Terminology of head of the east coast of the Malay Peninsula in the South China lateral-line and neuromast organs follow Miller (1988). The Sea. The inland water fishes of Tioman comprises mainly type specimens examined are deposited in the National of diadromous, secondary division freshwater forms and these Museum of Marine Biology & Aquarium, Pingtung, Taiwan appear to be concentrated mainly in the lower reaches and (NMMBP) and the Raffles Museum of Biodiversity Research, estuaries (Alfred, 1966; Ng et al., 1999). There are only six National University of Singapore (ZRC). Other comparative
237 Chen & Tan: New species of Stiphodon from Pulau Tioman materials of congeners are deposited in the Biological subterminal. Eye medium to large (17.3-43.4 % HL), Laboratory, Imperial Household, Tokyo (BLIH), NSMT dorsolateral in position. Upper jaw teeth tricupsid; lower jaw (followed Leviton et al., 1985), and the Pisces collection of teeth conical. Labial teeth present in lower jaw. Tongue ISC (ISP). The collection localities are shown in Fig. 1. All margin rounded. Anterior nostril a short tube, posterior nostril abbreviations for meristic and morphometric characters a round hole. Gill-opening restricted, extending vertically follow Miller (1988) and Chen et al. (1999). and not beyond the rear edge of opercle. Isthmus broad. Vertebral count 10 + 16 = 26. Morphometric data listed in Table 1, and meristic counts in Table 2. Stiphodon aureorostrum, new species (Figs. 2-5) Fins. Ð First dorsal fin ray count VI, second dorsal fin ray count I/9; anal fin ray count I/10; pectoral fin ray count 15- All material examined from Pulau Tioman, Pahang, Peninsular 16 (modally 15); pelvic fin ray count I/5+I/5 (frequency Malaysia. distribution of fin-ray counts listed in Table 2). First dorsal fin with filamentous fin rays with fourth fin ray longest in Holotype. Ð Male (52.0 mm SL)(ZRC 46412), Juara: Sungai male; first dorsal fin rays about equal in female. Distal tip Keliling, coll. I-S. Chen, H. H. Tan & K. K. P. Lim, 17-18 Jul.2001. of longest first dorsal fin ray extending to the bases of third Paratypes. Ð 2 females (55.1-60.3 mm SL)(ZRC 46413), same to fifth fin rays of second dorsal fin when depressed; but the locality data as holotype; 2 males and 2 females (38.9-52.5 mm rear margin of first dorsal fin nearly reaching second dorsal SL)(NMMBP 1984), same locality data as holotype; 2 females (52.0- fin origin in female. Posterior fin rays of second dorsal fin 58.9 mm SL)( ZRC 45409), Juara: Sungai Keliling, coll. H. H. Tan, and anal fin long, reaching the procurrent rays in adult male 25 Jun.1999; 1 male (41.2 mm SL)( ZRC 45410), Juara: Sungai when depressed, but respective fin rays in female not Keliling, coll. P. K. L. Ng et al., 24 Jun.1999. elongated. Origin of anal fin inserted below origin of second dorsal fin in male; the origin of anal fin inserted below first Non-type material. Ð1 female (43.1 mm SL)(ZRC 45408), Juara: branched fin ray of second dorsal fin in female. Pectoral fin Sungai Keliling. large, the rear tip reaching nearer the vertical of anus than the rear margin of pelvic fin in male; but not extending to the Diagnosis. Ð Stiphodon aureorostrum differs from all its vertical midline between rear tip of pelvic fin and anus. congeners by the unique combination of features: I/9 second Caudal fin elliptical, rear margin rounded. dorsal fin rays, I/10 anal fin rays, 15-16 pectoral fin rays, 35-37 longitudinal scale rows, 4-11 predorsal scales, first Scales. Ð Body covered with moderately large ctenoid scales, dorsal fin with prolonged filamentous rays; pectoral fin with but the anterior portion before midline of first dorsal fin with 8-10 vertical rows of greyish black spots, body with narrow relatively small cycloid scales extending to predorsal origin 9-10 dark grey bars in male; golden longitudinal band on of nape. Predorsal squamation with the extension of bifid anterior and lateral aspects of snout on both sexes in life. pattern (shown in Fig. 5A), the extension representing the Stiphodon aureorostrum also represents the largest species sexual dimorphism in which the coverage of scales toward ever collected in the Indo-Pacific region. One female and beyond the vertical line of anterior terminal pore of specimen (ZRC 46413) measures 60.3 mm SL. posterior oculoscapular canal in female but never reaching the point in male. Longitudinal scale row 33-37; transverse Description. Ð Body subcylindrical and somewhat square- row 9-10; scale rows between first dorsal fin origin and like in cross section, compressed posteriorly. Head somewhat anterior dorsal pectoral fin origin 11-16; predorsal scale row angular and snout semi-circular in dorsal view. Snout tip 4-6 in male, 7-11 in female; maximal extension of predorsal overhanging the upper lip. Mouth slightly oblique and
Fig. 1. Location of Pulau Tioman. Map of Pulau Tioman showing Fig. 2. In-situ photograph of a small group of Stiphodon distribution of Stiphodon aureorostrum (solid symbol) and S. aureorostrum (centre area of image) grazing on gravel in Sungei atropurpureus (hollow symbol). Keliling (taken on 18 July 2001).
238 THE RAFFLES BULLETIN OF ZOOLOGY 2005 scale series 11-14 in male, 19-22 in female. Abdominal Male: Head dorsum yellowish-brown with dark brown median squamation before anus extending to the rear edge patches, golden-yellow patch on area above eye. Anterior of pelvic fin base. Head including snout, cheek, opercle; tip of snout with golden-yellow longitudinal stripe to front pectoral fin base and pre-ventral region naked. of eye, bordered dorsally and ventrally with brown. Eye with golden iris. Region behind eye tinged with golden-yellow, Head lateral-line system (Fig. 5B) bordered ventrally with brown to opercle edge. Body dorsum Cacals: Pair of pores σ, σ’, and λ of anterior oculoscapular dark brown with posterior edge of scales black, forming a canal on dorsal side of snout; pore s’ on terminal of anterior reticulate pattern. Body dorsum with one gold bar anterior branch of head canal near the anterior nostril; pore σ near to first dorsal fin, two gold bars at first dorsal fin base, four the posterior nostril. Single pore k in interorbital region; no gold bars at second dorsal fin base. Body yellowish-brown pore w. Anterior oculoscapular canal with lateral pores α and with up to 10 dark brown bars, distance between bars ρ. Posterior oculoscapular canal with terminal pores θ and progressively more narrow towards posterior. Body with one τ. Preopercular canals with both terminal pores γ and ε, but brown bar anterior to first dorsal fin, two bars below first lacking pore δ. dorsal fin, six bars below second dorsal fin and one bar at caudal peduncle. Ventrum light brown. Pectoral fin light Head sensory papillae: The infraorbital papillae present as yellowish with 10 to 12 curved thin brown bars, base with 6 short transverse rows shown in detail as Fig. 5B. Row 6 gold spot. Pelvic fin hyaline. First dorsal fin base yellow separated into row 6s and 6i. The row b and d present. Rows with dark brown patches, distal part reddish with white os, ot, and oi well separated on opercle. Row os separated iridescent edge. Second dorsal fin yellowish with dark brown into four divisions. patches, distal part reddish with white iridescent edge. Anal fin dull reddish-brown with thin white edge. Caudal fin base Life coloration. Ð See Figs. 3A-C for life coloration of male with golden bar, whitish spot at base of upper caudal rays; and female specimens. upper and lower bands of caudal fin light reddish with white edge, middle section blackish with six to eight thin white bars, thin white margin. A Female: Head coloration similar to male, except golden stripe is faint. Body dorsum similar to male. Body with nine brown bars, one bar anterior to first dorsal fin, two bars below first dorsal fin, one bar in between dorsal fins, four bars below second dorsal fin and two bars at caudal peduncle. Ventrum light brown. Coloration of pectoral, pelvic, anal and caudal fins similar to male. First and second dorsal fins with black rays, hyaline interradial membrane, subdistal reddish band with white margin. B Preserved coloration. Ð See Figs. 4A-B for preserved coloration of male and female specimens. Body light creamy
A
B C
Fig. 3. Stiphodon aureorostrum, new species: A, Life male holotype, Fig. 4. Stiphodon aureorostrum, new species: A, Preserved male (ZRC 46412, 52.0 mm SL); B, Close-up of male’s head; C, life holotype (ZRC 46412, 52.0 mm SL); B, Preserved female paratype female paratype (ZRC 46413, 60.3 mm SL). (ZRC 46413, 60.3 mm SL).
239 Chen & Tan: New species of Stiphodon from Pulau Tioman
yellow, longitudinal stripe absent in male, a broad longitudinal, dark brown stripe present in lateral midline of A female. Ten narrow dark brown cross-bars in body side of male which much narrower than its inter-space, four bars before second dorsal fin origin, six bars below second dorsal fin base to caudal peduncle; nine to ten light brown or indistinct bars in female. Upper half of lateral side with a longitudinal, blackish brown line behind eye to upper tip of caudal fin base. Dorsal part from nape to caudal fin origin with nine dark blotches, and scales on the blotches usually with distal dark brown spot. Ventral side of body creamy yellow. Head creamy yellow; greyish brown in snout to interorbital region, anterior cheek and opercle in male; a short black bar below eye, a horizontal, brown line surrounding lower part of snout to opercle. Interorbital region to dorsal part of snout with three pairs of dark brown spots in female. Upper lip greyish brown and lower lip whitish. First dorsal B fin whitish with dark brown rays and posterior membrane behind fifth fin ray dusky brown in male; whitish with brown fin rays in which the first having two basal spots in female. Second dorsal fin greyish with darker rays in male; whitish in having three longitudinal rows of dark brown spots in female. Pelvic fin with dark grey membrane with whitish rays in male; whitish with a distal, narrow, longitudinal greyish black stripe in female. Pectoral fin whitish with a prominent longitudinal dark bar in female but indistinct in male; with ten to twelve vertical rows of brownish black dots on its fin rays. Caudal fin with six to eight vertical rows of black stripes in male, but usually separated as spots in posterior rows in female. Upper distal area of caudal fin whitish and lower one third dark grey in male.
Etymology. Ð From the combination of the Latin aureus, golden, and rostrum, snout. This is in allusion to the golden longitudinal band on the anterior and lateral aspects of snout on both sexes in life. Used as a noun in apposition.
Distribution. Ð Stiphodon aureorostrum is known only from the freshwater streams of Pulau Tioman, Peninsular Malaysia. It is found in the lower reaches of Sungai Keliling, Juara Bay, eastern Tioman; however an individual was observed by ISC in the lower reaches of Sungai Paya on the western side of the island. This species appears to be rare in the type locality, which seems to be predominated by Glossogobius celebius (Gobiidae).
Field observations. Ð Stiphodon aureorostrum was observed in-situ (Fig. 2) grazing on the gravel substratum in Sungai Keliling, usually in a group of 8 to 10 individuals, usually with one or two Glossogobius celebius or Lutjanus argentimaculatus (Lutjanidae) following closely behind. This species appears to prefer areas of slow to moderate water flow, whereas its sympatric congener, S. atropurpureus seems to prefer faster flowing water amongst large rocks. Stiphodon aureorostrum can also be found in areas of faster water flow but in smaller groups of 3 to 4 individuals. The genus Stiphodon was not found further upstream, possibly due to competition with Puntius lateristriga (Cyprinidae). Nocturnal Fig. 5. A. Head squamation drawing (female; scale bar = 1 mm); observations were attempted. Stiphodon aureorostrum were B. Head pore pattern drawing (male; lateral, ventral and dorsal views, scale bar = 1 mm). not seen, only Clarias leiacanthus (Clariidae) was active.
240 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 1. Morphometric data of Stiphodon aureorostrum.
Holotype male specimens female specimens (n=5) (n=6) SL (mm) 52.0 41.2-55.1 38.9-60.3 % SL range average range average head length 23.4 22.4-23.4 22.9 21.1-22.5 21.6 snout to first dorsal fin length 34.0 32.9-35.5 34.2 33.9-36.3 34.5 snout to second dorsal fin length 55.8 54.7-57.8 56.0 55.2-58.0 56.5 snout to anus length 52.4 49.8-52.4 51.1 50.0-57.6 53.5 snout to anal fin origin length 56.1 53.6-56.2 54.9 55.9-59.0 57.4 prepelvic length 23.3 22.7-23.4 23.1 19.9-23.6 21.1 caudal peduncle length 19.1 18.4-19.6 18.9 16.7-20.0 18.4 caudal peduncle depth 10.6 10.4-12.2 11.2 10.2-11.5 10.8 first dorsal fin base length 18.1 16.4-20.1 17.7 15.3-19.6 17.0 second dorsal fin base length 27.2 26.6-28.8 27.8 23.5-24.8 24.2 anal fin base length 28.2 27.6-31.3 29.2 25.6-26.9 26.3 pectoral fin length 21.8 20.6-24.0 22.3 18.3-22.5 20.8 caudal fin length 27.7 23.1-29.5 25.7 20.3-23.1 21.5 pelvic fin length 15.4 14.3-16.8 15.3 13.3-16.1 14.7 body depth at pelvic fin origin 13.8 13.5-14.9 14.1 12.5-15.3 13.6 body depth at anal fin origin 14.4 14.2-16.8 15.5 14.8-17.5 16.0 body width at anal fin origin 13.8 13.1-15.5 14.2 10.0-16.4 12.3 distance between pelvic fin origin and anus 28.8 28.8-30.0 29.4 30.6-33.9 32.3 % HL snout length 41.8 35.4-41.8 37.8 18.8-43.1 37.1 eye diameter 21.2 17.3-21.2 20.0 21.4-43.4 26.1 cheek diameter 24.1 23.2-26.2 24.1 21.1-31.4 24.7 postorbital length 40.0 40.0-50.5 44.3 37.7-49.8 45.5 maximal head width 73.6 68.8-73.6 70.8 61.9-76.7 71.3 head width at upper margin of gill-opening 58.5 56.4-59.6 58.4 55.0-65.9 61.1 fleshy-int 43.6 39.3-43.8 42.1 38.8-46.0 41.8 lower jaw length 40.0 36.7-40.0 38.6 35.8-42.7 38.6 % caudal peduncle length caudal peduncle depth 55.6 55.6-65.5 59.0 55.9-64.5 58.8
Comparison with related species. Ð Among its congeners, sized Ryukyu endemic species Ð S. imperiorientis Watson & the new species Stiphodon aureorostrum most resembles S. Chen, 1998, in sharing the spotted pectoral fin and distinct multisquamus Wu & Ni, 1985, from Hainan Island. They sexually dimorphic predorsal scale pattern. However, S. have similar pattern of spotted pectoral fin and several aureorostrum can be distinguished from S. imperiorientis by meristic features, but S. aureorostrum differs from S. the following features: 1) larger body size (vs. smaller); 2) multisquamus in the following characters: 1) larger body size first dorsal fin with third spine longest in males (vs. fourth (vs. smaller); and more elongated first dorsal fin rays in males dorsal spine longest); 3) with more numerous (about 12) and (vs. less elongate); 2) predorsal squamation with more scale broader dark bars on body in male (vs. fewer and narrower extension to anterior region (vs. fewer scale extension); 3) dark bars); 4) golden stripe across snout in life male (vs. snout body more slender and elongated (vs. less slender and and cheek iridescent blue); 5) caudal fin red in upper distal elongated); 4) absence of any blue on head and the lateral region in male (vs. no pattern); and 6) distinct dark bars on side in stead of golden snout in former male (vs. distinct blue body of female (vs. absence of bars). background); and 5) narrower dark transverse bars in former male (vs. iridescent blue). Remarks. Ð There is still a need to investigate the true diversity of the giant Stiphodon species found in the hill Stiphodon aureorostrum is also rather similar to the large stream habitats of different river basins draining into the South
241 Chen & Tan: New species of Stiphodon from Pulau Tioman
Table 2. Frequency of selected meristic counts of Stiphodon aureorostrum.
pectoral longitudinal scale row transverse predorsal scales fin rays scale row 15 16 33 34 35 36 37 9 10 4 5 6 7 8 9 10 11 male 9 1 1 3 4 2 - 3 2311----- female 10 2 - - 5 4 3 4 2 - - - 1 1 2 1 1 scales between origin of first dorsal fin maximal extension of predorsal scale series and upper margin of pectoral fin base 11 12 13 14 15 16 11 12 13 14 15 16 17 18 19 20 21 22 male 1 2 2 - - - 1 1 12 ------female - - 1 2 1 2 ------2 2 1 1 Male (n=5), female (n=6)
China Sea. There are most probably several unnamed species Chen, I.-S. & L.-S. Fang, 1999. The freshwater and estuarine fishes in this region (Chen, in preparation). of Taiwan. National Museum of Marine Biology & Aquarium, Taiwan. 287 pp. (In Chinese). Chen, I-S., H. L. Wu & K. T. Shao, 1999. A new species of Comparative material Rhinogobius (Teleostei: Gobiidae) from Fujian Province, China. Ichthyological Research, 46: 171-178. Stiphodon imperiorientis Watson & Chen, 1998 Ð Holotype: Herre, A. W. C. T., 1927. Gobies of the Philippines and the China NSMT.P 48063, 48.1 mm SL; Japan: Ryukyu Islands, Okinawa; Sea. Monograph of the Bureau of Science, Manila Monograph, Iriomote Island, Nakama River, Yaemama-gun; 2 Sep.1986. 23: 1-352, frontispiece, pls. 1-30. Paratypes: BLIH 19811202, 42.7 mm SL, same locality as above Leviton, A. E., R. H. Gibbs, E. Heal & C. E. Dawson, 1985. (10 Jul.1981); BLIH 1986400, 43.4 mm SL, same locality as above Standards in herpetology and ichthyology: Part 1. Standard (2 Nov.1986). symbolic codes for Institutional resource collections in herpetology and ichthyology. Copeia, 1985: 802-832. Stiphodon multisquamus Wu & Ni, 1985 Ð ISP- uncatalogued, 2 ex., 38.6-45.3 mm SL; China: Guangdong Province, Hainan Island: Lim, K. K. P. & H. H. Ng, 1999. Clarias batu, a new species of Linshuei River basin, hill stream near Paoting; I-S. Chen, 1 Aug catfish (Teleostei: Clariidae) from Pulau Tioman, Peninsular 1996. Malaysia. In: Sodhi, N. S., H. S. Yong & P. K. L. Ng (eds.), The biodiversity of Pulau Tioman, Peninsular Malaysia. Raffles Bulletin of Zoology, Supplement No. 6: 157-167. ACKNOWLEDGEMENTS Miller, P. J., 1988. New species of Corygobius, Thorogobius and Wheelerigobius from West Africa (Teleostei: Gobiidae). We thank NUS for funding ISC from the SPRINT program; Journal of Natural History, 22: 1245-1262. Peter K. L. Ng and Abdul Latiff, for collecting the first Ng, H. H., H. H. Tan & K. K. P. Lim, 1999. The inland fishes of specimen of Stiphodon from Pulau Tioman; Kelvin Lim, for Pulau Tioman, Peninsular Malaysia. In: Sodhi, N. S., H. S. Yong helping with field work. We are very grateful for R. Watson & P. K. L. Ng (eds.), The biodiversity of Pulau Tioman, for providing his many comprehensive papers for Indo-Pacific Peninsular Malaysia. Raffles Bulletin of Zoology, Supplement No. 6: 169-187. Stiphodon and valuable discussion for this genus. We also extend our thanks to the two referees for constructive Watson, R. E. & I-S. Chen, 1998. Freshwater gobies of the genus comments for enriching this manuscript. This work had been Stiphodon from Japan and Taiwan (Teleostei: Gobiidae: Sicydiini). Aquarium Journal of Ichthyology and Aquatic partially supported by research grant RP3982324 to Peter K. Biology, 3: 55-66. L. Ng from the National University of Singapore and also partially supported by NSC 2001-2002 research grants in Watson, R. E. & M. Kottelat, 1995. Gobies of the genus Stiphodon Taipei and the West Pacific research project from Academia from Leyte, Philippines, with descriptions of two new species (Teleostei: Gobiidae: Sicydiinae). Ichthyological Exploration Sinica to ISC. of Freshwaters, 6: 1-16. Weber, M., 1895. Fische von Ambon, Java, Thursday Island, dem Burnett-Fluss und von der Süd-Küste von Neu-Guinea. In: LITERATURE CITED Zoologische Forschungsreisen in Australien und dem malayischen Archipel; mit Unterstützung des Herrn Dr. Paul Alfred, E. R., 1966. Observations on the fauna of Pulau Tioman von Ritter ausgeführt. Jahren 1891-1893 von Dr. Richard and Pulau Tulai.-8. Fishes of stream drainages. Bulletin of the Semon. 5: 259-276. National Museum, Singapore, 34: 97-103. Wu, H. L. & Y. Ni, 1985. Suborder Gobioidei. In Anonymous Chen, I.-S. & K. T. Shao, 1996. A taxonomic review of the gobiid (ed.), The Freshwater and Estuaries Fishes of Hainan Island. fish genus, Rhinogobius Gill, 1859 from Taiwan, with Guangdong Science and Technology Press, Guangzhou. Pp. 259- descriptions of three new species. Zoological Studies, 35: 200- 314. (In Chinese). 214.
242 THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 243-249 Date of Publication: 31 Dec.2005 © National University of Singapore
AMBLYCEPS CARINATUM, A NEW SPECIES OF HILLSTREAM CATFISH FROM MYANMAR (TELEOSTEI: AMBLYCIPITIDAE)
Heok Hee Ng Fish Division, Museum of Zoology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, Michigan 48109-1079, USA.
ABSTRACT. Ð Amblyceps carinatum, a new species of hillstream catfish from the middle Irrawaddy River drainage in Myanmar is described here. It can be distinguished from congeners in having a unique combination of the following characters: the adipose fin in the form of a long, low ridge (length of adipose-fin base 37.5Ð 44.6% SL), a smooth posterior edge of the pectoral spine, poorly developed pinnate processes along the median caudal-fin rays, a deeply forked caudal fin, body depth at anus 13.1Ð15.0% SL, head depth 10.6Ð 13.6% SL, eye diameter 9.4Ð11.8% HL vs. 8.3), 39Ð40 vertebrae, and a uniform body coloration. The valid species of Amblyceps are listed. The status of A. apangi and A. arunachalensis are discussed; the former species is considered valid and the latter a junior synonym of A. mangois. Amblyceps mucronatum is also recorded from the Irrawaddy River drainage for the first time.
KEY WORDS. Ð Siluriformes, Sisoroidea, Irrawaddy River.
INTRODUCTION Among the thirteen species, two are known to occur in Myanmar or in river drainages that flow through Mynamar Amblyceps Blyth, 1858, is a group of catfishes typically (Ng & Kottelat, 2000), viz. A. murraystuarti and A. inhabiting fast flowing hillstreams or faster-flowing stretches mucronatum. Recent collections have uncovered a third of larger rivers, and is distributed throughout South and species, which is described in this study as Amblyceps Southeast Asia. Its members are diagnosed by an epiphyseal carinatum, new species. commissure of the supraorbital sensory canals not passing through the epiphyseal bar, fifth ceratobranchial expanded MATERIAL AND METHODS posteromedially, the presence of pinnate processes along the median caudal-fin rays, and the presence of triangular Measurements were made point to point with dial calipers fenestrae immediately medial to both inner and outer and data recorded to tenths of a millimeter. Counts and mandibular barbels (Chen & Lundberg, 1995). Externally, measurements were made on the left side of specimens Amblyceps species are easily distinguished by the presence whenever possible. Subunits of the head are presented as of the pinnate processes along the median caudal-fin rays percentage proportions of head length (% HL). Head length (although these processes may be poorly developed in some and measurements of body parts are given as percentage species), a prominent cup-like skin flap above the base of proportions of standard length (% SL). Measurements follow the pectoral spine, adipose fin largely separate from caudal Ng & Kottelat (2000), and institutional abbreviations follow fin and, with three exceptions, the presence of a deeply forked Eschmeyer (1998). caudal fin. Following Ng & Kottelat (2000) and Ng (2001), eleven valid species of Amblyceps are recognized, viz. A. TAXONOMY mangois (Hamilton, 1822), A. laticeps (M’Clelland, 1842), A. caecutiens Blyth, 1858, A. murraystuarti Chaudhuri, 1919, Amblyceps carinatum, new species A. foratum Ng & Kottelat, 2000, A. mucronatum Ng & (Fig. 1) Kottelat, 2000, A. platycephalus Ng & Kottelat, 2000, A. serratum Ng & Kottelat, 2000, A. tenuispinis Blyth, 1860, A. Material examined. Ð Holotype - UMMZ 245588, 34.7 mm SL, variegatum Ng & Kottelat, 2000 and A. macropterus Ng, Myanmar: Kachin, Irrawaddy drainage, hillstreams at Tonpan 2001. A twelfth species, A. apangi Nath & Dey, 1989, is village, on road from Myitkyina to Tanai, coll. Hain Zin & Hlaing Hlaing Oo, 20 Feb.2005. recognized as valid in this study.
243 Ng: New species of Amblyceps from Myanmar
Paratypes Ð UMMZ 245490, 3 ex., 25.8Ð36.0 mm SL, data as for Head depressed, with expanded cheek muscles imparting holotype. slightly swollen appearance in front of opercular region. Anterior and posterior nares separated only by base of nasal Diagnosis. Ð Amblyceps carinatum can be distinguished from barbel. Gill openings broad, extending from immediately congeners in having the adipose fin in the form of a long, ventral to post-temporal to isthmus. Dorsal surface of head low ridge (vs. blade-like, length of adipose-fin base 37.5Ð covered with thick, tuberculate skin. Eye ovoid, horizontal 44.6% SL vs. 13.6Ð33.5; Fig. 2). It further differs from axis longest; located entirely in dorsal half of head and congeners on the Indian subcontinent with a deeply forked subcutaneous. caudal fin (A. mangois, A. tenuispinis, and A. macropterus) in having poorly developed (vs. well developed) pinnate Barbels in four pairs. Maxillary barbel long and slender, processes along the median caudal-fin rays (Fig. 1 in Ng & extending to middle of pectoral-fin rays. Nasal barbel slender, Kottelat, 2001). Amblyceps carinatum can be further extending to opercle. Inner mandibular-barbel origin close distinguished from A. apangi, A. laticeps and A. murraystuarti to midline, extending to middle of pectoral-fin base. Outer in having a deeply forked (vs. truncate or emarginate) caudal mandibular barbel originatesing posterolateral of inner fin, and from A. caecutiens in having a deeper body (13.1Ð mandibular barbel, extending to middle of pectoral-fin rays. 15.0 vs. 9.8Ð10.7) and larger eyes (9.4Ð11.8% HL vs. 2.0Ð 3.4). It further differs from A. foratum in having larger eyes Mouth terminal, with lower jaw very slightly longer than (9.4Ð11.8% HL vs. 5.5Ð8.5), from A. platycephalus in having upper; anterior edge barely visible dorsally. Lips fleshy and a deeper head (10.6Ð13.6% SL vs. 8.3), and body (13.1Ð slightly fimbriated, with very well developed rictal fold 15.0% SL vs. 11.5), from A. serratum in having a smooth present below anterior border of eye. Oral teeth small and (vs. serrated) posterior edge of the pectoral spine and more villiform, in irregular rows on all tooth-bearing surfaces. vertebrae (39Ð40 vs. 34Ð36), and from A. variegatum in Premaxillary teeth in single broad semilunate band. Dentary having larger eyes (9.4Ð11.8% HL vs. 5.3Ð6.9) and a uniform teeth in two narrow crescentic bands separated at midline. (vs. mottled) body coloration. Dorsal fin located above anterior one-third of body, with I,6 Description. Ð Biometric data in Table 1. Head depressed, (4) rays; fin margin convex; spine very short and straight, body cylindrical. Anus and urogenital openings located at encased in thick skin and with fluid-filled cavity imparting vertical through middle of adpressed pelvic fin. Skin tuberculate. Lateral line midlateral, apparently extending to base of caudal fin; visible externally anterior to, but not posterior to, middle of dorsal-fin base. Only individual laterosensory pores visible externally posterior to middle of dorsal-fin base. Vertebrae 19+20=39 (1), 20+19=39 (1), or 20+20=40* (2).
Fig. 2. Adipose fins of: a. Amblyceps carinatum, holotype, UMMZ 245588, 34.7 mm SL; b. other Amblyceps species (A. mucronatum, Fig. 1. Amblyceps carinatum, holotype, UMMZ 245588, 34.7 mm UMMZ 245497, 41.2 mm SL illustrated). Arrows indicate origin SL, Myanmar: hillstreams at Tonpan village. of adipose fin and scale bar represents 10 mm.
244 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 1. Biometric data for Amblyceps carinatum (n=4)
Holotype Range Mean±SD Standard length 34.7 25.8Ð36.0 %SL Predorsal length 32.3 30.4Ð32.3 31.5±0.79 Preanal length 63.4 63.4Ð66.1 64.3±1.24 Prepelvic length 49.9 47.7Ð52.5 50.6±2.27 Prepectoral length 21.3 18.7Ð22.1 20.7±1.45 Length of dorsal-fin base 14.4 13.4Ð15.5 14.6±0.90 Length of anal-fin base 21.6 19.8Ð23.3 21.3±1.56 Pelvic-fin length 11.5 11.1Ð13.4 11.8±1.08 Pectoral-fin length 19.3 17.7Ð19.3 18.3±0.73 Caudal-fin length 28.5 25.0Ð28.5 26.5±1.80 Length of adipose-fin base 39.8 37.5Ð44.6 40.8±2.98 Dorsal to adipose distance 10.7 7.8Ð10.7 9.1±1.48 Length of caudal peduncle 15.6 14.5Ð16.7 15.6±0.90 Depth of caudal peduncle 14.4 12.7Ð14.4 13.4±0.79 Body depth at anus 15.0 13.1Ð15.0 13.7±0.90 Head length 24.5 24.5Ð26.4 25.2±0.86 Head width 17.9 16.3Ð17.9 17.1±0.68 Head depth 12.7 10.6Ð13.6 12.2±1.27 %HL Snout length 32.9 29.4Ð36.2 33.2±2.88 Interorbital distance 31.8 31.8Ð40.6 34.2±4.35 Eye diameter 11.8 9.4Ð11.8 10.5±1.01 Nasal barbel length 70.6 70.6Ð78.1 74.6±3.12 Maxillary barbel length 111.8 111.8Ð129.4 118.2±7.71 Inner mandibular barbel length 76.5 46.9Ð85.9 68.3±16.88 Outer mandibular barbel length 94.1 84.1Ð111.8 96.0±11.54
somewhat bulbous appearance along entire length. Adipose Dorsal and anal fins brown for proximal half, distal half fin long and low, anterior origin at vertical through pelvic- hyaline. Pectoral and pelvic fins hyaline. Adipose fin brown, fin origin. Caudal fin strongly forked, with upper lobe longer with hyaline distal margin. Caudal fin with brown fin rays than lower (but less than 1.5 times length of lower lobe) and and hyaline interradial membrane. Nasal and maxillary i,8,7,i (4) principal rays. Procurrent rays symmetrical and barbels brown at base, off white for almost entire length, extending considerably anterior to fin base. Anal-fin base mandibular barbels off white. ventral to adipose-fin origin at distance about half length of anal-fin base. Anal fin with straight anterior margin and Distribution. - Known only from the middle part of the straight or slightly concave posterior margin, spanning Irrawaddy River drainage in Myanmar (Fig. 3). posterior half of adipose-fin base and with v,9 (1), v,10* (2) or vi,10 (1) rays. Pelvic-fin origin midway between verticals Etymology. - From the Latin carina (keel), in reference to through posterior end of dorsal-fin base and anal-fin origin. the long, low adipose fin of this species. Pelvic fin with convex margin and i,5 (4) rays; tip of adpressed fin not reaching anal-fin origin. Pectoral fin narrow, with I,6* (3) or I,6,i (1) rays; posterior fin margin strongly convex. DISCUSSION Pectoral spine short and straight, with smooth anterior and posterior margins. Spine encased in thick skin, with fluid- The shape of the adipose fin diagnoses A. carinatum from all filled cavity imparting somewhat bulbous appearance along other congeners. It is the only character that reliably proximal half. distinguishes it from the sympatric A. mucronatum. Although A. carinatum seemingly has a longer head (24.5Ð26.4% SL Coloration. - In 70% ethanol: Dorsal and lateral surfaces of vs. 20.3Ð22.6) and shorter caudal peduncle (14.5Ð16.7% SL head, and body brown, fading to off white on ventral surfaces. vs. 18.3Ð21.8) than A. mucronatum, the differences are
245 Ng: New species of Amblyceps from Myanmar obscured by the size disparity of the material between the be determined. Therefore, it cannot be demonstrated two species examined (no specimen above 40 mm SL for A. conclusively that the available material of A. carinatum are carinatum and no specimen under 40 mm SL for A. adults and that size is a useful character in distinguishing A. mucronatum). Therefore, it is difficult to discount ontogeny carinatum from A. mucronatum. as accounting for the biometric differences observed. Biplots of caudal peduncle length and head length against SL (Fig. Recent studies of Amblyceps taxonomy (Ng & Kottelat, 2000; 4) graphically illustrate this point. Until a larger series of Ng, 2001) have not mentioned two nominal species described material can be studied to exclude the effects of ontogeny in by Nath & Dey (1989): A. apangi Nath & Dey, 1989 (type explaining the biometric differences, the lengths of the head locality: Dikrong River; the Dikrong is a tributary of the and caudal peduncle are not used in diagnosing the two Brahmaputra River) and A. arunachalensis Nath & Dey, 1989 species here. (type locality: Dikrong River). Specimens of an Amblyceps species with a truncate caudal fin from the Brahmaputra River The same argument (i.e. ontogeny) can be applied to discount drainage closely matches the description of A. apangi, and using the size and shape of the adipose fin as a diagnostic is hereby referred to this species (a photograph of A. apangi character, but there is enough circumstantial evidence to is provided in Fig. 6). Amblyceps apangi is most similar to suggest its utility as a diagnostic character. In species where A. murraystuarti, which is known from the upper Irrawaddy a large series (spanning the interval between the sizes of both River drainage. Both species differ from all other congeners the series of A. carinatum and A. mucronatum) is available in having truncate (vs. deeply forked or emarginate) caudal (A. mangois, A. laticeps, A. murraystuarti, and A. foratum), fins and more vertebrae (45Ð48 vs. 34Ð41). Amblyceps apangi the size and, in particular, the general shape and relative length differs from A. murraystuarti in having a longer head (20.2Ð of the adipose fin does not change much during ontogeny (as 24.6% SL vs. 18.3Ð20.5) and more vertebrae (47Ð48 vs. 45Ð shown in Fig. 5 using A. mangois as an example), thus making 46). In specimens above ca. 50 mm SL, A. apangi has a this character useful in diagnosing species of Amblyceps. wider head (13.8Ð15.2% SL vs. 10.6Ð12.7; Fig. 7), although this difference is not clearly evident between individuals of The gonads of two paratypes of A. carinatum were examined. both species below this size. Furthermore, the live colors of The largest paratype (UMMZ 245490, 36.0 mm SL) is a female in which the ovaries contained immature oocytes corresponding to the stage defined as initial maturation by dos Santos et al. (2004). The gonads of the other paratype (UMMZ 245490, 28.3 mm SL) were too small for its sex to
Fig. 4. Scatterplots of: a. length of caudal peduncle (CPL) and b. Fig. 3. Collecting locality for the type series of Amblyceps head length (HL) plotted against standard length for Amblyceps carinatum. carinatum and A. mucronatum.
246 THE RAFFLES BULLETIN OF ZOOLOGY 2005 the two species are different, with A. apangi being bronze (in which the type locality of A. mucronatum is located) yellow and A. murraystuarti gray. (Rainboth, 1991; 1996), this pattern of distribution is unusual. It is possible that the populations of A. mucronatum in the I was unable to examine material clearly referable to A. Mekong, Salween and Irrawaddy River drainages represent arunachalensis, but I was unable to find any distinct more than one species, but I was unable to find distinct differences between material identified as A. mangois from characters to separate the populations. A thorough study with the Brahmaputra River drainage and both the original more material is necessary to completely resolve this question description of A. arunachalensis, and the photograph of this and until such a study is possible, I consider the Mekong, species provided in Nath & Dey (2000). Both A. Salween and Irrawaddy populations of A. mucronatum arunachalensis and A. mangois share a caudal fin with greatly conspecific for the time being. unequal lobes, with the upper lobe much longer than the lower (upper lobe about 1.5 times length of lower). This character Comparative material. Ð Amblyceps mangois: UMMZ is otherwise only seen in A. tenuispinis, another Indian 2445888, 1 ex., 29.4 mm SL, India: West Bengal, Dolong species, but both A. arunachalensis and A. mangois have River in the vicinity of Shildanga, 26°23'13.0"N deeper bodies than A. tenuispinis (11.9Ð13.5% SL for A. 89°14'31.0"E; UMMZ 244680, 3 ex., 17.2Ð30.0 mm SL, mangois vs. 9.7Ð11.6 for A. tenuispinis). The body depth of India: West Bengal, Schutunga River at Ansole, 26°22'24.0"N A. arunachalensis is 15.6Ð16.0% SL as calculated from data 89°11'17.0"E; UMMZ 244752, 8 ex., 28.8Ð47.1 mm SL, West in the original description (which is unusually high for Bengal, Raidak I River at Shipra, just outside Buxa Tiger Amblyceps species, as all other congeners have body depths Reserve, 26°31'12.0"N 88°42'32.0"E; UMMZ 244760, 10 ex., under 15% SL), but is only 13.6% SL as calculated from the 26.6Ð51.3 mm SL, India: West Bengal, Jorai and Raidak I illustration in the original description and 12.4% SL as Rivers, 26°31'N 89°47'E; UMMZ 244866, 15 ex., 29.0Ð43.0 calculated from the illustration in Nath & Dey (2000). Given mm SL, India: West Bengal, Tista River at Tista Barrage, the probable inaccuracy of the data in the original description 26°45'10.0"N 88°34'11.0"E. and the overall similarity in physiognomy of the two nominal species (comparing A. mangois with the photographs of A. Amblyceps laticeps: UMMZ 208806, 15 ex., 17.5Ð42.1 mm arunachalensis in Nath & Dey, 2000), A. arunachalensis is SL, Bangladesh: Chittagong, Sylhet district, Piyain Gang here considered a junior synonym of A. mangois. River at Songram Punji, 400 m downstream of Indian border, 25°11'N 92°1'E; UMMZ 244765, 1 ex., 41.5 mm SL, India: Amblyceps mucronatum has been previously reported from West Bengal, market at Barobisha, 26°40'N 89°40'E; UMMZ the Mekong and the Salween River drainages (Ng & Kottelat, 244865, 1 ex., 37.6 mm SL, India: West Bengal, Tista River 2000), and the occurrence of this species in the Irrawaddy at Tista Barrage, 26°45'10.0"N 88°34'11.0"E. River drainage is reported for the first time in this study. Given that the freshwater fish fauna of the Irrawaddy River Amblyceps caecutiens: ZRC 43717, 4 ex., 39.2Ð56.3 mm SL, drainage is much more similar to that of the Ganges- Myanmar: Bago. Brahmaputra system than it is to the Mekong River drainage Amblyceps tenuispinis: ZRC 1563, 6 ex., 46.5Ð62.1 mm SL, India: Uttar Pradesh, Naini Tal district, Gola River at Kathgodam.
Amblyceps murraystuarti: ZSI F9736/1, holotype, 72.4 mm SL, Myanmar: Putao plains; NRM 40944, 5 ex., 90.8Ð128.3 mm SL, Myanmar: Kachin, Ma Kyaww Wa Chaung and its tributary Nan Hto Chaung in Putao, 27°19'44.0"N 97°22'36.0"E; UMMZ 245489, 6 ex., 34.4Ð81.9 mm SL, Myanmar: Kachin, Myitkyina district, hillstreams at Tonpan village, on road from Myitkyina to Tanai.
Amblyceps apangi: BMNH 1932.4.22.2Ð5, 3 ex., 28.1Ð30.8 mm SL; RMNH 16093, 2 ex., 38.6Ð51.6 mm SL, India: West Bengal, Sevoke stream, Tista Valley; CAS-SU 69784, 1 ex., 42.8 mm SL, India: West Bengal, Kalimpong Duars and Siliguri Terai; UMMZ 244754, 7 ex., 27.8Ð81.0 mm SL,
Fig. 5. Adipose fins of Amblyceps mangois: a. UMMZ 244680, 17.2 mm SL and b. UMMZ 244760, 50.8 mm SL, showing little Fig. 6. Amblyceps apangi, UMMZ 244754, 44.6 mm SL, India: change in relative length and general shape. West Bengal, Raidak I River at Shipra.
247 Ng: New species of Amblyceps from Myanmar
India: West Bengal, Raidak I River at Shipra, just outside Po mouth at confluence Nam Xone, 18°58"N 102°28"E. Buxa Tiger Reserve, 26°31'12.0"N 88°42'32.0"E. Amblyceps variegatum: ZRC 43719, holotype, 62.9 mm SL; Amblyceps foratum: ZRC 43718, holotype, 63.9 mm SL; ZRC CMK 14560, 8 paratypes, 43.5Ð60.9 mm SL, Thailand: 38733, 14 paratypes, 19.8Ð54.1 mm SL, Malaysia: Kanchanaburi, Nam Kung, upstream of Ban Huai Pak Khung, Terengganu, Sungai Brang just outside Sekayu waterfall park, 14°37'20.0"N 98°31'24.0"E. 4°57'51.0"N 102°57'46.0"E. Amblyceps macropterus: UMMZ 187411, holotype, 47.4 mm Amblyceps mucronatum: ZRC 41323, holotype, 42.9 mm SL, SL, Pakistan: Baluchistan, Las Bela district, Hab River at Thailand: Loei, Huang River tributary at 17°16'0.9"N Hinidan. 101°8'29.2"E; CMK 5027, 4 paratypes, 45.3Ð56.0 mm SL, Thailand: Loei, Nam Man, 2 km upstream of Amphoe Dan Sai, 17°16'N 101°9"E; CMK 5539, 1 paratype, 72.9 mm SL, ACKNOWLEDGMENTS Thailand: Tak, Huay Khokhe, approximately 6 km S of 60 km mark on TakÐMae Sot highway; CMK 14667, 1 paratype, I am grateful to Tin Win for help in obtaining material and 61.0 mm SL, Thailand: Tak, Mae Nam Charao at and the following for permission to examine material under their upstream of Ban Makham Porn, 16°58'33.0"N 98°39'4.0"E; care: Darrell Siebert (BMNH), David Catania (CAS), Maurice UMMZ 245497, 11 ex., 41.2Ð66.5 mm SL; Myanmar: Kottelat (CMK), Sven Kullander (NRM), Martien van Oijen Yangon, lower Kanazou Njaun Chaung, tributary of (RMNH), Douglas Nelson (UMMZ), Kelvin Lim (ZRC) and Ngamoyeik Chaung, at Gyongon village, 27 km from Hlegu A. K. Karmakar (ZSI). Funding from the Carl L. and Laura on YangonÐBago road. C. Hubbs Research Fellowship from the Museum of Zoology, University of Michigan and the All Catfish Species Inventory Amblyceps platycephalus: ZRC 42745, holotype, 60.2 mm (NSF DEB-0315963) are gratefully acknowledged. SL, Thailand: Mae Hong Son, Nam Mae Sa-Nga at Ban Yang Top Sole, immediately downstream of Nam Tok Pa Sua, 19°29'0.0"N 97°57'50.0"E. LITERATURE CITED
Amblyceps serratum: UMMZ 234708, holotype, 35.5 mm SL, Blyth, E., 1858. Report of Curator, Zoological Department, for May, Cambodia: Stung Treng, shallow channel across S end of 1858. Journal of the Asiatic Society of Bengal, 27: 267Ð290. Kaoh Han, 14 km NE of Stung Treng, 13°37"N 106°3"E; Blyth, E., 1860. Report on some fishes received chiefly from the UMMZ 235399, 1 ex., 43.4 mm SL, Laos: Champasak, Sitang River and its tributary streams, Tenasserim Provinces. Mekong River at Ban Hang Khone, just downstream of Khone Journal of the Asiatic Society of Bengal, 27: 138Ð174. Falls; UMMZ 238843, 1 ex., 36.0 mm SL, Laos: Attapeu, Chaudhuri, B. L., 1919. Report on a small collection of fish from Xe Kamen at crossing on road to Pleiku, 14°54"N 107°9"E; Putao (Hkamti Long) on the northern frontier of Burma. Records UMMZ 240006, 1 ex., 38.7 mm SL, Laos: Vientiane, Nam of the Indian Museum, 16: 271Ð287, Pl. 22.
Fig. 7. Dorsal views of heads of: a. Amblyceps apangi, UMMZ 244754, 81.0 mm SL; b. A. murraystuarti, UMMZ 245489, 81.1 mm SL.
248 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Chen, X.-P. & J. G. Lundberg, 1995. Xiurenbagrus, a new genus Nath, P. & S. C. Dey, 2000. Fish and Fisheries of North Eastern of amblycipitid catfishes (Teleostei: Siluriformes), and India (Arunachal Pradesh). Narendra Publishing House, Delhi. phylogenetic relationships among the genera of Amblycipitidae. 217 pp. Copeia, 1995(4): 780Ð800. Ng, H. H., 2001. Amblyceps macropterus, a new species of dos Santos, J. E., N. Bazzoli, E. Rizzo & G. B. Santos, 2004. amblycipitid catfish (Osteichthyes: Amblycipitidae) from Reproduction of the catfish Iheringichthys labrosus (Lütken) Pakistan. Ichthyological Exploration of Freshwaters, 12: 201Ð (Pisces, Siluriformes) in Furnas reservoir, Minais Gerais, Brazil. 204. Revista Brasileira de Zoologia, 21: 193Ð200. Ng, H. H. & M. Kottelat, 2000. A review of the genus Amblyceps Eschmeyer, W. N., 1998. Catalog of Fishes. California Academy (Osteichthyes: Amblycipitidae) in Indochina, with descriptions of Sciences, San Francisco. 2905 pp. of five new species. Ichthyological Exploration of Freshwaters, Hamilton, F., 1822. An account of the fishes found in the river 11: 335Ð348. Ganges and its branches. Archibald Constable, Edinburgh and Rainboth, W. J., 1991. Cyprinid fishes of Southeast Asia. In: Hurst, Robinson, London. 405 pp. Winfield, I. & J. S. Nelson (eds.) Cyprinid fishes: systematics, M’Clelland, J., 1842. On the fresh-water fishes collected by William biology and exploitation. Chapman & Hall, London. Pp. 156Ð Griffith, Esq., F. L. S. Madras Medical Service, during his travels 210. under the orders of the Supreme Government of India, from Rainboth, W. J., 1996. The taxonomy, systematics, and 1835 to 1842. Calcutta Journal of Natural History, 2: 560Ð589, zoogeography of Hypsibarbus, a new genus of large barbs Pls. 15, 18, 20, 21. (Pisces, Cyprinidae) from the rivers of southeastern Asia. Nath, P. & S. C. Dey, 1989. Two new fish species of the genus University of California Publications in Zoology, 129: 1Ð199. Amblyceps Blyth from Arunachal Pradesh, India. Journal of the Assam Science Society, 32: 1Ð6.
249
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 251-255 Date of Publication: 31 Dec.2005 © National University of Singapore
GLYPTOTHORAX EXODON, A NEW SPECIES OF RHEOPHILIC CATFISH FROM BORNEO (TELEOSTEI: SISORIDAE)
Heok Hee Ng Fish Division, Museum of Zoology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, Michigan 48109-1079, USA.
Ike Rachmatika Division of Zoology, Research and Development Center for Biology, Indonesian Institute of Sciences, Gedung Widyasatwaloka, Jalan Raya Jakarta km. 46, Cibinong, Bogor 16911, Indonesia.
ABSTRACT. Ð Glyptothorax exodon, a new species of rheophilic catfish from Borneo is described here. It can be distinguished from Sundaic congeners except G. platypogonides and G. siamensis in having a slender body (13.8Ð15.8% SL vs. 17.3Ð20.9). It differs from G. platypogonides in having a deeper caudal peduncle (6.8Ð7.3% SL vs. 5.7Ð6.7) and more exposed premaxillary teeth when the mouth is closed and from G. siamensis in having a larger eye (10.4Ð13.2% HL vs. 7.5Ð9.1) and a mottled color pattern lacking distinct pale stripes (vs. a uniform dark brown with distinct pale midlateral and mid-dorsal stripes).
KEY WORDS. Ð Rheophilic catfish, Kalimantan Barat, Glyptothorax, Kapuas River.
INTRODUCTION MATERIAL AND METHODS
Glyptothorax is the most diverse and the most widely Measurements were made point to point with dial calipers distributed sisorid catfish group, with 86 nominal species and data recorded to tenths of a millimeter. Counts and found in Asia Minor (in the Tigris and Euphrates River measurements were made on the left side of specimens drainages) westwards to East Asia (to the Yangtze River whenever possible. Subunits of the head are presented as drainage) and southwards to Southeast Asia. They typically proportions of head length (HL). Head length and inhabit fast flowing hillstreams or faster-flowing stretches of measurements of body parts are given as proportions of larger rivers, and are distinguished by their distinctive thoracic standard length (SL). Measurements follow Ng & Dodson adhesive apparatus, comprising an elliptical field of folded (1999), and institutional abbreviations Eschmeyer (1998). longitudinal pleats of skin. They are additionally diagnosed by a detached distal portion of the premaxilla and long and thin lateral arms of the vomer that extend underneath the entire TAXONOMY length of the articular process of the lateral ethmoid (de Pinna, 1996). Despite recent phylogenetic studies (de Pinna, 1996), Glyptothorax exodon, new species the monophyly of the group has yet to be rigorously tested. (Fig. 1)
During ichthyological surveys in the Kapuas River drainage Bagarius bagarius (non Hamilton) Vaillant, 1902: 72. in western Borneo (Roberts, 1989; Ng & Rachmatika, 1999), Glyptothorax platypogonoides (non Bleeker) Roberts, 1989: 136, specimens identified as G. platypogonides (Bleeker, 1855) Fig. 104. were collected. Comparison of this material with Sumatran Glyptothorax aff. platypogonoides Ng & Rachmatika, 1999: 182, Fig. 8B. material of G. platypogonides (also collected during recent surveys; Tan & Ng, 2000) revealed that the Bornean material Material examined. Ð Holotype - MZB 9940, 57.5 mm SL, Borneo: represented a distinct undescribed species, which is described Kalimantan Barat, Sungai Tekelan, coll. I. Rachmatika, 5 Sep.1997. below as Glyptothorax exodon, new species.
251 Ng & Rachmatika: New species of Glyptothorax from Borneo
Paratypes Ð BMNH 19823.29.190 (1), 63.2 mm SL; CAS 49419 Head depressed and narrow, triangular when viewed laterally (1), 59.0 mm SL, RMNH 28907 (1), 57.2 mm SL, Borneo: and with acute snout margin when viewed from above. Snout Kalimantan Barat, mainstream of Sungai Pinoh, 20Ð60 km upstream prominent. Anterior and posterior nares large and separated from Nangapinoh, coll. T. R. Roberts & S. Wirjoatmodjo, 22Ð26 only by base of nasal barbel. Gill openings broad, extending Jul.1976; RMNH 7845 (1), 60.2 mm SL, Borneo: Kalimantan Barat, from immediately ventral to posttemporal to isthmus. Bony Nanga Raun, coll. J. Büttikofer, 1897; MZB 9412 (2), 55.1–60.0 mm SL, Borneo: Kalimantan Barat, Embaloh, Sungai Jaket, coll. I. elements of dorsal surface of head covered with thick, Rachmatika et al., 19 Sep.1997. tuberculate skin. Eye ovoid, horizontal axis longest; located entirely in dorsal half of head. Orbit with free margin. Diagnosis. Ð Glyptothorax exodon be distinguished from Sundaic congeners except G. platypogonides and G. siamensis Barbels in four pairs. Maxillary barbel long and slender, in having a slender body (13.8Ð15.8% SL vs. 17.3Ð20.9). It extending to middle of pectoral-fin base. Nasal barbel slender, differs from G. platypogonides in having a deeper caudal extending to midway between its base and anterior orbital peduncle (6.8Ð7.3% SL vs. 5.7Ð6.7) and more exposed margin. Inner mandibular-barbel origin close to midline, premaxillary teeth when the mouth is closed (Fig. 2) and from extending to two thirds of distance between its base and that G. siamensis in having a larger eye (10.4Ð13.2% HL vs. 7.5Ð of pectoral spine. Outer mandibular barbel originating 9.1) and a mottled color pattern lacking distinct pale stripes posterolateral of inner mandibular barbel, extending to base (vs. a uniform dark brown with distinct pale midlateral and of pectoral spine. mid-dorsal stripes). Mouth inferior, premaxillary tooth band almost wholly Description. Ð Morphometric data in Table 1. Head exposed when mouth is closed. Oral teeth small and villiform, depressed, body moderately compressed. Dorsal profile rising in irregular rows on all tooth-bearing surfaces. Premaxillary evenly from tip of snout to origin of dorsal fin, then sloping teeth in single broad semilunate band. Dentary teeth in two gently ventrally from there to end of caudal peduncle. Ventral narrow crescentic bands separated at midline. profile flat to anal-fin base, then sloping gently dorsally from there to end of caudal peduncle. Anus and urogenital openings located at vertical through middle of adpressed pelvic fin. Skin prominently tuberculate, with ovoid tubercles particularly visible on sides of body. Lateral line complete and midlateral. Vertebrae 17+18=35 (1), 17+19=36 (2), 18+18=36 (2) or 18+19=37 (2).
Fig. 2. Ventral views of heads of: a. Glyptothorax exodon, RMNH Fig. 1. Glyptothorax exodon, paratype, RMNH 28907, 57.2 mm 28907, paratype, 57.2 mm SL; b. G. platypogonides, ZRC 41588, SL; Borneo: Sungai Pinoh. Dorsal, lateral and ventral views. 56.0 mm SL. Scale bar represents 5 mm.
252 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Dorsal fin located above anterior third of body, with I,6 (7) band in middle. Anal fin with brown base and brown rays; fin margin convex; spine short and gently curved. melanophores forming V-shaped subdistal mark on anterior Adipose fin with anterior margin concave and posterior half of fin; rest of fin hyaline. Adipose fin brown to brownish margin angular. Caudal fin strongly forked, with equal upper gray, with hyaline distal margin. Caudal fin brown to and lower lobes and i,7,8,i (7) principal rays. Procurrent brownish gray , tips and posterior margins of lobes hyaline. rays symmetrical and extending only slightly anterior to fin Maxillary barbels brown to brownish gray dorsally, beige base. Anal-fin base ventral to adipose-fin origin. Anal fin ventrally. with straight anterior margin and straight or slightly concave posterior margin ; with iv,8 (2) or iv,9 (5) rays. Pelvic-fin Distribution. Ð Known from the Kapuas River drainage in origin immediately posterior to vertical through posterior end western Borneo (Fig. 3). of dorsal-fin base. Pelvic fin with slightly convex margin and i,5 (7) rays; tip of adpressed fin not reaching anal-fin Etymology. Ð From the Greek exo- (out) and odontos (tooth), origin. Pectoral fin with I,8 (7) rays; posterior fin margin in reference to the exposed premaxillary teeth. slightly concave; anterior spine margin smooth, posterior margin with 10Ð12 serrations. Thoracic adhesive apparatus Habitat. Ð Glyptothorax exodon is found in swift, clear present, located on belly and consisting of logitudinal pleats streams with a substrate consisting of gravel, cobbles or large of skin in narrow elliptical field. Adhesive apparatus with rocks (Roberts, 1989). Other fish found in this habitat include median depression on posterior half and extending from just Barbonymus collingwoodi, Crossocheilus spp., behind gill opening to level of base of posteriormost pectoral- Cyclocheilichthys repasson, Epalzeorhynchos kalopterus, fin ray. Garra borneensis, Paracrossocheilus spp., Puntius spp., Rasbora spp., Schismatorhynchos heterorhynchus, Tor Coloration. Ð In 70% ethanol: Dorsal and lateral surfaces of tambroides, Gastromyzon contractus, Nemacheilus head, and body brown to brownish gray, fading to beige on kapuasensis, Glyptothorax major, Hemibagrus cf. nemurus, ventral surfaces. A dark, diffuse midlateral stripe running Doryichthys martensii, Mastacembelus spp. and Auriglobus along lateral line; lateral line without such pigmentation in amabilis. some individuals, causing it to show up as paler stripe within darker stripe. Body with several long, elliptical pale spots: first above lateral line and spanning region between dorsal DISCUSSION and adipose fins, second on anterior half of caudal peduncle above lateral line and third on anterior half of caudal peduncle Given the wide distribution of Glyptothorax species and the below lateral line. Dorsal fin with brown base and broad large number (87) of nominal species, it would not be realistic subdistal triangular brown mark. Pectoral fins hyaline, with to compare G. exodon with all congeners without making it brown melanophores forming a diffuse broad, transverse band difficult for the reader to follow. However, Southeast Asian subdistally. Pelvic fin hyaline, with faint transverse brown Glyptothorax species have restricted distributions (HHN, unpub. data), which means that species found in a particular region are not likely to occur outside of it; it is therefore more meaningful to restrict our comparisons of G. exodon to only those species that occur in Sundaic Southeast Asia.
The identity of the Sundaic Southeast Asian species of Glyptothorax remains confused (Roberts, 1989; Kottelat &
Fig. 4. Scatterplot of depth of caudal peduncle (CPD) plotted against Fig. 3. Distribution map of G. exodon. standard length for G. exodon and G. platypogonides.
253 Ng & Rachmatika: New species of Glyptothorax from Borneo
Table 1. Biometric data for Glyptothorax exodon (n=7)
Holotype Range Mean±SD %SL Head length 26.8 25.7Ð29.2 26.8±1.28 Head width 16.0 16.0Ð18.1 17.0±0.85 Head depth 14.8 13.8Ð15.1 14.3±0.56 Predorsal length 37.9 36.7Ð39.2 37.9±0.96 Preanal length 64.1 64.1Ð67.8 66.4±1.40 Prepelvic length 49.7 48.6Ð53.0 49.7±1.69 Prepectoral length 21.0 20.6Ð25.4 22.7±2.16 Body depth at anus 15.5 13.8Ð15.8 14.5±0.88 Length of caudal peduncle 18.1 18.0Ð20.6 19.3±1.07 Depth of caudal peduncle 6.8 6.8Ð7.3 7.0±0.21 Pectoral-spine length 20.2 16.2Ð20.3 18.7±1.51 Pectoral-fin length 25.0 22.1Ð25.0 23.7±1.06 Dorsal-spine length 17.9 15.0Ð20.0 16.7±1.95 Length of dorsal-fin base 13.0 10.6Ð14.9 12.8±1.43 Pelvic-fin length 16.5 15.2Ð16.5 15.7±0.46 Length of anal-fin base 14.8 14.2Ð15.3 14.9±0.46 Caudal-fin length 29.0 28.0Ð31.0 29.2±1.02 Length of adipose-fin base 13.7 11.0Ð14.0 13.0±1.13 %HL Snout length 45.5 44.1Ð47.0 45.7±0.97 Interorbital distance 26.0 23.6Ð26.6 25.3±1.21 Eye diameter 11.0 10.4Ð13.2 11.6±0.97 Nasal barbel length 19.5 14.9Ð19.9 17.6±2.07 Maxillary barbel length 111.0 91.8Ð111.0 98.7±9.31 Inner mandibular barbel length 37.0 29.8Ð39.1 34.3±4.46 Outer mandibular barbel length 63.0 47.8Ð66.9 54.8±8.11
Lim, 1995). Current literature lists only three species: G. Zakaria-Ismail & Lim, 1995) as G. platypogonides, but can major (Boulenger, 1894), G. platypogon (Valenciennes, 1840) be distinguished from it in having a fairly uniform color and G. platypogonides as valid. The group is more diverse pattern with distinct pale mid-dorsal and midlateral stripes than the literature implies and furthermore, many of the names (vs. highly mottled coloration without distinct pale stripes). have been misapplied (Kottelat & Lim, 1995). Since the Chu & Mo (1999) consider G. siamensis (a species described group is currently being revised elsewhere (Ng & Kottelat, from Peninsular Thailand) a junior synonym of G. dorsalis in prep.), it would not be appropriate to discuss the identities Vinciguerra, 1890, a species otherwise known only from the of all of the Sundaic Southeast Asian species. Therefore, we Irrawaddy and Salween River drainages. The first author follow the current literature in recognizing only the three has examined material of G. dorsalis from the Salween River species as valid for now, but we note that the Southeast Asian drainage and we conclude that they are not conspecific: G. members of the group as a whole can be divided into those dorsalis always possesses darker spots randomly distributed with deep bodies (17.3Ð20.9% SL) and those with slender on the head and body and a pale colored nuchal shield (vs. bodies (12.0Ð15.8% SL). Both G. exodon and G. dark spots and pale colored nuchal shield always absent in platypogonides belong to the latter group; this means that an G. siamensis). in depth discussion of the identities of all other deep bodied Sundaic Southeast Asian species can be deferred for now. The differences in biometrics between G. exodon and G. platypogonides are not solely due to ontogeny. A biplot of Additionally, we consider G. siamensis Hora, 1923 to be valid caudal peduncle depth (Fig. 4) against SL shows that the member of this group of slender bodied species. Glyptothorax regression lines are significantly different (ANCOVA; siamensis has often been misidentified in the literature (e.g. P<0.00000005).
254 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Comparative material. Ð Glyptothorax major: BMNH Singapore and the All Catfish Species Inventory (NSF DEB- 1892.9.2.59, 1 syntype, 107.6 mm SL; BMNH 1892.10.7.26, 0315963) is gratefully acknowledged. 1 syntype, 74.2 mm SL; BMNH 1893.3.6.173Ð178, 5 syntypes, 31.0Ð84.8 mm SL; Borneo: Sarawak, Senah, Tagora and Baram Rivers. CAS 49418, 1 ex., 89.7 mm SL; Borneo: LITERATURE CITED Kalimantan Barat, Sungai Tamang, small forested stream flowing into Sungai Pinoh, opposite mouth of Sungai Kelawi. Chu, X.-L. & T.-P. Mo, 1999. Sisoridae. In: Chu, X.-L et al. (eds.) FMNH 68922, 1 ex., 96.1 mm SL; Borneo: Sarawak, Kapit Fauna Sinica. Osteichthyes. Siluriformes. Science Press, Beijing. district, Mengiong River drainage, Sungai Tekalit and Nanga Pp. 114Ð181. (In Chinese). Tekalit camp. De Pinna, M. C. C., 1996. A phylogenetic analysis of the Asian catfish families Sisoridae, Akysidae, and Amblycipitidae, with Glyptothorax platypogon: RMNH 2998, 3 syntypes, 61.0Ð a hypothesis on the relationships of the neotropical Aspredinidae 67.0 mm SL: Java. MZB 8516 35 ex., 24.5Ð65.0 mm SL; (Teleostei, Ostariophysi). Fieldiana: Zoology (New Series), 84: Java: Bogor, Cisarua, Desa Tugu Jembatan Utara, Ciliwung. iÐiv + 1Ð83. UMMZ 155696, 1 ex., 59.0 mm SL; Java: Cisokan. Eschmeyer, W. N., 1998. Catalog of Fishes. California Academy of Sciences, San Francisco. 2905 pp. Glyptothorax platypogonides: ANSP 27285, 1 ex., 77.5 mm Kottelat, M. & K. K. P. Lim, 1995. Freshwater fishes of Sarawak SL; Sumatra: Sumatera Barat, Tamah Datar, Batusangkar. and Brunei Darussalam: a preliminary annotated checklist. RMNH 6912, 4 ex., 53.63Ð67.1 mm SL; RMNH 15289, 3 Sarawak Museum Journal, 48: 227Ð256. ex., 65.6Ð73.5 mm SL; Sumatra: Sumatera Selatan, Lahat. Ng, H. H. & J. J. Dodson, 1999. Morphological and genetic UMMZ 243339, 3 ex., 40.4Ð71.7 mm SL; ZRC 41588 15 descriptions of a new species of catfish, Hemibagrus chrysops, ex., 43.9Ð78.2 mm SL; Sumatra: Sumatera Barat, from Sarawak, East Malaysia, with an assessment of Pulaupunjung market at Sungaidareh. UMMZ 243329, 1 ex., phylogenetic relationships (Teleostei: Bagridae). Raffles Bulletin 59.6 mm SL; Sumatra: Sumatera Barat, market at Kiliranjao. of Zoology, 47(1): 45Ð57. Ng, H. H. & I. Rachmatika, 1999. The catfishes (Teleostei: Glyptothorax siamensis: CAS 132671 (1), 92.8 mm SL; Siluriformes) of Bentuang Karimun National Park, West Malaysia: Selangor, hills west of Ginting Sempak, 29km E Kalimantan, Indonesia. Raffles Bulletin of Zoology, 47(1): 167Ð of Kuala Lumpur. USNM 109609 (1), 61.5 mm SL; Thailand: 183. Nakhon Si Thammarat. ZRC 2412 (2), 83.0Ð89.2 mm SL; Roberts, T. R., 1989. The freshwater fishes of Western Borneo Malaysia: Selangor, 18 km from Kuala Lumpur along (Kalimantan Barat, Indonesia). Memoirs of the California Bentong Road. Academy of Sciences, 14: xii + 210 pp. Tan, H. H. & H. H. Ng, 2000. The catfishes (Teleostei: Siluriformes) of central Sumatra. Journal of Natural History, 34: 267Ð303. ACKNOWLEDGMENTS Vaillant, L., 1902. Résultats zoologiques de l’expédition scientifique The authors thank the following for permission to examine Néerlandaise au Bornéo central. Poissons. Notes from the Leyden material under their care: Dominique Didier (ANSP), Darrell Museum, 24: 1Ð166. Siebert (BMNH), David Catania (CAS), Mary Anne Rogers Zakaria-Ismail, M. & K. K. P. Lim, 1999. The fish fauna of Tasik (FMNH), Martien van Oijen (RMNH), Lynne Parenti Temengor and its tributaries south of Banding, Hulu Perak, (USNM), and Kelvin Lim (ZRC). Funding from the Raffles Malaysia. Malayan Nature Journal, 48: 319Ð332. Museum of Biodiversity Research, National University of
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 257-263 Date of Publication: 31 Dec.2005 © National University of Singapore
A NEW SPECIES OF RHACOPHORUS (ANURA: RHACOPHORIDAE) FROM GUNUNG GADING, SARAWAK
Indraneil Das Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia. Email: [email protected]
Alexander Haas Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. Email: [email protected]
ABSTRACT. Ð A new species of rhacophorid of the genus Rhacophorus is described from Gunung Gading National Park, Sarawak, western Borneo. Rhacophorus gadingensis, new species, is compared with congeners from Borneo and other parts of southeast Asia. The new species is diagnosable in showing the following combination of characters: SVL 29.5 mm in the larger of the two specimens known; snout obtuse, slightly projecting beyond mandible; head wider than long; fingers and toes webbed; lacking nuptial pads (in males) and a dermal flap along forearm; a weak supracloacal fold; spinose calcar present; skin not co-ossified to forehead, median lingual process absent; supratympanic fold weak; dorsum brown, with a dark interorbital bar; dark brown blotches on dorsum and flanks with blue blotches.
KEY WORDS. Ð Rhacophorus, new species, Rhacophorus gadingensis, Rhacophoridae, systematics, Borneo.
INTRODUCTION We report here a new species of Rhacophorus, which does not match any of the described species of the genus from The genus Rhacophorus Kuhl & van Hasselt, 1822 (Anura: Borneo and from any other part of southeast Asia, and is here Rhacophoridae), as defined by Brown & Alcala (1994), is described as new to science. The species is allocated to known to contain over 60 nominal species of which at least Rhacophorus for showing the following characters considered 41 occur in southeast Asia alone (Frost, 1985; Harvey et al., diagnostic for the genus, according to the concept of Brown 2002; Inger et al., 1999; Iskandar & Colijn, 2000; Ohler et and Alcala (1994; see also, Inger, 1966; Wilkinson & Drewes, al., 2000; Vasudevan & Dutta, 2000). The members of the 2000; Wilkinson et al., 2002): fingers webbed; tips of digits genus are distributed from southern China, Sri Lanka & disk-like, with circummarginal grooves; vomerine teeth southwestern and northeastern India south to Indo-China and present; and eyes large. For a critique of other concepts of Indo-Malaya (Frost, 1985). Of these, as many as 14 species the genus, where character states are not explicitly stated (e.g., have been reported from Borneo (Inger & Stuebing, 1997; Dubois, 1986; 1992; Iskandar & Colijn, 2000), see Harvey Inger & Tan, 1996). et al. (2002).
The herpetofauna of Gunung Gading, a granite massif in Sarawak, western Borneo, was subject to the attention of the MATERIALS AND METHODS Italian nobleman, the Marquis Giacomo Doria of Genoa (1840Ð1913) and the botanist Odoardo Beccari (1843Ð1920), The holotype and paratype were collected at ca. 0200-0230 in the 1860s, incidental to their botanical collections. hrs. photographed in life, fixed in formalin ca. four hours Herpetological material collected during this expedition were after collection, and subsequently washed in water and described in a series of papers by the German zoologist, transferred to 70% ethanol about two weeks after collection. Wilhelm Carl Hartweg Peters (1815-1883) of the The following measurements were taken with MitutoyoTM dial Zoologisches Museum Humboldt-Universität, Berlin. No vernier callipers (to the nearest 0.1 mm) ca. four months after new rhacophorids, however, were described on the basis of collection: snout-vent length (SVL, from tip of snout to vent); these collections. tibia length (TBL, distance between surface of knee to surface
257 Das & Haas: New Rhacophorus from Gunung Gading of heel, with both tibia and tarsus flexed); head length (HL, Paratype: Adult male (ZRC)(field number ID-7832), paratopotype; distance between angle of jaws and snout-tip); head width same data as for holotype. (HW, measured at angle of jaws); head depth (HD, greatest transverse depth of head, taken posterior of the orbital Diagnosis. Ð A small (SVL 29.5 mm in the larger of two region); eye diameter (ED, horizontal diameter of the eyes); specimens) species of Rhacophorus, diagnosable from interorbital distance (IO, least distance between upper congeneric species in showing the following combination of eyelids; internarial distance (IN, distance between nostrils); characters: snout obtuse, slightly projecting beyond mandible; eye to snout distance (E-S, distance between anterior-most head wider than long; fingers and toes webbed; lacking nuptial point of eyes and tip of snout); eye to nostril distance (E- pads (in males) and dermal flap along forearm; a weak N, distance between anterior-most point of eyes and nostrils); supracloacal fold; spinose calcar present; skin not co-ossified axilla to groin distance (A-G, distance between posterior to forehead, median lingual process absent; supratympanic edge of forelimb at its insertion to body to anterior edge of fold weak; dorsum brown, with a dark interorbital bar; dark hind limb at its insertion to body); body width (BW, greatest brown blotches on dorsum and flanks with blue blotches. width of body); and tympanum diameter (TD, vertical and horizontal). Colour notes were taken from Fujichrome Velvia 50 ASA 35 mm slide transparency film, and Description of holotype (adult female). Ð A small species of compared with colour swatches of F. B. Smith (1975; 1981). Rhacophorus, SVL 29.5 mm; body elongate, with a narrow Sex was determined through dissection to examine the waist; head broader than long (HW/HL ratio 1.9); snout slopes gonads. slightly beyond mandible in lateral view and obtuse in dorsal view; nares oval, laterally positioned, nearer tip of snout than Comparative materials examined are listed in Appendix I. to orbit of eye (E-N/E-S ratio 0.71); internarial distance Sources of additional data on character states and distribution slightly less than distance from anterior margin of eye to of congeneric species of Rhacophorus include the following nostril (IN/E-N ratio 0.97); eye large (ED/HL ratio 0.56); works: Alcala & Brown (1998), Berry (1975), Boulenger pupil vertical; its diameter greater than eye to nostril distance (1882; 1912), Bourret (1942), Brown & Alcala (1994), (ED/E-N ratio 1.55); skin not co-ossified to forehead; pinal Hoffmann (1995), Inger (1954; 1966), Inger & Stuebing ocellus absent; interorbital width greater than upper eyelid (1989; 1997), Iskandar (1998), Malkmus et al. (2002), width (IO/UE ratio 1.82); canthus rostralis distinct; loreal Manthey & Steiof (1998), Ohler et al. (2000), Taylor (1962), region vertical; maxillary teeth present; a weak ‘W’- shaped van Kampen (1923) and Ziegler & Köhler (2001). Museum notch (= symphysial knob) on anterior edge of mandible; abbreviations, where available (indicated with an asterisk), mouth extends to posterior corner of eye; tongue follow Leviton et al. (1985). These include: subtriangular, smooth, bifid apically, lacking a median lingual process, free posteriorly for a third its length; choanae located Natural History Museum, London, U.K. (BMNH*) close to anterior of palate; vomerine ridge reduced, in two Zoological Museum of the Department of Wildlife and oblique series, beginning at inner front edge of choana, and National Parks, Kuala Lumpur, Malaysia (DWNP) closer to choanae than to each other; pupil horizontal; no rictal Forest Research Institute Malaysia, Kepong, Malaysia gland at posterior corner of mouth; tympanum distinct; its (FRIM) horizontal diameter slightly larger than the vertical diameter, Philippines National Museum, Manila, Philippines (PNM*) and a third of the eye diameter; supratympanic fold weak; no Sarawak Biodiversity Centre, Semenggoh, Sarawak, Malaysia (SBC) dermal flap along forearm; weak supracloacal fold and Sarawak Museum, Kuching, Sarawak, Malaysia (SM*) postcloacal tubercles absent; dorsum, including upper eyelids Sabah Parks Zoological Museum, Gunung Kinabalu Park, and upper surfaces of limbs smooth; abdomen and inner side Sabah, Malaysia (SP) of thighs with weak glandular structures. Zoological Museum of the Department of Biology, Universiti Brunei Darussalam, Bandar Seri Begawan, Brunei Darussalam (UBD) Raffles Museum of Biodiversity Research, National University of Singapore, Singapore (ZRC; the abbreviation used in Leviton et al., 1985, is USDZ*).
Specimens with ID-field numbers are being accessioned with this collection, apart from the types of the present species.
SYSTEMATICS
Rhacophorus gadingensis, new species (Figs. 1Ð2)
Material examined. Ð Holotype: Adult female (ZRC 1.11532)(field number ID-7831) from Gunung Gading (03¼42'2"N 115¼32'06"E), Lundu Division, Sarawak, East Malaysia (Borneo), coll. I. Das, Fig. 1. Lateral view of holotype of Rhacophorus gadingensis, new 13 Aug.2004. species, in life (ZRC 1.11532).
258 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Arm moderately long and thin; fingers long and thin, webbing and rest of venter unpigmented yellow-cream, flanks of body on finger I, none; on finger II, none (inner) and to basal with sky blue (# 168D) blotches enclosing irregular brown subarticular tubercle (outer); on finger III, to basal subarticular (# 129) areas; upper surfaces of limbs with light brown cross- tubercle (inner) and to distal subarticular tubercle (outer), and bars; and lower surfaces of thighs and tibia yellow-cream on finger IV, to distal subarticular tubercle; relative length coloured; digits of arms and webbing pale yellow cream; those of fingers: 3 > 4 > 2 > 1; finger tips dilated, with of the feet, including webbing, greyish-yellow; pupil black; circummarginal grooves; width of disk on finger III greater surrounding iris gem ruby (# 110), lightening to buff (# 124) than tympanum diameter; subarticular tubercles prominent, peripherally. rounded, numbering one on first and second fingers, two on third and fourth fingers; palmar tubercles indistinct; hind Measurements (in mm; holotype, followed by paratype in limbs relatively long and thin, meeting each other but not parentheses). Ð SVL 29.5 (23.7); HL 8.1 (7.1); HW 9.6 (8.3); overlapping when folded right angle to body; toes long and HD 5.0 (4.0); BW 7.8 (5.5); TBL 16.7 (13.3); TD (vertical) thin; webbing on toe I, to base of disk; toe II, to distal 1.6 (1.3); TD (horizontal) 1.5 (1.9); ED 4.5 (3.7); UE 3.3 subarticular tubercle (inner) and base of disk (outer); toe III, (3.2); IN 2.8 (3.0); IO 6.0 (5.1); E-S 4.1 (3.8); E-N 2.9 (2.0); to distal subarticular tubercle (inner) and base of disk (outer); E-T 0.6 (0.6); A-G 15.8 (12.5). toe IV, to distal subarticular tubercle (both outer and inner); and toe V, to base of disk; relative length of toes (longest to Variation. Ð The paratype, a male, is smaller in size than the shortest): 4 > 3 > 5 > 2 > 1; toe tips rounded; subarticular female holotype. It lacks nuptial pads or external vocal tubercles weakly developed, rounded, numbering one on first pouches. and second toes; two on third and fifth toes; and three on fourth toe; inner and outer metatarsal tubercles absent; disks Etymology. Ð The new species is named for Gunung Gading, on fingers wider than those on toes; heel bears a weak the type locality of the new species. postaxial fold, with a spinous calcar proximally. Ecological notes. Ð The holotype and paratype were found Colour. In life, tawny (# 38) dorsally, with a thin antique on vegetation, ca. 3 m and 1 m, respectively, overhanging brown interorbital band that extends across the eyelids; large granite boulders at the edge of a torrential forest stream, > blotches of the same color on torso, including a transversely 8 m in width. They were collected ca. 0200-0230 h. The widened one at nuchal region, two paired oval ones on call and larval stages of the new species remain unknown. midbody and a single median one on sacral region; throat The following species of frogs were found sympatric with the new species: Chaperina fusca, Limnonectes laticeps, L. leporinus, L. kuhlii, Meristogenys poecilus, Rana hosii, R. picturata, R. raniceps, Staurois guttatus, Nyctixalus pictus, Philautus tectus, Polypedates colletti, Rhacophorus appendiculatus, R. pardalis, Rhacophorus kajau, Leptolalax hamidi, Megophrys nasuta and Ansonia spinulifer.
COMPARISONS
The only congeners from Borneo with a calcar are Rhacophorus angulirostris (Ahl, 1927) (Borneo and possibly Sumatra), which differs from the new species in showing a short, pointed snout; two outer fingers webbed to pads; flanks and inner surfaces of legs yellow with several large spots; and colour variable but with one or several subocular light spots (see Malkmus, 1993; 1995); R. baluensis Inger, 1954 (a Borneo endemic), which differs from the new species in its smooth dermal ridge from outer fingers to elbow, and another from outer toe to heel; and dorsum light or dark brown, with either dark transverse marks or irregular light blotches; R. dulitensis Boulenger, 1892 (Peninsular Malaysia, Sumatra, Borneo), which shows a sharp, pointed snout; toes webbed to base of disks; edge of webbing of outer toes red; and dorsum green, with or without small white dots; and R. pardalis Günther, 1858 (Philippines, Borneo, Sumatra), which differs from the new species in showing a head longer than broad; tympanum half eye diameter; nuptial pad on finger I; iris orange; and webbing on hand and feet orangish-red. Fig. 2. Hand and feet of the holotype of Rhacophorus gadingensis, Additionally, in the last two nominal species, the appendage new species, (ZRC 1.11532). Markers = 5 mm. at the heel is a rounded narrow or broad flap.
259 Das & Haas: New Rhacophorus from Gunung Gading
The presence of a calcar at the heel separates Rhacophorus brown with cross-bands; R. bifasciatus van Kampen, 1923 gadingensis, new species, from the following Bornean (Sumatra), tympanum half to two-thirds eye diameter; toes congeners (additional characters at variance to that seen in (except toe IV) webbed to base of disks; and dorsum purple the new species follow): R. appendiculatus (Günther, 1859) with two irregular white dorsolateral bands; R. bimaculatus (Thailand, Peninsular Malaysia, Sumatra, Borneo, southern (Peters, 1867) (Mindanao in the Philippines), tympanum half Philippines), head longer than wide; tympanum half eye eye diameter; head with a small, dermal process; dorsum diameter; a cutaneous fringe along edge of jaws and along, green, with a large white or blue subocular spot; R. inner edge of tarsus and forearm; and dorsum green with bipunctatus Ahl, 1927 (southern China, northeastern India, darker variegations; R. cyanopunctatus (Manthey & Steiof, Myanmar, Thailand and Vietnam), webbing on toes (except 1998) (Thailand, Vietnam, Peninsular Malaysia, Sumatra, toe IV) to base of disks; dermal fringe along forearm and Borneo); vomerine teeth rows absent; fingers with narrow tarsus; and dorsum green or grey, with black spots; R. dermal fringe; hind limbs long, overlapping when folded right bisacculus Taylor, 1962 (northeastern India, Thailand), a row angle to body; a white spot below eye; flanks and inner side of tubercles on undersurface of arm and another on tarsus; of lower thigh and shank dark brown with blue spots; and large nuptial pads; and dorsum brown with a darker, dorsum sandy brown; R. fasciatus Boulenger, 1895 (Borneo), subtriangular area on occiput; R. calcaneus Smith, 1924 vomerine teeth in two long oblique series; fingers and toes (Vietnam), webbing on outer fingers not reaching base of webbed to base of disks; and dorsum pale reddish-brown, disks; toes webbed to base of disks; a glandular supratympanic with brown specklings and dark brown cross-bars; R. gauni fold; and dorsum pale green with numerous pale yellow spots; (Inger, 1966) (Borneo), a conical dermal tubercle in middle R. catamitus Harvey et al., 2002 (Sumatra), snout slopes to of upper eyelid; vomerine teeth absent; and a row of tubercles a low, swollen rostral tubercle; webbing on hand reduced to on edge of lower arm; R. everetti Boulenger, 1894 (Palawan, basal tubercles of fingers I and II; a row of tubercles present Borneo), a small inner metatarsal tubercle; dorsal surface with along antebrachium and tarsus; tubercle present at tibio-tarsal widely scattered tubercles with pointed projections; and articulation; dorsum of body shagreened with low tubercles; dorsum green red and yellow variegations; R. harrissoni Inger supratympanic fold thick; and dorsum brown with dark cross- & Haile, 1960 (Borneo), toes webbed to base of disks; dorsum bars; R. dorsoviridis Bourret, 1937 (Vietnam), tympanum half reddish-brown, flanks greenish-yellow; SVL of males 50-56 eye diameter; webbing on toe IV to penultimate subarticular mm; of females 60-70 mm; R. kajau Dring, 1983 (Borneo), tubercle; and dorsum green; R. duboisi Ohler et al., 2000 a narrow dermal ridge along arm and along hind limb; dorsum (Vietnam), SVL to 65.7 mm; dorsum with small granules; green, with small white spots; and webbing black; R. dermal ridge along toe V; canthus rostralis with a brown band; nigropalmatus Boulenger, 1895 (Thailand, Peninsular and dorsum green and brown, with spots arranged in bands; Malaysia, Sumatra, Borneo), snout rounded, not projecting; venter grey-white with small grey spots; R. edentulous Müller, tympanum half to two-thirds eye diameter; webbing reaches 1894 (Sulawesi), dorsum green, unpatterned or with black disk of finger IV as a broad sheet; dorsum green; webbing spots; R. exechopygus Inger et al., 1999 (Vietnam), outer on hand and feet marked with black; R. reinwardtii (Schlegel, fingers webbed to base of disks; weakly crenulated dermal 1837) (Borneo, Java, Thailand, Peninsular Malaysia and ridges on forearm and tarsus; and dorsum grey to brown, Vietnam), tympanum three-fourths eye diameter; fingers and uniform or with obscure dark blotches at back of head; R. toes webbed to disks; cutaneous fringe along forearm; and georgii Roux, 1904 (Sulawesi), head as long as broad; fingers dorsum green, with a black axillary spot; large black area in two-third webbed; a small inner metatarsal tubercle; and webbings of fingers and toes; and R. rufipes Inger, 1966 dorsum pale green with lighter spots; R. margaritifer (Borneo), tympanum three-fourths eye diameter; webbing on (Schlegel, 1837) (Java), tympanic annulus somewhat hand and feet orangish-red. indistinct; numerous white tubercles present below vent; and dorsum greyish-white (in preservative; live coloration The new species from Gunung Gading is compared with other unknown), finely dotted with black R. maximus Günther, 1859 congeners from southeast Asia, listing only opposing suite (southern China, northeastern India, Thailand), tympanum of characters. Rhacophorus achantharrhena Harvey et al., greater than half eye diameter; fingers and toes webbed to 2002 (Sumatra), male SVL to 40.55 mm; thick smooth dermal disks; large inner metatarsal tubercle; and dorsum blue-green flaps along margins of forearm and tarsus; dorsum weakly or violet; R. modestus Boulenger, 1920 (Sumatra), head as spiculate in males; supratympanic fold thick and conspicuous; broad as long; snout rounded; tympanum half to two-thirds and dorsum green; R. annamensis Smith, 1924 (southern eye diameter; and dorsum grey or brown, with a white streak Vietnam), snout rounded; tympanum half eye diameter; or a series of white spots along flanks; R. monticola webbing on fingers and toes reaching the base of disks; and Boulenger, 1896 (Sulawesi), tympanum half eye diameter; dorsum with irregular black, red or green spots; R. baliogaster finely shagreened above; and dorsum greyish-green, with Inger et al., 1999 (Vietnam), dorsum dark brown, with small, scattered yellow spots; R. notater Smith, 1924 (Vietnam), irregular dark spots; ventral surface of head and body white snout rounded; tympanum indistinct, about half eye diameter; with black spots; and females with a distinct rostral cone; R. toes webbed to base of disks; a feeble dermal ridge along barisani Harvey et al., 2002 (Sumatra), male SVL to 53.16 fore- and hind limbs; and dorsum pale grey with large, mm; nares about equidistant between eye and snout-tip; low irregular blotches; R. orlovi Ziegler and Köhler, 2001 dermal ridge or tubercles along ulna and tarsus; dermal flap (northern Vietnam), calcar absent; webbing complete on feet; with long triangular calcar present at tibio-tarsal articulation; and dorsum reddish-brown; R. poecilonotus Boulenger, 1920 dorsum diffusely covered with rounded tubercles; and dorsum (Sumatra), head strongly depressed; tympanum two-thirds eye
260 THE RAFFLES BULLETIN OF ZOOLOGY 2005 diameter; and toes webbed to base of disks; R. prominanus Jamili Nais, Maklarin Lakim and Paul Yambun (SP), Smith, 1924 (Thailand, Peninsular Malaysia, Sumatra), Kamariah Abu Salim, Joseph K. Charles, David Edwards and tympanum subequal to eye diameter; toes webbed to disks; Helen Pang Yoke Yew (UBD) and Kelvin Kok Peng Lim, and dorsum green, with or without dark spots; R. Peter Kee Lin Ng and Chang Man Yang (ZRC). Finally, we rhyssocephalus Wolf, 1936 (Sangihe Island) large size, SVL thank Robert F. Inger and Tzi-Ming Leong for helpful to 85 mm; vomerine teeth separated from each other by a comments on the manuscript. gap almost as broad as length of row; snout rounded; snout longer than eye diameter; tympanum slightly over half eye diameter; subarticular tubercles distinct; heels overlap when LITERATURE CITED hind limbs are placed at right angle to body; skin finely granular, specially at posterior of head and on scapular region; Ahl, E., 1927. Zur Systematik der asiatischen Arten der several distinct bone crests on dorsal surface of head, Fröschgattung Rhacophorus. Sitzungsberichte der Gesellschaft including two short, V-shaped ones above tympanum that Naturforschender Freunde zu Berlin, 1927: 35-47. diverge distally; three more distinct crests form a U-shape in Ahl, E., 1931. Anura III. Das Tierreich 55. Walter de Gruyter & interorbital region; and dorsally light violet, with dense, dark Co. XVI + 477 pp. marbling; R. robinsoni Boulenger, 1903 (Thailand), Alcala, A. C. & W. C. Brown, 1998. Philippine amphibians. An tympanum two-thirds eye diameter; vomerine teeth touching illustrated fieldguide. Bookmark, Inc., Makati City. xii + 116 inner edge of choanae; toes webbed to base of disks; a strong pp. supratympanic fold; and dorsum pinkish-brown or pale Berry, P. Y., 1975. The amphibian fauna of peninsular Malaysia. brown, bluish-grey on flanks; R. taroensis Smith, 1940 Tropical Press, Kuala Lumpur. x + 130 pp. (Myanmar), head depressed; tympanum adjacent to orbit of Boulenger, G. A., 1882. Catalogue of the Batrachia Salientias. eye and subequal in size; toes webbed to base of disks; a Ecaudata in the collection of the British Museum. Second dermal fringe on outer side of forearm and hind limbs; and edition. British Museum, London. xvi + 503 pp., 30 pls. dorsum green with small black spots; R. turpes Smith, 1940 Reprinted 1966, Wheldon & Wesley, Codicote & Verlag J. (Myanmar), head as broad as long, depressed; toes webbed Cramer, Weinham. to base of disks; supratympanic fold glandular; and dorsum Boulenger, G. A., 1912. A vertebrate fauna of the Malay Peninsula pale pinkish-brown, with small black spots in males; and R. from the Isthmus of Kra to Singapore including the adjacent verrucosus Boulenger, 1893 (Myanmar, Thailand and islands. Reptilia and Batrachia. Taylor and Francis, London. Vietnam), SVL to 57 mm; body relatively robust; vomerine xiii + 294 pp. teeth in two oblique rows from anterior-median margin of Bourret, R., 1942. Les batraciens de l’Indochine. Mémoires de choana, the rows only a little shorter than their lengths; snout l’Institut Océanographique de l’Indochine, Hanoi. 547 pp., pls. as long or slightly longer than interorbital distance; toe V > IÐIV. III; tarsal fold present, formed by a broken line of skin folds; Brown, W. C. & A. C. Alcala, 1994. Philippine frogs of the family tibio-tarsal articulation with spur-like flap; skin dorsally with Rhacophoridae. Proceedings of the California Academy of numerous warts, especially on flanks; tip of snout with small Sciences, 48(10): 185-220. flap; and dorsum brownish-grey above, with lighter and Dubois, A., 1986. Miscellanea taxomonica batrachologica (I). darker lichenous or bark-like blotches. Alytes, 5(1Ð2): 7-95. Dubois, A., 1992. Notes sur la classification des Ranidae (Amphibiens Anoures). Bulletin Mensuel de la Société Linnéene ACKNOWLEDGEMENTS de Lyon, 61(10): 305-352. Frost, D. R., (ed.) 1985. Amphibian species of the world. A We would like to thank Datuk Cheong Ek Choon, Controller taxonomic and geographical reference. Allen Press, Inc., and of National Parks and Nature Reserves, Sarawak Forestry Association of Systematics Collections, Lawrence. (iv) + 732 Corporation, for permit (No. 5/2004, dated 06.07.2004) to pp. conduct research on the amphibians of Gunung Gading Harvey, M. B., A. J. Pemberton & E. N. Smith, 2002. New and National Park, and for export of specimens (permit number poorly known parachuting frogs (Rhacophoridae: Rhacophorus) 04635). We are also grateful to our respective institutions, from Sumatra and Java. Herpetological Monograph, 16: 46Ð the Institute of Biodiversity and Environmental Conservation, 92. Universiti Malaysia Sarawak and Biozentrum Grindel und Hoffmann, P., 1995. Untersuchungen zur Anurenfauna des Mt. Zoologisches Museum, Universität Hamburg, for supporting Kinabalu (Borneo) Teil II: Familien Ranidae, Rhacophoridae. our research on the amphibian fauna of Borneo, which was Sauria, Berlin 17(3): 9-17. supported by a grant from Volkswagen-Stiftung, Germany Inger, R. F., 1954. Systematics and zoogeography of Philippine (grant I/79 405). The second author acknowledges the Amphibia. Fieldiana Zoology, 33: 183-531. Economic Planning Unit, at the Prime Minister’s Office, Inger, R. F., 1966. The systematics and zoogeography of the Kuala Lumpur, for issuing him a research permit. Curators Amphibia of Borneo. Fieldiana Zoology, 52: 1-402. Reprinted of the following institutions permitted us to examine 1990, Lun Hing Trading Company, Kota Kinabalu. comparative material under their care: Colin John McCarthy Inger, R. F., N. Orlov & I. Darevsky, 1999. Frogs from Vietnam: (BMNH), Lim Boo Liat (DWNP), Norsham Suhaina binti a report on new collections. Fieldiana Zoology (New Series), Yaakob (FRIM), Arvin Cantor Diesmos (PNM), Eileen Yen 92: iÐiv + 1Ð46. and Margarita Naming (SBC), Charles Leh Moi Ung (SM),
261 Das & Haas: New Rhacophorus from Gunung Gading
Inger, R. F. & R. B. Stuebing, 1989. Frogs of Sabah. Sabah Parks Wilkinson, J. A., R. C. Drewes & O. L. Tatum, 2002. A molecular Trustees, Kota Kinabalu. 132 + iv pp. phylogenetic analysis of the family Rhacophoridae with an Inger, R. F. & R. B. Stuebing, 1997. A field guide to the frogs of emphasis on the Asian and African genera. Molecular Borneo. Natural History Publications (Borneo) Sdn Bhd/Science Phylogenetics and Evolution, 24:265-273. and Technology Unit, Sabah, Kota Kinabalu. x + 205 pp. Wolf, S., 1936. Revision der Untergattung Rhacophorus. Bulletin Bahasa Malaysia edition, 1999, Panduan lapangan katak-katak of the Raffles Museum, 12: 137-217. Borneo. Natural History Publications (Borneo) Sdn Bhd/Jabatan Ziegler, T. & J. Köhler, 2001. Rhacophorus orlovi, sp. n., ein neuer Muzium Sabah, Kota Kinabalu. x + 225 pp. Ruderforsch aus Vietnam (Amphibia: Anura: Rhacophoridae). Inger, R. F. & F. L. Tan, 1996. Checklist of the frogs of Borneo. Sauria, Berlin, 23(3): 37-46. Raffles Bulletin of Zoology, 44(2):551-574. Iskandar, D. T., 1998. Amfibi Jawa dan Bali. Puslitbang Biologi- LIPI and GEF- Biodiversity Collections Project, Bogor. xviii APPENDIX I + 117 pp., 26 pls. English edition, 1998, The amphibians of Java and Bali. Research and Development Centre for Biology- List of comparative material examined LIPI and GEF- Biodiversity Collections Project, Bogor. xix + 117 pp., 26 pls. Rhacophorus angulirostris (Ahl, 1927): Gunung Kinabalu, Sabah, Iskandar, D. T. & E. Colijn, 2000. Preliminary checklist of southeast Malaysia (IDÐ7309; PNM 7951); 16th mile, Crocker Range Asian and New Guinean herpetofauna. I. Amphibians. Treubia, National Park, Sabah, Malaysia (IDÐ7388; SP 20110; ZRC 31(3)(Suppl.): 1Ð133. 1.9085Ð86); Gunung Trus Madi, Sabah, Malaysia (SP 02913). Leviton, A. E., S. C. Anderson, R. H. Gibbs, E. Heal & C. E. Dawson, R. appendiculatus (Günther, 1859): Gn. Podam, Bau, Sarawak, 1985. Standards in herpetology and ichthyology. Part I. Malaysia (SBC A.00297); Gunung Gading, Sarawak, Malaysia Standard symbolic codes for institutional resource collections (PNM 7977); Niah National Park, Sarawak, Malaysia (ZRC in herpetology and ichthyology. Copeia, 1985(3): 802-832. 1.9021); Sungei Seran, Labang, Sarawak, Malaysia (ZRC Liem, (D.) S. S., 1970. The morphology, systematics and evolution 1.2721); Linumumsut Lake, Maliau Basin, Sabah (SP 02817). of the Old World treefrogs (Rhacophoridae and Hyperoliidae). R. baluensis Inger, 1954: Reked Maligan, Church Camp, Gunung Fieldiana Zoology, 57: 1-145. Murud, Sarawak, Malaysia (ZRC 1.9029Ð32); Gunung Malkmus, R., 1993. Zur Zeichnungsvariabilität und zum Kinabalu, Sabah, Malaysia (IDÐ7727Ð28; 7751). Totstellverhalten des Ruderfroeisches Rhacophorus R. bipunctatus Ahl, 1927: Genting Highlands, Selangor, Peninsular angulirostris Mocquard, 1890. Sauria, Berlin, 15(3): 35-38. Malaysia (ZRC 1.8061Ð65; ZRC 1.9090); Cameron Highlands, Malkmus, R., 1995. Kopfseitenflecken bei beiden Geschlechtern Pahang, Peninsular Malaysia (ZRC 1.4077). von Rhacophorus angulirostris Ahl, 1927 (Amphibia: R. cyanopunctatus (Manthey & Steiof, 1998): Lakum Forest Rhacophoridae). Salamandra, 31: 245-246. Reserve, Raub, Pahang, Peninsular Malaysia (ZRC 1.10533); Malkmus, R., U. Manthey, G. Vogel, P. Hoffmann & J. Kosuch, Sungei Kinchin, Rompin, Pahang, Peninsular Malaysia (ZRC 2002. Amphibians & reptiles of Mount Kinabalu (North 1.1684Ð85); Tawau Hills Park, Sabah, Malaysia (ID 8277). Borneo). Koeltz Scientific Books, Königstein. 424 pp. R. dulitensis Boulenger, 1892: Batu Apoi, Brunei Darussalam (UBD Manthey, U. & C. Steiof, 1998. Rhacophorus cyanopunctatus sp. 550), Tasek Merembun, Brunei Darussalam (UBD 658; ZRC n. (Anura: Rhacophoridae), ein neuer Flugfrogsch von der 1.3162); Tasek Merembun, Brunei Darussalam (UBD 658Ð59); Malaiischen Halbinsel, Sumatra und Borneo. Sauria, Berlin, Baram, Sarawak, Malaysia (ZRC 1.900); Lower Segama, Lahad 20(3): 37-42. Datu, Sabah, Malaysia (SP 20377); Tawau Hills Park, Sabah, Ohler, A.-M., O. Marquis, S. Swan & S. Grosjean, 2000. Amphibian Malaysia (SP 00637; 01264). biodiversity of Hoang Lien Nature Reserve (Lao Cai Province, R. everetti Boulenger, 1894: Palawan, Philippines (BMNH northern Vietnam) with description of two new species. 94.6.3.126Ð27; syntypes of R. everetti); Kinabalu, Sabah, Herpetozoa, 13(1/2): 71-87. Malaysia (BMNH 94.6.3.126; syntype of Rhacophorus everetti Smith, F. B., 1975. Naturalist’s color guide. Parts I and II. macrosceles); Gunung Kinabalu, Sabah, Malaysia (IDÐ7720; th American Museum of Natural History, New York. Part I: 8 pp. 7814Ð7817); 16 mile, Crocker Range National Park, Sabah, + 18 colour swatches; Part II: xiii + 229 pp. Malaysia (IDÐ7422). Smith, F. B., 1981. Naturalist’s color guide. Part III. American R. fasciatus Boulenger, 1895: Gunung Mulu, Sarawak, Malaysia Museum of Natural History, New York. (iv) + 37 pp. (IDÐ7784); Gunung Penrissen, Sarawak, Malaysia (ZRC 1.901). Taylor, E. H., 1962. The amphibian fauna of Thailand. University R. gauni (Inger, 1966): Kilapan, Crocker Range National Park, of Kansas Science Bulletin, 63(8): 265-599; errata (= 1 p). Sabah, Malaysia (SP 02177); Tawau Hills Park, Sabah, Malaysia (SP 00645); Maliau Basin, Sabah (SP 20244; 20281). van Kampen, P. N., 1923. The Amphibia of the Indo-Australian Archipelago. E. J. Brill, Leiden. xii + 304 pp. R. harrissoni Inger & Haile, 1960: Batu Apoi, Brunei Darussalam (UBD 214). Vasudevan, K. & S. K. Dutta, 2000. A new species of Rhacophorus (Anura: Rhacophoridae) from the Western Ghats, India. R. kajau Dring, 1983: Gunung Gading, Sarawak, Malaysia (IDÐ Hamadryad, 25: 1: 21-28. 7598); Batu Apoi, Brunei Darussalam (UBD 39, 216, 474, 482, 491, 492, 569; ZRC 1.3154; ZRC 1.8663). Wilkinson, J. A. & R. C. Drewes, 2000. Character assessment, genus level boundaries, and phylogenetic analyses of the family R. margaritifer (Schlegel, 1837): Pengalengan, Java, Indonesia (ZRC Rhacophoridae: a review and present day status. Contemporary 1.902). Herpetology, (2):1-24. URL: www.cnah.org/ch/ch/2000/2/ R. nigropalmatus Boulenger, 1895: Batu Apoi, Brunei Darussalam index.html (UBD 366); Silensing, Pahang, Peninsular Malaysia (ZRC
262 THE RAFFLES BULLETIN OF ZOOLOGY 2005
1.1068); Lata Berkoh, Taman Negara, Pahang, Peninsular R. prominanus Smith, 1924: Cat Snake Trail, middle elevation of Malaysia (ZRC 1.10160); Tambunan, Gunung Trus Madi, Genting Highlands, Selangor, Malaysia (FRIM 0072; 0260Ð61); Sabah, Malaysia (SP 0223). Genting Highlands, Selangor, Peninsular Malaysia (ZRC R. pardalis Günther, 1858: Salleh Trail, Sungei Relau, Taman 1.9044–45); Fraser’s Hill, Pahang, Peninsular Malaysia (ZRC Negara, Pahang, Peninsular Malaysia (DWNP A.1029Ð30); 1.3273); Sungei Relau, Taman Negara, Pahang, Peninsular Interpretive Trail, Sungei Relau, Taman Negara, Pahang, Malaysia (DWNP A.1008). Peninsular Malaysia (DWNP A.1013Ð14); Poring, Sabah, R. reinwardtii (Schlegel, 1837): Sungei Relau, Taman Negara, Malaysia (IDÐ7351); Gunung Meraja, Bau, Sarawak, Malaysia Pahang, Peninsular Malaysia (DWNP A.0995); Sukabumi, Java, (SBC A.00008, SBC A.00237); Gunung Podam, Bau, Sarawak, Indonesia (ZRC 1.3640–44); also, “Java” (ZRC 1.5273–75); Malaysia (SBC A.00296, A.00300); Pa’Umor, Bario, Kelabit Interpretive Trail, Sungei Relau, Taman Negara, Pahang, Highlands, Sarawak, Malaysia (IDÐ7483Ð84); Lambir Hills, Peninsular Malaysia (DWNP A.0995). Sarawak, Malaysia (IDÐ7470Ð71); Kuala Belalong Field Studies R. tunkui Kiew, 1987: Panti Bunker Trail, Johor, Peninsular Malaysia Centre, Temburong District, Brunei Darussalam (ZRC 1.8664Ð (ZRC 1.10363Ð74); Taman Negara, Pahang, Peninsular 68); Inambacon Caves, Bohol, Philippines (ZRC 1.10831); Malaysia (ZRC 1.10162). Mount Maquiling, Laguna, Luzon, Philippines (ZRC 1.10822).
263
THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 265-270 Date of Publication: 31 Dec.2005 © National University of Singapore
A NEW SPECIES OF POLYPEDATES (ANURA: RHACOPHORIDAE) FROM GUNUNG MURUD, SARAWAK (NORTHWESTERN BORNEO)
INDRANEIL DAS Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia. Email: [email protected]
ABSTRACT. Ð A new species of rhacophorid of the genus Polypedates is described from Gunung Murud, Sarawak State, north-western Borneo. Polypedates chlorophthalmus, new species is compared with congeners from Borneo and other parts of southeast Asia. The new species is diagnosable in showing the following combination of characters: SVL 62.1 mm in the unique holotype; snout rounded, not projecting beyond mandible; head slightly wider than long; canthus rostralis sloping; no dermal flap along forearm; supracloacal fold absent; skin not co-ossified to cranial bones, median lingual process absent; supratympanic fold distinct; dorsum brown, with a thin dark gray line at back of forehead, nearly converging at the axilla, lacking dark lines; lower flanks and anterior edge of thighs with dark blotches; throat with dark pigmentation; and iris bright green.
KEY WORDS. Ð Polypedates, new species, Polypedates chlorophthalmus, Rhacophoridae, systematics, new species, Borneo.
INTRODUCTION other part of south-east Asia, is here described, based on the recent collections from Gunung Murud. The species is The genus Polypedates (Anura: Rhacophoridae), as defined allocated to Polypedates for showing the following characters by Brown and Alcala (1994), is known to contain 16 nominal considered diagnostic for the genus, according to the concept species, of which 10 occur in southeast Asia (Glaw et al., of Liem, 1970 (see also Inger, 1966): fingers free of webbing; 2000; Frost, 1985, also the internet version, Frost, 2004; tips of digits disk-like, with circummarginal grooves; Iskandar & Colijn, 2000). The members of the genus are vomerine teeth present; eyes large, with horizontal pupil; distributed from southern China, Sri Lanka and southwestern dermal tarsal and anal folds absent; and dorsum dark brown. and northeastern India south to Indo-China and Indo-Malaya Although the genus Polypedates has been considered (Frost, 1985). Of these, four species have been reported from synonymous with Rhacophorus by some authorities, based Borneo (Inger & Stuebing, 1997; Inger & Tan, 1996). on morphology (e.g., Dubois, 1986; 1992) or acoustic data (Matsui & Wu, 1994), phylogenies of the group, based on a The herpetofauna of Gunung Murud, the highest mountain reanalysis of data in Liem (1970) show support for the validity in Sarawak (2,423 m ASL), has been poorly sampled. The of the genus (see Channing, 1989; Wilkinson & Drewes, present day knowledge of the fauna derives from the 2000; Wilkinson et al., 2002). collections made by the Swedish naturalist and adventurer, Eric Georg Mjöberg (1882–1938), Curator of the Sarawak Museum, in 1922. Mjöberg’s herpetological collections were MATERIAL AND METHODS published by Smith (1925), and Mjöberg (1925) himself described this expedition in detail, enumerating collection The holotype was collected at ca. 2100 h, photographed in localities. We commenced long-term field work in Gunung life, fixed in formalin ca. 4 h after collection and subsequently Murud, Sarawak, since 2003, with the goal of inventorying washed in water and transferred to 70% ethanol about 8 weeks the herpetological diversity of the mountain range, collecting after collection. The following measurements were taken with a number of novelties in the process. MitutoyoTM dial vernier calipers (to the nearest 0.1 mm), 4 months after collection: snout-vent length (SVL, from tip of A new species of Polypedates, which does not match any of snout to vent); tibia length (TBL, distance between surface the described species of the genus from Borneo or from any of knee to surface of heel, with both tibia and tarsus flexed);
265 Das: New species of Polypedates from Gunung Murud head length (HL, distance between angle of jaws and snout- sloping; no dermal flap along forearm; supracloacal fold tip); head width (HW, measured at angle of jaws); head depth absent; skin not co-ossified to cranial bones, median lingual (HD, greatest transverse depth of head, taken posterior of the process absent; supratympanic fold distinct; dorsum brown, orbital region); eye diameter (ED, horizontal diameter of eye); with a thin dark gray line at back of forehead, nearly interorbital distance (IO, least distance between upper eyelids; converging at the axilla, lacking dark lines; lower flanks and internarial distance (IN, distance between nostrils); eye to anterior edge of thighs with dark blotches; throat with dark snout distance (E-S, distance between anterior-most point of pigmentation; and iris bright green. eyes and tip of snout); eye to nostril distance (E-N, distance between anterior-most point of eyes and nostrils); axilla to Description of the holotype (adult female). Ð A medium groin distance (A-G, distance between posterior edge of species of Polypedates, SVL 62.1 mm; body elongate, with forelimb at its insertion to body to anterior edge of hindlimb a narrow waist; head slightly broader than long (HW/HL ratio at its insertion to body); body width (BW, greatest width of 1.04); snout elongate, rounded at the tip, not exceeding level body); and tympanum diameter (TD, vertical and horizontal). of mandible in lateral view and vertical in dorsal view; nares Color notes were taken from Fujichrome Velvia 50 ASA 35 oval, laterally positioned, nearer tip of snout than to eye (E- mm slide transparency film, and compared with color N/E-S ratio 0.74); internarial distance greater than distance swatches of F. B. Smith (1975; 1981). Sex was determined from anterior margin of eye to nostril (IN/E-N ratio 0.72); through dissection to examine the gonads. eye large (ED/HL ratio 0.39); pupil horizontal; its diameter greater than eye to nostril distance (ED/E-N ratio 1.07); Comparative materials examined are listed in Appendix I. canthal ridge distinct; skin of forehead free, not co-ossified Sources of additional data on character states and distribution to the nasal, sphenethmoid or frontoparietal elements of the of congeneric species include the following works: Alcala cranium; pineal ocellus present; interorbital width greater than and Brown (1998), Ahl (1931), Berry (1975), Boulenger upper eyelid width (IO/UE ratio 2.22); canthus rostralis (1882; 1912), Bourret (1942), Hoffmann (1995), Inger (1966), sloping; loreal region vertical; maxillary teeth present; a weak Inger & Stuebing (1989; 1997), Iskandar (1998), Malkmus ‘W’- shaped notch (= symphysial knob) on anterior edge of et al. (2002), Taylor (1962), Wolf (1936), and et al. (1991). mandible; mouth extends to posterior corner of eye; tongue Museum abbreviations, where available (indicated with an subtriangular, smooth, bifid apically, lacking a median lingual asterisk), follow Leviton et al. (1985). These include: process, free posteriorly for a third its length; choanae located Forest Research Institute Malaysia, Kepong, Malaysia close to anterior of palate; vomerine ridges large, in two (FRIM) oblique series, beginning at inner front edge of choana, and Philippines National Museum, Manila, Philippines (PNM*) separated by a distance ca. 0.5 of their own length; pupil Sarawak Biodiversity Centre, Semenggoh, Sarawak, Malaysia horizontal; no rictal gland at posterior corner of mouth, but (SBC) 2Ð3 small tuberculate structures present; tympanum distinct; Sabah Parks Zoological Museum, Gunung Kinabalu, Sabah, its vertical diameter greater than horizontal diameter; Malaysia (SP) supratympanic fold distinct, extending from the posterior Biology, Universiti Brunei Darussalam, Bandar Seri corner of eyelid to above insertion of forearm; no dermal flap Begawan, Brunei Darussalam (UBD) along forearm or tarsus; supracloacal fold absent; postcloacal Raffles Museum of Biodiversity Research, National tubercles absent; dorsum, including upper eyelids and upper University of Singapore, Singapore (ZRC; the abbreviation surfaces of limbs smooth; abdomen and inner side of thighs used in Leviton et al., 1985, is USDZ*). Specimens with IDÐ with weak glandular structures. field numbers are being accessioned with this collection, apart from the types of the present species; and Fore-and upper arm short and thick; fingers long and thin, Zoological Survey of India, Kolkata, India (ZSI*). lacking webbing; relative length of fingers (longest to shortest): 3 > 4 > 2 > 1; finger tips dilated, with circummarginal grooves; width of disk on finger III less than SYSTEMATICS tympanum diameter; subarticular tubercles prominent, rounded, numbering one on first, second and fourth fingers, Polypedates chlorophthalmus, new species two on third finger; palmar tubercles indistinct; hind limbs (Figs. 1-3) relatively long and thin, meeting each other and overlapping slightly when folded right angle to body; toes long and thin; Material examined. Ð Holotype Ð ZRC 1.11531 Ð Sarawak webbing on toe I to top of subarticular tubercle; toe II to distal Biodiversity Centre Zoological Museum (field number IDÐ8017) subarticular tubercle (inner) and base of disk (outer); toe III from an unnamed stream ca. 0.8 km NE of Samling Camp at to distal subarticular tubercle (inner) and base of disk (outer); Ravenscourt, 04¼ 05'14.1"N 115¼ 28' 42.7" E, Lawas Division, toe IV distal subarticular tubercle, reaching base of disk as Sarawak; 1,351 m ASL, at middle elevations of Gunung Murud, a narrow sheath (both outer and inner); and toe V to slightly East Malaysia (Borneo), coll. I. Das, 16 Oct.2003. Adult female. below base of disk; relative length of toes (longest to shortest): Diagnosis. Ð A medium-sized (SVL 62.1 mm) species of 4 > 5 > 3 > 2 > 1; toe tips rounded, with circummarginal Polypedates, diagnosable from congeneric species in showing grooves; subarticular tubercles well developed, rounded, the following combination of characters: SVL 62.1 mm in numbering one on first and second toes (Fig. 3); two on third the unique holotype; snout rounded, not projecting beyond and fifth toes; and three on fourth toe; large, elongated inner mandible; head slightly wider than long; canthus rostralis metatarsal tubercle; outer metatarsal tubercle absent; disks
266 THE RAFFLES BULLETIN OF ZOOLOGY 2005 on fingers wider than those on toes; heel lacks postaxial fold Etymology. Ð The new species is named for its remarkable or calcar. green iris, from the Greek for green-eyed.
Colour. Ð In life, the dorsum is russet (# 34), with numerous Ecological notes. Ð The holotype was found on vegetation, smoke gray (# 44) blotches; a vandyke brown (# 121) line ca. 0.8 m above a granite boulder at the edge of a forest stream. at back of forehead, commencing from posterior of upper It was a gravid female, containing large and small eyelid, converging medially a little ahead of the posterior end unpigmented ova, assumed to represent multiple clutches. The of the suprascapula, but failing to meet each other; lips barred males, call and larval stages of the new species remain with walnut brown (# 221B) and yellow ochre (# 123C); unknown. The following species of anuran amphibians were flanks with a few vandyke brown (# 121) blotches; the iris found sympatric with the new species: Ansonia sp., of eye apple green (# 61); pupil black; tympanic fold on the Meristogenys whiteheadi and Rhacophorus angulirostris. inferior aspect edged with walnut brown (# 221B); tongue Because of the relative inaccessibility of the site, follow-up yellowish-pink, unpigmented; bars on limbs and webbing visits to collect additional specimens have not been possible. vandyke brown (# 121); and venter pale horn color (# 92). In preservative, gray-brown dorsally, with a darker gray line Comparisons. Ð Anuran eye coloration is known to be of in scapular region; dark lips barred with pale gray; fore and systematic importance (see review in Glaw & Vences, 1997), hind limbs gray brown with darker bars; webbing pale gray; and two south-east Asian anuran species has been described throat and pectoral regions with extensive gray mottling; recently on the basis of iris coloration - Philautus flanks of body with small, gray-black blotches; undersurfaces erythrophthalmus Stuebing & Wong (2000) and of limbs grayish-yellow with dark gray variegations; digits Leptobrachium sp. Ohler et al., 2004 (both known only from of arms and legs yellowish-gray. the respective holotypes, from Sabah, Malaysia [Borneo] and Laos, respectively). In possessing bright green iris, the new Measurements (in mm). Ð SVL 62.1; HL 19.8; HW 20.6; species from Gunung Murud can be separated from all other HD 11.1; BW 19.7; TBL 38.0; TD (vertical) 3.9; TD south-east Asian species of Polypedates. Additionally, (horizontal) 2.9; ED 7.7; UE 5.5; IN 5.2; IO 12.2; E-S 9.7; Polypedates chlorophthalmus, new species, can be E-N 7.2; E-T 2.3; A-G 31.9. differentiated from P. colletti (Boulenger, 1890) (range: southern Thailand, Peninsular Malaysia, the Natuna Islands, Sumatra and Borneo), on the basis of the following characters
Fig. 1. Lateral view of holotype of Polypedates chlorophthalmus, new species in life (SBC ZRC 1.11531).
Fig. 2. Close-up of head of the holotype of Polypedates Fig. 3. Head (3.1), hand (3.2) and feet (3.3) of holotype of chlorophthalmus, new species ZRC 1.11531, showing bright green Polypedates chlorophthalmus, new species ZRC 1.11531. Markers iris. Marker = 5 mm. = 5 mm.
267 Das: New species of Polypedates from Gunung Murud diagnostic of the latter species: snout acute; canthal ridge and toe IV with two phalanges free of web; and P. mutus indistinct; fingers basally webbed; webbing on toe IV to distal (Smith, 1940) (southern China, Myanmar), head longer than subarticular tubercle on outer edge and to beyond median wide; dorsum brown a dark interorbital bar and spots or four subarticular tubercle on inner edge; lips unbarred, but with longitudinal stripes; a dark hour-glass shaped mark on scattered dark pigments; and a distinct X-shaped mark on dorsum; iris yellow; and outer edges of thighs with bright dorsum. Polypedates leucomystax (Gravenhorst, 1829) (as yellow blotches. currently understood, comprises a complex of sibling species, see Narins et al., 1998; north-eastern India and southern China, south to Indo-Malaya and Indo-China, to the ACKNOWLEDGEMENTS Philippines and Timor), differs from the new species in possessing head longer than broad; snout tip subacute, Field work was supported by an Intensification of Research exceeding level of mandible; canthus rostralis vertical; iris in Priority Areas Grant (number: 08-02-09-10007-EA0001) dark brown; dorsum typically with four narrow, dark stripes from the Federal Government of Malaysia. The Universiti or plain or with mottled coloration in individuals from the Malaysia Sarawak and the Institute of Biodiversity and Bornean population; lips lacking dark bars, and throat and Environmental Conservation, Universiti Malaysia Sarawak lower flanks and anterior edge of thighs lacking dark provided laboratory and other facilities. I am grateful to my pigmentation. Polypedates macrotis (Boulenger, 1891) colleagues at the Institute of Biodiversity and Environmental (Peninsular Malaysia, Sumatra, the Mentawai Archipelago, Conservation and the Faculty of Resource Science and Borneo, the Natuna Islands, and islands of the southern Technology, Fatimah Abang, Andrew Alek Tuen, Mustafa Philippines), differs from the new species in showing snout Abd Rahman and Mohammad Tajuddin Abdullah, for their that projects beyond mandible; a dark broad temporal stripe; support during the field work and for their friendship. lips unbarred; iris yellowish-gray; skin of forehead ossified Collection permit was provided by the Sarawak Biodiversity to frontoparietal bones in adult females; and two to four, Centre, permit number SBC-RP-0070-ID. Export permit white, postcloacal tubercles present. The last Bornean species (number 04635) was provided by the Sarawak Forest of the genus, P. otilophus (Boulenger, 1893) (Borneo and Department. Curators of the following institutions permitted Sumatra), is diagnosable from the new species from Murud me to examine comparative material under their care: Lim in being larger (SVL in adult females to 97 mm); has a serrated Boo Liat (DWNP), Norsham Suhaina binti Yaakob (FRIM), crest over the tympanum; skin co-ossified to bony elements Arvin C. Diesmos (PNM), Eileen Yen and Margarita Naming of cranium; spiny protuberances at posterior of jaws (in (SBC), Jamili Nias, Maklarin Lakim and Paul Yambun (SP), adults); webbing on feet reaches median subarticular tubercle; Kamariah Abu Salim, Joseph K. Charles, David Edwards and a calcar on heel and frequently, on elbow; dorsum of body Helen Pang Yoke Yew (UBD), Kelvin Kok Peng Lim, Peter and hind limbs with numerous thick and thin dark longitudinal Kee Lin Ng and Chang Man Yang (ZRC), and J. R. B. Alfred lines; throat unpatterned; and iris yellow. and Shyamal Kumar Chanda (ZSI). Finally, I thank Rafe Brown and Robert Inger for comments on the manuscript. The new species from Gunung Murud is compared with all other known congeners from south-east Asia, listing only opposing suite of characters: LITERATURE CITED Polypedates dennysii (Blanford, 1881) (southern China and Myanmar, assigned to Rhacophorus by Wilkinson et al., Ahl, E., 1931. Anura III. Das Tierreich 55. Walter de Gruyter & 2002), iris yellow; SVL to 120 mm; toes webbed to base of Co., Berlin. XVI + 477 pp. digits; and dorsum green, with isolated pale blotches on dorsal Alcala, A. C. & W. C. Brown, 1998. Philippine Amphibians. An surface and flanks; P. dugretei David, 1871 (southern China illustrated fieldguide. Bookmark, Inc., Makati City. xii + 116 and Vietnam: preliminary molecular data presented by Yang pp. & Lin, 1997 support the placement of this nominal species Berry, P. Y., 1975. The Amphibian Fauna of Peninsular Malaysia. in the genus Rhacophorus, although Wilkinson & Drewes, Tropical Press, Kuala Lumpur. x + 130 pp. 2000, retain it in Polypedates, pending further studies), iris Boulenger, G. A., 1882. Catalogue of the Batrachia Salientia s. yellowish-brown; dorsum green with reddish-brown blotches; Ecaudata in the Collection of the British Museum. Second toe IV with two phalanges free of web; and fingers basally edition. British Museum, London. xvi + 503 pp., 30 pls. webbed; P. feae Boulenger, 1893 (southern China, Myanmar, Reprinted 1966, Wheldon & Wesley, Codicote & Verlag J. Thailand and Vietnam), SVL to 125 mm; head as long as Cramer, Weinham. wide; dorsum green; and fingers with webbing; P. insularis Boulenger, G. A., 1912. A Vertebrate Fauna of the Malay Peninsula Das, 1995 (Great Nicobar Island, India), dorsum typically from the Isthmus of Kra to Singapore including the Adjacent with an hour-glass marking; iris yellow; toe webbing to base Islands. Reptilia and Batrachia. Taylor and Francis, London. of disks of all toes, except toe IV; vomerine teeth separated xiii + 294 pp. by a distance equal to their own; and skin on forehead Bourret, R., 1942. Les Batraciens de l’Indochine. Mémoires de completely free; P. megacephalus Hallowell, “1860” 1861 l’Institut Océanographique de l’Indochine, Hanoi. 547 pp., Pls. (revived from the synonymy of P. leucomystax by Matsui et IÐIV. al., 1986; see also Inger et al., 1999: southern China, Thailand, Brown, W. C. & A. C. Alcala, 1994. Philippine frogs of the family Laos and Vietnam), iris yellow; dorsum brown with a dark Rhacophoridae. Proceedings of the California Academy of hour-glass mark on scapular region; a dark canthal region; Sciences, 48(10): 185Ð220.
268 THE RAFFLES BULLETIN OF ZOOLOGY 2005
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Treubia, Yang, D.-T. & Y.-S. Lin, 1997. Molecular phylogenetics and 31(3)(Supplement):1Ð133. biogeography of the genus Rhacophorus in Taiwan. American Society of Ichthyologists and Herpetologists, 77th Annual Meeting, Seattle, Washington (Abstract).
269 Das: New species of Polypedates from Gunung Murud
APPENDIX I Samarahan, Sarawak, Malaysia; SP 01189-93, Hutan Simpan, Mandamail, Pitas, Kudat, Sabah, Malaysia; UBD 05, 07Ð08, List of comparative material examined 73, Tasek Lama, Bandar Seri Begawan, Brunei Darussalam; UBD 552, Kampung Sungei Damit, Tutong, Brunei Darussalam; UBD 505 and 689, Batu Apoi, Temburong District, Brunei Polypedates colletti Ð DWNP A.0996, Interpretive Trail, Sungei Darussalam; UBD 429, Jalan Menggis Dua, Bandar Seri Relau, Taman Negara, Pahang, Peninsular Malaysia; IDÐ7932, Begawan, Brunei Darussalam; UBD 580, 588, Bukit Patoi, Gunung Santubong, Sarawak, Malaysia; PNM 7959 and 7964, Temburong, Brunei Darussalam; ZRC 1.903, Ayer Rajah Bako National Park, Sarawak, Malaysia; IDÐ7508Ð09, Long Campus, Singapore; ZRC 1.1347, Hindhide Drive, Singapore; Asap, Belaga, Sarawak, Malaysia; IDÐ7495, Niah National Park, ZRC 1.1498, Jurong, Singapore. Sarawak, Malaysia; UBD 494 and 536, Batu Apoi, Temburong District, Brunei Darussalam; UBD 657, Tasek Merembun, Polypedates macrotis— DWNP A.0994, Interpretive Trail, Sungei Brunei Darussalam. Relau, Taman Negara, Pahang, Peninsular Malaysia; DWNP A.1022Ð24, behind staff quarters, Sungei Relau, Taman Negara, Polypedates insularis Ð ZSI A8731 (holotype), ZSI A8732Ð34 Pahang, Peninsular Malaysia; PNM 7955, Gunung Gading (paratopotypes), ca. 2 km E mouth of Galathea River, Galathea National Park, Sarawak, Malaysia; IDÐ7555, Gunung Beremput, National Park, Great Nicobar, India; ZSI A8575, Campbell Bay, Sarawak, Malaysia; IDÐ7588, Gunung Gading National Park, Great Nicobar, India; ZSI A8735Ð36 (paratypes), Shompen Hut, Sarawak, Malaysia; IDÐSWAK 19, Lambir Hills National Park, Great Nicobar, India; ZSI A8737Ð40 (paratypes), ca. 2 km E Sarawak, Malaysia; IDÐ7773, Pulau Balambangan, Sabah, Kopen Heat, Great Nicobar, India. Malaysia; SP 00920, Danum Valley Field Centre, Lahad Datu, Polypedates leucomystax Ð DWNP-A.0978Ð80, Sungei Relau, Sabah, Malaysia; SP 20189, Ulu Padas, Long Pasia, Sabah, Taman Negara, Pahang, Peninsular Malaysia; PNM 7970, Malaysia; UBD uncat. (Chris Reading coll.), Batu Apoi, Gunung Penrissen, Sarawak, Malaysia; SBC A.00050, Gn. Temburong District, Brunei Darussalam. Ropih, Bau, Sarawak, Malaysia; SBC A.00169, Gn. Umbut, Bau, Polypedates otilophus — SBC A.00022, Gn. Apin, Bau, Sarawak, Sarawak, Malaysia; SBC A.00176Ð77, SBC A.00253, SBC Malaysia; SBC A.00170, Gn. Umbut, Bau, Sarawak, Malaysia; A.00274, Gn. Tabai, Bau, Sarawak, Malaysia; SBC A.00199Ð ZRC 1.3923, Gunung Buri, Sarawak, Malaysia; ZRC 201, Gn. Doya, Bau, Sarawak, Malaysia; SBC A.00235, Gn. uncatalogued, Lambir Hills National Park, Sarawak, Malaysia; Meraja, Bau, Sarawak, Malaysia; SBC A.00298, Gn. Podam, SP 02725, Tawau Hills Park, Sabah, Malaysia; SP 02425, Mahua Bau, Sarawak, Malaysia; ZRC Ð 1.11090, Gunung Gading Camp, Crocker Range National Park, Sabah, Malaysia; UBD National Park, Sarawak, Malaysia; ZRC 1.7672, 1.7709, Kota 537, Batu Apoi, Temburong District, Brunei Darussalam.
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 277-279 Date of Publication: 31 Dec.2005 © National University of Singapore
THE FORMER STATUS OF THE WHITE-SHOULDERED IBIS PSEUDIBIS DAVISONI ON THE BARITO AND TEWEH RIVERS, INDONESIAN BORNEO
Erik Meijaard The Nature Conservancy, J. Kuranji no. 1, Vorvoo, 75123, Samarinda, East Kalimantan, Indonesia Email: [email protected] (Corresponding author)
S. (Bas) van Balen Department of Animal and Plant Sciences, Sheffield University, Alfred Denny Building, Western Bank, Sheffield, England, UK Email: [email protected]
Vincent Nijman Zoological Museum, University of Amsterdam, PO Box 94766, 1090 GT Amsterdam, the Netherlands Email: [email protected]
ABSTRACT. Ð Analysing historic species ranges is important for assessing the population trend and conservation status of threatened species. Understanding the spatial variation in threats to these species requires an analysis of regional differences in historic and current status, and the underlying causes of their decline. Historic literature can help in this process. Here we provide a translation of field notes by Salomon Müller during his 1836 travels in SE Kalimantan. We focus on the notes regarding White-shouldered Ibis Pseudibis davisoni (Hume, 1875), one of Southeast Asia’s most critically endangered birds. Indonesian records for this species from the few last decades are mostly restricted to the Mahakam River in East Kalimantan, Indonesian Borneo. Müller recorded the species almost daily along the middle reaches of the Barito River in Central Kalimantan, and provided details on its food and morphology. Müller’s data suggest that in the 19th century a healthy population of White-shouldered Ibis was present along the Barito River. We speculate on the connection of this population with the Mahakam one and the cause of its apparent extinction.
KEY WORDS. Ð bird, Borneo, endangered species, historic range, historic literature.
INTRODUCTION Collar et al. (2001) provided lengthy and detailed descriptions of the historic conservation status of many threatened A proper assessment of population trends and threats to Southeast Asian birds. One of the most critically endangered species is needed to understand the processes that cause birds of Southeast Asia, and one that has undergone a dramatic population declines. Understanding the historic situation decline over the course of the 20th century, is the White- provides important collateral data. What we need to know is shouldered Ibis Pseudibis davisoni. Once distributed over whether a particular species was historically rare, or whether large parts of Southeast Asia, from Myanmar and Vietnam large changes in the species’ status have occurred in relatively in the north to Peninsular Malaysia and Borneo in the south, recent times. Knowledge of this points to potentially limiting it is nowadays confined to pockets of riverine and swamp factors. For instance, a species that has traditionally been rare forest in Myanmar, Vietnam and Indonesian Borneo. At is probably limited by ecological factors (e.g., a species that present, the largest single population is confined to the only feeds in a particular rare habitat type). Species that have Mahakam River in East Kalimantan where the species is recently declined are probably affected by ‘modern’ threats regularly recorded (Sözer & van der Heijden, 1997; Collar like large-scale deforestation, human disturbance, or forest et al., 2001,VN & EM, pers. obs.). Other Bornean records fires. Especially for species that range over large areas it is are from the Seruyan River, a tributary of the Barito River important to assess their historic status on a region by region in 1984 (Holmes & Burton, 1987), the northern Barito basis. This way, regional differences between the historic (‘Barito Utara’) in 1979 (J. T. Marshall in Smythies, 1999), status, current status, and local causes of decline can and possible sightings, in 1974, on the Banjarmasin airport, contribute to understanding the processes that reduce at the mouth of the Barito River, and in Binuang just northeast population numbers of the species. of Banjarmasin (Holmes & Burton, 1987). Older Barito
277 Meijaard et al.: Former status of White-shouldered Ibis in southwest Borneo records date from Purukcahu in the upper reaches of the river [page 226] ‘Along the borders [of the Teweh River up to (Chasen & Kloss, 1931) in 1909, and the always unspecified Kampung Silo] we often saw little groups of sooty-black record from 1836 by S. Müller. Collar et al. (2001) provided ibises.’ a detailed assessment of the historic status of the Asian mainland populations of White-shouldered Ibis, but gave few Note 66: ‘The unfeathered parts of the specimen that we shot details on its historic status in Borneo (NB: A number of were coloured as follows: Skin of head and chin sooty-black; localities on their map have been mixed up). the anterior side of the naked neck is bluish-white, grading posteriorly into sky-blue; bill is lead-coloured grading into Salomon Müller (1804-1864) was a German taxidermist from purple near the basal part; legs are a dirty light-red, although Heidelberg, who, as a member of the Natuurkundige in older birds this seems to change to a darker, more purplish Commissie voor Nederlandsch Indië travelled in July Ð hue.’ December 1836 through southeast Kalimantan. Müller collected birds, amongst others seventy taxa new to Borneo (Smythies and Davison, 1999), but also some plants on the DISCUSSION Troe and Palandau tributaries of the Karau River (note that Karau is the Punan and Bakumpai general word for stork, Holmes (1991) discussed the status of the White-shouldered Sözer & van der Heyden 1997) a tributary of the Barito (Van Ibis in Kalimantan and noted for the Barito River that the Steenis-Kruseman, 1950). He observed the White-shouldered species has always been rare. This perception is Ibis and collected several specimens. As his detailed records understandable given that the last confirmed record from this are written in Dutch, we consider it worthwhile reproducing river dates back to 1984. Also more recently, the White- those sections relevant to understanding the biology of the shouldered Ibis has not been recorded by ornithologists who White-shouldered Ibis. We provide a translation of Müller’s surveyed the river and its tributaries over the last two decades original Dutch text. In addition, we use this note as a reminder (e.g. Wilkinson et al., 1991; van Balen & Prentice, 1997; see of the importance of historic literature for the conservation Collar et al., 2001). However, as appears from his field notes, of many species that remain virtually unknown. Where in Müller’s time it was commonly observed along the Barito necessary, we provide details between square brackets. We River, especially in its middle reaches. For the Mahakam determined geographic coordinates in the text by comparing River in the early 1990s, Sözer & Nijman (2005) reported an Müller’s detailed map with a recent digital map of the region. encounter rate of 0.6 Ð 2.5 birds per 100 km, with a large variation between sites. Maximum encounter rates at some sites could be as high as 16 birds per 100 km, with birds SALOMON MÜLLER’S ACCOUNT being observed almost daily. If we consider that Müller could travel some 20-30 km a day, the historic abundance of White- [pages 198-199] ‘Near Becebang [three small islands in the shouldered Ibis along the Barito River could be as high as Barito River, south of Muara Kalahian, 1¼42'36"S that along some stretches of the Mahakam River in the 1990s. 114¼49'48"E some 250 km from the mouth the Barito River], Although admittedly highly speculative, given the presence and from there on almost daily, we occasionally saw a sooty- of White-shouldered Ibis along the middle reaches of the brown ibis with white shoulder feathers. It is known among Barito and Teweh Rivers (c. 250 km in length), the total local people as Burung Karau. Usually, we observed groups population along this river could easily have been a hundred of two, and rarely of three or four birds together. Their individuals or more. frequent, slightly mournful call is rather loud, but despite this they are very secretive. Only after many failed attempts I The Barito River population is spatially separated by that of managed to approach a few birds by stalking them through the Mahakam River basin by two narrow mountain ranges: forest, across the [Barito] river from Tanjung-Jawa village the Müller range in the upper reaches of both rivers, and a [1¼36'00"S 114¼49'48"E]. There, I shot one bird from a pair low range between the Teweh and Lahai Rivers, tributaries feeding among Pekumpai-grass [Panicum stagninum Retz]. of the Barito River, and Pelajan River, a tributary of the This male bird was typified like several other male and female Mahakam River (Smyhties, 1981; Sandy, 1986). Straight line birds that we obtained later by the somewhat wrinkled skin distances between the river basins are locally less than 25 on the head and the nipple-shaped appendices on the neck. km, and possibly the ibises of these basins in Müller’s days Despite the presence of these features, which may only appear formed one single population. during mating season, I do not consider this species to be different from Ibis papillosa [=Black ibis Pseudibis papillosa Why did the species disappear from the Barito and Teweh from mainland Asia], which was described and depicted by Rivers? There are two obvious candidates for its causes, not Mr. Temminck in his Planches Coloriees (see note 66). mutually exclusive: human disturbance by hunting, and by Although we have not observed this ibis on any other island deforestation. But have these factors been much more of the Indonesian archipelago it was not rare at this latitude prevalent along the Barito River during the 19th and early and further up the Barito and Tewei [=Teweh, a tributary of 20th century than along the Mahakam River where the species the middle Barito River, branching off at Muara Teweh still occurs? Before answering that question we focus on the (0.87¼S 114.93¼E)] Rivers, and even north of the equator. In key ecological requirements of White-shouldered Ibis. its stomach we found remains of worms and larvae of aquatic Müller’s information suggests that the species is most insects.’ common along the strongly meandering middle section of
278 THE RAFFLES BULLETIN OF ZOOLOGY 2005 the Barito, where many oxbows occur. Similarly, along the ACKNOWLEDGMENTS Mahakam River, the species is most common along the meandering middle section of that river. These swamp areas We thank Nigel Collar for his insightful review of the are prone to frequent flooding and probably provide key submitted manuscript. We thank the Bogor library of the habitats like seasonally flooded grasslands, lakes and pools, Indonesian Institute of Sciences (LIPI) for lending us a copy and marshes and muddy streams (Collar et al., 2001). of Müller’s book.
Information from the mid-1850s (Schwaner, 1853) suggests that the middle Barito River was surrounded by mostly LITERATURE CITED continuous forest. Dry grasslands were restricted to some minor parts of the region and areas of shifting cultivation van Balen, S. & C. Prentice, 1997. Birds of the Negara River Basin, rapidly converted back to shrubland and secondary forests South Kalimantan, Indonesia. Kukila, 9: 81-107. (Schwaner, 1853). Panicum stagninum grasslands, i.e. the Bock, C., 1881. The head-hunters of Borneo. London. vegetation in which Müller shot his specimen, were restricted Boschwezen, 1931/1932. Begroeingskaart. Vluchtig opgenomen to swampy riverside flatlands. Knapen (2001), however, door de Onderafdeeling (Kantoor Bandjermasin) van het reports that the area of the lower and middle Barito was Boschwezen in 1931/1932. Scale 1: 100,000. rapidly developed after the 1850s following a dramatic shift Chasen, F. N. & C. B. Kloss, 1931. Additions to the list of Bornean from pepper to rice growing. Human population densities birds. Bulletin of the Raffles Museum, 4: 124. along the middle Barito River were very low in the 1850s Collar, N. J., J. V. Andreev, S. Chan, M. J. Crosby, S. Subramanya with only 1,905 inhabitants recorded along a 200 km stretch & J.A. Tobias (eds.), 2001. Threatened birds of Asia: the of the middle Barito River (Schwaner, 1853: 105), but these BirdLife International red data book. Cambridge: BirdLife grew rapidly following the expansion of rice growing. By International. about 1880, most of the lower and middle Barito area was Holmes, D. A., 1991. Note on the status of white-shouldered ibis ‘terribly devastated’ because of swidden cultivation and in Kalimantan. Kukila, 5: 145-148. almost no primary forest was left (Knapen, 2001). Holmes, D. A. & K. Burton., 1987. Recent notes on the avifauna of Kalimantan. Kukila, 3: 2-32. Unlike the Barito River, forest loss along the Mahakam River is much more recent, and those sections where White- Knapen, H., 2001. Forests of fortune? The environmental history shouldered Ibis are found remain forested. When the of Southeast Borneo, 1600-1880. KITLV Press, Leiden. Norwegian naturalist Bock (1881) travelled up the Mahakam Müller, S., 1857. Reis in het zuidelijk gedeelte van Borneo gedaan River in 1878, he reported that most of the area around the in het jaar 1836. Pages 129-326 in S. Müller, editor. Reizen en lakes [0º33’S 116º87'E] and upstream from there was still onderzoekingen in den Indischen Archipel gedaan op last der Nederlandsche Indische Regering tusschen de jaren 1828 en completely forested. A Dutch map from the early 1930s 1836. Vols. 1 and 2. Koninklijk Instituut voor de Taal-, Land- (Boschwezen, 1931/1932) shows that the area around the en Volkenkunde van Nederlandsch-Indië, Amsterdam, The Mahakam Lakes and at least 120 km upstream from there Netherlands. was still covered in dense forests. In fact, most of these swamp Sandy, I. M., 1986. Atlas Indonesia (sixth edition). Jakarta: PT forests were still intact up until the major fires of the early Dhasawarna & Universitas Indonesia. 1980s, as shown on satellite imagery. As such, deforestation and rapidly increasingly human population densities is much Schwaner, C. A. L. M. 1853. Borneo. Beschrijving van het stroomgebied van den Barito en reizen langs eenige voorname more recent along the Mahakam than the Barito River. rivieren van het zuid-oostelijke gedeelte van dat eiland in de jaren 1843-1847. 2 Vols. van Kampen, Amsterdam, the Loss of riverine forest will not only have led to a loss of Netherlands. breeding sites, but also to secondary effects such as a change Smythies, B. E. & G. W. H. Davison, 1999. The birds of Borneo. of the depositional character of river habitats (with finer 4th Edition. Kota Kinabalu: Natural History Publications substrates and more eroded banks), decrease in water clarity (Borneo). (as result of an increase in erosion) and an increase in Sözer, R. & A. J. W. J. van der Heijden, 1997. An overview of the temperature regime (as a result of loss of over-hanging trees) distribution, status, and behavioural ecology of the White- which may interfere with the birds’ ability to find food shouldered ibis in East Kalimantan. Kukila, 9: 126-140. (Nijman & Sözer, 2005). Furthermore, loss of forest would Sözer, R. & V. Nijman, 2005. Effects of ENSO-induced forest fires have led to an increased accessibility to nesting and feeding and habitat disturbances on abundance and spatial distribution sites, possibly leading to increased egg and chick collecting of an endangered riverine bird in Borneo. Animal Conservation, and hunting. 8: 27-31. van Steenis-Kruseman, M. J., 1950. Malaysian plant collectors and collections. Djakarta: Noordhoff-Kolff. Wilkinson, R., G. Dutson, B. Sheldon, Darjono & Y. R. Noor, 1991. The avifauna of the Barito Ulu region, Central Kalimantan. Kukila, 5: 99-116.
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THE RAFFLES BULLETIN OF ZOOLOGY 2005 53(2): 281-286 Date of Publication: 31 Dec.2005 © National University of Singapore
EAT IN OR TAKE AWAY Ð SEED PREDATION AND REMOVAL BY RATS (MURIDAE) DURING A FRUITING EVENT IN A DIPTEROCARP RAINFOREST
Konstans Wells Department of Experimental Ecology, University of Ulm, Albert-Einstein Allee 11, D-89069 Ulm, Germany Email: [email protected]
Robert Bagchi Department of Animal and Plant Science, University of Sheffield, Western Bank, Sheffield S10 2TN, U.K. Email: [email protected]
Abstract. Ð Seed predators are considered important in the evolution of mast fruiting. Mast events in turn provide an abundant food resource for consumers, but only for a limited period of time. In this study seed removal experiments, feeding trials and small mammal trapping were used to determine the potential of forest rats (Muridae) as predators and dispersers of two tree species, Lithocarpus gracilis (Fagaceae) and Parashorea malaanonan (Dipterocarpaceae) during a minor fruiting event in a dipterocarp rainforest at Danum Valley (Sabah, Malaysia) in Sept/Oct 2004. Seeds of both species were exploited by rodents at similar frequencies. Nearly all seeds exploited were removed from the feeding stations. Seeds placed near burrows of Maxomys rajah were moved into the burrows. Gnawed seeds collected in fruit fall traps (8% of total collected) indicated that Parashorea seeds were also exploited in the canopy. Partially eaten seeds often retained their ability to germinate. Captive rats ate a range of dipterocarp and other hard-shelled seed and feeding on an individual seed could take up to 27 minutes. Trap success for rats, but not tree shrews (Tupaiidae) and civets (Viverridae), decreased during the fruiting season. This supports the idea that this group benefits from mast fruiting. Rats are evidently important predators of the seeds of these tree species, both before and after dispersal. However, due to the removal of seeds before eating, long consumption times and the ability of partially consumed seeds to germinate, they may also function as secondary dispersal agents.
KEY WORDS. Ð Seed dispersal, scatter hoarding, predator satiation, Parashorea, Lithocarpus, Maxomys.
INTRODUCTION Several hypotheses have been suggested to account for the evolution of mast fruiting (Kelly, 1994). Of these, the most Fruit production in tropical forests is highly variable through widely accepted suggests that synchronous fruiting evolved time and often occurs in synchronised “mast” events (Curran to satiate seed predators allowing greater per capita et al., 1999; Leighton & Leighton, 1983; Smythe, 1986). In recruitment (Janzen, 1971, 1974; Kelly, 1994; Curran & southeast Asian dipterocarp rainforests, mast fruiting has been Leighton, 2000; Kelly & Sork, 2002; Sakai, 2002). well reported as a community-wide event including seeds of Dipterocarp seeds (we use the term ‘seed’ rather than ‘fruit’ the Dipterocarpaceae but also other species with different fruit for dipterocarp and lithocarp seeds that comprise a nutritious phenology (Ashton et al., 1988; Curran & Leighton, 2000; endosperm essential for the germination, except if fleshy fruits Sakai, 2002). However, the degree of synchrony between trees are included in the argumentation) are an easily accessible in fruiting may range from a few individuals of one tree and energy-rich food resource with thin testa that are attractive species to a diverse community of various taxa and fruit types. to Sus barbatus (bearded pigs) and possibly to several rodents. Several species that participate in mast events also produce Reproductive success of this vulnerable seed type is increased fruit at other times, with some fruit produced throughout the by predator satiation via masting and also by rapid year. The pay off for any specific pattern of synchrony and germination of the lipid-rich seeds within one week (Curran quantity in fruiting will depend on 1) interactions with the & Webb, 2000). Abundant fruit crops are simultaneously abiotic environment (Kelly, 1994; Kelly & Sork, 2002; Sakai, produced by Lithocarpus (stone oaks), comprising another 2002); 2) fruit characteristics (e.g. size, protection; Janzen, important element of dipterocarp forests (Cannon & Manos, 1971); 3) interactions with other fruiting species (Curran & 2003). Lithocarpus fruit every year, and so are not considered Leighton, 2000) and 4) dynamic interactions with the masting species (Sakai et al., 1999). The structure of hard- consumer and disperser communities (Kelly, 1994; Kelly & shelled nuts in a fused cupule should provide protection Sork, 2002). against predation of these seeds. Meanwhile the long
281 Wells & Bagchi: Seed predation and removal by rats germination period of one to nine months (Ng, 1991) both collected from the forest floor. A thin thread with a coloured increases the exposure to consumers and makes them suitable flag was attached to each seed, winded between fruit and for caching. Caching by rodents appears likely though not sepals/cupules, respectively, and 55 experimental stations yet fully proven (see Corlett, 1998) and may further increase consisting of one seed of each species where arranged linearly resource utilisation in time. 2-3 m apart in two areas (505 station nights in total). Further seed stations (three per burrow) were established near burrows However, seed fates and the diversity of seed consumers are of Maxomys rajah (Thomas.) (n = 2) that where located little investigated for Oriental tropical rainforests (see Blate beforehand. Each station was monitored daily for up to 11 et al., 1998; Wong et al., 1998; Sodhi et al., 2003), whereas days. Disappeared or damaged seeds were replaced upon each seed predation and dispersal by diverse rodent assemblages check and the germination of non-removed seeds was has been frequently recorded for Neotropical forests (e.g. recorded. We considered seeds found less than 50 cm away Fleming, 1979; Asquith et al., 1997; Hoch & Adler, 1997; from the original position as displaced, seeds not found within Brewer & Rejmanek, 1999; Charles-Dominique et al., 2003). 50 cm as removed. The selective removal of only single seeds, Rats (Muridae) are abundant and omnivorous forest dwellers remains of gnawed off sepals or cupules or cracked seeds in various strata of dipterocarp rainforests (Harrison, 1962; with incisor marks were considered to indicate rodents as the Wells et al., 2004), yet their role as seed predators and most likely predators. A removed seed could have either been dispersers have been little investigated (but see Blate et al., predated or dispersed, but we were not able to distinguish 1998; Curran & Leighton, 2000). Their effective utilisation between the two. of three-dimensional space in the forest suggest great potential for seed predation in space and time due to arboreal foraging A juvenile Maxomys cf. rajah/surifer and a female adult and possible caching. However, as rats are also exposed to Maxomys whiteheadi (Thomas.) were kept in captivity for carnivore predators during feeding, their seed handling and three to four days for feeding observations before release at consumption time is also subject to constraints. The the place of capture. Fruit of various species (Neoscortechinia constitution and nutritional value of different seeds types should determine the handling time, proportion of sp. (Euphorbiaceae), Shorea fallax, S. leprosula, S. parvifolia, consumption of a single seed, and whether seeds are removed. P. malaanonan, Dryobalanops lanceolata (all Different seed characters and distributions should thus Dipterocarpaceae), L. gracilis (Fagaceae), Aglaia elliptica specifically influence resource exploitation and resulting seed (Meliaceae), four species of Ficus sp. (Moraceae) and fate, and hence may be an important proximate determinant Tetrastigma sp. (Vitaceae)) were offered simultaneously of satiation strategies. Here we report on a seed fate study every evening. Feeding was observed continuously for the and feeding observations in order to estimate the role of rats first hour of each trial, after which the rat was left undisturbed as predator and dispersal agents on two seed types except for periodic cleaning and provisioning with clean (Parashorea and Lithocarpus) that were both commonly water. During the first hour, the time spent eating fruit of available during a dipterocarp fruiting event in a Bornean each species exploited and any movement of fruit were rainforest. recorded. At the end of every trial, the number of fruit of each species consumed was noted. Damaged or consumed fruit were then replaced for the start of the next trial. Damaged MATERIALS AND METHODS fruit were placed on moist tissue paper to assess their viability. Seeds were checked for the presence of a radical every two This study was conducted at the Danum Valley Conservation days for a period of one month after collection. Area (4°57.800'N 117°48.173'E), Sabah, Malaysia. The conservation area includes more than 43,000 ha of lowland To assess rodent predation on dipterocarp seeds in the canopy, evergreen rainforest and is surrounded by 973,000 ha of seed damage was recorded for seeds gathered with seed traps selectively logged forest. The vegetation is dominated by the placed near fruiting P. malaanonan. We established seed traps Dipterocarpaceae in the canopy and Euphorbiaceae in the in at distances of 2, 10, 20, and 30 meters along a transect understorey (Newberry et al., 1992; Marsh & Greer, 1992). line at ten separate trees with large fruit crops. Seeds from A partial dipterocarp mast-fruiting event occurred in the traps were gathered every four days for a period of 2 September Ð November 2004. The principle species involved months. Seeds were classed as undamaged, potentially viable were Parashorea malaanonan (Merr.), Shorea leprosula (more than half of the seed remaining) or predated (seeds (Miq.), and S. parvifolia (Dyer.). Fruits were also available were more than half consumed or cracked into pieces). from several species that fruit more regularly including Potentially viable seeds were put on moist tissue paper in the Lithocarpus gracilis (Korth.) Soepadmo (Fagaceae), several laboratory to monitor germination ability as outlined above. Ficus species (Moraceae), several Euphorbiaceae and Rubiaceae among others. P. malaanonan and L. gracilis fruits To compare the trap success of rats and other small mammals were particularly abundant. between the fruiting season in September Ð October 2004 and other seasons, locally made cage traps (28 x 14 x 14 cm) A seed choice and removal experiment was conducted with baited with ripe banana were placed for 16 consecutive days two types of seeds: P. malaanona (mean weight = 5.0 g ± on the forest floor in March and December 2003 and in 0.10, n = 292) and L. gracilis (mean weight = 3.1 g ± 0.7, September Ð October 2004, comprising a total trapping effort n = 90). Seeds with no signs of predation or germination were of 5,967 trap nights. During the seed fall period, P.
282 THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 1. The fate of seeds exploited by rodents at 55 feeding stations (505 station nights in total).
Parashorea Lithocarpus Eaten at station 3 0 Displaced 1 7 Removed 5 4 Total 911
Table 2. Proportion of consumed seeds and ‘explored’ seeds that were removed or found with gnawed off wings or radicals in a feeding choice experiment. Data are pooled for all trials. Maximum handling times [min.] of seeds are given in brackets. Feeding trials lasted for four nights for M. rajah/surifer and three nights for M. whitheadi with four fruits of each species offered.
Species Family M. cf. rajah/surifer M. whiteheadi Consumed Explored Consumed Explored Neoscortechinia forbesii Euphorbiaceae 0.81 0 0.08 0.08 Lithocarpus gracillus Fagaceae 0.13 0.38 0.17 0.25 Dryobalanops lanceolata Dipterocarpaceae 0 0 0 0 Parashorea malaanonan Dipterocarpaceae 0.50 (27) 0.25 0.08 0.66 Shorea fallax Dipterocarpaceae 0.25 (10) 0 0.33 0 Shorea leprosula/ parvifolia Dipterocarpaceae 0.41 (4) 0.24 0.91 0 Aglaia elliptica Meliaceae 0 0 0 0 Ficus sp. (4 species) Moraceae 0 0 0 0 Tetrastigma sp. Vitaceae 0 0 0 0 malaanonan and L. gracilis seeds were placed in some traps Both M. rajah (n=1) and Leopoldamys sabanus (Thomas.) to confirm their consumption by small mammals. For a (n=1) consumed the P. malaanonan and L. gracilis seed diversity estimate we used the Shannon-Wiener H’ index and provisions placed in traps they were caught in. We were not compared H’ between different trapping periods with 10,000 able to test further species. runs by use of the “Species diversity and richness” software (Henderseon & Seaby, 1998). A total of 424 seeds were recovered from the seed traps. Out of these, 390 seeds (92 %) were undamaged. A further 18 (4 %) seeds were less than half consumed and potentially still RESULTS viable, while 16 (4 %) seeds were more than half consumed or cracked into pieces with no further germination ability. Rodents appeared to frequently utilise P. malaanonan and L. Of the 18 half consumed seeds collected, two germinated in gracilis seeds as a food resource. At the experimental feeding the laboratory. stations, nine P. malaanonan (16%) and 11 L. gracilis (20%) seeds were exploited (Table 1), most probably by rodents. Maxomys cf. rajah/ surifer and M. whiteheadi kept in captivity Consumption of exploited seeds near the place of encounter both fed on fruit of all dipterocarps tested (except D. was low. Only two P. malaanonan seeds were completely lanceolata), L. gracilis and Neoscortechinia sp.. Fruits of the eaten near (within 50 cm) the feeding station. The radicals four Ficus species, A. elliptica and Tetrastigma sp. were of another two P. malaanonan seeds were gnawed off. The rejected by both species (Table 2). During every feeding trial, wings and attached flags were gnawed off the other P. the rats fed on a mixed fruit diet without finishing the available malaanonan seeds before removal to an unknown fate. No amount of any single fruit resource. Dipterocarp seeds were exploited P. malaanonan seeds recovered were viable. While moved a short distance from their original position, the sepals the removal rate was similar for L. gracilis, seeds were more gnawed off and feeding on a single seed lasted for up to 27 frequently displaced (distances between 10-30 cm) and two minutes on one P. malaanonan seed. Seeds from L. gracilis of these displaced seeds were found buried. No L. gracilis were frequently hidden under leaves or in corners of the cage seeds were eaten in situ. while consumption was less than on the available dipterocarp seeds. In most cases, seeds were only partially eaten, and some Seeds placed in the vicinity of Maxomys rajah burrows were half-consumed dipterocarp seeds were still able to germinate. moved inside them, indicated by the flags with threads leading down the holes. We recorded four removals of P. malaanonan Overall, 188 small mammals (18 species) were captured seeds, one removal of a L. gracilis seed and one simultaneous during the three trapping periods (Table 3). The number of removal of two P. malaanonan and one L. gracilis from near species trapped (range eight to 13 species) and diversity the burrows. (Shannon-Wiener H’, range 1.57 to 1.97, p < 0.05 for all
283 Wells & Bagchi: Seed predation and removal by rats
Table 3. Trap success per 1,000 trap nights for the taxonomic groups of small mammals captured during the three trapping periods. Trap nights per period are given in brackets.
Family No. of species trapped No. of individuals per 1,000 trap nights March 2003 Dec 2003 Sept/Oct 2004 (1915) (2025) (2027) Muridae 10 32.37 27.65 4.93 Sciuridae 2 2.09 2.47 0 Tupaiidae 4 9.40 3.95 9.86 Viverridae 2 0.52 0 1.97 Total 18 44.38 34.07 16.76 Shannon-Wiener H’ 1.97 1.84 1.58 comparisons, Table 3) were lowest during the fruiting season Beck & Terborgh, 2002). Therefore, rats can be expected to in Sept/Oct 2004. The capture of certain species varied forage on scattered seeds or at the edges of seed shadows in between trapping periods. During the fruiting season in Sept/ a wide variety of habitat patches that are less attractive or Oct 2004, trap success (individuals/ per trap night) was inaccessible to larger mammals like bearded pigs (Sus significantly lower for rats (Pearson’s Chi Square: x2 = 19.88, barbartus). Small arboreal species, e.g. the omnivorous and df = 2, p < 0.05). An increased (though not significant) trap abundant tree rat Niviventer cremoriventer (Miller.) (mean success was recorded for tree shrews (Tupaiidae) and civets weight of adults approximately 69 ± 13 g, n = 142), are agile (Viverridae) during the fruiting event, indicating that the trap enough to traverse thin distal branches of less than only 2- success for rats and other small mammal is influenced by the 3 mm in diameter (K. Wells, pers. obs.), where they might seasonal availability of abundant fruit resources. explore fruits at an early developmental stage. The small size of rodent seed predators is not only of significance for patterns of seed exploitation on the forest floor but also in three- DISCUSSION dimensional space. Our results in combination with these traits further encourage the idea that seed utilisation by rodents is Our results confirm that both terrestrial rats of the genus much more extended in space (e.g. arboreal exploitation) and Maxomys and the semi-arboreal L. sabanus feed on time (e.g. predation on early developmental stages and dipterocarp and lithocarp seeds (see Curran, 1994; Blate et caching) compared to larger predators. al., 1998; Curran & Leighton, 2000). Much of the literature dealing with dipterocarp fruiting concerns regional mast According to the interfamilial predator satiation hypothesis events (e.g. Curran & Leighton, 2000; Sakai, 2002). Curran (Curran & Leighton, 2000) resident seed consumers, like and Leighton (2000) suggest that bearded pigs, the most rodents, prefer non-dipterocarp seed resources. As a result, important seed predators in their study, are satiated at the when seeds of other species are abundant, dipterocarps suffer regional rather than local scale. This mechanism is dependent much less predation by residents. However, in this study, on pigs foraging together in large groups across large areas resident seed predators appeared to prefer dipterocarp seeds. to feed on dipterocarp seeds (Curran & Webb, 2000), This may have partly been due to the paucity of other small however, a behaviour that was not observed during the fruiting hard shelled fruit during this period (except for dipterocarps event described here and is probably only triggered by and Lithocarpus). Low trap success of common rodents regional masting events. Recent studies in Sabah record some during the fruiting season and a lack of such a change for seed survival for a few dipterocarp species even in non-mast small mammals probably incapable of consuming dipterocarp years (R. Bagchi, C. Philipson, unpublished data). While seeds (tupaiids and viverrids) are also consistent with this bearded pigs were responsible for some seed predation (R. conclusion. Bagchi, pers. obs.) rodents, which are much less mobile though omnipresent, appeared to be the main seed predators The exploitation of seeds offered at seed stations revealed in the study presented here. The predators and hence the scale no quantitative difference between the exploitation of at which predator satiation occurs may differ in non-mast Parashorea and Lithocarpus seeds. The majority of seeds years or in different localities. Overall, seed predation by exploited from both species were removed from the feeding rodents may exert a very different selective force on fruiting station. However, seed handling seems to differ as displaced patterns to that exerted by pigs. seeds of Parashorea were consumed, whereas some displaced lithocarp seeds were buried. The differences in morphology Small seed predators (e.g. rats and squirrels) may be more and germination time between the two seed types probably efficient at foraging for scattered seeds as a single seed will affect the fate of removed seeds, for example the probability provide proportionally more energy than for larger species of consumption or caching (Forget, 1991; Jansen & Forget, (see MacArthur & Pianka, 1966; Smythe, 1986). Collectively, 2001). Dipterocarp seeds germinate very rapidly making them foraging small seed predators are expected to effectively unsuitable for scatterhoarding. In contrast, L. gracilis seeds discover most resource patches (Janson & Emmons, 1990; germinate after between 173-178 days (Ng, 1991) making
284 THE RAFFLES BULLETIN OF ZOOLOGY 2005 them more suitable for caching and delayed consumption. for identifying fruits. K.W. appreciates grants form the The records of buried seeds as well as the seed storing German Academic Exchange Service (DAAD) and academic behavior of captive rats make scatterhoarding of Lithocarpus support from Elisabeth K.V. Kalko and Martin Pfeiffer. R.B. seeds by rodents likely. So far, there is very little information received a studentship from the Natural Environmental available on the extent of caching by rodents and its possible Research Council (NERC) and is grateful to Malcolm Press impacts on seed dispersal in Oriental rainforests (Leighton and Julie Scholes for academic supervision and field & Leighton, 1983; Corlett, 1998). Despite the differences assistance from Adzley Madran. We thank Navjot S. Sodhi between the seed types, both seeds may be dispersed and an anonymous reviewer for constructive comments on inadvertently by rodents moving them. Because Lithocarpus an earlier draft of the manuscript. seeds are more likely to be cached they will be more likely to be forgotten and survive. LITERATURE CITED Movement of dipterocarp seeds into burrows or other places is most likely a strategy to consume the seed in a protected Adler, G. H., 1998. Impacts of resource abundance on populations place rather than a scatterhoarding activity. Given the of a tropical forest rodent. Ecology, 79(1): 242-254. potentially long time required for complete consumption of Ashton, P. S., T. J. Givnish & S. Appanah, 1988. Staggered flowering Parashorea seeds, a rat would expose itself to a high risk of in the Dipterocarpaceae: new insights into floral induction and predation if it consumed the seed at the point of discovery. the evolution of mast fruiting in the aseasonal tropics. American This movement of seeds will add to the handling time prior Naturalist, 132(1): 44-66. to seed consumption, and this may increase the likelihood of Asquith, N. M., S. J. Wright & M. J. Clauss, 1997. Does mammal local predator satiation. community composition control recruitment in neotropical forests? Evidence from Panama. Ecology, 78(3): 941-946. Studies of caching may also help our understanding of the Beck, H. & J. Terborgh, 2002. Groves versus isolates: how spatial impact that resource availability has on the population aggregation of Astrocaryum murumuru palms affects seed dynamics of seed consumers. Cached seeds, while they extend removal. Journal of Tropical Ecology, 18(2): 275-288. the availability of the resource for hoarding species, are Blate, G. M., D. R. Peart & M. Leighton, 1998. Post-dispersal unavailable to other species. Frugivorous mammals that have predation on isolated seeds: a comparative study of 40 tree been studied in detail show seasonal patterns in population species in a Southeast Asian rainforest. Oikos, 82(3): 522-538. size and reproductive activity, supporting the idea that Brearley, F. Q., S. Prajadinata, P. S. Kidd, J. Proctor & Suriantata, temporal or limited resource availability is a major force in 2004. Structure and floristics of an old secondary rain forest in structuring these populations (Terborgh, 1986; Wolff, 1996; Central Kalimantan, Indonesia, and a comparison with adjacent Adler, 1998; Schnurr et al., 2002). We might therefore expect primary forest. Forest Ecology and Management, 195(3): 385- 397. that seed hoarders will show less fluctuation in their population sizes than other species. Brewer, S. W. & M. Rejmanek, 1999. Small rodents as significant dispersers of tree seeds in a Neotropical forest. Journal of Vegetation Science, 10(2): 165-174. At the moment we little understand the interactions between rodent seed predators and their food sources. The absence of Cannon, C. H. & P. S. Manos, 2003. Phylogeography of the large mammals and human disturbance affect small mammal Southeast Asian stone oaks (Lithocarpus). Journal of Biogeography, 30(2): 211-226. abundances and seed predation (Terborgh, 1986; Asquith et al., 1997; Wright et al., 2000; Caro, 2002; Wyatt & Silman, Caro, T. M., 2002. Factors affecting the small mammal community 2004). Further, variability in tree species composition among inside and outside Katavi National Park, Tanzania. Biotropica, 34(2): 310-318. forest patches and most severely in secondary forests affects seed availability and predation (Sodhi et al., 2003; Brearley Charles-Dominique, P., J. Chave, M. A. Dubois, J. J. De Granville, et al., 2004). Rats and squirrels are often the last mammals B. Riera & C. Vezzoli, 2003. Colonization front of the understorey palm Astrocaryum sciophilum in a pristine rain to remain in disturbed forest fragments (Corlett, 1998). forest of French Guiana. Global Ecology and Biogeography, Anthropogenic change could therefore increase the 12(3): 237-248. importance of rodent seed predation quite drastically. The Corlett, R. T., 1998. Frugivory and seed disperal by vertebrates in potential effect this may have on tree recruitment is the Oriental (Indomalayan) region. Biological Reviews of the unpredictable but is likely to be considerable. Cambridge Philosophical Society, 73: 413-448. Curran, L. M. 1994. The ecology and evolution of mast-fruiting in Bornean Dipterocarpaceae: a general ectomycorrhizal theory. ACKNOWLEDGMENTS Ph.D. Thesis. Princeton University. Curran, L. M., I. Caniago, G. D. Paoli, D. Astianti, M. Kusneti, M. We thank the Economic Planning Unit Malaysia for research Leighton, C. E. Nirarita & H. Haeruman, 1999. Impact of El permits and are indebted to Glen Reynolds (The Royal Nino and logging on canopy tree recruitment in Borneo. Science, Society) and Jadda Suaimi (Yayasan Sabah) for the friendly 286(5447): 2184-2188. hospitality and support at Danum Valley. Data on Parashorea Curran, L. M. & M. Leighton, 2000. Vertebrate responses to seeds were collected in collaboration with Christopher spatiotemporal variation in seed production of mast-fruiting Phillipson, Sam Phillips, Eleanor Slade and Jerome Dipterocarpaceae. Ecological Monographs, 70(1): 101-128. Villeneuva. We are grateful to Bernadus “Mike” Bala Ola
285 Wells & Bagchi: Seed predation and removal by rats
Curran, L. M. & C. O. Webb, 2000. Experimental tests of the Newberry, D. M., E. J. F. Campbell, Y. F. Lee, C. E. Ridsdale & spatiotemporal scale of seed predation in mast-fruiting M. J. Still, 1992. Primary lowland dipterocarp forest at Danum Dipterocarpaceae. Ecological Monographs, 70(1): 129-148. Valley, Sabah, Malaysia - structure, relative abundance and Fleming, T. H., 1979. Do tropical frugivores compete for food? family composition. Philosophical Transactions of the Royal American Zoologist, 19: 1157-1172. Society of London Series B-Biological Sciences, 335(1275): 341- 356. Forget, P. M., 1991. Scatterhoarding of Astrocaryum paramaca by Proechymis in French Guiana: comparison with Myoprocta Ng, F. S. P., 1991. Manual of Forest Fruits, Seeds and Seedlings. exilis. Tropical Ecology, 32: 155-167. Volume 1. Forest Record No. 34. Forest Research Institute of Malaysia (FRIM). 400 pp. Harrison, J. L., 1962. The distribution of feeding habits among animals in a tropical rain forest. Journal of Animal Ecology, Sakai, S., 2002. General flowering in lowland mixed dipterocarp 31(1): 53-63. forests of South-east Asia. Biological Journal of the Linnean Society, 75(2): 233-247. Henderson, P. A. & R. M. H. Seaby, 1998. Species diversity and richness 2.3. Pisces Conservation Ltd. Sakai, S., K. Momose, T. Yumoto, T. Nagamitsu, H. Nagamasu, A. A. Hamid & T. Nakashizuka, 1999. Plant reproductive Hoch, G. A. & G. H. Adler, 1997. Removal of black palm phenology over four years including an episode of general (Astrocaryum standleyanum) seeds by spiny rats (Proechimys flowering in a lowland dipterocarp forest, Sarawak, Malaysia. semispinosus). Journal of Tropical Ecology, 13(1): 51-58. American Journal of Botany, 86(10): 1414-1436. Jansen, P. A. & P. M. Forget, 2001. Scatterhoarding rodents and Schnurr, J. L., R. S. Ostfeld & C. D. Canham, 2002. Direct and tree regeneration. In: Bongers, F., P. Charles-Dominique & P. indirect effects of masting on rodent populations and tree seed M. Forget (eds.), Dynamics and plant-animal interactions in a survival. Oikos, 96(3): 402-410. neotropical rainforest. Kluwer Academic Publishers, The Netherlands. Pp. 275-288. Smythe, N., 1986. Competition and resource partitioning in the guild of neotropical terrestrial frugivorous mammals. Annual Review Janson, C. H. & L. H. Emmons, 1990. Ecological structure of the of Ecology and Systematics, 17: 169-188. non-flying mammal community at Cocha Cashu biological station, Manu National Park, Peru. In: Gentry, A. H. (ed.) Four Sodhi, N. S., K. S. H. Peh, T. M. Lee, I. M. Turner, H. T. W. Tan, Neotropical Rainforests. Yale University Press, New Haven. D. M. Prawiradilaga & Darjono, 2003. Artificial nest and seed Pp. 314-338. predation experiments on tropical southeast Asian islands. Biodiversity & Conservation, 12: 2415-2433. Janzen, D. H., 1971. Seed predation by animals. Annual Review of Ecology and Systematics, 2: 465-492. Terborgh, J., 1986. Community aspects of frugivory in tropical forests. In: Estrada, A. & T. H. Fleming (eds.), Frugivores and Janzen, D. H., 1974. Tropical blackwater rivers, animals and mast seed dispersal. W. Junk, Dordrecht, The Netherlands. Pp. 371- fruiting by the Dipterocarpaceae. Biotropica, 6(2): 69-103. 384. Kelly, D., 1994. The evolutionary ecology of mast seeding. Trends Wells, K., M. Pfeiffer, M. B. Lakim & K. E. Linsenmair, 2004. Use in Ecology & Evolution, 9(12): 465-470. of arboreal and terrestrial space by a small mammal community Kelly, D. & V. L. Sork, 2002. Mast seeding in perennial plants: in a tropical rain forest in Borneo, Malaysia. Journal of why, how, where? Annual Review of Ecology and Systematics, Biogeography, 31(4): 641-652. 33: 427-447. Wolff, J. O., 1996. Population fluctuations of mast-eating rodents Leighton, M. & D. R. Leighton, 1983. Vertebrate responses to are correlated with production of acorns. Journal of Mammalogy, fruiting seasonality within a Bornean rain forest. In: Sutton, S. 77(3): 850-856. L., T. C. Whitmore & A. C. Chadwick (eds.), Tropical Rain Wong, T. C. M., N. S. Sodhi & I. M. Turner, 1998. Artificial nest Forest: Ecology and Management. Blackwell Science, Oxford, and seed predation experiments in tropical lowland rainforest UK. Pp. 181-196. remnants of Singapore. Biological Conservation, 85: 97-104. MacArthur, R. H. & E. R. Pianka, 1966. On optimal use of a patchy Wright, S. J., H. Zeballos, I. Dominguez, M. M. Gallardo, M. C. environment. American Naturalist, 100(916): 603-609. Moreno & R. Ibanez, 2000. Poachers alter mammal abundance, Marsh, C. W. & A. G. Greer, 1992. Forest-land use in Sabah, seed dispersal, and seed predation in a neotropical forest. Malaysia: an introduction to Danum Valley. Philosophical Conservation Biology, 14(1): 227-239. Transactions: Biological Sciences, 335(1275): 331-339. Wyatt, J. L. & M. R. Silman, 2004. Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities. Oecologia, 140(1): 26-35.
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SURVEY ON BIRDS OF PREY AND OWLS (FALCONIFORMES AND STRIGIFORMES) ON JAVA SEA ISLANDS: CORRECTION AND ADDITIONS
Vincent Nijman Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, the Netherlands. Email: [email protected]
ABSTRACT. Ð In Southeast Asia the short-eared owl Asio flammeus is a northern migrant and is normally not recorded south of Singapore and, rarely, northern Borneo. The occurrence of short-eared owl in the Kangean archipelago, Java Sea, has been noted in several publications, including a recent one in this journal (Nijman, Raffles Bull. Zool. 2004, 52(2): 647-651). Kangean would represent the southernmost locality for the species and the first for Indonesia. The contention that short-eared owl does occur on Kangean probably all traces back to Hartert (Novit. Zool. 1902, 9: 419-441) who listed Strix flammea javanica in his list of birds of Kangean. Strix flammea is a synonym for the Barn owl Tyto alba and not of Asio flammeus. The description provided by Hartert clearly point to a barn owl and not to a short-eared owl. There are no indications that short-eared owl has ever been recorded on Kangean, and only two species of owl are present in the archipelago, i.e. barn owl and Sunda scops-owl Otus lempiji.
KEY WORDS Ð Aves, avifauna, Bawean, Indonesia, migration, zoogeography.
Recently I (Nijman, 2004) published the results of a survey (Smythies 1999). The occurrence of short-eared owls on on the birds of prey (Falconiformes) and owls (Strigiformes) Kangean would represent the southernmost record of the of the island of Bawean in the Java Sea. Some raptors (most species, and indeed the first record for Indonesia. notably the Brahminy kite Haliastur indus and white-bellied sea-eagle Haliaetus leucogaster) were inexplicably absent Both my trip and that of de Iongh et al. (1982) were made from the island and the data from the owls were rather scant. in the boreal summer (June and August, respectively), and In the discussion section of my paper I, therefore, made a no short-eared owls were recorded. Short-eared owls have comparison of the raptor community of Bawean with three not been recorded by other ornithologists visiting the islands, other archipelagos in the Java Sea and stressed the need for i.e. Vorderman (1893), Hoogerwerf (1962), or A.P. Setiadi a more detailed analysis of the distribution patterns of owls (pers. comm.). De Iongh et al. (1982: 51) however, do list in the region. One of the archipelagos included in the the species in their appendix 3 and this was the basis for comparison was Kangean situated some 300 km east of inclusion of the species in my paper. De Iongh et al. (1982) Bawean and 125 north of the island of Bali on the edge of refer to Hartert (1902) as the source for the occurrence short- the Sunda Shelf. It comprises some 30 islands, the main island eared owl on Kangean but contrary to proper practise I did after which the archipelago is named measuring 487 km2. not check this publication. Neither Wells (1999) nor For Kangean I tabulated five species of Falconiformes and MacKinnon (1990) do provide a source for the occurrence of three Strigiformes, referring to a report by de Iongh et al. short-eared owl on the Kangean Islands. (1982) and my own unpublished data from a trip I made to Kangean in 1997. One of the owls listed is the short-eared owl Asio flammeus. The occurrence of this species on the Hartert (1902) described the avifauna of Kangean based on Kangean Islands was also noted by Wells (1999) adding that a collection made by Ernst Prillwitz. It is not clear when the it is a rare vagrant to the Java Sea. Likewise, MacKinnon collection was made, but from the general lack of (northern) (1990) reported the short-eared owl for Kangean Islands, migrants in the species list we can assume that it did not occur again as a rare winter migrant. during the boreal winter. Hartert (1902: 428) includes Strix flammea javanica in his list of birds of Kangean. Strix The short-eared owl has a wide, largely Holarctic distribution flammea is a synonym for the Barn owl Tyto alba and not of and East Asian populations migrate down south as far as Asio flammeus. From the description provided it is clear that Singapore (November to March: Wells, 1999) and, rarely, indeed a barn owl has been collected and not a short-eared Sarawak (November) and Brunei (January), northern Borneo owl.
287 Correction and additions: Nijman Birds of prey and owls of Java Sea Islands
As such there is no evidence for the occurrence of short-eared LITERATURE CITED owl on Kangean, and the only owls that occur on the island are the barn owl and Sunda scops-owl Otus lempiji. de Iongh, H. H., B. van Helvoort, B., S. H. Atmosoedirdjo, S. Sutanto, 1982. An ecological survey of the Kangean Island Archipelago in Indonesia. Privately published, IJselstein, ACKNOWLEDGEMENTS Hoogerwerf, A., 1962. Some ornithological notes on the smaller islands around Java. Ardea 50(3-4): 180-206. I wish to thank Neville Kemp for drawing the attention to Köning, C., F. Weick, F. J.-H. Becking, 1999. Owls. A guide to the the short-eared owl record from Kangean allowing me to set owls of the world. Pica Press, Sussex. the record straight, and to Bas van Balen for additional MacKinnon, J., 1990. Field guide to the birds of Java and Bali. information. Also thanks to the Indonesian Institute for Gadjah Mada University Press, Yogyakarta. Sciences (LIPI) and the Directorate General of Forest Nijman, V., 2004. Survey on birds of prey and owls (Falconiformes Protection and Nature Conservation (PHKA) for granting and Strigiformes) on Bawean, Java Sea, with records of three permission to conduct research in Indonesia. My research in species new to the island. Raffles Bulletin of Zoology 52(2): 647- Indonesia is funded by P.A. Hens Memorial Fund, Martina 651. de Beukelaar Foundation, and J.C. van der Hucht Fund. Smythies, B. E., 1999. The birds of Borneo, 4th revised edition. Natural History Publications, Kota Kinabalu. Vorderman, A. G., 1893. Bijdrage tot de kennis der vogels van den Kangean-Archipel. Natuurkundig Tijdschrift voor Nederlandsch-Indië, 52(4): 181-208. Wells, D. R., 1999. The birds of the Thai-Malay Peninsula. Vol. 1 Non-passerines. Academic Press, London.
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BOOK REVIEW
Fishes of Bitung. species identification guide for fishery purpose style). Each Northern Tip of species is listed alphabetically and followed by the common Sulawesi, Indonesia. English and Indonesian names. A diagnosis of the species Edited by Seishi Kimura providing fin ray counts and lateral scale count is provided, and Keiichi Matsuura, with a short write-up. This is accompanied with a well taken 2003. Ocean Research picture of a freshly preserved specimen. Institute, The University of Tokyo, Tokyo, iv + Although this guide illustrates mainly freshly preserved fish, 244 pages. it would also provide a good guide for recreational divers to the region, especially at Lembeh Straits, which is a world This is a richly illustrated class muck diving site. medium sized volume covering 78 families and All in all, this publication is a good guide for every local and 584 species. It provides foreign fisheries biologist, ichthyologist and diver to have a good coverage of the on their book shelf. shallow coastal water marine fish, obtained by field collection and market purchases in 2000. This is the second bilingual book in English and Reference Bahasa Indonesian edited by the second author (the first being “Field Guide to Lombok Island” – Matsuura et al., 2000). Matsuura, K., O. K. Sumadhiharga & K. Tsukamoto (eds.) 2000. Field guide to Lombok Island. Identification guide to This guide does not cover all the fishes known from Bitung. marine organisms in seagrass beds of Lombok Island, The emphasis of this guide is on the less colourful non coral Indonesia. Ocean Research Institute, University of Tokyo, reef fishes. This is to promote a better understanding of the Tokyo, viii + 449. true diversity of tropical fishes. The intention of the authors is for this guide to be used for study, research and education by not only ichthyologists and fisheries scientists but also by university students and local government administrators. Tan Heok Hui Department of Biological Sciences Each family of fish has a short introduction, followed by a National University of Singapore section on similar families occurring in the area (ala FAO Kent Ridge 119260, Republic of Singapore
289