THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 1-10 Date of Publication: 28 Feb.2006 © National University of

THE EURICANIA MELICHAR (: ) FROM

Chang-Qing Xu, Ai-Ping Liang* and Guo-Mei Jiang Institute of Zoology, Chinese Academy of Sciences, 19 Zhongguancun Road, Beijing 100080 People’s Republic of China Email: [email protected] (*Corresponding author)

ABSTRACT. – Two new of Euricania Melichar (Hemiptera: Ricaniidae), E. brevicula, new species, and E. longa, new species, are described from China. Four previously recorded species, E. ocellus (Walker), E. facialis Walker, E. clara Kato and E. xizangensis Chou & Lu are redescribed and illustrated. A key to all the Chinese species in the genus is provided.

KEY WORDS. – Hemiptera, Ricaniidae, Euricania, new species, redescription, China.

INTRODUCTION Generic diagnosis. – Head including eyes broader than pronotum. Frons oblique, broader than long, with central, The Ricaniidae is one of the smaller families of the sublateral and lateral carinae. Frontoclypeal suture arched. superfamily Fulgoroidea, currently containing about 400 Vertex broad and narrow, with a carina between eyes. described species in over 40 genera (Metcalf, 1955; Chou et Pronotum narrow, with a central carina. Mesonotum narrow al., 1985). The family is mainly distributed in the Afrotropical, and long, with 3 carinae: central carina straight; lateral carinae Australian and Oriental regions, with some species in the inwardly and anteriorly curved, converging closely together Palaearctic Region. The ricaniid fauna of China is very poorly on anterior margin, each bifurcating outwardly near middle known. About 32 species are recorded from China (Fennah, in a straight longitudinal carina to or near anterior margin. 1956; Chou et al., 1985). Much basic taxonomic work on this group is needed in China. Tegmina transparent, moderately ampliate, somewhat elliptical; apical area with a pterostigma; apical margin about The genus Euricania (Hemiptera: Ricaniidae) was established as long as anal margin; with 3 longitudinal veins arising from by Melichar (1898). Until now 43 species were described in basal cell; R1 and R2 with common stem; M veins with the genus from the Oriental, Australian and Palearctic regions. common stem; cross veins on disc enclosing a number of The most recent studies of genus Euricania were by Yang irregular cellular areas on basal half; with 2 transverse lines (1989) and Chou & Lu (1985, 1977), four Euricania species formed by cross veins beyond middle, the inner line much were recorded in China. All these species were described angularly waved. Wings short, with 2 cross veins beyond without genital characters except E. ocellus (Walker) (Yang, middle and several longitudinal veins forked near apex. Hind 1989). In this paper, we describe two new species of Euricania tibiae with 2 lateral spines and 6 apical spines. Male genitalia and redescribe and illustrate the four previously known symmetrical; phallobase with 2 spinose processes at each species from China. The specimens used in this study are apical angle, ventral process twisted, directed cephalad at apex deposited in the Institute of Zoology, Chinese Academy of with basal part incised; genital styles with dorsal and ventral Sciences, Beijing, China (IZCAS), the Raffles Museum of Biodiversity Research, National University of Singapore, margins nearly parallel. Female genitalia symmetrical, bursa Singapore (RMBR), the Collection of Hebei copulatrix with 2 isometric pouches: first pouch with well University, Baoding, China (HU) and the Shanghai Institute visible cell and sclerotized ornamentation; second pouch with of Entomology, Chinese Academy of Sciences, Shanghai, numerous pores; gonaplac with teeth at inner side of apical China (SIE). margin.

Distribution. – Australian, Oriental and Palaearctic regions.

Genus Euricania Melichar, 1898 Key to species of Euricania from China

Euricania Melichar, 1898: 393 (Type species: Euricania ocellus 1. Tegmina without transverse fasciae (Fig. 29) .. E. clara Kato Walker, 1851). – Tegmina with transverse fasciae (Figs. 3, 16, 42, 55, 68) .....2

1 Xu et al.: Euricania from China

2. Tegmina with ring-shaped fasciae (Figs. 16, 55) ...... 3 Kuiqi, 100-200m, coll. unknown, 2 Aug.1955; 1 female, Jianyang, – Tegmina without ring-shaped fasciae (Figs. 3, 42, 68) ...... 4 Huangkeng, Guilin, 300m, coll. S. Q. Jiang, 4 Aug.1955; 1 female, 3. Male aedeagus with dorsal spinose process longer than half Shaowu, Chengguan, 150m, coll. F. J. Pu, 28 Jun.1955; 2 males, 1 length of phallobase (Fig. 59)...... E. ocellus Walker female, ZHEJIANG: Hangzhou, Lingyin, coll. Y. Chou, 19 Jul.1963; – Male aedeagus with dorsal spinose process shorter than half 2 males, Wuyanling, coll. B. Zhen, 20 Jul.1983; 6 males, 4 females, length of phallobase (Fig. 20)...... E. brevicula, new species JIANGXI: Poyang, coll. unknown, 15 Jun.1930; 2 males, Nanchang, 4. Male aedeagus with dorsal spinose process longer than half Meiling, coll. P. Y. Zhu, 4 Jun.1987; 1 male, 1 female, HENAN: length of phallobase (Fig. 46)...... E. longa, new species Jigongshan, 500m, H. X. Wu, 14 Jul.2001; 7 males, 2 females, – Male aedeagus with dorsal spinose process shorter than half Jigongshan, coll. Y. L. Wang & M. N. Wang, 17 Aug.1990; 1 male, length of phallobase (Fig. 7, 72) ...... 5 SHANXI: Wugong, coll. Y. J. Liu, 17 Jul.1954 (all in IZCAS except 5. In profile view, male aedeagus dilated near base (Fig. 74) ..... 2 males, 1 female in RMBR)...... E. xizangensis Chou & Lu Japan: 1 male, 1 female, Kyoto, coll. K. Eki, 22 Aug.1931; 1 male, – In profile view, male aedeagus not dilated near base (Fig. 9). Tokyo: Aoyama, coll. M. Higashino, 9 Aug. 1938; 1 female, Kyoto, ...... E. facialis Melichar coll. Y. Yie, 20 Sep.1935 (IZCAS).

Length (incl. teg.). – Male 9.5-10.5 mm, female 10.5-11.0 Euricania facialis Melichar, 1898 mm. (Figs. 1-13) Description. – General colour brown. Head (Fig. 1) (including Euricania facialis Melichar, 1898: 259; Distant, 1906: 385; Chou eyes) narrower than pronotum. Frons (Fig. 2) brown; central et al., 1985: 74. carina longer than half length of frons; sublateral carinae shorter than central carina at about 1:1.4; lateral carinae light Material examined. – China: 1 male, FUJIAN: Jianyang, brown; carina between eyes light brown. Clypeus yellow, Huangkeng, Guilin, 290-310m, coll. F. J. Pu, 4 Aug.1960; 3 males, ecarinate. Vertex yellow, wider at anterior margin than long Jianyang, Huangkeng, Changxian, 340-400m, coll. C. L. Ma 23 in middle line at about 1:16, without central carina, posterior Jul.1960; 1 male, Jianyang, Guilin, coll. Y. G. Li, 14 Jul.1979; 3 males, 1 female, Wuyi, coll. Z. X. Cui, 14 Jul.1984; 2 males, 2 corner with light brown macula, occipital margin carinate, females, San’gang, coll. S. Z. Wang, 15 Jul.1984; 1 male, Fuzhou, brown. Pronotum brown, narrow, wider at widest part than

Figs. 1-13. Euricania facialis Melichar: 1, vertex, pronotum and mesonotum, dorsal view; 2, frons and clypeus; 3, tegmina; 4, wing; 5, male genitalia, left side; 6, male anal segment, dorsal view; 7, aedeagus, dorsal view; 8, aedeagus, ventral view; 9, aedeagus, left side; 10, female genitalia, left side; 11, female anal segment, dorsal view; 12, gonophysis VIII, right side; 13, pregenital sternite. Scale bars: Figs. 1, 2, 10 = 1.25mm; Figs. 3, 4 = 3.2mm; Fig. 5 = 0.635mm; Figs. 6, 7-9, 11 = 0.4mm; Figs. 12, 13 = 0.5mm.

2 THE RAFFLES BULLETIN OF ZOOLOGY 2006 long in middle line about 1:6.5, punctated beside central Jiuxianshan, coll. S. Z. Wang, 14 Jul.1979; 4 males, 3 females, carina; anterior margin carinate. Mesonotum and mesopleura GUANGXI: Bobai, coll. J. J. Wang, 18 Jun.1985; 2 males, brown. GUANGDONG: Meixian, Yinnashan, coll. P. Lin, 4-7 Aug.1963 (all in IZCAS except 2 males, 1 female in RMBR). Tegmina (Fig. 3) with costal marginal fascia brown, with a Length (incl. teg.). – Male 9.0-10.0 mm, female 10.5-11.5 mm. yellow spot near middle; apical and inner margins with brown fasciae; disc with a transverse, brown fascia; common stalk Description. – Head (incl. eyes) (Fig. 14) wider than of R1 and R2 longer than base cell. Wings (Fig. 4) small, pronotum. Frons (Fig. 15) brown, longer than half length of costal margin and inner margin with narrow, brown fasciae; frons; sublateral carinae shorter than central carina at about apical margin with a broad, brown fascia. 1:1.2; lateral carinae brown. Carinae between eyes brown. Clypeus yellow, ecarinate or indistinctly carinate. Vertex (Fig. Legs yellowish, fore femora with 3-4 lateral spines; middle 14) brown, wider at anterior margin than long in middle line coxa with one basal spine; middle femora with 3-5 lateral about 15:1, with light brown maculae at each posterior corner; spines. Abdomen brown or dark brown. Tegula dark brown. without central carina; occipital margin with a dark carina. Eyes grey. Ocelli golden. Pronotum brown and narrow, wider at widest part than long in middle line about 5.5:1; punctated beside central carina; Male genitalia. – Anal segment (Figs. 5, 6) large, oblong in anterior margin carinate. Mesonotum and mesopleura brown. dorsal view (Fig. 6), longer than wide at middle at about 1.5:1, lateral margins almost straight, apical margin slightly Tegmina (Fig. 16) with costal marginal fascia brown, with a concave, basal margin strongly convex, ventral margin in yellow spot near middle, posterior outer corner of the spot profile (Fig. 5) strongly convex; anal style set in middle but with a rather small white spot; apical margin and inner margin close to apex. Aedeagus with connective symmetrical. each with a brown fascia; disc with 2 transverse brown Phallobase (Fig. 7) with dorsal process short, shorter than fasciae: the outer one nearly straight and connected with apical half length of phallobase in middle line about 1:3, directed fascia near posterior corner; the inner one forming a ring in cephalad and laterad; lateral process shorter than dorsal middle, with median part of the ring white; common stalk of process at about 1:2. In ventral view, phallobase (Fig. 8) R1 and R2 longer than base cell. Wings (Fig. 17) rather small, constricted medially, apex concave medially. In profile view costal margin and inner margin with narrow brown fascias, (Fig. 9), phallobase not dilated near base. Genital styles (Fig. apical margin with a broad brown fascia. 5) longer than wide at middle about 3.6:1. Legs yellow; fore femora with 2-3 lateral spines; middle Female genitalia. – Anal segment (Fig. 10) rather small, femora with 3-4 lateral spines. Abdomen brown. Eyes brown ventral margin in profile slightly convex; anal style set beyond or piceous. Ocelli yellowish. middle, in dorsal view (Fig. 11), apical margin concave, ventral margin strongly concave, lateral margins convex. Male genitalia. – Anal segment in dorsal view (Fig. 18) Gonophysis VIII (Fig. 12) shoe-shaped, basal part and anterior trapeziform, longer than wide at middle about 1.5:1; lateral part in a right angle, anterior part dilated apically. Pregenital margin convex, apical margin slightly concave, basal margin sternite (Fig. 13) with anterior and ventral margins nearly slightly convex; ventral margin in profile (Fig. 19) strongly straight medially. Bursa copulatrix as in fig. 10. convex; anal style set in middle close to apex. Aedeagus with connective symmetrical. Phallobase (Fig. 20) with dorsal Distribution. – China (Fujian, Zhejiang, Jiangxi, Henan, process short, directed first cephalad then laterad at apex, Shanxi, Taiwan); Japan. shorter than phallobase in middle line about 1:3; lateral process shorter than dorsal process at about 1:2. In ventral Remarks. – This species is similar to E. longa, new species, view (Fig. 21), phallobase constricted near base, apex concave but can be separated from the latter by its short spinose process medially. In profile view (Fig. 22), phallobase slightly dilated of the male phallobase (Fig. 7) and its long sublateral carinae near base. Genital styles longer than wide at middle about of the frons (Fig. 2). 3:1.

Host plants. – Silver chain, orange, tea plant, mulberry. Female genitalia. – Anal segment (Fig. 23) rather small, ventral margin slightly convex; anal style set beyond middle, in dorsal view (Fig. 24), apical margin slightly concave, Euricania brevicula, new species ventral margin almost straight, lateral margin convex. (Figs. 14-26) Gonophysis VIII (Fig. 25) shoe-shaped, basal part and anterior part in a right angle. Pregenital sternite (Fig. 26) with anterior Material examined. – Holotype - Male, China: Fujian, Fuzhou, margin produced forward medially, ventral margin with Kuiqi, 100-200m, coll. unknown, 2 Aug.1955 (IZCAS). median portion concave. Paratypes: China, 2 males, 9 females, same data as holotype; 7 males, 1 female, Fuzhou, Kuiqi, coll. unknown, 3 Aug.1955; 2 males, Distribution. – China (Fujian, Guangxi, Guangdong). 27 females, Fuzhou, Kuiqi, coll. unknown, Jul.1955; 1 female, Fuzhou, Kuiqi, coll. unknown, 8 Aug.1955; 8 males, 3 females, Remarks. – This species is similar to E. ocellus, but can be

3 Xu et al.: Euricania from China separated from the latter by its long sublateral carinae of frons males, 18 females, Wugong, coll. Y. Chou, 12 Aug.1956 (all in (Fig. 15) and shorter dorsal spinose process of the male IZCAS except 4 males, 2 females in RMBR). phallobase (Fig. 20). Length (incl. teg.). – male 10.0-11.0 mm, female 12.0-13.0 Etymology. – The species name is derived from the Latin mm. breviculus, referring to the short dorsal spinous process of the male phallobase. Description. – Head (incl. eyes) (Fig. 27) wider than pronotum. Frons (Fig. 28) brown or dark-brown, central carina longer than half length of frons; sublateral carinae Euricania clara Kato, 1932 shorter than central carina at about 1:1.4; lateral carinae light (Figs. 27-39) brown. Carinae between eyes light brown. Clypeus yellow or light brown, ecarinate. Vertex (Fig. 27) brown or piceous, Euricania clara Kato, 1932: 228; Chou & Lu, 1977: 314; Chou et wider at anterior margin than long in middle line about 15:1, al., 1985: 70. posterior corner with macula, bright yellow; carinate sublaterally, without central carina; occipital margin carinate Material examined. – China: 45 males, 7 females, BEIJING: Xijiao, and piceous. Pronotum brown or piceous, narrow, wider at 50m, coll. Y. R. Zhang, 8 Aug.1951; 38 males, Beijing, coll. Z. B. widest part than long in middle line about 6:1, punctated Zhang, 16 Sept.1936; 14 males, 7 females, Beijing, coll. unknown, beside central carina; anterior margin carinate. Mesonotum 13 Aug.1935; 3 males, 38 females, Tiantan, 50 m, coll. unknown, piceous. Mesopleura brown or dark brown. 27 Aug.1938; 5 males, Wanshengyuan, coll. L. Y. Wang, 21 Jun.1948; 15 males, Xijiao, Yiheyuan, coll. Q. Zhou, 18 Aug.1956; 3 males, 4 females, Sanbao, coll. S. B. Liao, 19 Aug.1964; 4 males, Tegmina (Fig. 29) with costal marginal fascia brown or 2 females, Xiangshan, coll. C. Q. Xu, 14 Jul.2001; 1 male, GANSU: piceous, with a yellow spot near middle, posterior corner of Wenxian, coll. unknown, Jul.1988; 3 males, 4 females, the yellowish spot with one rather small white spot; apical SHANDONG: Pingyi, Wanshougong, coll. C. Q. Xu, 2 Oct.2002; margin with or without fascia, inner margin with a narrow 1 male, SHANXI: Qinling, Laoyu, coll. unknown, 8 Jul.1951; 14 brown fascia; disc without transverse fasciae; common stem

Figs. 14-26. Euricania brevicula, new species: 14, vertex, pronotum and mesonotum, dorsal view; 15, frons and clypeus; 16. tegmina; 17, wing; 18, male anal segment, dorsal view; 19, male genitalia, left side; 20, aedeagus, dorsal view; 21, aedeagus, ventral view; 22, aedeagus, left side; 23, female genitalia, left side; 24, female anal segment, dorsal view; 25, gonophysis VIII, right side; 26, pregenital sternite. Scale bars: Figs. 14, 15, 23 = 1.25mm; Figs. 16, 17 = 3.2mm; Figs. 18, 20-22, 24 = 0.4mm; Fig. 19 = 0.635mm; Figs. 25, 26 = 0.5mm.

4 THE RAFFLES BULLETIN OF ZOOLOGY 2006 of R1 and R2 longer than basal cell. Wings (Fig. 30) with Gonophysis VIII (Fig. 38) tadpole-shaped; basal part and costal margin and apical margin without fasciae, inner margin anterior part in an acute angle; teeth of anterior part normal, with a narrow, light brown fascia. not dilated. Median part of pregenital sternite (Fig. 39) with anterior margin and ventral margin nearly straight. Bursa Legs yellow; fore femora with 3-4 lateral spines; middle copulatrix as in fig. 36. femora with 3-4 lateral spines. Abdomen brown or dark brown, dorsum dark brown or piceous. Tegula brown or dark Distribution. – China (Beijing, Shandong, Shanxi, Gansu); brown. Eyes grey. Ocelli bright yellow. Japan.

Male genitalia. – Anal segment in dorsal view quadrate (Figs. Host plants. – Mulberry, silver chain, medlar. 31, 32), lateral margin convex; apical margin slightly concave, basal margin slightly convex, ventral margin in profile almost straight; anal style set about in middle close to apex. Aedeagus Euricania longa, new species with connective symmetrical. Phallobase (Fig. 33) with dorsal (Figs. 40-52) process short, directed dorsocephalad, shorter than half length of phallobase in middle line at about 1:3; lateral process Material examined. – Holotype - male, China: Yunnan, shorter than dorsal process at about 1:2. In ventral view (Fig. Xishuangbanna, Meng’a, 1050-1080 m, coll. S. Y. Wang, 19 34), phallobase not constricted medially, apex truncate. In Aug.1958. profile view (Fig. 35), phallobase slightly dilated near base. Paratypes: 19 males, 21 females, China: YUNNAN: Genital styles (Fig. 31) longer than wide in middle at about Xishuangbanna, Meng’a, 1050-1080 m, coll. S. Y. Wang, 19 3.4:1. Aug.1958; 2 males, 9 females, Xishuangbanna, Menglun, 650 m, coll. F. J. Pu, 15 Aug.1958; 1 male, 6 females, Xishuangbanna, Female genitalia. – Anal segment (Fig. 36) rather small, Xiaomengyang, 850 m, coll. S. Y. Wang, 30 Aug.1957; 3 males, 4 ventral margin almost straight; anal style set beyond middle. females, Xishuangbanna, Mengla, 620-650 m, coll. Y. R. Zhang,

Figs. 27-39. Euricania clara Kato: 27, vertex, pronotum and mesonotum, dorsal view; 28, frons and clypeus; 29, tegmina; 30, wing; 31, male anal segment, dorsal view; 32, male genitalia, left side; 33, aedeagus, dorsal view; 34, aedeagus, ventral view; 35, aedeagus, left side; 36, female genitalia, left side; 37, female anal segment, dorsal view; 38, gonophysis VIII, right side; 39, pregenital sternite. Scale bars: Figs. 27, 28, 36 = 1.25mm; Figs. 29, 30 = 3.2mm; Figs. 31, 33-35, 37 = 0.4mm; Fig. 32 = 0.635mm; Figs. 38, 39 = 0.5mm.

5 Xu et al.: Euricania from China

12 Jul.1959; 48 males, 37 females, Xishuangbanna, Mengzhe, (Fig. 43) rather small, costal and inner margins with narrow 870 m, coll. F. J. Pu, 2 Sept.1958; 4 males, 1 female, HUNAN: brown fasciae, apical area with a broad, brown fascia. Dayong, coll. unknown, Dec.1986 (all in IZCAS except 4 males, 2 females in RMBR). Legs yellow, median part of hind femora black; fore femora with 2-4 lateral spines; middle femora with 3-5 lateral spines. Length (incl. teg.). – male 9.0-10.0 mm, female 11.0-12.0 Tegula brown. Eyes piceous. Ocelli golden. mm. Male genitalia. – Anal segment in dorsal view (Figs. 44, 45) Description. – Head (incl. eyes) (Fig. 40) wider than trapeziform, longer than wide at middle at about 1.4:1, lateral pronotum. Frons (Fig. 41) dark-brown or piceous, sublateral margin slightly convex, apical margin slightly concave, basal carinae indistinct, shorter than central carina at about 1:3.4; margin slightly convex, ventral margin in profile view (Fig. lateral carinae light brown. Carina between eyes light brown. 44) strongly convex; anal style set in middle close to apex. Clypeus yellow, ecarinate. Vertex (Fig. 40) brown, without Aedeagus with connective symmetrical. Phallobase (Fig. 46) central carina, wider at anterior margin than long in middle with dorsal process long, directed cephalad and laterad, longer line about 14:1, macula present in posterior corner, light than half length of phallobase in middle line at about 1:2; brown; occipital margin carinate, piceous. Pronotum brown, lateral process shorter than dorsal process at about 1:3. In narrow, wider at widest part than long in middle line about ventral view (Fig. 47), phallobase constricted medially; apex 6:1, punctated beside central carina; anterior margin carinate. concave medially. In profile view (Fig. 48), phallobase dilated Mesonotum and mesopleura piceous. basally. Genital styles longer than wide at middle about 2.7:1.

Tegmina (Fig. 42) with costal marginal fascia brown, with a Female genitalia. – Anal segment (Fig. 49) rather small; anal yellow spot near middle; apical margin and inner margin with style set beyond middle, in dorsal view (Fig. 50), apical and broad brown fasciae; disc with one transverse brown fascia; lateral margin concave, ventral margin straight. Gonophysis common stalk of R1 and R2 longer than basal cell. Wings VIII (Fig. 51) shoe-shaped, basal part and anterior part in a

Figs. 40-52. Euricania longa, new species: 40, vertex, pronotum and mesonotum, dorsal view; 41, frons and clypeus; 42, tegmina; 43, wing; 44, male genitalia, left side; 45, male anal segment, dorsal view; 46, aedeagus, dorsal view; 47, aedeagus, ventral view; 48, aedeagus, left side; 49, female genitalia, left side; 50, female anal segment, dorsal view; 51, gonophysis VIII, right side; 52, pregenital sternite. Scale bars: Figs. 40, 41, 49 = 1.25mm; Figs. 42, 43 = 3.2mm; Fig. 44 = 0.635mm; Figs. 45, 46-48, 50 = 0.4mm; Figs. 51, 52 = 0.5mm.

6 THE RAFFLES BULLETIN OF ZOOLOGY 2006 right angle. Apical teeth of gonophysis VIII normal, not Material examined. – China: 13 males, 10 females, GUANGXI: dilated. Median part of pregenital sternite (Fig. 52) with Guilin, Liangfeng, 200m, coll. C. G. Wang, 11 Jul.1963; 2 males, anterior margin nearly straight, ventral margin mesially Guilin, coll X. K. Yang, 12 Jul.1992; 9 males, 10 females, Guilin, convex. Bursa copulatrix as in fig. 49. Yanshan, 200m, coll. Y. S. Shi, 10 Jul.1963; 1 male, 1 female, Longzhou, Nonggang, 15 Jun.2000, coll. J. Yao; 1 male, 1 female, Longzhou, Nonggang, coll. C. D. Zhu, 15 Jun.2000; 1 male, Distribution. – China (Yunnan, Hunan). Longzhou, Sanlian, coll. W. Z. Li, 15 Jun.2000; 2 males, 1 female, Longzhou, Nonggang, 330m, coll. J. Chen, 15 Jun.2000; 2 males, Remarks. – This species is similar to E. facialis Melichar, Yangshuo, 150m, coll. S. Y. Wang, 20 Jul.1963; 1 male, 2 females, but can be separated from the latter by its rather long dorsal Baishou, 25 Jun.1959; 3 males, Lingtian, coll. Z. L. Wu & X. L. spinose process of male phallobase (Fig. 46) and its short Lu, 3 Jun.1984; 4 males, 4 females, : Baoting, 80m, coll. sublateral carinae of frons (Fig. 41). C. Q. Li, 16 May1960; 5 males, 4 females, Shuiman, 640m, coll. S. F. Li, 30 May1960; 2 males, Lehui, coll. K. R. Huang, 06 May1954; Etymology. – The species name is derived from the Latin 1 male, GUANGDONG: Lianxian, coll. Y. W. Zhang, 28 Jun.1965; longus, referring to the long dorsal spinous process of the 2 males, Nada, 25 Apr.1954, coll. K. R. Huang; 1 male, Lechang, Pingshi, coll. Y. W. Zhang, 1 Jul.1965; 2 males, HUNAN: Changsha, male phallobase. Yuelushan, coll. unknown, 2 Aug.1957; Changsha: Yuelushan, 1 male, 1 female, coll. unknown, 15 Jul.1955; 1 male, Hunan, coll. unknown, no date; 27 males, 10 females, Chenzhou, coll. Y. L. Euricania ocellus (Walker, 1851) Zhang & Y. H. Chai, 18-26 Jul.1985; 3 males, 11 females, Hengshan, (Figs. 53-65) coll. Y. L. Zhang & Y. H. Chai, 10 Aug.1985; 9 males, 2 females, Changsha, Yuelushan, coll. Z. R. Lei, 30 Aug.1991; 2 males, Pochazia ocellus Walker, 1851: 429. GUIZHOU: Lilang, coll. Y. Z. Du, 10 Jul.1994 (all in IZCAS except Euricania ocellus - Melichar, 1898: 259; Distant, 1906: 385; Chou 4 males, 2 females in RMBR). : 1 male, Tonkin, Hoa-Binh, & Lu, 1977: 314; Yang, 1989: 201. coll. unknown, no date (IZCAS).

Figs. 53-65. Euricania ocellus (Walker): 53, vertex, pronotum and mesonotum, dorsal view; 54, frons and clypeus; 55, tegmina; 56, wing; 57, male genitalia, left side; 58, male anal segment, dorsal view; 59, aedeagus, dorsal view; 60, aedeagus, ventral view; 61, aedeagus, left side; 62, female genitalia, left side; 63, female anal segment, dorsal view; 64, gonophysis VIII, right side; 65, pregenital sternite. Scale bars: Figs. 53, 54, 62 = 1.25mm; Figs. 55, 56 = 3.2mm; Fig. 57 = 0.635mm; Figs. 58, 59-61, 63 = 0.4mm; Figs. 64, 65 = 0.5mm.

7 Xu et al.: Euricania from China

Length (incl. teg.). – Male 10.5-11.0 mm, female 11.0-12.0 Host plants. – Silver chain, orange, tea plant, tung, and castor- mm. oil plant.

Description. – General colour brown. Head (incl. eyes) (Fig. 53) wider than pronotum. Frons (Fig. 54) with central carina Euricania xizangensis Chou & Lu longer than half length of frons; sublateral carinae shorter (Figs. 66-78) than central carina at about 1:3; lateral carinae yellow. Carina between eyes brown. Clypeus brown, central carina indistinct. Euricania xizangensis Chou & Lu, 1981: 225; Chou et al., 1985: Vertex (Fig. 53) brown, wider at anterior margin than long 70. in middle line about 17:1, with light brown macula in each posterior corner, carinate centrally; occipital margin carinate, Material examined. – China: 4 males, 2 females, XIZANG: Motuo, piceous. Pronotum narrow, wider at widest part than long in 800-1100m, coll. G. D. Ren, 17 Aug.2003 (HU); 1 female, Muotuo, middle line about 13:1; anterior margin carinate. Mesonotum Maniwong, 930m, coll. F. S. Huang, 25 Aug.1974 (IZCAS); 2 males, and mesopleura brown or piceous. 2 females, Muotuo, coll. G. T. Jin, 13 Jul.1980 (SIE).

Tegmina (Fig. 55) with costal marginal fascia brown, with a Length (incl. teg.). – Male 10.5-11.0 mm, female 11.0-11.5 yellow spot near middle, with a very small white spot at outer mm. lower angle of the yellowish spot; apical and inner margins with brown fasciae; disc with 2 transverse fasciae: outer fascia Description. – Head (incl. eyes) (Fig. 66) wider than nearly straight and connected with apical fascia near posterior pronotum. Frons (Fig. 67) piceous, sublateral carinae longer corner, inner fascia forming a ring in middle with central part than half length of central carina at about 1:2; lateral carinae of the ring white; R1 and R2 with a long common stem; apex with basal part black and apical part light brown. Carina round, common stem closed to claval suture. Wings (Fig. 56) between eyes brown with median part black. Clypeus brown, rather small, apical margin with a brown fascia, costal and central carina light brown. Vertex (Fig. 66) piceous, wider at inner marginal fasciae very narrow, brown. anterior margin than long in middle line about 12.8:1, posterior corner without macula or with indistinct macula; Legs yellow; fore femora with 2-3 lateral spines; middle carinate laterally, without central carina; occipital margin femora with 4-5 lateral spines. Abdomen brown or dark carinate, piceous. Pronotum piceous, narrow, wider at widest brown. Tegula brown or dark brown. Eyes piceous, ocelli part than long in middle line about 6:1; punctated beside rufous. central carina; anterior margin carinate. Mesonotum and mesopleura piceous. Male genitalia. – Anal segment in dorsal view (Figs. 57, 58) longer than wide at middle about 1.5:1, lateral margin convex, Tegmina (Fig. 68) with costal marginal fascia piceous, with basal margin slightly convex; in profile view with ventral a yellow or yellowish spot near middle, posterior corner of margin convex; anal style set about in middle, close to apex. the spot with one very small white spot; apical area piceous; Aedeagus stout. In dorsal view (Fig. 59), phallobase with apical margin and inner margin with piceous fasciae; disc dorsal process very long, directed first cephalad then laterad with one transverse piceous fascia which is interrupted or at apex, longer than half length of phallobase in middle line disappeared near claval suture; common stem of R1 and R2 at about 1:2; lateral process shorter than dorsal process at longer than basal cell, common stem closed to claval suture. about 1:5, directed laterad at apex. In ventral view, phallobase Wings (Fig. 69) with apical margin with a piceous fascia, (Fig. 60) not constricted medially, apex concave medially. costal margin and inner margin without fasciae. In profile view, phallobase (Fig. 61) dilated basally. Genital styles (Fig. 58) longer than wide at middle about 1.7:1. Legs brown; fore femora with 2-4 lateral spines; middle femora with 2-5 lateral spines. Abdomen piceous. Tegula dark Female genitalia. – Anal segment (Fig. 62) rather small, brown or piceous. Eyes piceous, ocelli rufous. ventral margin slightly convex; anal style set at middle, apical margin in dorsal view (Fig. 63) almost straight, ventral margin Male genitalia. – Anal segment (Figs. 70, 71) in dorsal view concave, lateral margin almost straight. Gonophysis VIII (Fig. longer than wide at middle about 1.5:1, lateral margin convex, 64) tadpole-shaped, basal part and anterior part in an acute apical margin truncate, basal margin almost straight; ventral angle. Pregenital sternite (Fig. 65) with anterior margin and margin in profile (Fig. 70) strongly convex; anal style set ventral margin nearly straight medially. Bursa copulatrix (Fig. about in middle, close to apex. Aedeagus with connective 62) second pouch slightly bigger than first pouch. symmetrical. Phallobase (Fig. 73) with dorsal process short, directed laterad, shorter than half length of phallobase in Distribution. – China (Guangxi, Hainan, Hunan, Guizhou, middle line about 1:3.3; lateral process shorter than dorsal Taiwan); Bengal, Sikkim, , Japan, Vietnam. process at about 1:1.3. In ventral view (Fig. 74), phallobase constricted medially, apex concave medially; lateral process Remarks. – This species is similar to E. brevicula, but can shorter than phallobase in middle line about 1:2.6. In profile be separated from the latter by its shorter sublateral carinae view (Fig. 75), phallobase dilated near base. Genital styles of frons (Fig. 54) and very long dorsal spinose process of the (Fig. 70) longer than wide at middle about 3:1. male phallobase (Fig. 59).

8 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Female genitalia. – Anal segment (Fig. 75) rather small, from the Chinese Academy of Sciences (grant no. A2903077, ventral margin in profile slightly convex; anal style set beyond to APL), the National Natural Science Foundation of China middle; in dorsal view (Fig. 76), apical margin straight, ventral (grant no. 30370187) and the National Science Fund for margin nearly straight, lateral margins convex. Gonophysis Fostering Talents in Basic Research (NSFC-J0030092). VIII (Fig. 77) shoe-shaped, basal part and anterior part in a right angle, teeth of anterior part dilated apically. Pregenital sternite (Fig. 78) with anterior margin produced forward LITERATURE CITED medially. Bursa copulatrix with second pouch smaller than first pouch. Chou, I. & J. S. Lu, 1977. On the Chinese Ricaniidae with descriptions of eight new species. Acta Entomologica Sinica, Distribution. – China (Xizang). 20: 314-322. Chou, I. & J. S. Lu, 1981. Homoptera: Fulgoroidea. In: Chen, S.X. Remarks. – This species is similar to E. longa and E. facialis, (ed.), of Xizang. The Series of the Expedition to the but can be separated from E. longa by its shorter dorsal Qinghai-Xizang Plateau. Vol. 1. Sciences Press, Beijing. Pp. spinose process of male phallobase (Fig. 72) and longer 225-226. sublateral carinae of frons (Fig. 67) and from E. facialis by Chou, I., J. S. Lu, J. Huang & S. Z. Wang, 1985. Homoptera its dilated basal part of male aedeagus (Fig. 74), distinct carina Fulgoroidea. Economic Insects Fauna of China. Fasc. 36. of clypeus (Fig. 67) and the whole black body. Sciences Press, Beijing. Distant, W. L., 1906. The Fauna of British India, including Ceylon and Burma. Rhynchota 3 (Heteroptera-Homoptera). London: ACKNOWLEDGEMENTS Taylor & Francis. 503 pp. Fennah, R. G., 1956. Fulgoroidea from southern China. Proceedings This work was supported by the Hundred Talent Program of the California Academy of Science, (4)28: 441-527.

Figs. 66-78. Euricania xizangensis Chou & Lu: 66, vertex, pronotum and mesonotum, dorsal view; 67, frons and clypeus; 68, tegmina; 69, wing; 70, male genitalia, left side; 71, male anal segment, dorsal view; 72, aedeagus, dorsal view; 73, aedeagus, ventral view; 74, aedeagus, left side; 75, female genitalia, left side; 76, female anal segment, dorsal view; 77, gonophysis VIII, right side; 78, pregenital sternite. Scale bars: Figs. 66, 67, 75 = 1.25mm; Figs. 68, 69 = 3.2mm; Fig. 70 = 0.635mm; Figs. 71-74, 76 = 0.4mm; Figs. 77, 78 = 0.5mm.

9 Xu et al.: Euricania from China

Kato, M., 1932. Notes on some Homoptera from south Manchuria, Metcalf, Z. P., 1955. General Catalogue of the Homoptera. Fasc. collected by Mr. Yukimichi. Kontyû, 5(5): 216-229. IV. Fulgoroidea. Part 16. Ricaniidae. North Carolina State Matsumura, S., 1930. The Illustrated Thousand Insects of Japan. College, Raleigh, N.C., USA. Keiseisha, Tokyo. Wytsman, P., 1923. Genera Insectorum. Homoptera. 182: 120-173. Melichar, L., 1898. Monographie der Ricaniiden (Homoptera). Yang, C. T., 1989. Ricaniidae of Taiwan (Homoptera: Fulgoroidea). Annalen des K. K. Naturhistorischen Hofmuseums, 8(2-3): 197- Taiwan Museum Special Publication Series, 8: 201-202. 359. Walker, F., 1851. List of the Specimens of Homopterous Insects in the Collection of the British Museum, 2: 261-636.

10 THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 11-20 Date of Publication: 28 Feb.2006 © National University of Singapore

NEW SPECIES OF THE WATER STRIDER GENERA EOTRECHUS KIRKALDY AND RHYACOBATES ESAKI (HETEROPTERA: GERRIDAE) FROM VIETNAM

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

Chang Man Yang c/o Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore

ABSTRACT. – Three new species, Eotrechus vietnamensis, Rhyacobates anderseni, and Rhyacobates gongvo, are described from Vietnam. Both genera, Eotrechus and Rhyacobates, are first records for Vietnam. Rhyacobates abdominalis is also first reported from Vietnam.

KEY WORDS. – Vietnam, Gerridae, Eotrechus, Rhyacobates, new species, first record.

INTRODUCTION MATERIALS AND METHODS

The literature on aquatic heteropteran fauna of Vietnam is Specimens for this study were collected from Vietnam by rather scanty and scattered. The most comprehensive study the first author during the period from April to June 2003 on Gerridae is by Zettel & Chen (1996), who documented 40 and on 3 July 2004. Additional specimens of Eotrechus species in 18 genera. In this study, we described one new vietnamensis were provided by Dr. J. T. Polhemus (Colorado, species of Eotrechus and two new species of Rhyacobates, U.S.A). They were collected by sweeping a hand net over Both Eotrechus and Rhyacobates are recorded for the first the water surface (for Rhyacobates species) or around rock time from Vietnam. surfaces of forested streams (for Eotrechus species). All specimens were preserved in 75% alcohol except those were Species of the genus Eotrechus Kirkaldy, only found in dissected or mounted dry. Measurements of new species refer mountainous areas, inhabit hygropetric habitats on rock or to the holotype, the allotype, or the single winged specimen, soil kept wet by water seepage and stream, or on waterfall if no variations were given. For R. abdominalis, mean splash zone (Andersen, 1982, 1998). Hither to, there are eight measurements for appendages were given. The remaining described species (Andersen, 1982, 1998), two from southern measurements were given as ranges based on randomly China, two from northern , and four others found in selected specimens. All measurements were in millimetres. India, and . The new species Eotrechus Length of the body was measured from the anterior margin vietnamensis described in this study extends the known of the head to the posterior tip of the last abdominal tergum. distribution of the genus to Vietnam. Width of the body was measured across meso-acetabula.

The genus Rhyacobates Esaki inhabits fast-flowing The specimens were deposited in the following collections: mountainous streams. In their revision of the genus JTPC, Collection of J.T. Polhemus, Englewood, Colorado, Rhyacobates, Andersen & Chen (1995) recorded nine species U.S.A from Asia (China, Korea, Taiwan, Myanmar, and Thailand), NHMW, Natural History Museum Vienna, Austria all with rather restricted distribution. Only one species, R. ZMHU, Zoological Museum, Hanoi University of Science, malaisei Andersen & Chen, is found in China, Myanmar and Vietnam Thailand where these three countries meet. In this study, R. ZRC, Zoological Reference Collection, Raffles Museum of abdominalis Andersen & Chen, described from China is Biodiversity Research, National University of Singapore, recorded for Vietnam for the first time. Singapore

11 Tran & Yang: New Eotrechus and Rhyacobates from Vietnam

TAXONOMY Structural characteristics Apterous male (holotype): Head width across eyes 1.29; Eotrechus vietnamensis, new species interocular width 0.60; eye kidney-shaped on dorsal view, (Figs. 1-6, 26) length of eye 0.63. Antennae about 1.2x body length (6.99: 5.90), lengths of segments 1-4: 1.94: 1.55: 1.46: 2.04; first Material examined. – Holotype (apterous male) and allotype segment with 2-3 black spine-like hairs in apical part. (apterous female), Vietnam, Vinh Phuc province, Tam Dao National Pronotum, broader than long, slightly shorter than head length Park, suoi Thac Bac (near Doi Che), 21°27.005'N 105°38.771'E, (0.84: 1.16). Lengths of mesosternum and metasternum: 1.97 749 m, coll. A. D. Tran & Q. K. Hoang, 17 Jun.2003, TAD0351, and 0.37. Posterior margin of metasternum with a median (ZMHU). fringe of short black hairs (Fig. 3). Lengths of leg segments Paratypes: Vietnam: apterous 4 males, 3 females, same locality data (femur: tibia: tarsal segment 1: tarsal segment 2) as follows: as holotype; apterous 4 males, 1 female, Vinh Phuc province, Tam fore leg: 2.43: 1.89: 0.22: 0.32; middle leg: 6.90: 5.10: 0.61: Dao National Park, Suoi Mo, 21°27.103'N 105°38.872'E, 924m, coll. 0.49; hind leg: 7.20: 5.00: 0.73: 0.49. Fore femur (Fig. 2) A. D. Tran & Q. K. Hoang, 16 Jun.2003, TAD0350; apterous 3 simple, length about 6.57x maximum width (2.43: 0.37), males, 2 females, Vinh Phuc province, Tam Dao National Park, ventral surface with 2 black stout hairs (2-6 hairs in other 21 27.747'N 105 38.754'E, 1000m, coll. A. D. Tran & Q. K. Hoang, ° ° paratypes); fore tibia almost straight, with some long black 18 Jun.2003, TAD0354; apterous 1 male, macropterous 3 females, Vinh Phuc province, Tam Dao National Park, 21°27.439'N spine-like hairs on apical margin. Middle and hind femur 105°39.193'E, 1000m, coll. A. D. Tran & Q. K. Hoang, 18 Jun.2003, slender and slightly longer than the body, with scattered small TAD0355; apterous 5 males, 5 females, Vinh Phuc province, Tam brown spines. Claws stout, length 0.20. Abdomen relatively Dao National Park, Suoi Bua Lon, 600m, coll. A. D. Tran & Q. K. short, pregenital length 1.22, with patch of long yellow hairs Hoang, 19 Jun.2003, TAD0356; apterous 5 males, 3 females, Dien on median sterna 1-5. Sternum 7 about 2.25x length of two Bien province, Muong Phang, upstream and water fall of Muong preceding sterna combined (0.61: 0.27), posterior margin with Phang stream, 21 27.000'N 103 10.548'E, 1070m, coll. A. D. Tran, ° ° very deep rectangular median notch (about half the length of 28 Jul.2004, DY0419 (NHMW, ZMHU, and ZRC); apterous 1 male, Ha Tay province, Ba Vi National Park, small stream, 575 m, coll. sternum 7) (Fig. 3). Genital segments large, length 0.54, J. T. Polhemus, D. A. Polhemus & P. Nguyen, 4 Apr.2000, CL4391; pygophore broadly suboval, with a pair of disto-lateral apterous 1 male, 1 female, Ha Tay province, Ba Vi National Park, processes, each directed obliquely upwards, plate like, and seeping rock face on upper road, 895 m, coll. J. T. Polhemus, D. A. with slightly pointed apex in lateral view (Figs. 3-4). Polhemus & P. Nguyen, 4 Apr.2000, CL4391; apterous 6 males, 5 Parameres small and blunt. females, Lao Cai province, roadside seeps, 11 km NE Sa Pa, 22 23.42'N 103 52.92'E, 1220 m, coll. J. T. Polhemus, D. A. ° ° Apterous female (allotype): slightly bigger than the male. Polhemus & P. Nguyen, 7 Apr.2000, CL4396; apterous 2 females, Lai Chau province, Nam Ceung stream, 15 km N Lai Chau, Head width across eyes 1.37; interocular width 0.65; eye 22°08.67'N 103°11.55'E, 207 m, coll. J. T. Polhemus, D. A. kidney-shaped, length of eye 0.61. Antennae about 0.9x of Polhemus & P. Nguyen, 11 Apr.2000, CL4409; apterous 5 males, body length (6.20: 6.90), lengths of segments 1-4: 1.79: 1.41: 5 females, Lai Chau province, cascading tributary to Nam Na River, 1.16: 1.84; first segment with 2-3 black spines sub-apically. 12 km N Lai Chau, 22°07.325'N 103°11.50'E, 290 m, coll. J. T. Pronotum slightly shorter than head length (0.83: 1.16). Polhemus, D. A. Polhemus & P. Nguyen, 11 Apr.2000, CL4410 Lengths of mesosternum and metasternum: 2.24 and 0.51. (JTPC). Lengths of leg segments (femur: tibia: tarsal segment 1: tarsal segment 2): fore leg: 2.38: 1.79: 0.24: 0.39; middle leg: 6.80: Description. – Apterous male, length 5.5-5.9 (holotype 5.9), 4.80: 0.76: 0.61; hind leg: 6.90: 4.90: 0.83: 0.54. Fore femur width 1.85-2.07 (holotype 1.97) (n=20); apterous female simple, length about 7.0x maximum width (2.38: 0.34), length 6.3-7.0 (allotype 6.9), width 2.22-2.39 (allotype 2.32) (n=14); macropterous female length 7.2-7.8, width 2.20-2.44 ventral surface with five black stout hairs; fore tibia almost (n=3). straight with some long black spines on apical margin. Middle and hind femur slender and subequal to body length, scattered Colour. Apterous form, body mainly brown on dorsal side with small brown spines. Claws stout, length 0.20. Pregenital and pale yellow on ventral side (females with two brown abdominal venter 0.32x body length (2.24: 6.90). Venter of markings on mesosterno-pleura and median brown patches sterna 1-6 without patch of long hairs. Sternum 7 about 1.18x on metasterno-pleura); dorsal surface covered with silvery length of two preceding sterna together (0.60: 0.51), posterior or greenish pubescence. Head with one median yellow stripe; margin straight. Genital segments not concealed, proctiger antennae yellow to brown. Pronotum with five longitudinal round and slightly deflected (Fig. 6). yellow stripes on dorsal and lateral sides. Mesonotum mainly yellow with six longitudinal markings: two slender dark Macropterous female: similar to apterous female but slightly brown lateral stripes, two slender dark brown sub-lateral bigger in size. Fore wings distinctly surpassing abdominal stripes and two light brown markings in the centre. apex. Wing veins as shown in Fig. 5. Metanotum and abdominal tergum mainly dark brown. Pro-, meso- and meta-sternopleura with dense silvery and Macropterous male unknown. reflective pubescence. Legs: all coxae, trochanters and femora yellow (slightly darker in females), all tibiae and tarsi brown. Etymology. – This species is named after its country of origin. In macropterous form, anterior part of pronotum with five yellow stripes as in apterous form, posterior part totally Distribution & habitat. – Northern Vietnam (Vinh Phuc, Ba brown, wings brown. Vi, Lai Chau, Dien Bien, and Lao Cai provinces). Eotrechus

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Figs. 1-6. Eotrechus vietnamensis, new species. 1-4, male holotype: 1, lateral view of head and prothorax. 2, right fore leg. 3, abdomen, ventral view. 4, genital segments, lateral view. 5, fore wing of male. 6, female allotype, lateral view of abdomen. Scales bars 1-6: 1 mm.

13 Tran & Yang: New Eotrechus and Rhyacobates from Vietnam vietnamensis was collected on wet surfaces of rocks, cliffs Additional material. – 1 apterous female, China, Yunnan, covered with mosses at waterfalls or along forest streams at Xishuangbanna, Hill stream, 90km from Jinghong to Simao, different elevations (from 200-1200 m) of mountainous areas 1070 m, coll. H. H. Tan, 20 May.2000, THH0068 (ZRC). in northern Vietnam. Adults and nymphs were seen resting on almost vertical rock surfaces. When disturbed, they jumped Description. – Apterous male: length 6.0-6.1 (allotype 6.0), away or hid in shaded spaces beneath the rocks. They were width 1.83-1.85 (allotype 1.83) (n=5); apterous female: length found in the same habitat as Onychotrechus species. 6.8-7.0 (holotype 6.8), width 2.52-2.57 (holotype 2.52) (n=8); macropterous male: length 6.1-6.2, width 1.85-1.88 (n=2). Remarks. – In the key given by Andersen (1982), the male of E. vietnamensis could be keyed to E. brevipes by having Colour. Body almost black dorsally, covered with silvery plate-shaped processes in the pygophore, and the relatively pubescence. Head and pronotum black with semi-circular short middle and hind femura which are only slightly longer yellow markings. Antennae mainly dark brown to black, first than body. However, males of this species can be antennal segment with yellow basal half, last segment with distinguished form E. brevipes by the following characters: whitish area in distal two fifths. Male with longitudinal thin lateral processes of pygophore flattened, expanded distally yellow stripe on posterior half of mesonotum. Yellow stripe and without finger-like process; median emargination of on mesonotum of female broader than that of male. Venter sternum 7 much deeper than in all known Eotrechus species; bright yellow except anterior part of mesosterno-pleura black. fore trochanter without patch of spinules, fore femur of males All coxae, trochanters and femora yellow. Fore femur of both more slender, fore tibia without apical process. The female sexes with two brown longitudinal stripes on dorsal and of E. vietnamensis, could not be keyed to any described external side, more slender and lighter brown in the female. species because the genital segments are not concealed and Middle and hind femora yellow at base but brown or black the proctiger is not elongated but deflected. In lateral view, at distal end. All tibiae and tarsi dark brown to black. sternum 7 of the female is rather similar to that of E. sinensis but it has fewer stout hairs on fore femur and a lesser ratio of Structural characteristics sternum 7: sternum 5+6. Apterous female (holotype): Head width across eyes 1.22, eye length 0.54, interocular width 0.48. Lengths of antennal Male specimens from Dien Bien province and Lai Chau segments 1-4: 2.86: 0.68: 1.02: 0.73. Pronotum broader than province differed from those collected in locality of holotype long, length and width: 0.63 and 1.36. Mesonotum length in the following characters: fringe of short black hairs on 2.02. Posterior margin of metanotum with a median pointed posterior margin of metasternum less dense, hair patch on process (Fig. 7). Lengths of leg segments (femur: tibia: tarsal sterna 1-5 less dense, median notch of sternum 7 gradually segment 1: tarsal segment 2): fore leg: 2.86: 2.37: 1.33: 0.71; widened towards posterior margin, disto-lateral processes of middle leg: 8.50: 5.20: 2.24: 0.32; hind leg: 8.60: 3.10: 0.15: pygophore slightly thicker and less pointed, posterior margin 0.20. All coxae with rings of dark bristle-like hairs at apical of pygophore slightly produced on median. However, the margin. Fore trochanter with long hairs ventrally, hind presence of a hair fringe on metasternum, hair patches on trochanter with dense long hairs dorsally. Fore femur with sterna 1-5, and a very deep median notch of sternum 7 are nine long dark bristle-like hairs posteriorly and scattered with unique among Eotrechus species. Therefore, we treated these soft long yellow hairs on venter (Fig. 11). Middle tibia lined differences as intraspecific variations. with brownish hair-fringe ventrally. Hind femur with long yellow hairs on basal part. Abdomen relatively short, As the manuscript was being prepared, Dr. J. T. Polhemus pregenital abdomen length 1.27. Length of sterna 5 and 6: (Colorado, U.S.A) informed us that he had collected 0.30 and 0.40. Abdominal segment 7 long, length 0.80, raised specimens which he believed could be the same species as slightly upwards dorsally, bearing a pair of long projections E. vietnamensis. He kindly sent us part of his material (lot pointing outwards and downwards (Figs. 8-10); posterior number CL4391, CL 4396, and CL4410) to confirm its margin more or less straight with two short lateral processes identity. His specimens are conspecific with E. vietnamensis (Figs. 9 & 10). Abdominal segment 7 almost enclosing the as he had suspected, and as such, we have included all his genital segments. material as paratypes. Apterous male (allotype): Head width across eyes 1.16, interocular width 0.43, eye length 0.46. Lengths of antennal Rhyacobates anderseni, new species segments: 2.67: 0.68: 1.05: 0.71. Pronotum broader than long, (Figs. 7-16, 27) length and width: 0.59 and 1.22. Mesonotum length 1.76. Posterior margin of metanotum without a median process. Material examined. – Holotype (apterous female) and allotype (apterous male), Vietnam, Ha Tinh province, Vu Quang National Lengths of leg segments (femur: tibia: tarsal segment 1: tarsal Park, Khe Lim, 18°16.416'N 105°26.467'E, 291 m, coll. A. D. Tran, segment 2): fore leg: 2.73: 2.18: 0.83: 0.51; middle leg: 8.10: 24 Apr.2003, TAD0304 (ZMHU). 4.50: 2.02: 0.34; hind leg: 8.00: 2.52: 0.12: 0.17. Structure of legs similar to female except middle femur with small Paratypes: Vietnam: apterous 3 males, 6 females, same locality data spinules but not in a row, spinules more scattered on distal as holotype; apterous 1 male, 1 female, macropterous 2 males, Ha part. Abdomen short, pregenital abdomen length 1.37, about Tinh province, Vu Quang National Park, Song Con, Sao La station, 0.23x of body length. Sternum 7 length 0.43, posterior margin 18°15.386'N 105°27.442'E, 293 m, coll. A. D. Tran, 25 Apr.2003, TAD0307a (NHMW, ZMHU, and ZRC). straight with long hairs. Genital segments slightly curve

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Figs. 7-16. Rhyacobates anderseni, new species. 7-11, female holotype: 7, lateral view. 8-10, subgenital segments: 8, lateral view. 9, dorsal view. 10, ventral view. 11, right fore leg. 12-16, male: 12, proctiger. 13-14, left paramere, two different views. 15, vesical sclerites. 16, right fore wing. Scales bars 7-12, 16: 1 mm, 13-15: 0.1 mm.

15 Tran & Yang: New Eotrechus and Rhyacobates from Vietnam downwards. On lateral view, abdominal segment 8 with We examined a single female from Yunnan, China which is concave venter, dorsal margin straight. Venter of abdominal almost identical to the Vietnamese specimens except for slight segment 8 length 0.35. Pygophore large, ovate; proctiger with colour differences: head mainly yellow, larger yellow marking round angular projections on each side (Fig. 12). Parameres on pronotum, first and second antennal segments yellow, third falciform, slightly broad, long, not conspicuously setose (Figs. and fourth antennal segments light brown. 13-14). Vesical sclerites as illustrated in Fig. 15.

Macropterous male: Head width across eyes 1.15, eye length Rhyacobates gongvo, new species 0.46, interocular width of head 0.48. Head with long bristle- (Figs. 17-25, 28) like hair around eyes. Lengths of antennal segments 1-4: 2.70: 0.67: 1.00: 0.71. Posterior margin of metanotum with a Material examined. – Holotype (apterous female) and allotype median pointed process. Lengths of leg segments (femur: (apterous male), Vietnam, Lao Cai province, Sa Pa, Hoang Lien tibia: tarsal segment 1: tarsal segment 2): fore leg: 2.73: 2.21: National Park, Sin Chai, 22°20.421'N 103°48.844'E, 1366 m, coll. 0.78: 0.51; middle leg: 8.20: 4.61: 2.05: 0.32; hind leg: 8.30: A. D. Tran, 1 Jun.2003, TAD0337 (ZMHU). 2.47: 0.13: 0.16. Fore wings surpassing abdominal apex, wing Paratypes: Vietnam: de-alated macropterous 2 males, 3 females, vein shown in Fig. 16. Other characteristics similar to apterous same locality data as holotype (ZRC); apterous 3 males, 6 females, male. same locality as holotype, coll. A. D. Tran, 3 Jul.2004, TAD0415 (NHMW, ZRC). Macropterous female unknown. Description. – Apterous male: length 6.3-6.5 (allotype 6.5), Etymology. – This species is dedicated to the late Prof. N. width 2.2 (n=2). Apterous female: length 8.3, width 2.52. De- M. Andersen (Copenhagen) for his great contributions to the alated macropterous male: length 6.4, width 1.97 (n=2). De- research on semi-aquatic Heteroptera. alated macropterous female: length 7.5, width 2.44 (n=3).

Distribution & habitat. – Northern Vietnam and Yunnan, Colour. Body mainly black, covered with silvery pubescence. China. The type material was collected from moderate Head yellow with black markings dorsally. Antennae mainly flowing streams of a forest at an elevation of 300 m. The black, fourth antennal segment with whitish area in distal two specimen from Yunnan was collected from a stream above fifths. Pronotum with large sub-triangular yellow spot in 1000m in elevation. apterous female and smaller and ovate in apterous male (Fig. 28). Pronotum in macropterous form with ovate (in males) Remarks. – Andersen & Chen (1995) listed the generic or diamond-shaped (in females) yellow marking in anterior characteristics of the genus Rhyacobates as: (a) mesonotum part and yellow posterior margin. Mesonotum in apterous black with pale median stripe; (b) first antennal segment form black with longitudinal yellow stripe posteriorly, stripe longer than other three segment together, last antennal length about three fourths of mesonotum length. Metanotum segment with whitish area in distal two fifths; (e) middle and and dorsal abdomen black. Venter of female light yellow hind tarsi without claws; (f) male parameres moderate in size, except anterior 2/3 of mesosternum. Venter of male mainly falciform and not conspicuously setose; (h) middle femur black or dark brown, except acetabula and genital segments. without distinct row of black spines; (i) eighth abdominal All coxae and trochanters yellow. Fore femur yellow with segment of male reduced ventrally, shorter than seventh one brown dorsal stripe. Middle and hind femora yellow at segment; (j) posterior segments of female abdomen usually base, brown or black at distal part. All tibiae and tarsi dark curved dorsad (apterous form); (k) seventh abdominal brown or black. segment of female tube-like, long and completely enclosing the genital segments. Rhyacobates anderseni possesses all Structural characteristics the above characters except that the female abdomen is Apterous female (holotype): Head width across eyes 1.37, relatively short, not curved dorsad, and abdominal segments interocular 0.52, eye length 0.56. Lengths of antennal 1-6 combined are only slightly longer than abdominal segment segments 1-4: 3.54: 0.80: 1.10: 0.80. Pronotum broader than 7. This new species is also distinctly different from other long, length 0.68, width 1.46. Mesonotum length 2.19. known Rhyacobates species by having a very conspicuous Metanotum without spine on median. Lengths of leg segments median process on the posterior margin of metanotum in the (femur: tibia: tarsal segment 1: tarsal segment 2): fore leg: female. This process is also found in Andersenius and 3.50: 2.91: 1.63: 0.81; middle leg: 10.10: 5.60: 2.52: 0.37; Pleciobates. However, elongated hind coxae which are hind leg: 10.20: 3.10: 0.16: 0.20. All coxae with a ring of diagnostic for the genus Andersenius were not present in this long dark hairs at apical margin. Fore trochanter with some new species. It is also distinct from Pleciobates by the long yellow hairs ventrally (Fig. 21). Middle and hind colouration of mesonotum, general structure of male paramere trochanters without hairs. Fore femur slender, with a row of and endosoma (for Pleciobates spp., see Zettel & Chen, 1996), 6-8 long dark bristle-like hairs and some scattered long soft and the absence of a row of black spines on the male middle yellow hairs on venter. Middle femur without small spinules. femur. The dorsal projections of seventh abdominal segment Abdomen elongate and straight, posterior part of sternum 7 in the new species are rather similar to those of R. malaisei slightly depressed ventrally (Fig. 17). Abdomen relatively but differ in the curvature of the internal margin. long, pregenital segments length 3.15. Length of sterna 5 and

16 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Figs. 17-25. Rhyacobates gongvo, new species. 17-21, female holotype: 17, lateral view. 18-20, subgenital segments: 18, lateral view. 19, dorsal view. 20, ventral view. 21, right fore leg. 22-25, male: 22, proctiger. 23, vesical sclerites. 24-25, left paramere, two different views. Scales bars 17-22: 1 mm, 23-25: 0.1 mm.

17 Tran & Yang: New Eotrechus and Rhyacobates from Vietnam

6: 0.45 and 0.57. Sternum 7 length 0.95, not totally enclosing middle leg: 9.80: 5.60: 2.44: 0.38; hind leg: 10.00: 2.91: 0.16: genital segments, posterior ventral margin straight without 0.20. process. Dorsal processes of sternum 7 long, straight and flatten (Figs. 18-20). De-alated macropterous male: similar to apterous male. Head width across eyes 1.25, interocular width 0.48, eye length Apterous male (allotype): Head width across eyes 1.29, 0.48. Pronotum with longitudinal ridge along mid-line, interocular width 0.51, eye length 0.49. Lengths of antennal pronotum length: width 2.44: 1.78. Lengths of antennal segments 1-4: 3.25: 0.78: 1.22: 0.78. Pronotum short, length: segments 1-4: 3.30: 0.78: 1.19: 0.78. Lengths of leg segments width 0.66: 1.39. Mesonotum length 1.93. Metanotum without (femur: tibia: tarsal segment 1: tarsal segment 2): fore leg: median spine or process on posterior margin. Lengths of leg 3.35: 2.96: 1.11: 0.63; middle leg: 10.00: 5.40: 2.37: 0.36; segments (femur: tibia: tarsal segment 1: tarsal segment 2): hind leg: 10.40: 2.52: 0.15: 0.17. fore leg: 3.40: 2.91: 1.10: 0.61; middle leg: 9.80: 5.10: 2.24: 0.34; hind leg: 10.00: 2.39: 0.15: 0.17. Other characters Etymology. – The word “gong vo” in Vietnamese means similar to apterous female except venter of fore femur with water skaters, it is used here specifically as a noun in a row of 4 long, dark, bristle-like hairs at basal half. Middle apposition. femur with small spinules not placed in a row. Abdomen short and slightly curved downwards posteriorly. Pregenital Distribution & habitat. – Northern Vietnam, only known abdomen length 1.68, about 0.27x body length. Sternum 7 from the type locality. Specimens were collected from a fast length 0.51. Venter of abdominal segment 8 length 0.40. flowing rocky mountainous stream in open area. They were Genital segments length 0.88. Proctiger with small angular seen skating swiftly on the water surface and well projections on each side (Fig. 22). Parameres relative long camouflaged in the torrential environment by their silvery and slender (Figs. 24-25). Vesical sclerites as illustrated in pubescence. They were also found resting on shaded sides of Fig. 23. exposed surfaces of rocks in the stream. Both Rhyacobates gongvo and R. abdominalis were found in the same habitat. De-alated macropterous female: similar to apterous female except slightly smaller in size. Head width across eyes 1.32, Remarks. – Rhyacobates gongvo matches diagnostic eye length 0.49, interocular width 0.48. Pronotum length: characters of the genus Rhyacobates except for characters (j) width 2.63: 2.00. Lengths of antennal segments 1-4: 3.25: and (k) (see remarks for R. anderseni above): posterior 0.73: 1.11: 0.78. Lengths of leg segments (femur: tibia: tarsal segments of female straight, not curved dorsad; seventh segment 1: tarsal segment 2): fore leg: 3.40: 2.86: 1.54: 0.79; segment of female abdomen not completely enclosing the

Fig. 26. Habitus of Eotrechus vietnamensis, holotype. Fig. 27. Habitus of Rhyacobates anderseni, holotype.

18 THE RAFFLES BULLETIN OF ZOOLOGY 2006 genital segments. This species is relatively similar to R. Paratypes examined: apterous 1 male, 1 female, labelled “China: malaisei in the following characters: abdomen of female Guangdong Prov., Ruyang Nat. Res., Lao Peng Keng stream, 1100m, elongate, almost straight in lateral view, tube-like seventh 14 Aug.1990, coll. P.P. Chen, C9012” (NHMW). sternum shorter than two preceding sterna combined, seventh abdominal segment with long projections; male elongate with Descriptive notes. – The specimens of R. abdominalis from relatively short abdomen, eighth abdominal segment ventro- Vietnam were not significantly different from specimens laterally depressed; similar lengths of antennal segments. described by Andersen & Chen (1995) from China, except However, it could be separated from R. malaisei by the they are slightly bigger and the median process of sternum 7 following characters: proctiger of male without hook-shaped in the female is less pointed. The following measurements projection on each side; paramere long and slender; are of Vietnamese specimens: projections on abdominal segment 8 of female straight, not curved. Apterous male: length 6.8-7.0, width 2.10-2.20 (n=4). Head width across eyes 1.30, interocular width 0.63, eye length 0.51. Lengths of antennal segments 1-4: 3.63: 0.99: 1.13: 0.92. Rhyacobates abdominalis Andersen & Chen, 1995 Lengths of leg segments (femur: tibia: tarsal segment 1: tarsal segment 2): fore leg: 3.98: 3.15: 1.11: 0.69; middle leg: 10.80: Rhyacobates abdominalis Andersen & Chen, 1995: 58-59, Figs. 12- 5.65: 2.47: 0.48; hind leg: 11.00: 3.88: 0.13: 0.21. 15. Apterous female: length 9.9-10.0, width 3.50 (n=5). Head Material examined. – Vietnam: apterous 4 males, 5 females, de- width across eyes 1.55-1.58, interocular width 0.79, eye length alated macropterous 1 female, Lao Cai province, Sa Pa, Hoang Lien 0.63. Lengths of antennal segments 1-4: 4.59: 1.21: 1.31: 1.04. National Park, Sin Chai, 1300m, coll. A.D. Tran, 1 Jun.2003, Lengths of leg segments (femur: tibia: tarsal segment 1: tarsal TAD0337 (ZMHU, ZRC); apterous 6 males, 5 females, de-alated segment 2): fore leg: 4.86: 4.25: 2.45: 1.07; middle leg: 12.00: macropterous 2 males, 1 female, same locality, coll. A.D. Tran, 3 7.10: 3.69: 0.58; hind leg: 13.00: 5.75: 0.17: 0.25. Length of Jul.2004, TAD0415 (ZMHU, ZRC). sterna 5 and 6: 0.59 and 0.71. Sternum 7 length 1.51.

De-alated macropterous male: length 6.4-6.6, width 2.06-2.11 (n=2). Head width across eyes 1.29-1.32, interocular width 0.52, eye length 0.54. Lengths of antennal segments 1-4: 3.53: 1.01: 1.16: 0.97. Lengths of leg segments (femur: tibia: tarsal segment 1: tarsal segment 2): fore leg: 3.72: 3.19: 0.98: 0.67; middle leg: 10.30: 5.60: 2.40: 0.44; hind leg: 12.40: 3.82: 0.11: 0.25. Pronotum covers almost of mesonotum, mainly black with a median yellow mark anteriorly and pale- yellowish posterior margin.

De-alated macropterous female: length 9.4-9.6, width 3.14- 3.30 (n=2). Head width across eyes 1.49-1.55, interocular width 0.63-0.79, eye length 0.62-0.63. Lengths of antennal segments 1-4: 4.46: 1.21: 1.31: 1.02. Lengths of leg segments (femur: tibia: tarsal segment 1: tarsal segment 2): fore leg: 4.70: 4.03: 2.33: 1.02; middle leg: 12.70: 7.00: 3.54: 0.56; hind leg: 12.30: 5.75: 0.16: 0.27. Pronotum similar to macropterous male.

Distribution. – China and Vietnam.

Remarks. – This is the first record of this species for Vietnam. It was found in the same habitat as Rhyacobates gongvo.

ACKNOWLEDGEMENTS

We are grateful to Prof. Peter K. L. Ng (National University of Singapore) and Prof. Nguyen Xuan Quynh (Hanoi University of Science, Vietnam) for their enthusiastic support of our study on aquatic Heteroptera of Vietnam. We also thank Dr. H. Zettel (Natural History Museum Vienna, Austria) for valuable comments on the manuscripts and for the loan of paratypes of Rhyacobates abdominalis, Dr. J. T. Polhemus Fig. 28. Habitus of Rhyacobates gongvo, holotype.

19 Tran & Yang: New Eotrechus and Rhyacobates from Vietnam

(Colorado, U.S.A) for the loan of Eotrechus vietnamensis LITERATURE CITED specimens. We are indebted to the late Prof. N. M. Andersen (Zoological Museum, University of Copenhagen, Denmark) Andersen, N.M., 1982. Semiterrestrial water striders of the genera for his valuable comments on the manuscript. The first author Eotrechus Kirkaldy and Chimarrhometra Bianchi (Insecta, thanks Prof. Tran Ninh (Hanoi University of Science) who Hemiptera, Gerridae). Steenstrupia, 9(1): 1-25. facilitated his trip to Vu Quang National Park that led to the Andersen, N.M., 1998. Notes on the genus Eotrechus Kirdaldy discovery of Rhyacobates anderseni and Mr. Hoang Quoc (Hemiptera: Gerridae) with a cladistic analysis and a new species Khanh (Hanoi) for assistance in the field that led to the from China. Steenstrupia, 24: 1-8. discovery of Eotrechus vietnamensis. We also thank Dr. Andersen, N.M. & P-p. Chen, 1995. A taxonomic revision of the Lanna Cheng (University of California, San Diego) for Ptilomerine genus Rhyacobates Esaki (Hemiptera: Gerridae), linguistic review on the manuscripts. This study is supported with five new species from China and adjacent countries. by National University of Singapore, Research Grant number Tijdschrift voor Entomologie, 138: 51-67. R-154-000-222-112. Zettel, H. & P-p. Chen, 1996. Beitrag zur Taxonomie und Faunistik der Gerridae mit Neubeschreibungen der Gattung Andersenius gen.nov. aus der Unterfamilie Ptilomerinae und weiterer Arten (Insecta: Heteroptera: Gerridae). Entomologische Abhandlungen Staatliches Museum für Tierkunde Dresden, 57(6): 149-182.

20 THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 21-48 Date of Publication: 28 Feb.2006 © National University of Singapore

A PHYLOGENETIC REASSESSMENT OF THE RARE S. E. ASIAN FIREFLY GENUS PYGOLUCIOLA WITTMER (COLEOPTERA: LAMPYRIDAE: LUCIOLINAE)

L. A. Ballantyne School of Agricultural and Veterinary Sciences, Charles Sturt University, PO Box 588, Wagga Wagga, NSW 2678, Australia.

C. Lambkin CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia.

ABSTRACT. – Cladistic analysis supports the return of the rare Malaysian (Bornean) firefly genus Pygoluciola Wittmer to generic status; it is redescribed from the five species included, and keys to males and known females presented. The number of reliably identified specimens in published accounts increases from 21 (in five species) to 28. Females of P. guigliae and P. wittmeri are described including morphology of the female abdomen and aspects of reproductive system anatomy which may be of significance in female identification. Discussion addresses different preservation techniques relative to interpretation of taxonomic characters and conjecture about function, association of males and females, functional morphology of the terminal abdomen of both sexes, and the value of colouration characters. Luciola undulata (Pic) is reassigned to Atyphella Olliff.

KEY WORDS. – Pygoluciola, taxonomy, female morphology, female reproductive anatomy, colour characters, preservation methods, Atyphella undulata.

INTRODUCTION with male abdomen structure, indicated to them the possibility of a copulation clamp. The dried pinned specimens precluded The Luciolinae present a most interesting array of any investigation of internal anatomy. opportunities for observation and study. They are apparently exclusively flashing fireflies, and a species of Pteroptyx A recent collection of seven alcohol preserved specimens, undertakes the spectacular synchronous rhythmic flashing on which includes both males and females of two species the Selangor River at Kampong Kuantan, (Pygoluciola wittmeri and P. guigliae) permits an (Ballantyne, 2001). Their importance to the tourism industry investigation and reinterpretation of male and female terminal of other Asian countries is being belatedly recognised. abdomen morphology, and for the first time an investigation for this group of female reproductive anatomy. For the taxonomist the variety of male terminal abdomen modifications has excited speculation about their function Until this collection, which was killed, fixed and preserved (Ballantyne, 1987b), and some resolution (the copulation by immersion directly into 70% alcohol, the rare group was clamp in Pteroptyx valida Wing et al., 1983). Such external represented by five species known from published records abdominal structures were used to erect new genera (e.g. Pteroptyx Olivier, 1902, based on the deflexed elytral apices of 11 males and 10 females [Pygoluciola guigliae (male, and ‘trilobed’ abdominal V 7). These subdivisions of the female), P. hamulata (three males), P. kinabalua (five males, Luciolinae, as listed by McDermott (1966), are unsatisfactory eight females), P. stylifer (male, female) and P. wittmeri and not representative (Ballantyne & Lambkin, 2000, 2001). (male)] (Olivier, 1885; Ballantyne, 1968; Ballantyne & The early taxonomy was thus male based, there being no Lambkin, 2001; Wittmer, 1939). Unusual in the Luciolinae, distinctive features of the female abdomen until now. four of the five species are now known from both sexes. Rarity of occurrence of fireflies when they are obvious by That distinctive features could occur externally on the female their light and easy to net, usually relates to their occurrence abdomen of any Lucioline firefly was unsuspected until in difficult or dangerous terrain, or in largely uncollected Ballantyne & Lambkin (2001), describing L. [P.] kinabalua, territory. The recent collecting activities in are thus highlighted aspects of the female abdomen, which, coupled most welcome.

21 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Pygoluciola Wittmer, a rare group of fireflies with unusual Specimens were scored by LB from types, and specimens male terminal abdomen modifications, and restricted to the compared to types, with these exceptions, which were scored Malaysian states of Sabah and , was considered a from published work and/or identified specimens (authority subgenus of Luciola (McDermott, 1966; Ballantyne, 1968; for identification indicated): Luciola dejeani Ballantyne & Ballantyne & Lambkin, 2001). Ballantyne & Lambkin (2000) Lambkin, 2000; Curtos costipennis specimens identified by scored Luciola (Pygoluciola) stylifer Wittmer in a cladistic N. Ohba, in ANIC, Jeng et al. (1998); Luciola pedemontana analysis of 43 species of Luciolinae. Because of many specimens identified by F. Papi, in ANIC, Jeng et al. (2003); missing features their analysis did not give unequivocal Colophotia praeusta identified by LB in comparison with directions for its taxonomic position. The analysis was rerun specimens in Olivier collection at Museum National by Lambkin using L. (P.) kinabalua as a second exemplar d’Histoire Naturelle, Paris, specimens in National Museum for this subgenus, with no missing adult characters, and ‘?’ of Natural History in Washington, D. C.; ‘Sisiak’ undescribed for larval characters only (Ballantyne & Lambkin, 2001), specimens, code named ‘Luciola 12’ in Lloyd (1973), work but again did not yield unequivocal directions for the in progress, in LB’s collection to be deposited in ANIC; ‘Mt. taxonomy of this group, thus Ballantyne & Lambkin (2001) Missim’ is an undescribed species based on two males and continued to address Pygoluciola as a subgenus of Luciola two females from high elevation in New Guinea, in ANIC; as had McDermott (1966). The specimens described here, Pteroptyx malaccae Ballantyne, 2001; Lampyroidea syriaca together with a re-examination of the type species identified specimens in BMNH London; Atyphella (Pygoluciola stylifer Wittmer), permit a re-evaluation of their marginipennis work in progress (Ballantyne) (Ballantyne & findings. Lambkin, 2000).

The position of Pygoluciola is addressed with a third cladistic Abbreviations for taxonomic characters are: ASW, antennal analysis and revised. The genus is redescribed and keyed socket width; FS, antennal flagellar segments, referred to by from known males and females. Males and females of two number e.g. FS 3, 4; GHW, greatest head width; ML, median species of Pygoluciola are described or redescribed from lobe aedeagus; MN, mesonotal plates; MPP, median posterior features of freshly collected specimens, with detail of the projection of ventrite 7; MS, mesoscutellum; LL, lateral lobe terminal abdomen of each including aspects of the female aedeagus; PLP, posterolateral projections of ventrite 7; SIW, genitalia and reproductive tract. smallest interocular width; T7, T8, abdominal tergite seven, eight; V followed by a number, abdominal ventrite/s.

MATERIALS AND METHODS Abbreviations for repositories of specimens are: ANIC Australian National Insect Collection CSIRO, Taxonomic characters are based on Ballantyne & Lambkin Canberra (2001), which in turn were described in detail in Ballantyne BBM Bernice Bishop Museum, Hawaii & Lambkin (2000); changes or emendations to these BORN Borneensis, Sabah characters are described following the character description; CMG Museo Civico di Storia Naturale Giacomo characters are numbered thus: 9 (8) indicating that character Doria’, Genoa 9 in this paper was character 8 in Ballantyne & Lambkin BMNH Natural History Museum, London (2001). Characters and states are included in the descriptions RMNH National Museum of Natural History Naturalis, e.g. ‘head moderately exposed 4 (1)’ = character 4, state 1; Leiden ‘92 (0&1)’ indicates that character 92 is polymorphic, equal numbers of specimens show either states 0 or 1; 92(0, 1) indicates for a polymorphic character that state 1 is the TAXONOMY majority state. Visible male abdominal sternites are referred to by their actual number, which is one more than their visible Pygoluciola Wittmer, 1939 number (= ventrites). The ventral plate of the concealed 9th abdominal segment is referred to as a sternite. Ballantyne Pygoluciola Wittmer, 1939:21. (1968) refers the terminal abdominal tergite to tergite 7; Luciola (Pygoluciola) – McDermott, 1966: 115; Ballantyne, 1968: abdominal segmentation was reviewed in Ballantyne (1987a) 119; Ballantyne & McLean, 1970: 233; Ballantyne & Lambkin, and this tergite referred to abdominal segment 8. The degree 2000: 82; Ballantyne & Lambkin, 2001: 363. of head exposure/coverage (characters 4, 5) was expanded to Type species. – Pygoluciola stylifer Wittmer, 1939, monobasic indicate the differing mechanisms by which the head can be (RMNH). concealed (able to be retracted into a pronotal cavity vs. covered by the anterior expansion of the pronotum). Branham Redescription of genus. – Male. Body elongate slender, 2.9 and Wenzel (2003) interpreted this (on many more genera – 3.3 times as long as wide. than used here) as a capacity to retract between the hypomera, thus effectively characterising the size of a pronotal cavity. Pronotum Width/length 1.6 – 2.0; 1/5 – 1/7 as long as whole Distortion affects absolute dimensions (Ballantyne, 1987a), 1 body length; always wider across posterior /3 than anterior and most dimensions here are given as absolute and on a 1 /3; median anterior margin gently rounded, or slightly comparative basis e.g. GHW as a function of SIW. medianly emarginate, scarcely projecting beyond anterolateral

22 THE RAFFLES BULLETIN OF ZOOLOGY 2006 corners if at all; anterolateral corners defined, rounded and Abdomen with six ventrites belonging to segments two – narrowly obtuse or angulate acute, and may project anteriorly seven; (remnants of segment one may be represented by beyond anteromedian margin; lateral margins diverging cuticularisation in the intersegmental membrane anterior to posteriorly along their length, sometimes with some slight V2); posterior margin of V 3 and V4 not recurved. Light sinuousity at mid point; posterolateral corners broadly organs: occupying all of V 6 except sometimes for some rounded, acute, and projecting either a little beyond median irregular erosion along anterior margin; LO in V 7 occupying posterior margin or not at all, and delimited from rest of at least half the area of V 7, anterior margin often slightly posterior margin by shallow emarginations which do not emarginate and not reaching anterior margin of V 7; lateral follow the anterior margin of the elytral humeral angle; lateral and posterior margins of LO gently curved, posterior margin margin near posterolateral corners not indented; lateral margin may be gently emarginate, no margins reaching to sides or (viewed from beneath) with anterior ‘hypomeral’ area posterior margin of V7; no area posterior to LO in V7 greatly developed and distinct from the posterior area, which is expanded or swollen. Ventrite seven (V7): no hairy lobes or 2 flattened in posterior /3, slightly more widely so in posterior pointed projections along posterior margin of V7; half than in anterior area, such that the dorsal and ventral posterolateral processes absent, (MPP in pinned specimens surfaces are adpressed to each other in dried pinned specimens tends to be engulfed behind by the downturned apex of tergite and closely approach in ‘wet’ preserved specimens; dorsal 8); posterolateral corners rounded; dorsal face posterior to surface smooth, barely convex, except for depressed area of LO (i.e. MPP) lacking muscle attachments; no median median sulcus and narrow depressed area running along longitudinal groove, carina or dimple; MPP longer than wide, posterior margin; lacking pronounced ridges running or at least as wide as long, symmetrical, curving strongly obliquely from posterolateral areas to median area of disc, dorsally (at about a 90º angle) towards its apex which may convexity over the posterior margins of the eyes, and a be truncate and expanded, or flattened and gently or strongly conspicuous flattening of the dorsal surface in median emarginate, and in dried pinned specimens engages against posterior area; irregular small low tubercles present in the downturned apex of tergite eight; if the apex of the MPP posterolateral areas in four of five species; punctures small, is truncate and expanded, the flat posterior surface makes an shallow, fairly inconspicuous, some contiguous, some approximately 90º angle to the horizontal). Tergite seven: separated by their width. wider than long or about as wide as long, with strongly depressed lateral areas beneath which dorsoventral muscles Elytra punctation not conspicuously larger than pronotal attach, and an elevated rounded median area covering the punctation, not linear; apex not deflexed ; with four barely anterior prolongation of tergite eight and the aedeagal elevated apunctate interstitial lines, delimited at their sides complex; median posterior margin recurved. Tergite eight: by punctures which are linear along the side of the interstitial elongate, narrower than tergite seven, if viewed from above line; epipleuron and sutural ridge extending to rounded apex, is about twice as wide in anterior half as in posterior half with neither thickened in apical half; preapical sutural ridge (widest at point of lateral angulations), lateral margins not downturned; lateral margins not strongly explanate; narrowing posteriorly in posterior half, most of apex not epipleuron only slightly expanded; elytra at least five times visible from above, and strongly deflexed ventrally and in as long as wide, subparallel-sided or tapering posteriorly. dried pinned specimens enveloping the MPP of V7 from behind (in freshly killed specimens preserved in ethanol these Head in dorsal view is moderately exposed in front of structures do not approach closely); anterior margin of tergite pronotum; vertex shallowly to moderately depressed; lacking eight bifurcate into two elongate slender apically rounded posterolateral eye excavation; eyes moderately to widely projections; angulate margins at point where tergite eight separated above labrum; GHW 3 – 4.8 X SIW; labrum narrows posteriorly may be slightly asymmetrical; narrowed transverse (wider than long); antennal sockets separated by apex of tergite eight may be entire or gently emarginate; ASD up to twice but not three times ASW; frons vertex ventral surface in posterior (narrowed) half lacking median junction not defined; eyes moderately separated behind longitudinal grooves, lateral ridges, depressed troughs, flanges mouthparts ventrally. Mouthparts well developed, especially or any asymmetrical projections; ventral surface in anterior fleshy lobes of maxillae, and assumed functional; apical half with a median longitudinal groove separating two segment of labial palpi laterally flattened, with the longest, thickened surfaces in P. wittmeri; ventral surface in anterior inner edge bearing a number of slender finger like projections half lacking such developments, and lacking longitudinal which may differ from one palp to the other; apical segment ridges, flanges lateral ridges in P. guigliae, P. kinabalua and of maxillary palpi ovoid, not much longer than wide, and as P. stylifer. long as apical segment of labial palpi, with an acutely rounded tip. Antennae length two to three times GHW; 11 segmented, Aedeagal sheath (not extracted in P. stylifer, P. hamulata; all segments elongate slender (at least three times as long as significance of this structure not realised until Ballantyne, wide), simple, apical FS never conspicuously shortened or 1987a, b); about twice as long as wide, much narrower in club like, no FS produced laterally or flattened. anterior half; anterior half of sheath sternite symmetrical and very narrow, narrowed abruptly at about the mid point of its Legs femora and tibiae straight, not swollen, not curved in length, or slightly less; posterior half of sheath symmetrical, three of five species; tibiae of all legs curved in P. guigliae sheath sternite and tergite equally wide; tergite lacks swollen and P. stylifer; all basitarsi simple, not excavated on their lateral protuberances but lateral margins of tergite may be inner margins. MFC absent. visible very narrowly at sides of posterior half of V 9 (seen

23 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer only in dried pinned specimens of P. kinabalua); median stylifer, and Ballantyne & Lambkin (2001), without a re- anterior margin of tergite 9 evenly emarginate, lateral margins examination, erroneously keyed P. stylifer using the character anteriorly prolonged at the sides of the sheath sternite and of straight tibiae. This is corrected below. joining the narrowed anterior prolongation of this ventrite in two species, and visible from beneath; posterior margin of sternite narrowing gently, level with or projecting slightly List of species of Pygoluciola Wittmer posteriorly beyond tergite. 1. Pygoluciola guigliae (Ballantyne, 1968) Aedeagus Length/Width 2.5 – 4.0; LL and ML symmetrical, 2. Pygoluciola hamulata (Olivier, 1885) basal piece and dorsal base of LL asymmetrical; basal piece 3. Pygoluciola kinabalua (Ballantyne & Lambkin, 2001) in two narrow halves, not sitting symmetrically above ML, 4. Pygoluciola stylifer Wittmer, 1939 left half slightly more ventral than right; ML tapering to a 5. Pygoluciola wittmeri (Ballantyne, 1968) rounded acute apex, shorter than LL, sometimes just longer than half the length of the aedeagus; LL visible to either side of ML when viewed from above or below, separate along Key to species of Pygoluciola using males almost all their length dorsally, inner dorsal margins always 1. All tibiae curved; lateral margins of elytra tapering posteriorly divergent at least in basal fourth, and often closely ...... 2 approximate in apical half; apical half of LL subequal in – No tibiae curved; lateral margins of elytra usually sub-parallel width, apices no wider than preceding half, and sometimes sided ...... 3 out-turned; LL lacking leaf like lobes on their outer (lateral) 2. Apex of median posterior projection of tergite 8 no wider than margins, bearing hairs along at least half of their outer margin. rest and rounded; pronotum with large median darker area; lateral margins of pronotum slightly sinuate ...... P. guigliae Female macropterous, and assumed capable of flight; similar – Apex of median posterior projection of tergite 8 wider than rest to male differing in these aspects: pronotum about twice as and medianly emarginate; lateral margins of pronotum not wide as long, with lateral margins diverging posteriorly, but slightly sinuate ...... P. stylifer 3. Median posterior projection of abdominal ventrite 7 bifurcate median anterior margin often projecting beyond rounded at apex ...... 4 obtuse anterolateral corner; head of winged female form, – Median posterior projection of abdominal ventrite 7 not bifurcate smaller than that of male; abdomen with eight visible at apex ...... P. hamulata ventrites. Light organ occupying the whole of V 6 only; V7 4. Apex of median posterior projection of abdominal ventrite 7 with accessory external developments (transverse ridge, deeply emarginate, laterally ensheathing the downturned apex lateral mounds) in two of five species; tergite seven with of tergite 8 and projecting laterally beside it ...... P. wittmeri anteromedian mound in one of five species; V7 usually quite – Apex of median posterior projection of abdominal ventrite 7 deeply emarginate across posterior margin, V8 narrower than shallowly emarginate, not laterally ensheathing the downturned V7, sides tapering posteriorly and median posterior margin apex of tergite 8 and not projecting beside it..... P. kinabalua slightly indented. Female genitalia (known from two species) with styli longer than wide, parallel sided and apically rounded, and visible beyond end of V8; coxites not well Key to species of Pygoluciola using females differentiated, partly sclerotised and lateral margins 1. All tibiae curved ...... P. guigliae converging posteriorly; valvifers elongate, slender, well – No tibiae curved ...... 2 sclerotised and rod like, extending in front of coxites for about 2. Posterior margin of Ventrite 7 deeply emarginate; bearing a small 7 /10 of the total length of the complex. ridge anterior to median area of deepest emargination; ventrite 8 with anteromedian prolongation not any more sclerotised than Larva unknown. remainder of ventrite 8; tergite 7 with anteromedian area rounded and elevated, lateral areas not flattened ...... P. kinabalua Remarks. - Wittmer (1939) misinterpreted the abdominal – Posterior margin of Ventrite 7 shallowly emarginate, lacking segmentation and recorded seven ventrites. His third last an anteromedian ridge; ventrite 8 with anteromedian segment, a light organ, is actually V 6, and he interpreted the prolongation well sclerotised and visibly separated from remainder of ventrite 7; tergite 7 lacking a rounded and elevated light organ on V 7 as a separate segment surrounded at the anteromedian area, with lateral areas flattened .... P. wittmeri sides and behind by the last abdominal sternite. This is one segment, V 7, only. Ballantyne (1968) briefly redescribed Luciola (Pygoluciola) from four species, two of which were Pygoluciola guigliae (Ballantyne) known from three specimens only. There is inconsistency (Figs. 1, 6, 13-17, 19, 21, 27, 30, 32, 34, 35, 38-40) between that description and this because of the redefinition of abdominal segmentation by Ballantyne (1987a,b) (tergite Luciola (Pygoluciola) guigliae Ballantyne, 1968: 120 [Holotype seven there becomes tergite eight here; visible abdominal male British North Borneo, Bundu Tukan (BBM)]. sternites are referred to as ventrites and given their actual segmentation number), and certain other characters which are Diagnosis. – Both sexes with strongly curved tibiae on all redefined here (e.g. interstitial lines), and that redescription legs; males elongate, slender, 3.2 times as long as wide, with should be replaced by the current one. Ballantyne (1968) elytra tapering towards their apices, most similar to P. stylifer, did not distinguish the curved legs on the holotype of P. distinguished by the darker pronotal colour, the forwardly

24 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Figs. 1-17. Features of male Pygoluciola. 1, 6, 13-17: P. guigliae; 2: ‘Mt Missim’ male; 3, 7: P. kinabalua holotype male; 4, 10-12: P. stylifer holotype male; 5, 8-9: P. wittmeri. 1-5: dorsal, male pronota (left side shows colour pattern, right side sculpturing; solid line indicates extent of irregular tubercles, dotted lines retraction of fat body): 6,7: dorsal , whole body; aedeagi (8; 10 ventral; 9, 12 dorsal; 11, left lateral); 13: maxillary palp, ventral surface; 14: labial palp, ventral surface; 15-17: aedeagal sheath, dorsal, ventral, and lateral. These figures share scale lines: 1-5; 6-7; 8-14; 15-17. Figure legend: A, anus; BP, basal piece; EO, ejaculatory orifice; LL, lateral lobe; ML, median lobe; TS, tergite aedeagal sheath; VS, ventrite aedeagal sheath.

25 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Figs. 18-36. Features of female Pygoluciola: Figs. 18-27; Features of female Pygoluciola: Figs. 28-36. 18, 22, 26, 27-29, 31, 33, 36 (P. wittmeri); 19, 21, 27, 30, 32, 34, 35 (P. guigliae); 20, 23-25 (P. stylifer holotype male). 18, 19, 23, 28, 30, ventral terminal abdominal segments [29 stippled outline in V 7 = fat body, in 30 = lateral elevations of V7]; 20, 22, 29, dorsal, terminal abdominal segments; 21, vertical face of apex of MPP of ventrite 7, dorsal face uppermost; 24, posterior face apex of tergite 8; 25, 26, lateral terminal abdominal segments, ventrites uppermost [26 anus, stippled, protrudes through MPP of V7]; 27, dorsal face abdominal tergite 8; 31, 32, dorsal surface abdominal ventrite 7 [right side only; stippling = dorsally reflexed cuticle; in 32 dotted oval = position of elevation]; 33, 34, ventral face, abdominal ventrite 8; 35, 36, ventral female genitalia. These figures share scale lines: 18, 20, 22, 23, 28-30; 19, 25-27; 31-34; 35-36. Figure legend: AS, aedeagal sheath complex; C, coxite; LO light organ; ST, stylus; T7, T8 tergite 7, 8; V, valvifer, female genitalia.

26 THE RAFFLES BULLETIN OF ZOOLOGY 2006 projecting anterolateral pronotal angles, the slight narrowing margins are semitransparent until apex, which is very in the lateral pronotal margin, and the shape of the apex of narrowly darker brown (when elytron is viewed separate to abdominal tergite 8; female with posterior margin of V 7 underlying hind wing); elytral punctures very dark brown, deeply and broadly emarginate, with lateral projections interstitial lines pale brown and margined by a single row on apically acute and longer than broad, and smoothly raised each side of dark punctures; head between eyes, antennae elevations present in lateral areas; tergite 8 lacking any median and palpi dark brown except for pale under half of FS 4 – 9 mound; anterior prolongation of V9 not as long as posterior and inner half of apical segment of maxillary palpi; labrum (wider) portion of this ventrite, well sclerotised, and brown, semitransparent; ventral surface of pro and membranous separation from the rest of the ventrite in mesothorax light brown, of metathorax dark brown, darker posterior area only. in median area; legs 1, 2, dark brown except for pale brown 1 coxae and trochanters and ventral surface of basal /6 of Material Examined. – Three males, one female, Malaysia, femora; legs 3 dark brown except for pale trochanters and Sabah: 5.48°N 116.26°E, Mahua, 1045 m, coll. narrow basal area of femora; basal abdominal ventrites dark Mahadimenakbar bin Mohamed Dawood, 19 Mar.2004, 20.45 brown, V 6 pale semitransparent [underlying fat body shows – 22.00 hrs. (BORN). Specimens of both P. guigliae and P. through cuticle] along anterior margin where pale yellow light wittmeri were collected “at night by a river where some organ is broadly and shallowly emarginate; V7 pale individuals formed a group of about 10, perching and flashing semitransparent in areas not occupied by light organ, with on branches of trees. Some were collected while flying. They underlying fat body visible especially in anterior areas, and are quite common here” (pers. comm. Collector). That there a small, short, very dark median line just behind posterior was more than one species was not recognised at the site, margin of light organ; MPP pale, semitransparent; dorsal and it is not known whether one or both species aggregated surface of tergites 2 – 6 brown, dorsally reflexed margins of in the trees or whether one or both were taken in flight. No ventrites 2-5 brown; tergite 7 mottled brown in median (raised observations on light production patterns were made. area) mainly pale in lateral areas with a small median brown spot, posterior margin pale semitransparent; tergite 8 pale Redescription. – Male. Body 11.1 – 11.9 mm long; 3.2 times semitransparent with a slightly darker median line; dorsally as long as wide (Fig. 6). Pronotum yellow, semitransparent, reflexed margins of V6, V7 pale. marked extensively in dark brown in median area (Fig. 1); pubescence pale; underlying fat body clustered closely under Pronotum (Fig. 1) 2.8 – 3.0 mm (2) wide; 1.8 (2) - 1.9 mm 1 cuticle in all areas except around margin; MS, MN yellow; long; width/length 1.6; pronotum /6 as long as whole body; elytra pale brown semitransparent; both suture and lateral median anterior margin not indented, projecting only slightly

Fig. 37. Female reproductive system. A, diagrammatic representation of ventrite 8, part of female genitalia, bursa and spermatheca and gland [bursa is shown to side of spermatheca] Figure not to scale, dimensions represented relatively; B, internal aspect, hook in bursa; scale line is 1 mm. Figure legend: B, bursa (arrow indicates hook); O, median oviduct; S, spermatheca; P, gland; V8, ventrite 8; V, valvifers female genitalia.

27 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer beyond anterolateral corners and separated from them by paler brown markings on pronotum (extent is confused by rounded shallow emarginations; anterolateral corners retraction of underlying fat body in the coloured areas); angulate, (anterior and lateral margins subtend approximately ventral colouration as for male except for all legs which have a right angle between them); lateral margins divergent very dark brown apices of femora, dark brown tibiae and tarsi, 2 posteriorly with a slight narrowing at posterior half, then and pale golden brown coxae, trochanters and basal /3 of diverging evenly until posterolateral corner; posterolateral femora; basal abdominal ventrites moderately dark brown; corners angulate, acute, not projecting posteriorly as far as pale light organ in V6, and V7 and 8 shiny light brown; median posterior margin, which is entire, and separated from abdominal tergites brown except for golden brown tergite 8. it by shallow emarginations. Elytron (Fig. 6) 9.2 - 10 mm 2 1 long; lateral margins diverge in anterior /9 and converge in Pronotum 1.8 mm long; 3.3 mm wide; pronotum /7 as long 7 posterior /9; maximum width across elytra, measured in as whole body; median anterior margin not indented, not 2 anterior third, 4 mm; elytra 2.5- 3.0 mm wide (at /9 of their projecting either beyond or behind anterolateral corners and length from tip). separated from them by rounded shallow emarginations; anterolateral corners angulate obtuse; lateral margins Head gently depressed between eyes; GHW 1.7 - 1.8mm; divergent posteriorly with a slight narrowing at posterior half, SIW 0.5 - 0.6 mm; ASD = ASW or just slightly >; apical then diverging evenly until posterolateral corner; segments of labial palpi (Figs. 13, 14) with 3 teeth on each posterolateral corners angulate, acute, not projecting (1); 4 on R and 3 on L (1), 4 on each (1); antennal length posteriorly as far as median posterior margin, which is entire, almost 3 X GHW, pedicel slightly narrowed near base; with and separated from it by shallow emarginations. FS 1-6 subequal in length, about 4 times as long as wide, and FS 6 - 9 subequal in length slightly narrower than 1 - 6. Elytra 10.2 mm long; elytral lateral margins divergent in 2 1 anterior /7, subparallel in next /7, then convergent in apical 4 3 Legs with all tibiae curved, stouter at apices than at base /7; maximum width across elytra 4.3 mm (measured at /10 (maximum width of apex/maximum width at base of hind their length from front); maximum width across elytra 3.2 2 tibiae =1.2 - 1.7) and all femora stout, not curved (maximum mm (measured at /10 their length in front of apices). width hind femora 0.5 – 0.6 mm). Head completely concealed beneath pronotum at rest; GHW Abdomen (Figs. 19, 21) with V2 lacking faint median ‘carina’ 1.8 mm, SIW 0.6mm; ASD subequal to ASW; head barely between bases of coxae 3. Light organs: entire in V6 with a depressed between eyes; mouthparts well developed, broad emargination along anterior margin; entire in V7, with functional; apical segments of maxillary and labial palpi a shallow anteromedian emargination, not posteromedially shaped as for male, except the right apical segment of the emarginated. Ventrite 7: MPP 1.2 X longer than wide, not labial palpi has 4 elongate slender evenly spaced projections expanded at its tip when viewed from beneath; when viewed on the longer inner margin while the left palp has 4 with the 2 from side, tip expanded in posterior /3, such that the ventral middle two fused at their bases. margin of the MPP is continuous with the more anterior margin of V7, there being no change of angle, and the dorsal Legs with all tibiae curved, stouter at apices than at base surface expands such that the posterior half of the MPP is (maximum width of apex/maximum width at base of hind twice as deep as the anterior half; posterior margin of MPP tibiae = 1.3) and all femora moderately stout, not curved not emarginated dorsally (Fig. 21), rounded ventrally, with (maximum width hind femora 0.5 mm). ventral half of posterior half clothed in short fine hairs and dorsal half slightly depressed, paler than rest; dorsal surface Abdomen (Figs. 30, 32, 34) with posterior margin of V7 of anterior part of MPP with narrow short longitudinal evenly and quite deeply emarginate, posterolateral corners depression corresponding to external dark marking; apex of apically acute and longer than wide, projecting posteriorly tergite 8 (Fig. 27) narrows gradually towards tip, rounded, beyond the median posterior margin of V7 by more than their entire. width; ventral surface with posterolateral elongate rounded protuberances, but lacking any troughs, or ridges; anterior Aedeagus in the aedeagal sheath (Figs. 15 – 17) protrudes half of V8 narrowed, and linearly prolonged beneath V7, behind the posterior margin of the MPP of V7 in the median prolonged portion shorter than wider posterior portion of V8, line, lateral lobes splay to each side of the downturned tip of all of anterior prolongation well sclerotised; dorsal surface tergite 8. Aedeagus with ML much shorter than LL, which of tergite 7 evenly rounded. are widely divergent along their dorsal, inner margins in basal 2/7 of their length and closely approximate along their apical Genitalia similar to P. wittmeri (Figs. 35, 36), with some 4/7 with apices touching then diverging to each side; LL in sclerotisation between coxites, and valvifers long, slender and ‘wet’ preserved specimens white, plump (about as high as not diverging in anterior half. wide), becoming dorsoventrally flattened and darker after immersion in 20% KOH for about 20 minutes. Reproductive system (Fig. 37). The junction of the ‘vagina’, the median oviduct, and the duct leading to the bursa and Description. – Female. 12.0 mm long; body 4.3 mm wide spermatheca occurs just above the anterior prolongation of across widest point of closed elytra; macropterous, and V8 and the anterior tips of the valvifers of the female genital observed in flight; dorsal colouration as for male except for complex (2.5 mm from the posterior margin of V8). Muscles

28 THE RAFFLES BULLETIN OF ZOOLOGY 2006 run from the anterior narrow portion of abdominal V8 and Diagnosis. – Most similar to P. stylifer, distinguished by the attach onto the anterior portions of the valvifers. The non curved legs and the apex of abdominal tergite 8 being 1 presumed spermatheca is about /10 the size of the entire (that of stylifer is emarginate). spermathecal gland, both appear quite muscular and neither has any obvious internal cuticularisation. Paired anteriorly directed hooks, attached to the sides of the median duct [the Pygoluciola kinabalua (Ballantyne) bursa] at their bases only, occur just in front of the junction (Figs. 3, 7, 38-40) of the median oviduct to the vagina, and a half empty presumed spermatheca was found here. These hooks are Luciola (Pygoluciola) kinabalua Ballantyne & Lambkin, 2001: 371 above the narrowed anterior prolongation of V8. [Holotype male Malaysia, Sabah: Mt Kinabalu (BMNH)].

Remarks. – Ballantyne (1968) describe the pronotal colour Diagnosis. – Males elongate slender (Figs. 3, 7), about three on a pinned specimen as red-brown and the elytral with an times as long as wide, with elytra subparallel sided; one of apical brown area, and the single (pinned) female as lacking two species of Pygoluciola with the apex of the MPP of V7 curvature on the anterior tibiae. emarginate, gently so in this species; distinguished from P. wittmeri, which has a deeply emarginate apex; female with posterior margin of V7 deeply and broadly emarginate, with Pygoluciola hamulata (Olivier) a small transverse ridge just anterior to this emargination, (Figs. 38-40) lateral projections moderately broad and apically rounded; tergite 7 with strongly sclerotised median ‘mound’; lateral Luciola hamulata Olivier, 1885: 367; 1902: 80; 1907: 52; 1913: 59 margins of anterior prolongation of V8 not well sclerotised [Holotype male Borneo, Sarawak (CMG)]. nor well defined. Luciola (Luciola) hamulata – McDermott, 1966: 105. Luciola (Pygoluciola) hamulata – Ballantyne, 1968: 121

Figure 38. (A). The strict consensus of the 2992 MPT for 49 taxa of the Luciolinae found with phylogenetic analysis of 115 characters, excluding 10 colour and mimicry characters. (B). The majority rule consensus of the 2992 MPT for the Luciolinae, showing above the branch the percentage of MPT retaining that node. 153 of the MPT were compatible with the majority rule consensus.

29 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Figure 39. Semistrict consensus of 195 successively weighted trees. Percentage of successively weighted trees with this resolution shown above the branch. None of the MPT was compatible with the semistrict consensus successively weighted trees.

30 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Figure 40. Preferred MPT from 153 filtered MPT compatible with the majority rule consensus of the 2992 MPT, Part 1, without Atyphella clade. Unambiguous character and state changes were mapped using MacClade. Colour and mimicry characters, not used in the phylogenetic analysis, are enclosed in boxes. Bremer support values shown above nodes.

31 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Figure 41. Preferred MPT from 153 filtered MPT compatible with the majority rule consensus of the 2992 MPT, Part 2, Atyphella clade. Unambiguous character and state changes were mapped using MacClade. Colour and mimicry characters, not used in the phylogenetic analysis, are enclosed in boxes. Bremer support values shown above nodes.

32 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Pygoluciola stylifer Wittmer as wide and slightly shorter than scape); FS 6 – 9 more slender (Figs. 4, 10 – 12, 20, 23 – 25, 38-40) than preceding FS.

Pygoluciola stylifer Wittmer, 1939: 22 [Holotype male Borneo Legs with all tibiae curved. (RMNH). Labelled 1. hand written Pygoluciola stylifer Wittm.; printed det. W. Wittmer. 2. orange typed ‘Holotype’. 3. hand Abdomen (Figs. 20, 23 – 25) MPP of V7 expanded at its tip written ‘TYPUS’. 4. Printed ‘H. C. Siebers/M. O. Borneo Exp./ as described for P. guigliae; tergite 8 narrowed before its apex 120 M. bij L. Petak/1925 viii – ix.’]. which is expanded with posterior margin shallowly Luciola (Pygoluciola) stylifer – McDermott, 1966: 115; Ballantyne, 1968:119. emarginate.

Diagnosis. – Most similar to P. hamulata, distinguished by Aedeagus (Figs. 10 – 12) with ML about half as long as LL; the curved tibiae of all legs, and the emarginate apex of inner dorsal margins of LL strongly divergent in basal half, abdominal tergite 8. converging level with tip of ML and closely approximate until apical 1/6, where they diverge slightly. Redescription of Holotype Male. – Elongate slender, three times as long as wide; elytra tapering towards their apices. 11.2 mm long. Pronotum light golden brown with no apparent Pygoluciola wittmeri (Ballantyne) darker markings on the dorsal surface except for very narrow (Figs. 5, 8-9, 18, 22, 26-29, 31, 33, 36, 38-40) dark brown markings along anterior margin to either side of mid line; fat body is retracted along anterior and posterior Luciola (Pygoluciola) wittmeri Ballantyne, 1968: 121; Ballantyne & Lambkin, 2001: 371 [Holotype male, ‘British North Borneo’ margins and these areas appear paler brown; MS, MN paler (BBM)]. brown than pronotum; elytra light golden brown, slightly darker than the pronotum, and apical 1/6 a little darker than Diagnosis. – Male distinguished from P. kinabalua see above; rest; elytral punctures dark brown; head and maxillary palpi female with posterior margin of V7 shallowly emarginate and very dark brown, labial palpi mid brown; antennae dark posterolateral corners apically rounded and moderately broad; brown, left antenna (R is incomplete) with dorsal surface of ventral surface of V7, 8 smooth; anterior prolongation of V8 FS 6 – 9 paler brown than preceding segments, and very pale longer than posterior (wider) portion, and lateral margins well whitish yellow beneath; ventral surface of pro and meso sclerotised in posterior half. thorax light brown; legs 1 with coxae and femora light brown, tibiae darker brown, tarsi very dark brown; ventral surface Material examined. – One male, two females, Malaysia, Sabah, of metathorax dark brown ; legs 2 and 3 yellowish except for 5.48°N 116.26°E, Mahua, by the Mahua River, 1045 m, coll. dark brown tibiae and tarsi 2, and tarsi 3; basal abdominal Mahadimenakbar bin Mohamed Dawood, 19 Mar.2004, between ventrites dark brown, V6 creamy yellow (light organ), V7 2045 – 2200 hrs. Malaysian time (Borneensis). yellowish (light organ is torn from surface of V7); tergites 7 and 8 pale. Redescription. – Male. 10.1 mm long. Pronotum pale creamy yellow and semitransparent, marked faintly in pale brown in Pronotum (Fig. 4) 3.1 mm wide; 1.8 mm long; width/length posteromedian areas; pubescence pale; underlying fat body 1 = 1.7; pronotum /5 as long as whole body length; lateral clustered closely under cuticle in all areas except two margins smoothly divergent along their length; median anteromedian areas and narrowly across most of the anterior anterior margin rounded, not emarginate, projecting a little margin; MS, MN pale creamy yellow, fat body dense under beyond anterolateral corners, which are narrowly rounded and MN and restricted to posterior half of MS; elytra pale brown obtuse, but do not project anteriorly; posterolateral corners semitransparent; both suture and lateral margins are rounded, acute (60 - 70°), not projecting beyond posterior semitransparent until apex, which is very narrowly darker margin; dorsal surface of pronotum mainly fairly smooth and brown (when elytron is viewed separate to underlying hind flat except for slight depressions in median and anterolateral wing); elytral punctures very dark brown, interstitial lines areas and narrowly across the posterior margin, and with pale brown and margined by a single row on each side of irregular low elevations in posteromedian areas. dark punctures; head between eyes, antennae and palpi dark brown except for pale under half of FS 7 – 9 and inner half Elytra 9.4 mm long, apparently tapering toward their apices; of apical segment of maxillary palpi; labrum brown, 1 left elytron lateral margin divergent in anterior /4, subparallel semitransparent; ventral surface of pro and mesothorax light 1 1 sided in next /4, and convergent in posterior /2; right elytron brown, of metathorax very dark brown in median area with 1 similar but more strongly convergent in posterior /2 (this may metepisterna and metepimera paler brown; legs 1, 2 with pale be a post-mortem effect). creamy yellow coxae, trochanters and femora, except for brown inner margins of coxae and trochanters, and brown 2 Head gently depressed between eyes; GHW 1.9 mm; SIW apical /3 of femora; tibiae and tarsi of all legs dark brown; 0.5 mm; GHW/SIW 3.8; ASD subequal to ASW; apical legs 3 with ventral surface of coxae very dark brown except segments of both labial palpi with three elongate projections for narrow pale basolateral areas; trochanters and femora pale on their inner surface; antennae slightly longer than twice yellow; tibiae very light brown; tarsi dark brown, not as dark GHW; FS 1-5 subequal in length (about three times as long as tarsi 1, 2; V2 light brown with dusky cream lateral areas;

33 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

V3 with median brown area more extensive than that of 2, Description. – Female. 11.5 - 12.0 mm long; body 5.5 mm and small anterolateral paler areas which are outlined wide across widest point of closed elytra; macropterous, and posteriorly in dark brown; V4 moderately brown except for observed in flight; dorsal colouration as for male except for paler anterolateral areas; V5 similar, more darkly marked; extensive median dark brown markings on pronotum; ventral V6 wholly occupied with light organ; V7 pale semitransparent colouration as for male except for brown anterior ventrites in areas not occupied by light organ, with underlying fat body which are marked in posterolateral areas with small irregular visible especially in posterolateral areas, and a small very pale brown markings, pale light organ in V6, and V7 and 8 dark median triangular mark just behind posterior margin of shiny light brown; abdominal tergites light brown. light organ; MPP pale, semitransparent, fat body clustered in 2 1 anterior /3; dorsal surface of tergites 2 – 6 brown, darkening Pronotum 2.0 mm long; 4 mm wide; pronotum /6 as long as progressively towards tip of abdomen; dorsally reflexed total body; median anterior margin smoothly rounded, margins of ventrites pale; tergites 7 and 8 pale, 8 with a fine projecting a little beyond the rounded, obtuse anterolateral dark median line. angles which are not produced; lateral margins diverge posteriorly; median sulcus prominent, dark, and depressed Pronotum (Fig. 5) 1.6 mm long, 2.8 mm wide, width/length in its posterior half; dorsal surface narrowly depressed along 1 = 2; /7 as long as whole body; anterior margin rounded, posterior margin; pronotal surface smooth lacking any slightly indented in median line, not projecting beyond irregular tubercles; posterior area of median sulcus bearing a anterolateral corners and separated from them by slight cluster of long white posteriorly directed hairs; a line of hairs emarginations; anterolateral corners pointed, acute; lateral extends along posterior margin [hairs only visible when margins divergent posteriorly with no convergences; specimen dried]; punctures small mainly separated by their posterolateral corners rounded acute, not projecting beyond width. median posterior margin and separated from it by shallow emarginations; dorsal surface smooth, mainly flat; slight Elytra 10.5 - 11 mm long; elytra slightly convex sided lateral 2 irregularities of the surface on either side of the median area margins diverge along anterior half, subparallel for next /8 3 8 in posterior half of dorsal surface. then converge in posterior / .

Elytron 8.6 mm long; lateral margins subparallel for half their Head completely concealed beneath pronotum at rest; GHW 3 length, converging posteriorly in posterior /8. 2.0 mm, SIW 0.7mm; ASD < ASW; head barely depressed between eyes; mouthparts well developed, functional; apical Head shallowly depressed between eyes; GHW 1.7 mm; SIW segments of maxillary and labial palpi shaped as for male, 0.35mm;ASD > ASW but not 2 X ASW; antennal length except the apical segments of the labial palpi have three approximately twice GHW, with FS 1-6 subequal in length, elongate slender projections on their longer inner margin in about 4 times as long as wide, and FS 7-9 subequal in length one female, and four on left and three on right in second but slightly shorter than 1 - 6; apical segment of labial palpi female (the male has four on both). with 4 slender evenly spaced projections on both palpi. Abdomen (Figs. 28, 29, 31, 33) with posterior margin of V7 Legs with no segments swollen or curved; width of hind evenly and shallowly emarginate, posterolateral corners femora at widest area 0.4 mm. apically acute and moderately broad, projecting posteriorly beyond the median posterior margin of V7 by a little more Abdomen (Figs. 18, 22, 26, 27) with V2 with faint median than their width; ventral surface lacking any troughs, ‘carina’ between bases of coxae 3. Light organs: entire in elevations or ridges; anterior half of V8 acutely narrowed, V6; entire in V 7, with a shallow anteromedian emargination and linearly prolonged beneath V7, prolonged portion longer (not visible in this specimen unless abdomen is manipulated); than wider posterior portion of V8, lateral margins of anterior not posteromedially emarginated. Ventrite 7: with small prolongation well sclerotised in posterior half; dorsal surface anterolateral depressions but no obvious signs of strong D-V of tergite 7 laterally flattened, median area evenly and linearly muscle blocks; MPP much longer than wide, apically widely rounded; tergites 7, 8 and V 7, 8 more heavily sclerotised emarginate and curving dorsally where in the single male than preceding segments. examined the emargination engulfs the end of the alimentary canal; lateral arms of emarginate apex apically rounded, about Genitalia (Fig. 36) similar to P. guigliae, lacking any median as wide at base as at apex; median posterior margin of tergite sclerotisation between coxites, valvifers long slender and 7 rounded and projecting backwards over the anterior margin diverging in anterior half. of tergite 8; apex of tergite 8 narrowly and shallowly medianly emarginate. Reproductive system as described for P. guigliae; two females were dissected, in both the two structures arising anterior to Aedeagus (Figs. 8, 9) in the aedeagal sheath protrudes behind the bursa are well developed, thick walled and equal in size, the posterior margin of the LO in V7 in the median line and and assumed to be a gland and a spermatheca. the rectum is engulfed by the emarginate MPP. Aedeagus with ML a little shorter than LL, which are widely divergent Remarks. – The width between the high points on V8 of the along their dorsal, inner margins and closely approximate but female could accommodate the apex of the male MPP not contiguous at their apices. between them. In this alcohol preserved male the MPP and

34 THE RAFFLES BULLETIN OF ZOOLOGY 2006 tergite 8 apex are widely divergent, and the aedeagus, still Strict (Schuh & Farris, 1981), semistrict (Bremer, 1990), and partly contained in the aedeagal sheath, protrudes towards majority-rule consensus (Margush & McMorris, 1981) of the tip of tergite 8. In the (pinned) holotype the apex of tergite MPT were computed using PAUP*. The 50% majority rule 8 is contained between the apical emargination of the MPP. consensus shows all nodes that are found in more than 50% Because of its disposition in this specimen, there seems no of the trees. Groups are preserved in the majority rule reason to expect the aedeagus to be extracted laterally, despite consensus tree even if there are some MPT that support the downturned apex of tergite 8 and upturned apex of V 7. conflicting groups (Margush & McMorris, 1981). Bremer support (Bremer, 1988; Kallersjö et al., 1992; Bremer, 1994) values were calculated with TreeRot v.2 (Sorenson, 1999) PHYLOGENETIC ANALYSIS with 20 heuristic searches of the data. The command line (nchuck=10000 chuckscore=322 abortrep=yes) was added for Our earlier phylogenetic analyses of the Lampyridae each heuristic search to prevent prolonged searches of (Ballantyne & Lambkin, 2000, 2001) were expanded to suboptimal tree space. include 49 taxa, with the addition of another four species: Atyphella undulata, P. guigliae, P. hamulata, and P. wittmeri. Phylogenetic Results A full list of exemplar taxa is given in Table 1, the Data Analysis of 115 characters (excluding the colour and mimicry Matrix. One hundred and twenty-five morphological characters) in the data matrix in Table 1 produced 2992 MPT characters were scored comprising 315 states in the analysis. of length 312, consistency index (Kluge & Farris, 1969) 0.56, Most are binary, but 30 characters were coded as multistate consistency index without uninformative characters (Kluge characters each with three states, while characters 4, 5, 56, & Farris, 1969) 0.50, retention index (Farris, 1989) 0.75, and 65, 66, and 75 were coded with four states, characters 9, 11, rescaled consistency index (Farris, 1989) 0.42. The strict (Fig. 55, 73, and 103, with five states, character 2 with six states, 38A) and 50% majority rule (Fig. 38B) consensus trees were and character 98 with eight states. The six characters (97- calculated. Successive weighting of the 2992 MPT produced 100, 114-115) that describe colour, and four characters (101- 195 trees for which the semistrict consensus (Fig. 39) was 2, 116-7) that describe mimicry or colour patterns were not calculated and saved. A tree filter, created in PAUP*, found used in the phylogenetic analysis but mapped onto the none of the 2992 MPT was compatible with the semistrict resultant trees to examine patterns of evolution. consensus of the successively weighted trees. Therefore the successive weighting was unable to aid in choice of MPT. Character analysis However, a second filter found 153 of the 2992 MPT were Phylogenetic analyses presented here do not prejudge the compatible with the 50% majority rule consensus of the 2992 relative informativeness of characters (Larson & Dimmick, MPT. One of these 153 MPT (Figs. 40-41) was chosen as 1993; Kallersjö et al., 1999). All character changes were the preferred MPT, and Bremer supports calculated and weighted equally (Farris, 1990). Character polarity was shown above the nodes. Unambiguous character and state determined using outgroup comparison (Maddison et al., changes, including the 10 colour and mimicry characters not 1984; Nixon & Carpenter, 1993). Characters with more than used in the phylogenetic analysis, were mapped onto the one derived state were coded as multistate, not in binary form preferred MPT (Figs. 40-41) in MacClade (Maddison & (Meier, 1994), and treated as unordered (non-additive). Maddison, 2001). Cladistic analyses completed 1000 random step-wise addition heuristic searches, with TBR branch swapping, MULPARS, Characters and branches having maximum length zero collapsed to yield Characters and states are numbered to reflect their number in polytomies using PAUP*4.0b10 (Swofford, 2002). Ballantyne & Lambkin (2001) [e.g. 6(5) = character 6 here was character 5 previously], and are as described therein with Initial searches indicated that there were most parsimonious these exceptions which are described following the trees (MPT) of length 312, but searches tended to become appropriate character– 1. Modified: 2(2), 3(3), 4(4), 6(5), stuck in suboptimal tree space. Therefore 10000 random step- 15(14), 18(17), 22(21), 279260, 28(27), 40(39), 41(39), wise addition heuristic searches were completed, saving no 51(47), 71(63), 77(70), 93(82), 94(83), 98(86), 99(87), more than 3000 MPT of length greater than or equal to 313, 113(96); 2. New: 5, 48, 49, 52, 57-60, 78-81, 96, 101-2, 106- and abandoning the replicate if that limit was hit. 9, 111, 116, 117.

Successive approximations character weighting (successive Pronotum (1-10) weighting) is a method of choosing between MPT (Farris, 1. Hypomera: 1969; Carpenter, 1988). It is an iterative character weighting Open 0 procedure in which the weights are assigned, based on their Closed in front 1 observed level of homoplasy. Initial weights are derived from the MPT under equal weighting, and heuristic searches 2. Inclination of lateral pronotal margins: completed based on the Rescaled Consistency Index (Farris, Diverging along anterior half or more with some 1989). The procedure is repeated until the weights (and the convergence in posterior area which is not sinuate trees) remain stable over iterations. Successive weighting of (Ballantyne & Lambkin 2000, Fig. 1a-d, f, g, the MPT was computed using PAUP*. i-l) 0

35 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Diverging along all their length except for a slight 6(5). Posterolateral corners of pronotum (shape and sinuousity at about the mid point of the lateral angle): margin 1 Rounded obtuse (Ballantyne & Lambkin, 2000, Diverging along anterior half or more (widest Fig. 1 a-c, d, f-g, I, j-l) 0 1 across posterior margin) with posterior /4 sinuate Right angled or angulate obtuse (Ballantyne & (gentle convergence followed by gentle McLean, 1970, Fig. 4h) 1 divergence) 2 Rounded acute 2 Diverging along their length (Ballantyne & Pointed acute (Ballantyne, 1968, Fig. 130) 3 McLean, 1970, Fig. 4c) 3 States 2, 3 are combined and state 4 becomes state 3. Subparallel or wider across middle with slight convergence anteriorly and posteriorly 4 7(6). Posterolateral corners of pronotum: Converging posteriorly along their length Scarcely projecting beyond posterior margin nor (Ballantyne & Lambkin, 2000, Fig. 19a) 5 delimited by a deep emargination of the posterior New states 1 and 2 accommodate slight sinuousity of the lateral margin 0 margin; old state 1 becomes 4, and 3 becomes 5. In states 0-3 the pronotum is wider across the base than across the middle; in 4 it Projecting considerably beyond posterior margin may be wider across the middle than across the base; in 5 it is wider and often delimited by an emargination of the across the anterior margin than the base. Posterior margin (Ballantyne, 1968, Figs. 20, 33, 38, 51, 62, 70) 1 3. Anterolateral corners of pronotum: Projecting beyond posterior margin but not Obliterated (there is a smooth even transition from delimited by a wide pronounced emargination 2 the middle of the anterior margin to the lateral margins; no anterolateral corner is defined) (Lloyd 8(7). Lateral margin of pronotum near posterolateral & Ballantyne, 2003, Figs. 5-10,19) 0 corner: Broadly rounded, obtuse (Ballantyne & Lambkin, Not indented 0 2000, Fig. 1, 19a-e) (smooth fairly even transition Slightly indented 1 from middle of the anterior margin to the lateral The indentation is slightly irregular and small, contrasting with the margins, anterolateral corner discernable but not evenly sinuate posterior margin of character 2 state 2. well defined, rounded and widely obtuse) 1 Narrowly rounded, obtuse (corner defined, 9(8). Lateral margin of pronotum (viewed from rounded, and narrowly obtuse) 2 beneath): Angulate acute (corner defined, not smoothly Not flattened 0 rounded, acute 3 Flattened only in posterior half 1 Old state 2 is submerged into 3, and a new state 2. All of lateral margin narrowly flattened but not more so in posterior half 2 4 (4). Degree of head exposure in front of pronotum All of lateral margin narrowly to widely flattened (governed by retraction into pronotal cavity): but more so in posterior half of pronotum 3 Concealed but not retracted into the cavity beneath Lateral margin widely flattened along its length the pronotum (outgroup) 0 and anterior area as wide as or wider than posterior Greatly exposed (slightly retracted into the cavity area 4 beneath the pronotum) 1 Scarce to moderately exposed (moderately 10(9). Anterior margin of pronotum: retracted into the cavity beneath the pronotum) 2 Not explanate 0 Completely concealed (fully retracted into the Narrowly explanate 1 cavity beneath the pronotum) 3 See explanation following 5; a new state 1 is necessary. Elytron (11-18) 11(10). Punctation: 5. Degree of head exposure as covered by expanded Not conspicuously larger than pronotal anterior and lateral margins of the pronotum: punctation 0 Covered (Photuris)0Conspicuously larger than pronotal punctation 1 Not so covered 1 In the Luciolinae the head is concealed by retraction into the cavity 12(11). Apex: beneath the pronotum; and this concealment relates to the size of Not deflexed 0 both head and pronotal cavity; lateral pronotal margins even if Deflexed with apex rounded (Ballantyne & explanate do not contribute to this concealment. The head is either Lambkin, 2000, Fig. 20 j, k) 1 greatly exposed in front of the pronotum (when the head cannot be retracted within and beneath the pronotum), slightly exposed (when 13(12). Apex: at least some of the posterior section of the head is retracted within and beneath the pronotum at rest) or completely concealed (not Not deflexed 0 visible from above). In Photuris spp. (outgroup) the wide head Deflexed with sides A, B, C equal cannot be retracted into the pronotal cavity and both the explanate (Ballantyne & Lambkin, 2000, Fig. 20k) 1 anterior and lateral margins contribute to head concealment, from Deflexed with sides A, B longer than C above. Characters 4 and 5 make this distinction. (Ballantyne & Lambkin, 2000, Fig. 20j) 2

36 THE RAFFLES BULLETIN OF ZOOLOGY 2006

14(13). Development of epipleuron and sutural apex in 23(22). Proximity of antennal sockets: apical half of elytron: Contiguous 0 No thicker than rest 0 Separated by < 2 x ASW but not contiguous 1 Considerably thicker than anterior portions 1 Separated by at least 3 x ASW 2 Separated by > twice but not 3 x ASW 3 15(14). Interstitial line development as elevation similar to that of sutural ridge: 24(23). Frons-vertex junction: 4 well defined lines (1, 2, 3, 4 including the humeral carina, line 3 arising at or near the Not acute (Ballantyne & Lambkin, 2000, humerus) 0 Fig. 2 a, d, h) 0 3 well defined lines (lines 1, 2, and 3 or 4) 1 Acute (Ballantyne & Lambkin, 2000, Fig. 2 c, f, 2 well defined lines (lines 1, 2 closest to the g, I) 1 suture) 2 Only one line well defined 3 25(24). Median area of frons vertex junction: No lines well defined 4 Not elevated or indented 0 This character is amplified to reflect the development of all the Elevated and/or indented (Ballantyne & Lambkin, possible interstitial lines which may be visible (e.g. paler in colour 2000, Fig. 2f, g) 1 than rest, or often delimited by punctures along their sides), but not elevated comparable to the elevation of the sutural ridge, in which case they are classed as state 4. 26(25). Anterior margin of head: Not prolonged 0 16(15). Epipleuron: Prolonged in front of eye for about its width 1 Extending to apex of elytron 0 See Ballantyne, 1968, Figs. 132, 134. Extending past mid point of elytron but not to apex 1 27(26). Length width of labrum: Extending no further than mid point of elytron 2 Posterior margin of labrum not defined (clypeolabral suture absent) 0 17(16). Sutural ridge: Posterior margin of labrum defined (clypeolabral Extending to apex of elytron 0 suture present); labrum wider than long 1 Evanescent before elytral apex 1 Posterior margin of labrum, defined by clypeolabral suture present; labrum about as long 18(17). Margins: as wide 2 Parallel sided (may converge in posterior 1/3) 0 A new state 0 defines the situation where, in the absence of a Convex sided 1 clypeolabral suture, the posterior margin of the labrum cannot be accurately determined and it is not possible to measure L/W of labrum. Head, antennae, mouthparts (19-41) 19(18). Depression of vertex: 28(27). Clypeolabral suture: Minimal 0 Present, clypeus and labrum not fused, and junction Moderate - deep 1 between clypeus and labrum flexible 0 20(19). Approximation of eyes on ventral surface of head Present, clypeus and labrum fused, and junction (measured as eye separation taken just behind between clypeus and labrum inflexible 1 mouthparts/GHW measured ventrally): Absent, clypeus and labrum fused with no defined Wide separation(0.5 or greater) 0 junction between the two (Ballantyne, 1968, Figs. Close to moderate separation (0.4 or less) 1 132, 134) 2 Contiguous or almost so 2 John Lawrence (ANIC Canberra) determined the extent of the labrum and clypeus, and the position of the clypeolabral suture on specimens of Photuris and “Mt Missim’. The states are elaborated 21(20). Posterolateral eye excavation (Ballantyne, 1968, here; scoring remains the same as in Ballantyne & Lambkin (2001). Figs. 144, 147-150): Absent; if slightly developed not visible when head 29(28). Mouthparts: is retracted 0 Functional 0 Well developed and usually visible even when Non functional 1 head is evenly retracted 1 Non functional mouthparts have very small apical segments of labial A well developed posterolateral eye excavation is at least as and maxillary palpi, mandibles often do not cross in the median wide as long when viewed from the side. line and the antennal sockets are contiguous.

22(21). Antenna length: 30(29). Proximity of eyes above labrum (SIW/GHW): Longer than twice GHW 0 Close (1/6 - 1/15) 0 > GHW - 2 x GHW 1 Moderately separated > 1/6 1 Subequal to GHW 2 Widely separated 1/3 – 1/2 2 Amplification ‘much’ eliminated from state 0.

37 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

31(30). Antennal flagellar segment 1: 45(43). Curvature of femora 3: As long as or longer than pedicel 0 Not curved 0 Shorter than pedicel 1 Curved along their length (Ballantyne & Lambkin, 2000, Fig. 26h) 1 32(31). Antennal flagellar segment 1: Not expanded at its outer apex 0 46(44). Swelling of tibiae 3: Expanded at its outer apex 1 Not swollen 0 Swollen at least at their apices (Ballantyne & 33(32). Median area of antennal flagellar segment 1: Lambkin, 2000, Fig. 26h) 1 Not produced 0 Produced (Ballantyne & McLean, 1970 47(45). Curvature of tibiae 3: Fig. 18b) 1 Not curved 0 34(33). Flagellar segments 7-9: Curved (Ballantyne & Lambkin, 2000 Not conspicuously shorter than rest of FS 0 Fig. 26h) 1 Conspicuously shorter than rest of FS 1 48. Curvature of tibiae 2: 35(34). Number of segments: Not curved 0 11 0 Curved 1 < 11 1 49. Curvature of tibiae 1: 36(35). Flagellar segment 9: Not curved 0 Apically rounded 0 Curved 1 Apically pointed 1 Abdominal segments – ventrites (50 - 70) 37(36). Flagellar segments 2-8: 50(46). Ventrite 8: Not expanded 0 Present 0 Expanded at anterior apical angle 1 Absent 1 38(37). Antennal segments: 51(47). Light organ in ventrite 7: Not flattened 0 Entire (e.g. Ballantyne & Lambkin, 2000, Flattened 1 Fig. 4a) 0 39(38). Pedicel: Posterior medial division short - long (Ballantyne, Not produced at outer apex 0 1968, Fig. 42, 91) 1 Produced at outer apex 1 Bipartite (Ballantyne & Lambkin, 2000 Fig. 26a) 2 40(39). Shape of apical segment of labial palpi: State 2 was erroneously listed as 3 in Ballantyne & Lambkin (2001). Fusiform, or about as wide as long 0 Laterally flattened 1 52. Intrusion of swollen posterior half of ventrite 7 into light organ posterior margin: 41(39). Nature of inner margin of apical segment of labial Posterior half of ventrite 7 not swollen 0 palpi: Posterior half of ventrite 7 swollen but not reaching Entire (margin rounded, convex) 0 into posterior half of light organ 1 Dentate 1 Posterior half of ventrite 7 swollen and reaching Lunate 2 into emarginate posterior border of light organ 2 Character 39 has been split into two to reflect the shape of the apical segment of the labial palp and the nature of its inner margin. 53(48). Light organ in ventrite 7: Reaching sides and posterior margin of ventrite Legs (42-49) 70 42(40). Inner tarsal claw of each leg: Reaching sides but not posterior margin 1 Not split 0 Split 1 Not reaching sides or posterior margin 2

43(41). Metafemoral comb (Ballantyne, 1987a, Fig. 1j): 54(49). Size of light organs: Absent 0 Occupying at least half of the area of ventrite 7 or Present 1 more 0 Restricted to very small paired anterolateral 44(42). Femora 3 (Ballantyne & McLean, 1970, Fig. 4p): plaques that occupy less than 10% of the area of Not swollen 0 ventrite 7 (Ballantyne & McLean, 1970, Swollen 1 Fig. 18c) 1

38 THE RAFFLES BULLETIN OF ZOOLOGY 2006

55(50). Apex of MPP of ventrite 7: 63(55). Median longitudinal trough on ventral surface of MPP not developed 0 MPP: Apex truncate 1 Absent 0 Apex rounded 2 Present (Ballantyne & McLean, 1970, Apex gently emarginate (e.g. Ballantyne & Fig. 3 d, f) 1 Lambkin, 2000 Fig, 20 d, f, j, k, i) 3 Apex deeply emarginate (Ballantyne & Lambkin, 64(56). All of ventrite 7, especially in posterior half: 2000 Fig. 3 d, e, g, j; Ballantyne, 1968 Flat, not arched or swollen (e.g. Ballantyne & Fig. 13) 4 Lambkin, 2000, Fig. 4a) 0 Arched and often swollen (Ballantyne, 1968, Figs. 56(51). Length/ width of MPP of ventrite 7: 56, 89) 1 Not produced 0 About as long as broad or shorter; narrower than 65(57). Length of PLP of ventrite 7: half the width of ventrite 7 1 Not developed 0 About as long as broad; at least half as wide as ventrite 7 2 Slightly produced (Ballantyne & Lambkin, 2000, At least twice as long as wide 3 Fig. 26a) 1 Moderately produced, may extend beyond the tip 57. Relation of posterior margin of ventrite 7, to apex of the MPP 2 of tergite 8: Considerably produced (Ballantyne, 1987b Fig. 2 MPP developed 0 a, b) 3 MPP not developed, posterior margin of Ventrite 7 not enveloped by strongly ventrally deflexed 66(58). Width of PLP of ventrite 7: apex of tergite 8 1 Not developed 0 MPP not developed, posterior margin of ventrite Narrower than MPP (Ballantyne &Lambkin, 2000, 7 engulfed by the strongly ventrally deflexed apex Fig. 26a) 1 of tergite 8 2 As wide as MPP (Ballantyne & McLean, 1970, Fig. 9 a, b) 2 58. Dorsal inclination of MPP: Broader than MPP (Ballantyne, 1987b, Fig. 2 a, MPP not developed 0 b) 3 MPP developed, not strongly reflexed dorsally 1 MPP developed, strongly reflexed dorsally 2 67(59). Inclination of PLP of ventrite 7: Not developed 0 59. Relationship of MPP to tergite 8: Horizontal 1 MPP not developed 0 Oblique - vertical 2 MPP developed but not engulfed at its posterior See Ballantyne & McLean, 1970, Fig. 3 d, f; and Ballantyne, 1968, margin by the strongly downturned tergite 8 Figs. 11, 13. apex 1 MPP developed and engulfed at its posterior 68(60). Incurving hairy lobes along posterior margin of margin by tergite 8 apex 2 ventrite 7 (Ballantyne & Lambkin, 2000, Fig. 26a): Absent 0 60. Inclination of apex of tergite 8: Present 1 Apex of tergite 8 not strongly inclined ventrally 0 69(61). Pointed projection of ventrite 7 posterior margin Apex of tergite 8 strongly inclined ventrally 1 (Ballantyne & Lambkin, 2000, Fig. 26a): Character 52 is replaced by 57 – 60, to distinguish the situation where the apices of tergite 8 and the MPP of ventrite 7 are narrowed Absent 0 and approach each other by being strongly inclined (at approximately Present 1 a 90° angle) in either a ventral or dorsal direction; it eliminates species such as A. obsoleta, A. huonensis and ‘Mt. Missim’, where 70(62). Dimple on ventrite 7 (Ballantyne & Lambkin, both tergite 8 and ventrite 8 may curve slightly towards each other, 2000, Fig. 20k): but are not bent at a pronounced angle. Absent 0 Present 1 61(53). Median longitudinal carina in ventrite 7: Absent 0 Abdominal segments - tergites (71-76) Present (Ballantyne & McLean, 1970, Fig. 3 d, f) 1 71(63). Flanges on ventral face of tergite 8: Lacking flanges 0 62(54). Median longitudinal trough in ventrite 7: With flanges (Ballantyne & Lambkin, 2000, Fig. Absent 0 20l) 1 Present 1 State 2 is eliminated and state 1 modified.

39 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

72(64). Ventral face of tergite 8 with depressed lateral 78. Width of posterior area of ventrite 9 (posterior to troughs (Ballantyne & Lambkin, 2000 Fig. 26c, point of articulation with sheath tergite): d): As wide as tergite of sheath and symmetrical; if Absent 0 narrower than tergite of sheath then Present 1 symmetrical 0 About half as wide as tergite of sheath and 73(65). Posterior half of ventral face of tergite 8 with asymmetrical (right side of ventrite is evenly elongate longitudinal symmetrical developments emarginate along its length 1 margining a median longitudinal trough: Absent 0 79. Length of posterior area of ventrite 9 (posterior to point of articulation with sheath tergite): Fine transparent barely elevated ridges margining About as long as tergite of sheath 0 a median longitudinal trough 1 Shorter than tergite of sheath but not less than half Slightly thickened barely elevated ridges its length 1 margining a median longitudinal trough 2 Shorter than half the length of tergite of sheath2 Low barely elevated ridges present in posterior half of ventral surface only 3 80. Symmetry or asymmetry of posterior margin of Longitudinal raised well developed ridges sheath ventrite: delimiting a median longitudinal trough 4 Posterior margin of sheath ventrite entire, rounded 0 74(66). Ventral face of tergite 8 with asymmetrical Posterior margin of sheath ventrite projections (other than flanges), and/or transverse emarginate,symmetrical 1 ridges and/or hooks: Posterior margin of sheath ventrite emarginate, Absent 0 asymmetrical 2 Present (Ballantyne, 1987b, Fig. 2f) 1 Ballantyne (1987b) termed all the narrowed ventral projections of 81. Width of aedeagal sheath ventrite in anterior half: tergite 8 ‘flanges’. Flanges are here interpreted as the narrowed Broad; if narrowed then this is a gradual narrowing anterior projections often of lateral longitudinal ridges on the ventral anteriorly 0 surface of tergite 8. Abruptly narrowed, anterior prolongation much narrower than posterior half of ventrite 1 75(68). Width of tergite 8: About as wide as long 0 82(71). Length/width of aedeagal sheath: Very short 1 Never more than about 4 times as long as wide0 Much longer than wide and projecting Very long and narrow (about 7 times as long as considerably beyond MPP 2 wide) 1 Longer than wide but not projecting considerably beyond MPP 3 83(72). Lateral margins of aedeagal sheath: Lacking paraprocts (Ballantyne & Lambkin, 2000, 76(69). Width of posterior half of tergite 8: Fig. 21 c, d) 0 Not narrowed, or lateral margins converging With paraprocts (Ballantyne & Lambkin, 2000, posteriorly but not abruptly 0 Fig. 21 a, b) 1 Abruptly narrowed (Ballantyne, 1968, Figs. 17,110,112,114) 1 84(73). Length/width of tergite 9 of aedeagal sheath: About as long as wide 0 Aedeagal sheath (77- 84) Much wider than long 1 77 (70). Aedeagal sheath ventrite symmetry in posterior area i.e. posterior to point of articulation with the Aedeagus (85-96) sheath tergite, as distinct from symmetry or 85(74). Maximum width across lateral lobes/ maximum otherwise of posterior margin of ventrite 9: width of median lobe: Wide (4 – 6/1) (Ballantyne, 1968, Fig. 171) 0 Symmetrical (Ballantyne & Lambkin, 2000, Fig. Moderate (2/1) (Ballantyne & Lambkin, 2000, Fig. 21 b, d, e) 0 5) Asymmetrical in posterior area (Ballantyne & Narrow (less than 2/1) (Ballantyne & Lambkin, Lambkin, 2000, Fig. 21 i) 1 2000, Fig. 21 o, r, u; 26e) 2 Undescribed specimens ‘Sisiak’ were scored state 1 in Ballantyne & Lambkin, 2000; the posterior margin of the sheath ventrite is asymmetrical; state 1 is modified above and new characters 77-79 86(75). Inclination of apex of median lobe: address this inconsistency. Not curving ventrally 0 Curving ventrally (Ballantyne, 1968, Figs. 164, 168) 1

40 THE RAFFLES BULLETIN OF ZOOLOGY 2006

87(76). Extent of preapical ventral area of median lobe: 95(84). Lateral appendages of lateral lobes: Not produced 0 Absent 0 Produced and rounded 1 Present (McDermott & Buck, 1959, Produced and pointed (Ballantyne, 1968, Figs. 162, Fig. 62a-c) 1 164, 168) 2 96. Shape of basal piece of aedeagus: 88(77). Length of median lobe of aedeagus relative to Not hooded 0 lateral lobes: Hooded 1 Much shorter than LL 0 Subequal in length to lateral lobes or slightly Male colour patterns (97 – 102) longer 1 97(85). Colour of pronotum: Longer than LL but less than twice their Pronotum concolourous 0 length 2 Pronotum with dark markings 1 Much longer than LL i.e. more than twice their (separated) length 3 98(86). Colour of elytral margins compared to rest of elytra: 89(78). Separation of lateral lobes of aedeagus: Elytra concolourous (dark – light, pale brown) 0 Separated for > half their length 0 Separated for less than half their length (Ballantyne If elytron dark then only lateral margin pale 1 & Lambkin, 2000, Fig. 26 e-g) 1 Lateral and sutural margins pale at least in basal half 2 90(79). Width of lateral lobes of aedeagus: Lateral and sutural margins pale with base of Much wider and flatter at their apices than widest elytron dark 3 point of ML (Ballantyne & Lambkin, 2000, Fig. Lateral sutural and apical margins pale, base of 21 l, m) 0 elytron pale 4 About as wide at apices as widest point of ML If elytron pale then darker markings scattered at (Ballantyne & Lambkin, 2000 Fig. 5) 1 base and apex 5 Much narrower at apices than widest point of ML If elytron pale then dark markings at apex only6 (Ballantyne &Lambkin, 2000 Fig. 21 p,s,u,v; Fig. Elytra concolourous (Pale, usually yellow) 7 26 e-g) 2 States 1 and 7 are amplified to avoid confusion.

91(80). Separation of lateral lobes into broad basal section 99(87). Colour of interstitial lines 1, 2 as distinct from basal and narrowed widely separated apical section: elytron colour or colour of its margins: No such separation 0 No paler than dark area between lines 1 and 2 0 Present (Ballantyne & Lambkin, 2000, About as pale as this area or slightly paler 1 Fig. 21k) 1 Distinctly paler than this area so I lines appear as stripes 2 92(81). Aedeagal symmetry: Where interstitial lines traverse differing colours on the elytra this Aymmetrical 0 character is scored according to the dominant colour of these lines. Asymmetrical (Ballantyne & Lambkin, 2000, Fig. 26e, f) 1 100(88). Colour of terminal abdominal tergum: As dark as or darker than preceding terga 0 93(82). Fleshy leaf like lobes arising from lateral margins Pale (as pale as preceding terga or paler) 1 of lateral lobes: Absent 0 101. Dorsal colouration pattern: Present (Ballantyne & Lambkin, 2000, Fig. 21l, Dorsal colouration crypsis 0 n) 1 Dorsal colouration mimicry 1 This modification eliminates states 1 and 3, those species with various, non leaf like lobes supporting in some cases dense hair 102. Dorsal colouration pattern: bearing areas. Mimicry 0 Crypsis – background matching 1 94(83). Extent of ventral face of apices of lateral lobes: Crypsis – disruptive colouration 2 Extending to either side of ML and both visible In background matching, cryptic colour patterns randomly resemble from beneath (Ballantyne & Lambkin, 2000, Fig. the visual background in shape, size and colour at the time and place 5, Fig 21 k, l, m) 0 of highest predation risk (Endler, 1978). Because edges and Almost extending to sides of LL but both apices boundaries are the primary features used in visual recognition, a may not be visible in the same plane 1 mechanism often used to attain crypsis is disruptive coloration (Cott, Not extending to either side of ML and not visible 1957) where the organism’s shape, or even number, is obliterated from beneath except in basal half or less (Oxford & Gillespie, 1998). The blends into the background (Ballantyne & Lambkin, 2000, Fig. 21) 2 as the surface is broken into meaningless shapes or the attention of A new state 1 is included. the predator is drawn to nonessential patterns (Merilaita, 1998).

41 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Adult female morphology (103 – 113) 113(96). Head form: 103(89). Development of fore wings of female: Of same form as male 0 Fully developed (or covering all but two terminal Of winged female form (Ballantyne & Lambkin, abdominal segments) 0 2000, Fig. 6b) 1 Elytra longer than pronotum but shortened such Of wingless female form (Ballantyne & Lambkin, 1 2 that they cover approximately /2 - /3 of the 2000, Fig. 6d, e) 2 abdomen 1 A new state 0 is necessary. Elytra shorter than pronotum (more than half as long as pronotum) and contiguous in the median Adult female colour patterns (114 – 117) line 2 114(92). Colour of pronotum: Elytra shorter than half pronotal length and often Pronotum concolourous 0 contiguous or closely approaching in the median Pronotum with coloured markings 1 line 3 Elytra shorter than half pronotal length and widely 115(93). Colour of elytra: separated in the median line 4 Elytra concolourous 0 Macropterous gravid females may have one to two abdominal Elytra not concolourous 1 segments protruding beyond the elytral apices. 116. Dorsal colouration mimicry or crypsis: 104(90). Development of hind wings of female: Dorsal colouration crypsis 0 Fully developed 0 Dorsal colouration mimicry 1 Hind wings about 2/3 as long as macropterous state 1 117. Dorsal colouration pattern: Hind wings vestigial or absent 2 Mimicry 0 Crypsis – background matching 1 105(91). Extent of female light organ: Crypsis – disruptive colouration 2 Occupying ventrites 6 and 7 0 Restricted to ventrite 6 1 Larval Morphology (118 – 125) 118(97). Production of lateral margins of terga: 106. Developments of dorsal face of abdominal tergite Lateral margins not explanate (Ballantyne & 8: Lambkin, 2000, Fig. 22) 0 Tergite 8 lacking dorsal elevation 0 Lateral margins narrowly explanate especially at Tergite 8 with dorsal elevation 1 posterolateral corners (Ballantyne, 1968, Figs. 158- 160) 1 107. Developments of median ventral face of abdominal Lateral margins widely explanate (Ballantyne & ventrite 7: Lambkin, 2000, Figs. 12, 15) 2 Ventrite 7 lacking median ridge 0 Ventrite 7 with median ridge (Ballantyne & 119(98). Length/width of pronotum: Lambkin, Fig. 16) 1 Longer than wide 0 About as long as wide 1 108. Developments of lateral areas of ventral face of ventrite 7: 120(99). Nature of tergal margins: Ventrite 7 lacking lateral developments 0 Not ridged 0 Ventrite 7 with lateral developments 1 Ridged 1

109. Depressed lateral areas on ventrite 7 ventral face 121(100). Paired dorsal and ventral tubercles on protergum: (corresponding to dorsoventral muscles): Absent 0 Absent 0 Present (Ballantyne & Lambkin, 2000, Fig. 15a, Present 1 c) 1

110(94). Number of well defined elytral interstitial lines: 122(101). Shape of posterolateral corners of protergum: Four 0 Rounded (Ballantyne & Lambkin, 2000, Fig. 15 Less than four 1 a-c) 0 Acute (Ballantyne & Lambkin, 2000, 111. Elytral carina: Fig. 12 c) 1 Absent 0 Narrowly produced 2 Present 1 123(102). Margins of median line on terga 1-10: 112(95). Nature of pronotal punctures: Not ridged 0 Contiguous in at least lateral areas 0 Ridged 1 Not contiguous in any area 1

42 THE RAFFLES BULLETIN OF ZOOLOGY 2006

123(103). Size of punctures in anterior half of terga 2-10: it is unresolved in both the strict and majority rule consensus No larger than rest 0 (Fig. 38), and the preferred MPT (Fig. 40). The successively Larger than rest 1 weighted semi-strict consensus (Fig. 39) suggests that Curtos costipennis is closely related to Pygoluciola, as in our previous 125(104). Extent of posterolateral corners of tergum 11: analyses; however Curtos costipennis now falls between Not produced (Ballantyne & Lambkin, 2000, Fig. Pygoluciola and Atyphella. Future studies including more 7a, 15b) 0 members of Curtos will address the placement of this genus. Produced (Ballantyne & Lambkin, 2000, Fig. 12a- c, 15 a, c) 1 The clade of Atyphella, found in our previous analyses, is retained in the successively weighted semi-strict consensus (Fig. 39). However the majority rule consensus (Fig. 38B) DISCUSSION indicates that the clade Atyphella is present in only 75% of the MPT. The preferred MPT (Figs. 40, 41) shows the very Taxonomy low support values for the internal nodes of the clade. Future This study supports the return of Pygoluciola to generic status, studies including more members of Atyphella will address and the general trends shown in previous analyses (Ballantyne relationships in this genus. & Lambkin, 2000, 2001) while indicating directions for further resolution within the Luciolinae. Taxonomic features The basal (L. italica-L. cowleyi) and (Colophotia praeusta- that permit reliable identification of certain females occur on Pyrophanes beccarii) clades found in our previous analyses the external female abdomen and within the reproductive tract. are retained, not as sister clades but paraphyletic with low This is a useful development as Luciolinae taxonomy until support, in the strict and majority rule consensus (Fig. 38), now has been male based, and there have been no reliable the successively weighted semi-strict consensus (Fig. 39), and external or internal features of females to permit identification the preferred MPT (Fig. 40). to species. That the Luciolinae might be a monophyletic group united However there are also indications of problems with firefly by the possession of a reduced number of abdominal taxonomy that could have wider application to other soft segments, is under investigation. While it appears that it is bodied insects. The method and state of preservation confuses abdominal ventrite 8 in the male that is missing (Ballantyne interpretation of certain taxonomic characters, and influences 1987a, b), and this can be interpreted by counting the conjecture about the function of various structures. Certain abdominal spiracles to confirm segmentation, it has not been artefacts identified on the female abdomen are a consequence determined whether the ventrite is lost or fused with 7. of dehydration. The use of colour by early taxonomists in Branham and Wenzel (2001) established a monophyletic fireflies was widespread, but colour is of limited value in concept of the Lampyridae in which a Luciolinae clade was cladistic analyses. These aspects are explored below. represented by 8 species (5 of which were scored here). Although we continue to use the same set of species Phylogeny (Ballantyne & Lambkin, 2000, 2001), decisions about further Pygoluciola is clearly distinct and separate from Luciola. The revisions of taxonomic categories in the Luciolinae await genus Pygoluciola is found in all MPT, and therefore in the work in progress (Ballantyne, unpub.). However with few strict consensus (Fig. 38A), in all successively weighted trees exceptions, the same categories arise from this analysis as in (Fig. 39), and receives the highest Bremer support of any the previous two (Ballantyne & Lambkin 2000, 2001): group in this analysis of the Luciolinae (Fig. 39). 1. The type species of Luciola, L. italica (= pedemontana), The relationships between Pygoluciola and other taxa are less and L. dejeani, L. cowleyi, L. (Hotaria) parvula, clear. The majority rule consensus (Fig. 38B), like our Lampyroidea syriaca (the type species of Lampyroidea), previous analyses (Ballantyne & Lambkin, 2000, 2001), and Bourgeoisia hypocrita. Kawashima et al. (2003) indicates that ‘Mt Missim’ is more closely related to submerged Hotaria into Luciola without a cladistic Pygoluciola than any other taxa. ‘Mt Missim’ is more closely analysis. A distinctive aedeagal pattern of wide flaring related to Pygoluciola in 75% of the MPT, and 25% of the lateral lobes which are visible from beneath, and a small MPT would place ‘Mt Missim’ elsewhere. Close inspection leaf like lobe on their outer margins and aedeagal sheath of the character mapping on the preferred MPT (Figs. 40, broad and symmetrical. Both patterns are repeated in 41) shows that the undescribed species, ‘Mt. Missim’, is Luciola satoi Jeng et Yang, L. terminalis Olivier, and L. highly apomorphic, having many characters that are not praeusta Kiesenwetter (Jeng Yang & Lai, 2003). shared with any other taxon. Unlike our previous analyses (Ballantyne & Lambkin, 2000, 2001) the successively 2. Species of four existing genera: Pteroptyx (4 species), weighted semi-strict consensus (Fig. 39) suggests that ‘Mt Pyrophanes (1), Australian Luciola (4), and Colophotia Missim’ is more closely related to ‘Sisiak’ and the sub-genus praeusta. The aedeagus is elongate and slender, with the Pygatyphella. The placement of ‘Mt Missim’ remains apices of lateral lobes narrow and not visible from below, unresolved. and variously shortened. The aedeagal sheath is long , narrow and symmetrical. Ballantyne (1987a,b) The position of Curtos costipennis is also problematical, as distinguished 2 subgroups of Pteroptyx, (which are split

43 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

here), based on the presence/absence of lateral strong evidence for convergent evolution of similar coloration protuberances on the aedeagal sheath and the metafemoral in unrelated species producing a misleading signal. comb. For these reasons the six characters (97-100, 114-115) that 3. Atyphella Olliff, distinguished by Ballantyne & Lambkin describe colour, and four characters (101-2, 116-7) that (2000). The aedeagus has lateral lobes visible from describe mimicry or colour patterns were not used in the beneath and median lobe slightly longer than lateral lobes. phylogenetic analysis but mapped onto the preferred MPT The aedeagal sheath is rather broad, with the posterior (Figs. 40 - 41). The characters that describe mimicry or colour half asymmetrical. [A similar pattern of ‘exposed’ lateral patterns are clearly delineated by a box surrounding them on lobes, and an asymmetrical posterior half of the sheath the preferred MPT (Figs. 40 - 41). ventrite occurs in several aquatic Luciola from China (Jeng Lai & Yang 2003)]. In this analysis certain elements Most of the characters (97-100, 101-2, 114-115, 116-7) that are unresolved (three New Guinean species and A. costata describe colour or mimicry are apomorphic, and therefore from Australia) while ‘Sisiak’ groups with three Atyphella only found on branches leading to particular taxa (Figs. 40 – species distinguished by Ballantyne (1968) as Luciola 41). None of the colour or mimicry characters optimized on (Pygatyphella). Further analyses should include certain the preferred MPT is unique. Chinese and Japanese fireflies. The ‘Sisiak’-Pygatyphella clade is homoplasiously supported 4. Pygoluciola Wittmer. The aedeagal sheath has the by characters 102, 114, 117; Mt Missim-Pygoluciola clade anterior portion of the ventrite very narrow, the posterior by characters 98, 115; L. italica-L. cowleyi clade by character half very broad and symmetrical, a pattern not yet seen 115, and L. australis-Pyrophanes beccarii clade by characters elsewhere. 101-2 (see further discussion above).

Certain elements are still unresolved and should be addressed Few internal nodes are substantiated by colour or mimicry again – ‘Mt Missim’ (based on 4 specimens) is an undescribed characters, indicating that phylogenetically, there is little species from high elevation in New Guinea. The aedeagus grouping information and a great deal of homoplasy in these is similar to that of Atyphella, differing in the very narrow characters. Again there is evidence of convergent evolution median lobe which curves at its apex. ‘Mt Missim’ differs of similar coloration in unrelated species. from all other Luciolinae in lacking a clypeolabral suture. ‘Sisiak’ is based on undescribed New Guinean specimens, Methods of preservation and interpretation problems where the aedeagal sheath and aedeagus differ from that of Fireflies are soft bodied insects much given to distortion once Atyphella but repeated phylogenetic analyses still place it pinned. Collecting at least some into ethanol facilitates within Atyphella. dissection and manipulation (Ballantyne, 1987b). However, problems occur with colour interpretation (Ballantyne, 1987a, The problem of colour 2001), distortion of shape, determination of the extent of the Although morphological characters, especially those of the light organ, especially where the specimen was pinned after male terminal abdomen, have been used for grouping in the first immersion in ethanol, colour fading of larvae, and Luciolinae, early taxonomists, especially Ernest Olivier (see hardening of specimens preserved for a long period in ethanol McDermott, 1966), made much use of colour in describing (Ballantyne & Lambkin, 2000). new species, and colour still plays an important part. Species of are recognised by combinations of colour and Further problems arise here with ‘wet’ preserved specimens. pattern providing taxonomists with a means of identification The degree of explanation of the lateral pronotal margins, and categorisation (Brush, 1978). and determination of states for character 9 is more difficult to determine. The lateral lobes of the aedeagus in P. guigliae Colour is not included in many cladistic analyses as it is both are white and ‘plump’ (as wide as high) in the freshly difficult to quantify, and has traditionally been considered a preserved specimens, but when dehydrated appear character of little phylogenetic value. The poor regard for dorsoventrally flattened and slightly brown, as in the very colour characters in phylogenetic analysis stems, at least similar P. stylifer, suggesting they may have owed their partially, from the belief that colour is easily affected by contour to body fluids. Ballantyne (1968) used such features convergent evolution. Selection for crypsis in particular to help distinguish the aedeagi of P. guigliae and P. hamulata. habitat types leads to the convergent evolution of similar colours and colour patterns in unrelated species (Cott, 1957; A small hook on the inner margins of the lateral lobes in P. Oxford & Gillespie, 1998). Such phenotypic similarity reflects stylifer may be a result of dehydration; in the very similar, similar selection pressures, not shared ancestry, and in ‘wet’ preserved P guigliae, the same area is not pointed or phylogenetic analysis gives rise to homoplasy (Givnish & acutely developed, and is an area of heavier sclerotisation Sytsma, 1998) that can distort the inference of relationships. extending from the junction of the lateral lobes and running Previous quantitative phylogenetic studies (unpublished, ventrally to the ventral side of these lobes. Lambkin, 2001) combining a data set of over 200 colour characters with over 300 morphological characters in the The close approximation of the apices of abdominal tergite Exoprosopini (Diptera: Bombyliidae: Anthracinae) found 8 and ventrite 7 in dried specimens of Pygoluciola is not seen

44 THE RAFFLES BULLETIN OF ZOOLOGY 2006 in ‘wet’ specimens, where they may splay apart (see for it shows externally, as there is nothing about the female Functional Morphology below). Presumed distortions of the reproductive system to indicate any reinforcement against (pinned) female abdomen (seen in P kinabalua between the outside pressure. Any external pressure on the female bases of V7 and 8) are not seen in ‘wet’ specimens, and no abdomen of P. guigliae (which lacks any obvious external internal developments correspond with this apparent artefact, modifications), if such there is, would be exerted between which probably reflects, indirectly, the muscle attachments the elevations on V 7 and on the anterior half of V8 where it onto the anterior prolongation of ventrite 8 and the anterior projects under V7. This coincides internally at least with the portions of the valvifers (Ballantyne & Lambkin, 2001). attachment of muscles from the valvifers of the female genitalia to the anterior prolongation of V8, and above this, Female morphology and identification the hooks in the female reproductive system. These hooks Lucioline firefly taxonomy is male based until now, and in the duct, attached only at their bases, may be under the association of the sexes without taking a mating pair was, female’s control, and if erected they would probably prevent and in most cases still is, difficult (Ballantyne & McLean, any backward movement of a spermatophore once it had been 1970; Ballantyne, 1987a). Ballantyne (1988) associated inserted, particularly if pressure were exerted from outside females by taking them on the wing, flying with males where (LB squeezed the bursa accidentally during a dissection and no other species occurred. Ballantyne (1987b) described found that the spermatophore only moved forward). features of the female, none of which taken in isolation would have allowed reliable identification to species. Ballantyne However this would seem to be a female modification that & Lambkin (2000, 2001) advocated a more pragmatic on first glance, benefits the male. LB acknowledges Prof approach and made tentative identifications based on James Lloyd for the following suggestions that address the similarity of label data. possible function of the hooks in the female vagina: 1. If the copulating male sometimes removes ejaculate after depositing External abdominal modifications, seen in two species of sperm, because it is the presence of nutritive contributions Pygoluciola, fortuitously allow female identification. that persuade the female to copulate with him, he might However their actual function is unknown. The presence of reclaim and reuse the package to gain additional copulations hooks in the bursa of two species of female Pygoluciola, and with other females; 2. Males may remove sperm previously the existence of plates in the bursa of Pteroptyx valida female inserted by other males; if the female has invested time etc suggests the usefulness of these as taxonomic characters, and in selecting or waiting for a male and has accepted sperm an investigation of such plates in female Luciolinae is under from him, and then later is forced into copulation with another way (Ballantyne, unpub.). male, such hooks could reduce the chance that the second male could remove the spermatophore, thus negating her Functional morphology previous decision; 3. If another male tries to push or pry a In Pteroptyx valida, the female abdomen is held in a pincer copulating male away from a female, the intruder will have fashion between the deflexed elytral apices and the MPP of more difficulty in prising away the coupled male if both the the male ventrite 7 (‘copulation clamp’). No obvious external male and the female have a firm grasp on the spermatophore. morphology of the female abdomen hints at such an arrangement, but the valida female has cuticular plates in the The pronotal irregularities are seen in both sexes except for bursa that may prevent damage to the female during mating the single ‘wet’ preserved female of P. wittmeri. They may (Wing et al., 1983). They are absent in the Pygoluciola be a cluster of sense organs (no specimens were available for females dissected. EM study). Curved tibiae of all legs are seen in both sexes of one species. In certain Pyrophanes males, curved tibiae Ballantyne & Lambkin (2001) suggested a type of ‘copulation 3, and enlarged femora 3 are coupled with a metafemoral clamp could exist in P. kinabalua where the male abdomen comb, and taken together they may have some grasping can open posteriorly in a pincer like fashion, and may engage function (Ballantyne, 1987b: 174). The function of curved against the female, which has a transverse ridge on abdominal tibiae in Pygoluciola is unknown. ventrite 7, and a dorsal ‘mound’ on tergite 7. [Such splaying of the male terminal abdomen may nevertheless be nothing Ballantyne (1987b) suggested that the surface area of the more than a deterrent to another male trying to copulate with anterior prolongation of the aedeagal sheath ventrite, which an already coupled female (Ballantyne, 1987b); see also provides area for attachment of these muscles, could indicate Methods of Preservation above). That some clamping some of the complexity of movements the aedeagus can make, mechanism could occur in other Pygoluciola species is but in Pygoluciola the anterior prolongation of V9 although supported by two observations – first, that the male abdomen very narrow, has muscles from the base of the aedeagus in ‘wet’ preserved specimens described here shows a wide attaching all along its length, thus the width of this surface divergence between V7 and T 8. Secondly, the dorsoventral may not be a good indicator of the extent of this musculature. musculature is very strong, evidenced by the lateral depressions in abdominal tergite 7, the absence of light organ at the anterolateral corners of V7, where these muscles attach, ACKNOWLEDGEMENTS and where the sides of V7 may be very strongly sclerotised and ridge like. If some clamping mechanism exists, Ballantyne thanks Mahadimenakbar bin Mohamed Dawood (assuming a tail to tail mating position) the reinforcement for the opportunity to examine these specimens, Jenny

45 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Horsnall for invaluable help with assembling figures, and Farris, J. S., 1969. A successive approximations approach to Charles Sturt University for continued support. Lambkin character weighting. Systematic Zoology, 18: 374-385. thanks CSIRO Entomology and United States National Farris, J. S., 1989. The retention index and the rescaled consistency Science Foundation Partnerships Enhancing Expertise in index. Cladistics, 5: 417-419. Taxonomy program (DEB 9521825, 9977958)) for providing Farris J. S., 1990. Phenetics in camouflage. Cladistics, 6: 91-100. financial support. Givnish, T. & K. Sytsma, 1998. Homoplasy in molecular vs. morphological data: The likelihood of correct phylogenetic inference. In: Givnish, T. & K. Sytsma (eds.), Molecular LITERATURE CITED evolution and adaptive radiation. Cambridge University Press, New York. Pp. 55-101. Ballantyne, L. A., 1968. Revisional Studies of Australian and Haneda, Y., 1966. Synchronous Flashing of Fireflies in New Guinea. Indomalayan Luciolini (Coleoptera, Lampyridae, Luciolinae). Science Report of the Yokosuka City Museum, No. 12: 4-10. University of Queensland Papers, Department of Entomology, University of Queensland Press, II(6): 103-139. Jeng, M-L, J. Lai & P-S Yang, 2003. Lampyridae: a synopsis of aquatic fireflies with description of a new species. In: Jäch M. Ballantyne, L. A., 1987a. Further revisional studies on the firefly A. & L. Ji (eds.), Water Beetles of China. Volume 3. genus Pteroptyx Olivier (Coleoptera: Lampyridae: Luciolinae: Zoologische-Botanische Gesellschaft in Österreich and Weiner Luciolini). Transactions of the American Entomological Society, Coleopterologenverein, Wien. Pp. 539-562. 113: 117-170. Jeng, M.-L., P.-S. Yang, M. Satô, J. Lai, & J.-C. Chang, 1998. The Ballantyne, L. A., 1987b. Lucioline Morphology, Taxonomy and Genus Curtos (Coleoptera, Lampyridae, Luciolinae) of Taiwan Behaviour: A Reappraisal. (Coleoptera, Lampyridae). and Japan. Japanese Journal of Systematic Entomology, 4(2): Transactions of the American Entomological Society, 113: 171- 331-347. 188. Jeng, M.-L., P.-S. Yang, & J. Lai, 2003. Notes on the genus Luciola Ballantyne, L. A., 1988. The identities of Luciola australis (F.) (Coleoptera, Lampyridae, Luciolinae) of Taiwan. Special and L. guerini LaPorte (Coleoptera: Lampyridae). Journal of Bulletin of the Japanese Society of Coleopterologists, Tokyo, 6: the Australian Entomological Society, 27: 161-165. 247-262. Ballantyne, L. A., 2001. The bent winged fireflies of , Kawashima, I., H. Suzuki, & M. Satô, 2003. A Check-List of , Malaysia, Philippines and Thailand (Coleoptera: Japanese fireflies (Coleoptera, Lampyridae and Lampyridae: Luciolinae: Luciolini). Pteroptyx spp. of the Rhagophthalmidae). Japanese Journal of Systematic Polunin collection. Serangga, 6(1): 51-95. Entomology, 9(2): 241 – 261. Ballantyne, L. A. & C. Lambkin, 2000. The Lampyridae of Australia Kallersjö, M., J. S. Farris & V. A. Albert, 1999. Homoplasy (Coleoptera: Lampyridae: Luciolinae: Luciolini). Memoirs of increases phylogenetic structure. Cladistics, 15: 91-93. the Queensland Museum, 46(1): 15-94. Kallersjö, M., J. S. Farris, A. G. Kluge & C. Bult, 1992. Skewness Ballantyne, L. A. & M. R. McLean, 1970. Revisional studies on and permutation. Cladistics 8: 275-287. the firefly genus Pteroptyx Olivier (Coleoptera : Lampyridae : Luciolinae : Luciolini). Transactions of the American Kluge, A. & J. S. Farris, 1969. Quantitative phyletics and the Entomological Society, 96: 223-305. evolution of anurans. Systematic Zoology, 18: 1-32. Ballantyne, L. A. & C. Lambkin, 2001. A new firefly, Luciola Lambkin, C. L., 2001. Systematics and evolution of Australian (Pygoluciola) kinabalua, new species (Coleoptera: Lampyridae), exoprosopine (Diptera: Bombyliidae: Anthracinae) based from Malaysia, with observations on a possible copulation on molecules, morphology, and colour patterns. Ph.D. Thesis, clamp. Raffles Bulletin of Zoology, 49(2): 363-377. University of Queensland. Branham, M. A. & J. W. Wenzel, 2001. The evolution of Larson, A. & W. W. Dimmick, 1993. Phylogenetic relationships of bioluminescence in cantharoids (Coleoptera: Elateroidea). the salamander families: An analysis of congruence among Florida Entomologist, 84 (4): 565-586. morphological and molecular characters. Herpetological Monographs, 7: 77-93. Bremer, K., 1988. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42: 795- Lloyd, J. E., 1973. Fireflies of Melanesia: Bioluminescence, mating 803. behavior and synchronous flashing. Environmental Entomologist, 2(6): 991-1008. Bremer, K., 1990. Combinable component consensus. Cladistics, 6: 369-372. Maddison, D. R. & W. P. Maddison, 2001. MacClade Version 4.03. Sinauer Associates, Inc., Sunderland M. A. Bremer, K., 1994. Branch support and tree stability. Cladistics, 10: 295-304. Maddison, W., M. Donoghue & D. Maddison, 1984. Outgroup analysis and parsimony. Systematic Zoology, 33: 83-103. Brush, A., 1978. Avian Pigmentation. In: Brush, A. (ed.), Chemical Zoology. Volume 10. Aves. Academic Press, New York. Pp. Margush, T. & F. McMorris, 1981. Consensus n-trees. Bulletin of 141-163. Mathematical Biology, 43: 239-244. Carpenter, J. M., 1988. Choosing among multiple equally McDermott, F. A., 1966. Lampyridae (Luciolini, 98-118). In: Steel, parsimonious cladograms. Cladistics, 4: 291-296. W.O. (ed.) Coleopterorum Catalogus Supplementa. Volume 9 (Second Edition). W. Junk, Gravenhage. 149 pp. Cott, H., 1957. Adaptive Coloration in Animals. Methuen & Co., London. Meier, R., 1994. On the inappropriateness of presence/absence recoding for non-additive multistate characters in computerised Endler, J., 1978. A predator’s view of animal color patterns. In: cladistic analysis. Zoologischer Anzeiger, 232: 201-212. Hecht, M., W. Steere & B. Wallace (eds.), Evolutionary Biology. Volume 11. Plenum Press, New York. Pp. 319-364.

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Merilaita, S., 1998. Crypsis through disruptive evolution in an Oxford, G. & R. G. Gillespie, 1998. Evolution and ecology of spider isopod. Proceedings of the Royal Society of London. Series B. coloration. Annual Review of Entomology, 43: 619-643. Biological Sciences, 265: 1059-1064. Schuh, T. & J. S. Farris, 1981. Methods of investigating taxonomic Nixon, K. C. & J. M. Carpenter, 1993. On outgroups. Cladistics, congruence and their application in the Leptopodomorpha. 9: 413-426. Systematic Zoology, 30: 331-350. Olivier, E., 1885. Catalogue des Lampyrides faisant partie des Sorenson, M.D., 1999. TreeRot Version 2. (Sorenson, M.D., Boston collections du Musée Civique de Gênes. Annali del Museo University, Boston, MA.) Civico di Storia Naturale di Genova (2a)2: 333-374. Swofford, D., 2002. PAUP*: Phylogenetic Analysis Using Olivier, E., 1902. Catalogue des espèces de ‘Luciola’ et genres Parsimony (*and Other Methods). Version 4.0b.10. (Sinauer voisins décrits jusqu’à ce jour. Revue Scientfique du Associates, Sunderland, Massachusetts.) Bourbonnais et du centre de la France, 15: 69-88. Wing, S. R., J. E. Lloyd & T. Hongtrakul, 1983. Male competition Olivier, E., 1907. Coleoptera, Family Lampyridae (Luciolinae: 48- in Pteroptyx fireflies: Wing-cover clamps, female anatomy, and 56). In: Wytsman, P. (ed.), Genera Insectorum Bruxelles. P. mating plugs. Florida Entomologist, 66(1): 86-91. Wytsman Fasc. 53. 74 pp. Wittmer, W., 1939. Fauna Buruana. Coleoptera 7. Beitrag zur Olivier, E., 1913. The Lampyridae of Borneo. Sarawak Museum Kenntnis der indo-malayischen Malacodermata. Treubia, 17: Journal, 8: 55-60. 21-24 (Lampyridae).

47 Ballantyne & Lambkin: Phylogenetic reassessment of Pygoluciola Wittmer

Table 1. Data matrix describing 125 characters for 49 taxa, including polymorphic coding.

48 THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 49-57 Date of Publication: 28 Feb.2006 © National University of Singapore

THE GENUS PARAMEDETERA (DIPTERA: ) IN SINGAPORE, WITH A KEY TO THE ORIENTAL SPECIES

Patrick Grootaert Department of Entomology, Royal Belgian Institute of Natural Sciences, Vautierstreet 29, B-1000 Brussels, Belgium Temporary address: Raffles Museum of Biodiversity Research, Science Drive 2, National University of Singapore, Singapore 117600 Email: [email protected]

ABSTRACT. – The genus Paramedetera Grootaert & Meuffels, 1997 is reported for the first time from Singapore with five species. P. ankarum Grootaert & Meuffels, described from South Thailand, is recorded for the first time since its description. Four species new to science are described and illustrated: P. micropyga, new species, P. obscura, new species, P. chelata, new species and P. digitata, new species. A key is given for the thirteen Oriental species of the genus.

KEY WORDS. – Diptera, Dolichopodidae, Oriental, new species, Singapore.

INTRODUCTION At present fifteen species of Paramedetera Grootaert & Meuffels are known in the world, including the four new Paramedetera Grootaert & Meuffels, 1997 is a genus of species described here. P. sierraleonensis Grichanov, 1999 small flies that strongly resembles the cosmopolitan and very is the only Aftrotropical species up to now. All other species common genus Medetera. For a detailed generic diagnosis have been recorded from the Oriental or Australasian region. we refer to Grootaert & Meuffels (1997). Paramedetera has Despite intensive collections in the northern coastal plain of an ovoid third antennal segment with a latero-subapical arista. , only a single species, P. papuensis The ocellar tubercle is prominent and the long vertical bristles Grootaert & Meuffels, was found. The area of radiation of are inserted on an elevation. They possess four dorsocentral the genus seems to be with eleven species bristles of subequal size on the scutum. Rarely a very short recorded now, including the four new species described from anterior bristle is present that could make up for a fifth Singapore in the present paper. The number of records of the dorsocentral. The main characteristics to distinguish species and the number of new species found, indicate that Paramedetera from most Medetera are the straight veins there must be a lot more species to be described in Southeast r4+5 and M that run parallel to the wing tip. Paramedetera Asia The genus is also present in Central and South China species possess very few external characters to distinguish where two species have been described from the Henan and the species. There are slight differences in the colouration Guangxi provinces respectively: P. medialis Yang & Saigusa, of the ocellar, vertical and dorsocentral bristles, but the colour 2000 and P. jinxiuensis Yang & Saigusa, 2001. In addition ranges from black to pale brown or yellowish brown, we have seen at least two more species from Guangdong depending mainly on the incidence of the light. The halters province and Xishuangbanna in the south of the Yunnan can be whitish, pale greyish or occasionally brownish black. province. The females may have a specific melanization of the tergites. The basal tergites I, II or III may have unmelanized, triangular areas on both sides where the tergite is yellowish MATERIAL AND METHODS in contrast to the rest of the black tergites. These patterns are specific, but unfortunately only few females have been All specimens have been collected by sweep netting. They associated with males. A preparation of the male genitalia is are stored in 75% alcohol. A register number (e.g. 23086) generally necessary to identify the species. The shape of the refers to a record of the species in a database containing all aedeagus is complex and composed of a central tube with collection data. The numbers preceded by Si (e.g. Si303) refer lateral wings. These wings can bear small to huge prongs to the position of the tube in the collection. All holotypes and can have a rounded or pointed tip. In lateral view it may designated here are deposited in the Zoological Reference look like a ’s head. The shape of the hypandrium, if Collection at the Raffles Museum of Biodiversity Research, present, is diagnostic as is the position of the epandrial setae. Singapore (ZRC). Voucher specimens are conserved at the

49 Grootaert: Paramedetera of Singapore

Royal Belgian Institute of Natural Sciences, Brussels, Paratypes – 1 male, Nee Soon, 25 Nov.2003 (23086; coll. P. Belgium (RBINS). Grootaert, Si201); 1 male, Nee Soon, 3 Dec.2003 (23102, coll. P. Grootaert, Si218; coll. RBINS). The genitalia are figured with the ancestral ventral part (e.g. aedeagus) side up and their dorsal side (cerci) down. Description. – Male. Body length: 1.60 mm; wing length: 1.50 mm. Ocellar and vertical bristles pale yellowish. Palp The epandrium bears at both sides of its base, three epandrial black with a black apical bristle. Four dorsocentrals, preceded setae. They are grouped in a pair of lateral setae and a single by a short bristle (about a fifth length of anterior dc), 2 anterior medial seta. The outermost seta of the lateral pair is generally dc shorter than posterior 2. All bristles on scutum brownish. the largest. The epandrial setae can be inserted on a tubercle Scutellars pale brown. Propleural bristle long, pale. Halter or papilla that is either short or quite long. greyish-brown. Squama with a brown border, bearing 6 pale yellowish cilia.

SYSTEMATIC ACCOUNT Legs yellowish-brown, but all coxae and femora brown. Tibiae and tarsi paler, yellowish, 5th tarsomeres of all legs Paramedetera micropyga, new species dark brown. (Figs. 1-3) LI: Length of femur, tibia and tarsomeres (in mm): 0.42 : 0.40 : 0.21 : 0.12 : 0.08 : 0.06 : 0.07. Material examined. – Holotype - male: SINGAPORE: Nee Soon, LII: Length of femur, tibia and tarsomeres (in mm): 0.52 : 4 Dec.2002, swamp forest (22054, coll. P. Grootaert, Si303; ZRC). 0.55 : 0.31 : 0.17 : 0.12 : 0.07 : 0.06.

Figs. 1-3. Paramedetera micropyga, new species: 1, hypopygium ventral view; 2, aedeagus, ventral view; 3, hypopygium lateral view. ae: aedeagus; c: cercus; hy: hypandrium; les: lateral epandrial setae; mes: median epandrial seta.

50 THE RAFFLES BULLETIN OF ZOOLOGY 2006

LIII: Length of femur, tibia and tarsomeres (in mm): 0.56 : Paramedetera ankarum Grootaert & Meuffels, 1998 0.56 : 0.21 : 0.22 : 0.13 : 0.07 : 0.06. (Figs. 4-5)

Hypopygium (Figs. 1, 3). Aedeagus (Fig. 2) rather small, Material examined. – 1 male: SINGAPORE: Kent Ridge, Science hyaline, with a bird’s headlike tip, bearing a sharply pointed Park, 22 Nov.2003 (23082, coll. P. Grootaert, Si103; coll. ZRC). tip. Hypandrium triangular, a little erect, with a elongated tip. Median epandrial bristle the longest, on a long tubercle. Description. – Male. Body length: 1.25 mm; wing length: Lateral pairs of epandrial setae also on a tubercle. 1.38 mm. Ocellar and vertical bristles pale yellowish-brown. Palp black with a black apical bristle. Four dorsocentrals, 2 Female unknown. anterior dc shorter than posterior 2. All bristles on scutum yellowish brown. Scutellars pale brown. Propleural bristle long, pale. Halter white. Squama pale, with a brown border, Etymology. – The name micropyge refers to the rather small bearing 4 pale cilia. aedeagus. Legs paler than in the other species. Fore coxa anteriorly Differential diagnosis. – The aedeagus is rather small and slightly infuscated, posterior coxae completely yellowish the tips of the aedeagal wings are pointed in lateral view. brown. Femora only infuscated dorsally; tibiae and tarsi The inner rim of the wings does not bear a tooth-like yellow; 4th and 5th tarsomeres of all legs brown. projection, that distinguishes it from P. jixiuensis and P. LI: Length of femur, tibia and tarsomeres (in mm): 0.42 : borneensis. 0.40 : 0.21 : 0.11 : 0.08 : 0.06 : 0.07.

Figs. 4-5. Paramedetera ankarum Grootaert & Meuffels: 4, hypopygium ventral view; 5, hypopygium lateral view.

51 Grootaert: Paramedetera of Singapore

LII: Length of femur, tibia and tarsomeres (in mm): 0.49 : small fish), 23 April 2005 (25084, coll. P. Grootaert, Si742). 57 0.52 : 0.40 : 0.14 : 0.11 : 0.07 : 0.06. males, 34 females, same locality, 25 June.2005 (25189, coll. P. Grootaert, Si849, Si850; coll. RBINS). LIII: Length of femur, tibia and tarsomeres (in mm): 0.49 : 0.56 : 0.17 : 0.21 : 0.11 : 0.07: 0.06. Description. – Male. Body length: 1.55-1.60 mm; wing length: 1.48-1.50 mm. Ocellar and vertical bristles yellowish Hypopygium (Figs. 4-5): Wings of aedeagus with a bird’s brown. Palp black with a black apical bristle. Four headlike tip. Inner rim of the wings smooth. Hypandrium not dorsocentrals, 2 anterior dc shorter than posterior 2. All distinct. Epandrial setae inserted on short tubercle. bristles on scutum yellowish brown. Propleural bristle long, pale. Halter white. Squama pale, with a brown border, bearing Female unknown. 6 pale brown cilia.

Legs yellowish brown. Fore coxa brown anteriorly, posterior Paramedetera obscura, new species 4 coxae brown. Femora brown dorsally and yellow ventrally; (Figs. 6-7) tibiae and tarsi yellow; 4th and 5th tarsomeres of all legs brown. LI: Length of femur, tibia and tarsomeres (in mm): 0.54 : Material examined. – Holotype - male: SINGAPORE: Clementi 0.49 : 0.41 : 0.14 : 0.11 : 0.07 : 0.08. woods, 15 Mar.2005, drain (25014, coll. P. Grootaert, Si446; ZRC). LII: Length of femur, tibia and tarsomeres (in mm): 0.50 : 0.56 : 0.35 : 0.18 : 0.13 : 0.06 : 0.06. Paratypes – 4 males, 1 female, same data as holotype. 30 males, 28 LIII: Length of femur, tibia and tarsomeres (in mm): 0.56 : females, Clementi woods, major drain (streaming water containing 0.63 : 0.22 : 0.27 : 0.14 : 0.07 : 0.06.

Figs. 6-7. Paramedetera obscura, new species: 6, hypopygium ventral view; 7, hypopygium lateral view.

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Aedeagus almost black, with pointed apex in lateral view (Fig. Paramedetera chelata, new species 7). Hypandrium not distinct, represented by the broadly (Figs. 8-10) rounded margin of epandrium (Fig. 6). Epandrial setae inserted on short tubercle. Material examined. – Holotype - male: SINGAPORE: Nee Soon, 18 Mar.2005 (25018, coll. P. Grootaert, Si559; ZRC). Female. Body length: 1.87 mm; wing length: 1.75 mm. Larger than male. First tergite on both sides with a pale, narrow Paratype – 1 male, same provenance as holotype (coll. RBINS). triangular area at basal margin and a wider pale, triangular Description. – Male. Body length: 1.50 mm; wing length: area at apical margin. Tergite II with a very small, pale 1.45 mm. Ocellar and vertical bristles black. Palpi black with triangular at both sides of apical margin. Tergite III with a a black apical bristle. Four black dorsocentrals, 2 anterior dc large pale triangular area at both sides of apical margin. shorter than posterior 2. Propleural bristle long, pale. Halter black. Squama brown, with a brown border, bearing 6 pale Etymology. – The name obscura refers to the darkened, brown cilia. almost black aedeagus. Legs yellowish-brown. Fore coxa brown anteriorly, posterior Differential diagnosis. – It is so far the only species in the 4 coxae brown. Femora dorsally brown, ventrally yellow; genus that has a black aedeagus. However it should not be tibiae and tarsi yellow; 4th and 5th tarsomeres of fore and mid confused with the huge black prongs on the aedeagus of P. legs brown, apical 3 tarsomeres of hind legs brown. horrorifera.

Figs. 8-10. Paramedetera chelata, new species: 8, hypopygium ventral view; 9, tip of dorsal surstylus, cercus dorsally; 10, hypopygium lateral view.

53 Grootaert: Paramedetera of Singapore

LI: Length of femur, tibia and tarsomeres (in mm): 0.42 : its huge sharp prongs on the aedeagus. P. chelata new species 0.38 : 0.24 : 0.10 : 0.07 : 0.06 : 0.06. is a darker species than the others, with all bristling black LII: Length of femur, tibia and tarsomeres (in mm): 0.42 : and even black halters. 0.49 : 0.28 : 0.14 : 0.11 : 0.07 : 0.06. LIII: Length of femur, tibia and tarsomeres (in mm): 0.49 : 0.52 : 0.15 : 0.21 : 0.1 : 0.07 : 0.07. Paramedetera digitata, new species (Figs. 11-12) Hypopygium (Figs. 8-10). Aedaegus orange brown, with tip of wings rounded. Inner rim of the wings in ventral view, Material examined. – Holotype - male: SINGAPORE: Nee Soon, with two claw-like projections: one at tip and one near middle. 9 Mar.2005, swamp forest (25004, coll. P. Grootaert, Si384; ZRC). Hypandrium not distinct, represented by a small rounded projection. Median epandrial seta not on a tubercle. Description. – Male. Body length: 1.50 mm; wing length: 1.25 mm. Ocellar and vertical bristles brown. Palpi black with Female unknown. a black apical bristle. Four dorsocentrals, anterior 2 shorter than posterior 2. All bristles on scutum dark brown. Propleural bristle long, pale brown. Halter greyish-brown. Squama Etymology. – The name chelata refers to the large claw-like brown with long pale cilia. Legs yellowish-brown, but all projections on the inner rim of the wings of the aedeagus. coxae and femora brown. Tibiae and tarsi yellowish, 4th and 5th tarsomeres of all legs dark brown. Differential diagnosis. – The strong claw-like projections on the inner rim of the wings of the aedeagus are specific. LI: Length of femur, tibia and tarsomeres (in mm): 0.35 : Similar projections, but much smaller, are present in P. 0.35 : 0.19 : 0.07 : 0.06 : 0.04 : 0.06. sumatrensis Grootaert & Meuffels. The species P. jinxiuensis LII: Length of femur, tibia and tarsomeres (in mm): 0.38 : Yang & Saigusa and P. borneensis Grootaert & Meuffels have 0.42 : 0.21 : 0.10 : 0.07 : 0.04 : 0.06. also small projections on the inner rim of the wings. The LIII: Length of femur, tibia and tarsomeres (in mm): 0.42 : extreme is found in P. horrorifera Grootaert & Meuffels with 0.42 : 0.12 : 0.15 : 0.07 : 0.06 : 0.06.

Figs. 11-12. Paramedetera digitata, new species: 11, hypopygium ventral view; 12, hypopygium lateral view.

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Hypopygium (Figs. 11-12) rather elongate. Aedeagus orange, 8. Hypandrium absent (Fig. 20). Median epandrial setae on a short with a downwards pointed tip. Hypandrium triangular, a little tubercle (Papua New Guinea) ...... P. papuensis sunk between the rounded lobes of the epandrium that bears – Hypandrium indistinct (triangular) (Fig. 19). Median epandrial seta not on a tubercle; tip of wings on aedeagus pointed the basal epandrial seta. Epandrial setae all short, none on a downwards (Sumatra) ...... P. orientalis tubercle. Lateral to the epandrial setae is a black finger-like 9. Tip of aedeagus in lateral view like a bird’s head: rounded above projection of the epandrium. Dorsal surstylus with a curved and with a beak-like projection below, either directed up or down tip, large and wide in lateral view (not straight and stalk-like (Figs. 21, 22, 23, 24) ...... 10 as in the other species). – Tip of aedeagus simply pointed (Figs. 25, 26), not like a bird’s head in lateral view ...... 13 Female unknown. 10. Tip of beak-like projection turned up (Fig. 21, 24) ...... 11 – Beak-like projection pointing down (Figs. 22, 23) ...... 12 11. Lateral epandrial setae not tuberculate (Fig. 21) (China, Guangxi) Etymology. – The name digitata refers to the finger-like ...... P. jinxiuensis projections on the epandrium dorsad of the epandrial setae. – Lateral epandrial setae tuberculate (Figs. 1, 24) (Singapore) .. This is so far a unique character in Paramedetera...... P. micropyga 12. Median epandrial seta not on a tubercle (Fig. 22) (Thailand, Differential diagnosis. – A finger-like projection lateral to Singapore) ...... P. ankarum the epandrial setae is not present in other Paramedetera – Median epandrial seta on a short tubercle. Inner border of species. aedeagal wings with a tooth-like projection (Fig. 23) (Borneo, )...... P. borneensis 13. Aedeagus orange, tip of aedeagal wings sharply pointed, directed downwards; hypandrium triangular, pointed; basal epandrial seta Key to male Paramedetera from the Oriental and on a rounded lobe; a finger-like projection at each side of the Australasian fauna base of epandrium (Figs. 11, 12, 25) (Singapore) ...... P. digitata – Aedeagus black (Figs. 6, 26); hypandrium indistinct, broadly Since there are very little non-genital characters of diagnostic rounded, no finger-like projection at base of epandrium value, the key is largely based on the shape of male genital (Singapore) ...... P. obscura appendages. It is conceived as such that when the hypopygium is detached from the abdomen, most species can be identified. For unambiguous identification, the hypopygium should be GENERAL DISCUSSION removed, macerated and mounted for microscopic study. I suggest using mainly the composite Figs. 13-26. Most of the genera of the subfamily Medeterinae are arboreal. Paramedetera however is soil dwelling and found near 1. Aedeagus with large pointed, black prongs directed inwards (Fig. aquatic habitats. It is observed near riverbanks of streams 13) (Thailand) ...... P. horrorifera with solid substrates such as pebbles, boulders or rocks. That – Aedeagus usually without prongs; when prongs present, then is probably why Paramedetera is found in the open concrete small and pale ...... 2 drains cutting through parks in Singapore. We assume that 2. Tips of wings of aedeagus rounded in lateral view (Figs. 14-20) the larvae are semi-aquatic, or live in the mosses and algae ...... 3 splashed by the streaming water. The record of a few P. – Tip of aedeagal wings sharply pointed in lateral view (Figs. 21- papuensis specimens near a water hole is the only exception 26) ...... 9 3. Inner edge of at least one aedeagal wing (Fig. 15) with one or of Paramedetera found near stagnant water, though the other two tooth-like projections (Figs. 14, 15, 16) ...... 4 P. papuensis were collected on pebbles in a streambed. – Inner edge of aedeagal wings smooth, without tooth-like Paramedetera is tropical in distribution. It is not reported projection (Figs. 17-20) ...... 6 from Australia, though it is likely to be present in the northern 4. Hypandrium distinct; inner rim of aedeagal wings with small, rain forest since it is present in the northern coastal lowland tooth-like projections ...... 5 of Papua New Guinea (Grootaert & Meuffels, 1997). It is – Hypandrium not distinct; inner rim of aedeagal wings with two, also not known from the Neotropical Realm. A single species large, well separated tooth-like projections (Figs. 8, 14) is recorded from the Afrotropical Realm indicating a (Singapore) ...... P. chelata Gondwana link. When looking to the distribution of 5. In ventral view: inner rim (border) of aedeagal wings each with Paramedetera in the world, it seems that the centre of a single tooth close to tip; median epandrial seta on a long radiation is situated in Southeast Asia with eleven species, tubercle (Fig. 15) (China, Henan) ...... P. medialis – In ventral view: inner rim (border) of aedeagal wings with not taking into account the two South China species. pointed tip and a tooth-like projection close to the tip; Currently five species are recorded in Singapore. Only one hypandrium large triangular, with truncate tip (Fig. 16) species was previously known (Thailand), the others are new (Indonesia, Sumatra) ...... P. sumatrensis for science. The species accumulation curve and the number 6. Lateral epandrial setae on distinct tubercle ...... 7 of singletons and doubletons (seven species on 11, or 60% – Lateral epandrial setae not on a tubercle /or on a small elevation of the Southeast Asian species) suggest that a large number only ...... 8 of species remains to be described. Malaysia, Indonesia and 7. No hypandrium (Fig. 17) (Borneo, Brunei) ….. P. bruneiensis the Philippines are largely unexplored and are therefore likely – Hypandrium very short, triangular (Fig. 18) (Thailand) ...... to have the bulk of the undescribed species...... P. turschi

55 Grootaert: Paramedetera of Singapore

ACKNOWLEDGEMENTS Paramedetera borneensis Grootaert & Meuffels, 1998: Bull. Annls. Soc. r. belge Ent. 133 [1997]: 376. Type locality: The author thanks Dr. Peter Ng, director of Raffles Museum Brunei, Kuala Belalong (HT in RBINS). Distribution: Borneo, for Biodiversity Research, for the kind hospitality during his Brunei. sabbatical stay at NUS. Mr. K. L.Yeo assisted in the field. Paramedetera bruneiensis Grootaert & Meuffels, 1998: Bull. The authorities of National Parks are also thanked for issuing Annls. Soc. r. belge Ent. 133 [1997]: 378. Type locality: the Research permits to collect in the National Parks and for Brunei, Kuala Belalong (HT in RBINS). Distribution: Borneo, their interest in the research. He finally acknowledges a grant Brunei. in the Action 4 programme of the Belgian Science Policy. Paramedetera chelata, new species. Type locality: Singapore Dr. Marc Pollet and Dr. Isabella Van de Velde gave many (HT in ZRC). Distribution: Singapore. useful comments on the manuscript. Paramedetera digitata, new species. Type locality: Singapore (HT in ZRC). Distribution: Singapore. Paramedetera horrorifera Grootaert & Meuffels, 1998: Bull. LITERATURE CITED Annls. Soc. r. belge Ent. 133 [1997]: 383. Type locality: Thailand, Phang-Nga (HT in RBINS). Distribution: Thailand. Grichanov, I. Ya., 1999. A brief review of the Afrotropical fauna of Paramedetera jinxiuensis Yang & Saigusa, 2001: Bull. Inst. the subfamily Medeterinae (Diptera: Dolichopodidae) with r. Sci. nat. Belg., Entom. 71: 160. Type locality: China, descriptions of a new genus and new species. Bulletin de Guanxi province, Jinxiu (HT in CAU). Distribution: South l’Institut Royal des Sciences Naturelles de Belgique, Entomologie, 69: 87-112. China. Paramedetera medialis Yang & Saigusa, 2000: Insects of the Grootaert, P. & H. J. G. Meuffels, 1997. Dolichopodidae (Diptera) Mountains Funiu and Dabie regions: 200. Type locality: from Papua New Guinea XIV. Paramedetera, a new genus in China, Henan prov., Luoshan (HT in CAU). Distribution: the Medeterinae. Invertebrate Taxonomy, 11: 309-319. South China. Grootaert, P. & H. J. G. Meuffels, 1998 (1997). New Paramedetera Paramedetera micropyga, new species. Type locality: Grootaert & Meuffels, 1997 from (Diptera, Singapore (HT in ZRC). Distribution: Singapore. Dolichopodidae, Medeterinae). Bulletin et Annales de la Société royale belge d’Entomologie, 133: 375-388. Paramedetera orientalis (Hollis, 1964) [Cyrturella] (Grootaert & Meuffels, 1997: Invert.Taxon. 11: 315). Type Yang, D. & T. Saigusa, 2000. New species of Dolichopodidae from locality: Indonesia, Sumatra, Bukit Tingi (Fort de Kock) (HT Henan. Insects of the Mountains Funiu and Dabie regions: 189- 209. in ITZ). Distribution: Indonesia, Sumatra. Paramedetera obscura, new species. Type locality: Yang, D. & T. Saigusa, 2001 New and little known species of Singapore (HT in ZRC). Distribution: Singapore. Dolichopodidae from China (VIII). Bulletin de l’Institut Royal Paramedetera papuensis Grootaert & Meuffels, 1997: des Sciences Naturelles de Belgique, Entomologie, 71: 155-164. Invert.Taxon. 11: 310. Papua New Guinea, Madang prov. Boroi (HT in RBINS). Distribution: Papua New Guinea. Paramedetera sierraleonensis Grichanov, 1999: Bull. Inst. Catalogue of the Paramedetera species from the world r. Sci. nat. Belg., Entom., 69: 103. Type locality: Sierra Leone, Freetown (HT in Lund). Distribution: Sierra Leone. Paramedetera [genus] Grootaert & Meuffels, 1997: Paramedetera sumatrensis Grootaert & Meuffels, 1997: Invert.Taxon. 11: 309 (in subf. Medeterinae). Type species: Invert.Taxon. 11: 312. Type locality: Indonesia, Sumatra, Paramedetera papuensis Grootaert & Meuffels, 1997 Padang, Air terjun tiga tinkat lubuk hitam (HT in RBINS). (original designation) Distribution: Indonesia, Sumatra. Paramedetera turschi Grootaert & Meuffels, 1998: Bull. Paramedetera ankarum Grootaert & Meuffels, 1998: Bull. Annls. Soc. r. belge Ent. 133 [1997]: 380. Type locality: Annls. Soc. r. belge Ent. 133 [1997]: 385. Type locality: Thailand, Phang-Nga (HT in RBINS). Distribution: Thailand. Thailand, Ranong prov., Bok Kai waterfalls (HT in RBINS). Distribution: Thailand, Singapore.

56 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Figs. 13-26. Ventral (upper) and lateral view (lower) of aedeagal wings and hypandrium of Oriental and Australasian Paramedetera. 13, horrorifera; 14, chelata; 15, medialis; 16, sumatrensis; 17, bruneiensis; 18, turschi; 19, orientalis; 20, papuensis; 21, jinxiuensis; 22, ankarum; 23, borneensis; 24, micropyga; 25, digitata; 26, obscura.

57

THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 59-82 Date of Publication: 28 Feb.2006 © National University of Singapore

THE GENUS TEUCHOPHORUS (DIPTERA: DOLICHOPODIDAE) IN SINGAPORE

Patrick Grootaert Department of Entomology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1000 Brussels, Belgium Temporary address: Raffles Museum of Biodiversity Research, Science Drive 2, National University of Singapore, Singapore 117600 Email: [email protected]

ABSTRACT. – Thirteen species of Teuchophorus (Dolichopodidae: ) are reported from Singapore. Nine species are new to science and are described and illustrated. Only one species from the T. conspicuus – notabilis group was found: T. ornatulus Meuffels & Grootaert. Three species belong to the T. gratiosus group (former genus Mastigomyia Becker) that is supposed to be arboricolous: T. temasek, new species, T. neesoonensis, new species, and T. antennatus, new species. Four species belong to the T. pauper group: T. singaporensis, new species, T. meieri, new species, T. spinulosus, new species, and T. acuminatus, new species. T. pauper from Thailand is re-examined and illustrated. Three species belong to the T. pectinatus group and all live here in mangrove: T. simplicissimus Meuffels & Grootaert, T. limosus, new species, T. bulohensis, new species. Two species are considered incertae sedis and not assigned to a species group: T. krabiensis Meuffels & Grootaert, a species from coastal sandy beaches and T. stenostigma Meuffels & Grootaert, a species from stream beds in forest. A key is given for all Singaporean species. In addition, some records of species from Malaysia are given and a mangrove species from Pulau Tioman (Malaysia) related to Singaporean mangrove species is described as well.

KEY WORDS. – Diptera, Dolichopodidae, Teuchophorus, new species, Singapore, mangrove, swamp forest.

INTRODUCTION has only three representatives in Singapore. They are all found in mangrove habitats, including T. tiomanensis that is In 2003, Meuffels & Grootaert published 40 new species of described here from Pulau Tioman. Four representatives of Teuchophorus from the Oriental and Australasian region. the T. pauper group were found. This group is characterised Despite this huge input of new species, still more new species by the absence of the acrostichals and all were found in acidic were found after intensive sampling in a quite small area swampy habitats. like Singapore. Evenhuis & Grootaert (2002) reported two unidentified species of Teuchophorus from Singapore, but The typical microhabitat of Teuchophorus, being muddy and in the present paper I raise that number to thirteen species. sandy patches along small streams in forests, are very hard Nine species are new to science while the other four were to find in Singapore. Nevertheless, the nature reserves in the described by Meuffels & Grootaert (2003). Teuchophorus Central Catchment Area hold nine species. Moreover, four ornatulus, originally described from Thailand, is the only species are recorded in marine habitats: T. krabiensis is found Southeast-Asian species that was found here in Singapore exclusively on sandy beaches while T. simplicissimus and that belongs to the T. conspicuus-notabilis group. This species two newly described, related species are very abundant in group can be considered as a Gondwana connection since it mangrove habitats. has many representatives in the Afrotropical region described under the genus name Olegonegrobovia Grichanov, 1995. Remarkable is the presence of three new species of the rare MATERIAL AND METHODS T. gratiosus group (former genus Mastigomyia Becker, 1924), characterised by a very long third antennal segment. They Material was collected by sweep netting (5-13 October 2000, were found in the same habitat i.e. swamp forest and are 4-13 December 2002, 19 November-11 December 2003 and apparently arboricolous. The T. pectinatus group (Meuffels March-July 2005) and with Malaise traps (November- & Grootaert, loc. cit.), which is very diverse in New Guinea, December 2003 and a 6-month period in 2005: March-August

59 Grootaert: Teuchophorus of Singapore

2005). The Malaise traps were operated at following sites in TAXONOMY Singapore: Bukit Timah (secondary rain forest at place of sampling), Sime forest (rain forest), Central Catchment Area Teuchophorus Loew, 1857 (swamp forest), Sungei Bulo (mangrove) and Pulau Ubin (mangrove). The genus Teuchophorus belongs to the subfamily Sympycninae and is very closely related to the Sympycnus – The material is stored in alcohol in the collections of the Chaetogonopteron complex. Teuchophorus represents small Raffles Museum for Biodiversity Research, Singapore (ZRC: flies of about 1.5-4 mm (diagnosis see Meuffels & Grootaert, Zoological Reference Collection) and the Royal Belgian 2003). The third antennal segment is generally triangular with Institute of Natural Sciences, Brussels (RBINS). For each a dorsal arista. However, in the T. gratiosus group (former record a register number is given (register number in Database genus Mastigomyia Becker) the third antennal segment can linked to collection data such as locality, date, collector, range from 3.5 times to 5.5 times as long as wide. In these habitat). In addition, an inventory number (e.g. Si200) species the arista can be longer than the third antennal indicates where the specimens can be found in the wet segment, shorter or even completely absent. Females of the collection. T. gratiosus group have short, triangular antennae.

The terminology used in the descriptions is that of Meuffels Chaetotaxy on the mesoscutum is stable and is as follows: & Grootaert (2003) in order to conserve continuity in the The acrostichals are generally uniseriate in Teuchophorus and descriptions. The following abbreviations are used: a: anterior; rarely absent. In the T. pauper group the acr are always absent. acr: acrostichal bristles; ad: anterodorsal; av: anteroventral; Some species of the T. gratiosus group also lack acrostichals. dc: dorsocentral bristles; pd: posterodorsal; pv: posteroventral; Four to five dorsocentrals; one humeral with a hair beside, tp: posterior cross vein. two notopleurals, one posthumeral, one presutural, one sutural, one intraalar, two supraalars; scutellum with two strong marginals. Morphology of the male genital apparatus The first tarsomere of the hind leg is always shortened. Vein The epandrium (hypopygium or genital capsule) is composed m1+2 is turned up immediately after the connection with the of a spherical to pear-shaped capsule bearing a number of cross vein tp and not presenting a wing “boss” at some appendages. Dorsal (original position, before the rotation and distance of the cross vein like in the Sympycnus – flexion) are a pair of cerci. Their structure is simple in Chaetogonopteron-complex. Males have often a stigma, Teuchophorus. Sometimes the apical hairs are much longer being a darkened swelling on the costa. It is present in most than the rest (Fig. 44 in T. tiomanensis) or there is a long Palaearctic species, but in Singapore only T. stenostigma has apical bristle (Figs. 7, 8 in T. temasek). Two pairs of surstyli a slight thickening and darkening of the costa. The other are present: a dorsal and a ventral pair. The dorsal pair species do not display that character. generally bears a strong dorsal bristle with a few minute hairs on its tip. The ventral surstylus is generally larger than the dorsal and often bears a pair of long bristles at its ventral Key to male Teuchophorus of Singapore base. These bristles should not be confused with epandrial bristles. There is no pair of lateral epandrial lobes in 1. Third antennal segment more than 2.5 times as long as wide . Teuchophorus like in other groups such as Diaphorus, ...... 2 Nepalomyia, Dolichopus and others. However, one or more – Third antennal segment at most 1.5 times as long as wide, or epandrial setae are sometimes present (Fig. 8 in T. temasek). shorter ...... 4 2. Arista longer than third antennal segment; third antennal segment nearly 3.5 times as long as wide (Fig. 6) ...... The aedeagal complex is composed of three structures ...... T. temasek, new species (pg. 62) forming a tube: the hypandrium, the aedeagus and a pair of – Arista shorter than third segment or even spine-like; third appendages on the epandrium. The hypandrium is generally antennal segment more than 3.5 times as long as wide (Figs. 9, a single plate fused at its base with the epandrium, but the 12) ...... 3 hypandrium can bear two arms. On the dorsal wall of the 3. Arista shorter than base of third antennal segment. Third antennal epandrium there are generally one or two lobes that we call segment 5.5 times a long as wide (Fig. 12). Acr uniseriate .... here the dorsal epandrial lobes. The aedeagus is generally a ...... T. antennatus, new species (pg. 65) simple tube lying in the middle of a tube formed by the dorsal – Arista well developed but shorter than length of third antennal segment (Fig. 9). Third antennal segment 4.5 times as long as epandrial lobes and the hypandrium. wide. No acr ...... T. neesoonensis, new species (pg. 64) 4. Acr present ...... 5 Illustrations: The lateral view of the genitalia is always the – No acr present ...... 11 left side of the , because sometimes the hypandrium is 5. Mid femur ventrally with a pair of very long bristles (Fig. 2). asymmetrical (e.g. in T. pauper, T. singaporensis and the T. Mid tibia ventrally near middle with a long flattened bristle .. conspicuus group) so that right and left side are different...... T. ornatulus Meuffels & Grootaert (pg. 61) – Mid femur ventrally without long bristles; mid tibia without flattened bristles ...... 6

60 THE RAFFLES BULLETIN OF ZOOLOGY 2006

6. Third antennal segment about 1.5 times as long as wide; hind 14. Right arm of hypandrium much longer than left arm and tibia with some strong dark bristles in the preapical comb (Fig. wrinkled laterally (Fig. 20); ventral surstylus ventrally indented, 49) (sandy beach) ...... bearing two strong bristles (Thailand)...... T. krabiensis Meuffels & Grootaert (pg. 78) ...... T. pauper Meuffels & Grootaert (pg. 68) – Third antennal segment at most 1.2 times as long a wide, or – Right arm of hypandrium almost as long as left arm; tip of right shorter; hind tibia without stronger bristles in the preapical comb arm smooth (Fig. 19); ventral surstylus ventrally not indented ...... 7 ...... T. singaporensis, new species (pg. 66) 7. Wing costa slightly thickened between R1 and R2+3, but not a real stigma (Fig. 51) ...... T. stenostigma Meuffels & Grootaert (pg. 79) Species records and descriptions – Costa with at most a darkening of the area between R1 and R2+3 ...... 8 Group I. T. conspicuus-notabilis group 8. Hind tibia posteriorly near base with about 5 erect hairs (mangrove) ...... The T. conspicuus-notabilis group has several representatives ...... T. simplicissimus Meuffels & Grootaert (pg. 73) in the Afrotropical region. They were described by Grichanov – Hind tibia posteriorly without such hairs ...... 9 (1995) in the genus Olegonegrobovia. Future genetic work 9. Tip of cercus darkened and with long bristles ...... might reveal if recognition of Olegonegrobovia indeed ...... T. tiomanensis, new species (pg. 75) – Tip of cercus not darkened and without long bristles ...... 10 renders Teuchophorus paraphyletic. This species group has 10. Fore leg with a basal ventral spinule on first tarsomere. Surstyli also diversified in Southeast Asia with T. ornatulus, T. short, truncate; cerci longer than surstyli (Fig. 47) ...... ornatuloides and T. vexillifer (Meuffels & Grootaert, 2003), ...... T. bulohensis, new species (pg. 77) but I have seen four more undescribed species (one from – No spinule on fore metatarsus. Aedeagal complex contrastingly Northeast Thailand, one from Cambodia and two from South- black and large (Fig. 39)..... T. limosus, new species (pg. 74) China). This species group can be seen as a Gondwana-link 11. Mid femur ventrally with about 4-5 short spine-like bristles (Fig. between Southeast Asia and Africa. It is also the only 26) ...... T. spinulosus, new species (pg. 71) “Teuchophorus” present in the Afrotropics. – Mid femur ventrally without or only minute bristles...... 12 12. Third antennal segment about 1.5 times as long as wide, with a fine elongated tip (Fig. 28) ...... Teuchophorus ornatulus Meuffels & Grootaert, 2003 ...... T. acuminatus, new species (pg. 72) (Figs. 1-4) – Third antennal segment shorter than wide ...... 13 13. Hypandrium symmetrical (Fig. 23). Prosternum yellow, with a Meuffels & Grootaert, 2003: 150, Figs. 2-6. central brown spot...... T. meieri, new species (pg. 69) – Hypandrium asymmetrical (Fig. 18). Prosternum entirely brown Material examined. – SINGAPORE: 6 males, 3 females, Sime with a central black stripe ...... 14

Figs. 1-4. Teuchophorus ornatulus Meuffels & Grootaert, male: 1, antenna; 2, mid leg; 3, hind leg; 4, hypopygium lateral view.

61 Grootaert: Teuchophorus of Singapore forest, 24 Mar.2005 (25026, coll. P. Grootaert, Si561); 9 males, 19 Teuchophorus temasek, new species females, Sime forest, 15 Apr.2005 (25061, coll. P. Grootaert, Si715); (Figs. 5-8) 30 males, 37 females, Sime forest, 2 Jul.2005 (25191, coll. P. Grootaert, Si865); 1 male, Sime forest, 7 Sept.2005 (25334, coll. P. Material examined. – Holotype male: SINGAPORE: Nee Soon, Grootaert, Si1112). Central Catchment Area (Upper Peirce Reservoir), acidic swamp forest, Malaise trap, 3 Dec.2003 (23104, coll. P. Grootaert, Si39). MALAYSIA: 9 males, 2 females, Johor, Gunung Belumut, 22 Oct.2005, sandy riverbanks in oil palm plantation (25396, coll. P. Paratypes: SINGAPORE: 1 male, Nee Soon, 16 Mar.2005, Malaise Grootaert, Si1096); 7 males, 1 female, Gunung Belumut, 22 trap 1 in swamp forest (25015, coll. P. Grootaert, Si459); 1 female, Oct.2005, sandy and rocky riverbanks in primary rain forest (25397, Nee Soon, 16 Mar.2005, Mal. 2 (25016, coll. P. Grootaert, Si481); coll. P. Grootaert, Si1101). 8 males, 2 females, Nee Soon, 16 Mar.2005, Mal. 3, (25017, coll. P. Grootaert, Si496); 1 male, Nee Soon, 12 Aug.2005 (25291, coll. Diagnosis. – A medium-sized metallic green species with a P. Grootaert, Si949); 1 male, 1 female, Nee Soon, 19 Aug.2005 short antenna (shorter than wide) and wing without stigma (25298, coll. P. Grootaert, Si951); 5 males, 5 females, Nee Soon, and without “flag” (group of black bristles on hind margin of 16 Sept.2005 (25350, coll. P. Grootaert, Si1110, Si1111). wing). Mid femur ventrally with two very long bristles near base (Fig. 2). Mid tibia with a long, flattened ventral bristle Diagnosis. – A species of the T. gratiosus group with third beyond middle. Hind femur with only short ventral bristles. antennal segment nearly 3.5 times as long as wide and with arista longer than third segment. Acr absent. Distribution. – Singapore, Malaysia, South-Thailand (Trang prov.). Description. – Male. Body length: 1.5 mm; wing length: 1.65 mm. Head. Frons broad, with shining metallic green ground colour; Group V. T. gratiosus group face concolorous with frons, narrow, near middle as wide as an eye facet. Eye facets enlarged in front and below, small The T. gratiosus species group was erected by Meuffels and above. Palpi small, brown, with a black apical bristle. Rostrum Grootaert (2003) to group a number of species that were brown. Occiput shining metallic green. Chaetotaxy as usual; formerly considered to belong to the genus Mastigomyia a pair of long brown ocellars and verticals, 2 small Becker, 1922. Mastigomyia was synonymised with postocellars. Postocular cilia uniseriate, short, brown. Antenna Teuchophorus on the basis of the identical wing venation and (Fig. 6) with basal segments yellow, third segment brown, the inconsistent length of the third antennal segment and the very long, at least three times (3.2) as long as wide. Arista arista in males. Males have a very long third antennal segment, about 1.5 times as long as third segment. but the length of the arista is variable. In some species the Thorax. Mesoscutum and scutellum shining metallic green, arista is absent, or short, spine-like or the arista can be longer but yellowish-brown on sides (humeri, notopleural than the third segment. Females have a short third antennal depression, postalar calli). Pleurae brown in apical half, lower segment with long arista and cannot be separated from other half of mesopleura and sternopleura yellow. Long bristles on Teuchophorus species. This was one more reason to lump mesoscutum brown. Chaetotaxy as usual, but no acr, 5 dc. Mastigomyia with Teuchophorus. Future genetic work might Scutellum with 2 marginals, flanked each by a minute hair. prove if this synonymy was a useful action or not. Legs, including coxae, yellow. Fore leg. Coxa with very short, sparse brown hairs, and a Two species are known to occur in Southeast Asia: T. row of brown bristles towards apex. Femur with short, ventral gratiosus (Becker, 1924) (described from Taiwan and bristles in basal third, half as long as femur is wide. A weak recorded from Laos and Ryukyu Island, Japan) and T. posterior preapical present. Tibia about as long as femur, trangensis (Bickel, 1999) (Thailand). Here I describe three without bristles or serration. First tarsomere without basal additional species from Singapore, but more species are spinule. Length of femur, tibia and tarsal segments (in mm): expected since I have seen an undescribed species from 0.49 : 0.49 : 0.3 : 0.14 : 0.14 : 0.07 : 0.07. Northeast Thailand (Na Haeo, Loei province). Further species Mid leg. Coxa with long brown exterior bristle. Femur with in the T. gratiosus group are T. miricornis Meuffels & a brown preapical av and a preapical pv. Tibia with a weak Grootaert, 2003, from Papua New Guinea, T. amami (Bickel, brownish yellow pd on basal third; 2 ad and apical crown 1999) from Ryukyu Island (Japan) and T. anomalicerus with 2 somewhat longer bristles. Tarsomeres 4 and 5 (Hollis, 1964) from Nepal. unusually short. Length of femur, tibia and tarsal segments (in mm): 0.63 : 0.6 : 0.31 : 0.14 : 0.8 : 0.3 : 0.3. In contrast to most other Teuchophorus species which are Hind leg. Coxa with a brown exterior bristle (shorter than found on humid soils, the T. gratiosus-species are found by that on mid coxa). Femur slightly thickened, without any sweep netting in the vegetation or in Malaise traps and they bristle, except a preapical a and av. Tibia longer than femur, are not active at ground-level. Hence it is likely that the adults with 3 short brownish yellow pd. Length of femur, tibia and live in the vegetation and the canopy. tarsal segments (in mm): 0.7 : 0.84 : 0.14 : 0.17 : 0.1 : 0.07 : 0.06.

62 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Wing faintly brownish tinged, with brown veins. Costa not Discussion. – Teuchophorus temasek, new species, is closely thickened. r4+5 and m1+2 apically slightly diverging. Apical related to T. gratiosus but the latter has uniseriate acrostichal portion of Cu twice as long as tp. Halter yellow. Squama bristles and the arista very short. In T. temasek, the arista is with almost black border and black cilia. longer than the third antennal segment and there are no Abdomen brown; venter yellowish. Hairs and bristles acrostichal bristles. A synapomorphy are the long marginal brownish. First tergite with 5 pairs of long marginal bristles bristles on the first tergite. (outer most short). Hypopygium (Figs. 7-8) brown; cerci Teuchophorus temasek, new species, and T. neesoonensis, brown. Cercus with a long apical bristle (about as long as new species, are sister species. The main differences in the cercus is long). A pair of strong pedunculate epandrial bristles two species are the length of the third antennal segment and and a short basal epandrial (Fig. 7). Hypandrium a simple the arista. In T. temasek the third antennal segment is nearly plate. 3.5 times as long as wide with the arista longer than the third segment (about 1.5 times as long) and the basal antennal Female. Body length: 2.1 mm; wing length: 1.92 mm. segments are yellowish. In T. neesoonensis the third antennal Identical to male in most respects but third antennal segment segment is 4.5 times as long as wide and the arista is shorter short (Fig. 5): a little longer than wide. Face a little broader than the third segment (3/4 length of third segment). The basal than in male, a little wider than second antennal segment. antennal segments are brown. The structure of the hypopygium of both species is almost identical. Both have Etymology. – The species name temasek refers to the name two strong lateral epandrial bristles on a tubercle and a small of ancient Singapore, which means “city at the sea”. Used as median epandrial seta. A strong dorsal bristle is present on a noun in apposition. the dorsal surstylus. The cercus bears a very long apical

Figs. 5-8. Teuchophorus temasek, new species: 5, female antenna; 6, male antenna; 7, hypopygium ventral view; 8, hypopygium lateral view. D: dorsal; V: ventral.

63 Grootaert: Teuchophorus of Singapore bristle. Finally, T. temasek and neesoonensis differ from T. Diagnosis. – A species of the T. gratiosus group with third antennatus new species in having no acrostichal bristles and antennal segment nearly 4.5 times as long as wide and with the presence of a well-developed arista. T. antennatus has a arista shorter than third antennal segment (about 3/4 of length very short spine-like arista and distinct, uniseriate acrostichals. of third antennal segment). Basal antennal segments brown. It also lacks the very long apical bristles on the cercus and Acr absent. has only a single lateral epandrial bristle. Description. – Male. Body length: 1.6 mm; wing length: 1.5 Distribution and habitat. – Singapore. Swamp forest. mm. Head. Frons broad, with shining metallic green ground- colour; face concolourous with frons, narrow, near middle Teuchophorus neesoonensis, new species as wide as an eye facet. Eye facets enlarged in front and below, (Figs. 9-11) small above. Palpi small, brown, with a black apical bristle. Rostrum brown. Occiput shining metallic green Chaetotaxy Material examined. – Holotype male: SINGAPORE: Nee Soon, as usual; a pair of long brown ocellars and verticals, 2 small Central Catchment NR, acidic swamp forest, Malaise trap, 24 postocellars. Postocular cilia uniseriate, short, brown. Antenna Mar.2005 (25029, coll. P. Grootaert, Si600). (Fig. 9) with basal segments dark brown, third segment pale brown, very long, about 4.5 times as long as wide. Arista Paratypes: SINGAPORE: 2 males, Nee Soon, 26 Aug.2005, Mal 3, about 3/4 length of third segment. (25318, coll. P. Grootaert, Si1108); 1 male, Nee Soon, 7 Sept.2005, Mal. 2, (25332, coll. P. Grootaert, Si1051); 1 male, Nee Soon, 16 Thorax. Mesoscutum and scutellum shining metallic green, Sept.2005, Mal. 2, (25350, coll. P. Grootaert, Si1109). but yellowish-brown on sides (humeri, notopleural

Figs. 9-11. Teuchophorus neesoonensis, new species, male: 9, antenna; 10, hypopygium ventral view; 11, hypopygium lateral view.

64 THE RAFFLES BULLETIN OF ZOOLOGY 2006 depression, postalar calli). Pleurae brown on upper half, but sternite). Hairs and bristles brownish. First tergite with 3 pairs yellow on lower half. Long bristles on mesoscutum black. of long black bristles. First tergite with 6 long black marginal Chaetotaxy as usual, but no acr, 6 dc. Scutellum with 2 bristles. Hypopygium (Figs. 10-11) brown; cerci brown. marginals. Cercus with a long apical bristle (longer than cercus is long). Legs, including coxae, yellow. Dorsal surstylus with a very long dorsal bristle. A pair of Fore leg. Coxa with very short, sparse brown hairs, and a strong pedunculate epandrial bristles and a short basal row of brown bristles towards apex. Femur without ventral epandrial bristle (Fig. 10). Hypandrium a simple plate. bristles. A weak posterior preapical present. Tibia about as long as femur, without bristles or serration. First tarsomere Female unknown. without basal spinule. Length of femur, tibia and tarsal segments (in mm): 0.49 : 0.49 : 0.24 : 0.14 : 0.08 : 0.05 : Etymology. – The name neesoonensis refers to the type 0.05. locality that is probably one of Singapore’s most diverse Mid leg. Coxa with long brown exterior bristle. Femur with forests. a brown preapical av and a preapical pv. Tibia with a weak brownish pd on basal third; 2 ad and apical crown with 2 Discussion. – Teuchophorus neesoonensis, new species, is somewhat longer bristles. Tarsomeres 4 and 5 unusually short. very closely related to T. temasek, new species. For the Length of femur, tibia and tarsal segments (in mm): 0.56 : differences between the two species and the other species of 0.56 : 0.25 : 0.12 : 0.08 : 0.07 : 0.07. the T. gratiosus group, I refer to the discussion under T. Hind leg. Coxa with a brown exterior bristle (shorter than temasek, new species. The most obvious character is the that on mid coxa). Femur slightly thickened, without any length of the arista and the third antennal segment. bristle, except a preapical a and av. Tibia longer than femur, with 3 short brownish pd. Length of femur, tibia and tarsal Distribution and habitat. – Singapore. Swamp forest. segments (in mm): 0.77 : 0.77 : 0.17 : 0.19 : 0.08 : 0.05 : 0.05. Wing faintly brownish tinged, with brown veins. Costa not Teuchophorus antennatus, new species thickened. r4+5 and m1+2 apically slightly diverging. Apical (Figs. 12-14) portion of Cu 1.5 times as long as tp. Halter yellow. Squama with almost black border and black cilia. Material examined. – Holotype male: SINGAPORE: Sime forest, Abdomen brown; venter yellowish (except for brown 5th 2 Jul.2005, Mal. 1 (25189, coll. P. Grootaert, Si864).

Figs. 12-14. Teuchophorus antennatus, new species, male: 12, antenna; 13, hypopygium ventral view; 14, hypopygium lateral view.

65 Grootaert: Teuchophorus of Singapore

Paratypes: SINGAPORE: 1 male, Sime forest, 7 Sept.2005, Mal. 1 apical bristle. A single long, non-pedunculate lateral epandrial (25334, coll. P. Grootaert, Si1106); 1 male, Nee Soon, 16 Sept.2005, and a short basal epandrial bristle present. Dorsal surstylus Mal. 2 (25350, coll. P. Grootaert, Si1107). with a short dorsal bristle. Diagnosis. – A species of the T. gratiosus group. Third Female unknown. antennal segment 5.5 times as long as wide; base subrectangular. Arista spine-like, very short, about half as Discussion. – The new species is closely related to T. long as the subrectangular swelling of third antennal segment trangensis Bickel, 1999. Both have a very long third antennal is wide (by. Bristles on head and thorax brown. Acr brown, segment with a very short arista and the base of the third uniseriate and diverging. Four pairs of marginals on tergite antennal segment has a peculiar subrectagular broadening. 1. In T. trangensis the arista is somewhat longer than the width of the third antennal segment, whereas in the new species Description. – Male. Body length: 1.75 mm; wing length: the arista is shorter: about half as long as the width of the 1.75 mm. third segment. In T. trangensis both basal antennal segments Head. Frons broad, with shining metallic green ground- are yellow, while only the scape is yellow in the new species. colour; face concolorous with frons, narrow, near middle as The male genitalia of T. trangensis are not illustrated, but wide as an eye facet. Eye facets enlarged in front and below, Bickel states (l.c.) that the genitalia of T. trangensis are similar small above. Palpi small, brown, with a black apical bristle. to those of T. gratiosus. The latter has a very long aedeagus, Rostrum brown. Occiput shining metallic green. Chaetotaxy and two long, lateral epandrial setae on a tubercle. The new as usual; a pair of long brown ocellars and verticals, 2 small species has a short aedeagus with a sagged tip and only one postocellars. Postocular cilia uniseriate, short, brown. Antenna lateral epandrial seta. Although T. trangensis and the new (Fig. 12) with basal segment yellow, second segment pale species are morphologically very similar, we consider the brown, third segment brown, very long, about 5.5 times as morphological characters as sound enough to distinguish them long as wide, basally with a subrectangular swelling. Arista as separate species. For a further comparison with T. temasek very short, about half as long as base of third antennal and T. neesoonensis, I refer to the key and the descriptions segment. above. Thorax. Mesoscutum and scutellum shining metallic green, but brown on sides (humeri, notopleural depression, postalar Distribution and habitat. – Singapore. Swamp forest. calli). Pleurae brown on upper half, but yellow on lower half (sternopleuron, hypopleuron). Long bristles on mesoscutum dark brown. Chaetotaxy as usual. Acr distinct, uniseriate and Group VII. T. pauper group diverging, 6 dc. Scutellum with 2 marginals. Legs, including all coxae, yellow. Species without acrostichal bristles and without a basal Fore leg. Coxa with very short, brown hairs, and a row of spinule on tarsomere 1 of the fore leg. Males have no wing brown bristles at apex. Femur with minute ventral bristles, stigma. The legs are poorly bristled. The antennae are and a single longer preapical. A weak posterior preapical predominantly yellow and the third antennal segment is short. present. Tibia about as long as femur, without bristles or This group originally included two species: T. pauper serration. First tarsomere without basal spinule. Length femur, Meuffels & Grootaert, 2003 (Thailand, Indonesia, Sulawesi) tibia and tarsomeres (in mm): 0.58 : 0.49 : 0.28 : 0.14 : 0.084 and T. cteniuchus Meuffels & Grootaert, 2003 (Irian Jaya). : 0.056 : 0.056. Here I add T. singaporensis, new species, and T. meieri, new Mid leg. Coxa with long brown exterior bristle. Femur with species, that resemble very much to T. pauper in the somatic a short, brown preapical av and a strong preapical pv. Tibia characters. T. spinulosus, new species, and T. acuminatus, with a weak brownish pd on basal third; 3 weak pv and an new species, are added here provisionally on the basis of the apical crown of bristles. Length femur, tibia and tarsomeres absence of acr and the absence of a basal spinule on tarsomere (in mm): 0.63 : 0.63 : 0.28 : 0.112 : 0.07 : 0.07 : 0.056. 1 of the fore leg, but the colour and shape of antenna do not Hind leg. Coxa with a brown exterior bristle (shorter than really fit in the group. the one on mid coxa). Femur slightly thickened, without any bristle, except a preapical a and av. Tibia longer than femur, with 4 short brownish pd. Tarsus somewhat curved. Length Teuchophorus singaporensis, new species femur, tibia and tarsomeres (in mm): 0.77 : 0.91 : 0.07 : 0.224 (Figs. 15-19) : 0.112 : 0.084 : 0.07. Wing faintly brownish tinged, with brown veins. Costa not Material examined. – Holotype male: SINGAPORE: Sime forest, thickened. r4+5 and m1+2 apically slightly diverging. Apical 24 Mar.2005 (25026, coll. P. Grootaert, Si562, ZRC). portion of Cu twice as long as tp. Halter yellow. Squama with black border and black cilia. Paratypes: SINGAPORE: 1 male, 4 females, Nee Soon, 9 Mar.2005, Abdomen with brown tergites; first sternite yellow, following swamp forest (25006, coll. P. Grootaert, Si376); 3 females, Nee segments pale brown. Hairs and bristles brownish. Basal Soon, 9 Mar.2005 (25005, coll. P. Grootaert, Si393); 5 males, 5 tergite with 4 pairs of long black marginal bristles (outmost females, Sime forest, 24 Mar.2005 (25026, coll. P. Grootaert, Si562 pair shortest). Hypopygium (Figs. 13-14) brown; cerci brown. - male dissected); 2 females, Bukit Timah, 16 Mar.2005 (25021, Dorsal side of aedeagus near tip sagged. Cercus without long coll. P. Grootaert, Si515); 1 female, Pulau Ubin, Check Jawa, 26 Mar.2005 (25034, coll. P. Grootaert, Si536); 1 male, 4 females, Sime

66 THE RAFFLES BULLETIN OF ZOOLOGY 2006 forest, 15 Apr.2005 (25061, coll. P. Grootaert, Si716); 1 female, left arm and a longer right arm. In resting position aedeagus Kranji (beach forest), 27 Jul.2005 (25254, coll. P. Grootaert, Si927). shorter than hypandrium.

MALAYSIA: 2 males, 1 female, Pulau Tioman, Salang, 13 Jul.2005, Description. – Male. Body length: 1.38-1.50 mm; wing forest (25209, coll. P. Grootaert, Tio2); 4 males, Pulau Tioman, length: 1.25-1.30 mm. Salang, 13 Jul.2005 (25211, coll. P. Grootaert, Tio6); 7 males, Juara, 20 Jul.2005 (25246, coll. P. Grootaert, Tio128; extracted for DNA); Head. Frons and face with greenish black ground-colour, 8 males, 6 females, Johor, Gunung Belumut, 22 Oct.2005, riverbanks feebly shining. Frons broad. Face wide, at its narrowest point in palm oil plantation (25396, coll. P. Grootaert, Si1098). about as wide second antennal segment. Palpi yellowish brown, each with a short brownish yellow, sometimes dark THAILAND: 38 males, 29 females, Phang-Nga province, Thap Put, apical bristle. Rostrum yellowish brown. Occiput greenish 23 Oct.1997, (97107, coll. P. Grootaert) population of only T. black, feebly shining. Chaetotaxy as usual, all bristles singaporensis; 9 males, females (?), Phang-Nga province, Thap Put, brownish; 2 very short postocellars. Antenna (Fig. 15) short, 23 Oct.1997, (97107, coll. P. Grootaert) mixed population of T. yellow; second segment darkened on dorsal surface and singaporensis and T. pauper, so females unidentified; 24 males, 5 narrowly along apical margin; third segment darkened on females, Songkhla province, Ban Huai Mo, 30 Oct.1997, swamp forest (97150, coll. P. Grootaert, RBINS), pure population of T. about apical third. Third segment shorter than wide, with a singaporensis, no T. pauper found. short, blunt apex. Arista slightly more than three times as long as antenna; basal aristal segment slightly shorter than Diagnosis. – Medium-sized species with yellowish antennae. third antennal segment. No acr; 5 dc. Bristles on head, thorax and legs yellow to Thorax. Mesoscutum brown, with a greenish shine, becoming yellowish brown. Costa not thickened. Prosterna entirely paler towards sides; humeri, lower part of notopleura, and a brown with a central black stripe. Mesopleura almost entirely narrow border from wing root unto scutellum brownish brown. Hypandrium bifurcate, but asymmetric with a short yellow. Scutellum brownish with paler border. Mesopleura

Figs. 15-19. Teuchophorus singaporensis, new species, male: 15, antenna; 16, hind leg; 17, tarsomere 1 of hind leg; 18, hypopygium ventral view, with detail of tip of aedeagus and ventral epandrial lobe; 19, hypopygium lateral view. ae: aedeagus; del: dorsal epandrial lobe; hy: hypandrium; L: left; R: right.

67 Grootaert: Teuchophorus of Singapore almost completely brownish; sternopleura yellow; Prosterna T. singaporensis, new species, and T. pauper are very closely brown with a black central stripe. Chaetotaxy as usual, but related. Both have 5 equally long dorsocentrals that are there are no acr; all bristles brownish or brownish yellow. 5 yellowish-brown, prosterna entirely brown with a central dc. Scutellum with 2 marginals. No propleural bristle. black stripe and mesopleura almost entirely brown. The most Legs, including all coxae, yellow. The preapical posterior obvious difference is in the genitalia. Both, T. singaporensis comb on the basal tarsomere of the hind legs is contrastingly and T. pauper have an asymmetrical hypandrium while T. darkened (the other combs are pale). meieri has a symmetrical hypandrium. It is not possible to Fore leg. Coxa with short pale sparse hairs and a row of short differentiate T. pauper from T. singaporensis on the somatic brownish bristles towards apex. Femur with a very short hair- characters, but the structure of the male hypopygium is very like preapical pv. Tibia about as long as femur, without different: shape of the surstyli, hypandrium, dorsal epandrial bristles or serration. First tarsomere without ventral spinule. lobes and aedeagus. Length of tibia and tarsal segments (in mm): 0.4 : 0.2 : 0.09 : 0.07 : 0.06 : 0.07. T. meieri, new species, has 4 equally long, black dc, yellow Mid leg. Coxa anteriorly and exteriorly with very short prosterna with a small central black spot and only the upper hairlets. Femur with 2 minute preapical av and 1 minute third of the mesopleura brown, lower two thirds are yellow. preapical pv. Tibia a little longer than femur, with 2 small T. meieri has, seen in apical view, a large hypandrium with brown ad; 1 small pd near base; 2 small bristles in apical long, symmetrical arms. Both arms end in a large, bifid tip. crown. Length of tibia and tarsal segments (in mm): 0.65 : The ventral epandrial extensions are also symmetrical and 0.22 : 0.13 : 0.1 : 0.08 : 0.08. the tip, in lateral view, is very wide. The surstyli are rather Hind leg. Coxa with a very thin and short black exterior short in comparison to the long, digitiform surstyli in T. bristle. Femur (Fig. 16) with 1-2 preapical av and 1 preapical pauper and T. singaporensis. pv, all very short and weak. Tibia (Fig. 16) hardly longer than femur; 2 (sometimes 3) weak yellowish brown or black Distribution and habitat. – Thailand, Malaysia, Singapore. dorsal bristles on apical half, and in between them a row of Rain forest and swamp forest. slightly lengthened hairs. First tarsal segment slightly thickened; on its anterior apical rim a fringe of very short brownish hairlets. Length of tibia and tarsal segments (in Teuchophorus pauper Meuffels & Grootaert, 2003 mm): 0.6 : 0.1 : 0.2 : 0.1 : 0.07 : 0.075. (Figs. 20-21) Wing hyaline, slightly brownish tinged, with yellowish brown veins. Costa not thickened and not darkened. r4+5 and m1+2 Material examined. – THAILAND: 4 males, Phang-Nga, Sa apically very feebly, but distinctly diverging. Tp short, Nangmanora (type locality), 23 Oct.1997 (97104, coll. P. Grootaert); straight, oblique, much shorter than apical part of m3+4 (about 12 males, Phang-Nga province, Thap Put, 23 Oct.1997, (97107, coll. 2 : 5). Halter yellow. Squama small, yellowish with darkened P. Grootaert) mixed population of T. singaporensis and T. pauper so 3 females unidentified; 1 male, Satun province, Ban Du Son, 29 border, with brownish cilia. Oct.1997, forest with mainly bamboo, (97141, coll. P. Grootaert, Abdomen dorsally dark brown or brown, feebly shining, with RBINS); 119 males, 31 females, Songkhla province, Ban Khlong yellowish incisions. Venter and sides yellowish. Hairs and Kua, 29 Oct.1997, secondary rain forest (97142, coll. P. Grootaert, hindmarginal bristles on terga short, dark. Hypopygium (Figs. RBINS), pure population of T. pauper, no T. singaporensis present; 18 and 19) large, dark brown, partly blackish, especially the 2 males, 2 females, Satun province, 20 km north of La-Ngu, 1 asymmetrical hypandrium contrastingly black and bulging; Nov.1997, secondary rain forest (97155, coll. P. Grootaert, RBINS); right arm of hypandrium almost as long as left arm; tip 10 males, 1 female, Satun province, 21 km north of La-Ngu, 1 pointing downward and smooth. Aeadeagus rather thick. Two Nov.1997, secondary rain forest (97156, coll. P. Grootaert, RBINS). long dorsal epandrial lobes present that surpass the tip of the hypandrial arms, each bearing a small papilla. Diagnosis. – Medium-sized species with yellowish antennae. No acr; 5 dc. Bristles on head, thorax and legs yellow to Female. Body length 1.3-1.7 mm; wing length 1.2-1.5 mm. yellowish brown. Costa not thickened. Prosterna entirely Agrees in every aspect with the male. Face a little wider. brown with a central black stripe. Mesopleura almost entirely Legs as in male. Oviscapt with 6 short black acanthae on brown. Hypandrium bifurcate, but asymmetric with a very each hemitergite. short left arm and a longer right arm. The tip of the right arm is rugose in right lateral view. The tip of the aedeagus is long Discussion. – There are three very closely related species in effilate. The dorsal surstylus is dorsally near tip excavated the T. pauper group that should be compared: T. pauper and bears two long bristles at the base of the concavity. Meuffels & Grootaert, 2003, T. singaporensis, new species, and T. meieri, new species. All three species lack acrostichals, Description. – Male. Body length: 1.5 mm, wing length: 1.25 are of the same size, have yellowish brown antenna, no mm. peculiar bristling on legs and wing without stigma. Although Meuffels & Grootaert (2003) shows curiously bent all three species are small in size, and even if there is variation appendages below the hypandrium. Specimens of the type in colouration of the thorax and the bristles, they can easily locality (Sa Nangmanora near Phang-Na, Thailand) were re- be recognized in lateral view, by looking at the shape of the examined. In the original description (Fig. 120) the structure hypandrium and the surstyli. of the right hypandrial arm was misinterpreted. It is much

68 THE RAFFLES BULLETIN OF ZOOLOGY 2006 shorter and has a rugose (wrinkled) tip. The dorsal epandrial Description. – Male. Body length: 1.5 mm, wing length: 1.25 lobe is very long and bears two short, pointed appendages mm. (Fig. 20). Head. Frons and face with greenish blue black ground-colour, feebly shining. Frons broad. Face wide, at its narrowest point I have not found T. pauper in Singapore. Considering its almost as wide as second antennal segment is wide. Palpi distribution in South-Thailand and the mixed populations of brown, each with a short black apical bristle. Rostrum T. pauper and T. singaporensis, it is likely to occur in yellowish brown. Occiput greenish black, feebly shining. Singapore as well. Chaetotaxy as usual, all bristles black; 2 very short postocellars. Antenna (Fig. 22) short, yellowish brown; third Distribution and habitat. – South Thailand, Indonesia: segment hardly as long as wide, with a short, blunt apex. Sulawesi Utara. Rain forest and swamp forest. Arista slightly more than three times as long as antenna; basal aristal segment shorter than third antennal segment. Thorax. Mesoscutum dark brownish, shining, paler brownish Teuchophorus meieri, new species towards sides: humeri, lower part of notopleura, and a narrow (Figs. 22-24) stripe from wing root unto scutellum brownish yellow. Scutellum brown. Pleurae yellowish. Upper third of Material examined. – Holotype male: SINGAPORE: Nee Soon, 5 mesopleura brown, lower two thirds yellow, Sternopleura and Apr.2005, Malaise trap in swamp forest (25048, coll. P. Grootaert, hypopleura yellow. Bristles on thorax black. No acr; 4 dc. Si665). Scutellum with a pair of black marginals. A minute pale propleural bristle. Diagnosis. – Medium-sized species with yellowish antennae. Legs, including coxae, yellow (including posterior four No acrostichals. 4 black dc. Bristles on head, thorax and legs coxae). Tip of hind femora brown. black. Legs poorly bristled. Costa not thickened. Hypandrium Fore leg. Coxa with very short sparse hairs and a row of short symmetrical. Prosterna yellow, with a central brown spot. brownish bristles at apex. Femur with a very short hair-like Upper third of mesopleura brown, lower two thirds yellow. preapical pv. Tibia about as long as femur, without bristles

Figs. 20-21. Teuchophorus pauper Meuffels & Grootaert, male: 20, hypopygium ventral view, with detail of dorsal epandrial lobe (del), right hypandrial arm and tip aedeagus; 21, hypopygium lateral view. L: left; R: right.

69 Grootaert: Teuchophorus of Singapore or serration. No basal spinule on first tarsomere. Length of Venter and sides yellowish. Hairs and hindmarginal bristles femur, tibia and tarsal segments (in mm): 0.36 : 0.35 : 0.15 on terga extremely short, dark. Hypopygium (Figs. 23-24) : 0.08 : 0.07 : 0.06 : 0.07. moderately large, dark brown, partly blackish, with partly Mid leg. Coxa anteriorly and exteriorly with very short hairs, black appendages. Hypandrium with two long arms (Fig. 23). no exterior bristle. Femur with a minute preapical av. Tibia The tip of the arms are enlarged and bifid in lateral view (Fig. a little longer than femur, with 2 small brown ad (one basal, 24). In addition, the epandrium bears a pale, lateral lobe on one median); 1 small pd near base; 2 small bristles in apical both sides, partly covering the tips of the hypandrium (Fig. crown. Length of femur, tibia and tarsal segments (in mm): 24). (This structure is unique in Teuchophorus and probably 0.49 : 0.53 : 0.19 : 0.11 : 0.08 : 0.05 : 0.05. not homologous with the epandrial lobe in other groups). Tip Hind leg. Coxa with a very thin and short dark exterior bristle. of ventral epandrial lobes much enlarged; the lobes are Femur with 1-2 preapical av and 1 preapical pv, all very short symmetrical in ventral view. and weak. Tibia hardly longer than femur; 2 (sometimes 3) weak yellowish brown or black dorsal bristles on apical half, Female unknown. and in between them a row of slightly lengthened hairs. First tarsal segment slightly thickened; on its anterior apical rim a fringe of very short brownish hairs. Length of femur, tibia Etymology. – The new species is dedicated to my host Dr. and tarsal segments (in mm): 0.49 : 0.49 : 0.08 : 0.14 : 0.08 Rudolf Meier, Associate Professor at NUS and head of the : 0.07 : 0.07. Evolutionary Biology Laboratory. Wing hyaline, slightly brownish tinged, with brownish veins. Costa not thickened, no stigma but the costa between r1 and Discussion. – Teuchophorus meieri, new species, is very r2+3 darker brown (not black). r4+5 and m1+2 apically very similar to T. singaporensis and T. pauper in the absence of feebly diverging. Tp short, straight, oblique, much shorter acrostichal bristles, the similar size, general colouration, as than apical part of m3+4 (about 2 : 5). Halter yellow. Squama well as the poorly bristled legs. We refer to the discussion small, with darkened border, with few short brownish cilia. under T. singaporensis to distinguish T. meieri from T. Abdomen dorsally dark brown or brown, feebly shining,. singaporensis and T. pauper.

Figs. 22-24. Teuchophorus meieri, new species, male: 22, antenna; 23, hypopygium ventral view; 24, hypopygium lateral view. hy: hypandrium; del: dorsal epandrial lobe; el: epandrial lobe.

70 THE RAFFLES BULLETIN OF ZOOLOGY 2006

There are more species without acr especially in the T. Description. – Male. Body length: 1.63 mm; wing length: gratiosus group. Here in Singapore, there are T. temasek, new 1.63 mm. species, and T. neesoonensis, new species. The males of both Head. Frons and face with shining dark metallic green ground- have very long antennae. The females having short antenna, colour. Eyes almost touching on face. Palpus small, black, that are however at least as long as wide, are hard to with a small, black apical bristle. Rostrum brown. Occiput distinguish from T. pauper and T. meieri. T. spinulosus, new blackish green. Chaetotaxy as usual; no postocellars. species, and T. acuminatus, new species, also lack Postoculars uniseriate, short, black. Antenna (Fig. 25) brown; acrostichals, but have the third antennal segment about 1.5 third segment pointed triangular, 1.5 times as long as wide. times as long as wide. Arista slightly more than twice as long as antenna, shortly pubescent. Distribution and habitat. – Singapore. River banks in rain Thorax. Mesoscutum dark brown, paler at sides, but darker forest. toward tip; scutellum blackish. Pleurae largely brown; insertion of wing yellowish. Chaetotaxy as usual, but no acr. Legs and coxae yellow, but apical 1/4 of hind femora, and Teuchophorus spinulosus new species tarsi brown, hind tarsi darkest. Fore leg. Coxa anteriorly with (Figs. 25-27) very short black hairs, that become longer towards apex of coxa. Femur without bristles. Tibia slightly shorter than Material examined. – Holotype male: SINGAPORE: Sime forest, femur, without bristles or serration. First tarsal segment 24 Mar.2005 (25026, coll. P. Grootaert, Si560). without a basal, ventral spinule. Length of femur, tibia and tarsal segments (in mm): 0.56 : 0.42 : 0.19 : 0.07 : 0.06 : 0.06 Paratypes: SINGAPORE: 1 male, Sime forest, 24 Mar.2005 (25026, : 0.08. coll. P. Grootaert, Si560); 5 males, Sime forest, 1 Apr.2005 (25041, Mid leg. Coxa anteriorly with very short brown hairs. Femur coll. P. Grootaert, Si637); 1 male, Sime forest, 15 Apr.2005, sweeping (25061, coll. P. Grootaert, Si717); 1 male, Sime forest, 8 (Fig. 26) ventrally with about 4-5 short spine-like bristles Apr.2005 (25053, coll. P. Grootaert, Si736); 1 male, Sime forest, 1 being 3/4 of width of femur; a weak preapical pv, and a yet Jul.2005 Mal. 1 (25189, coll. P. Grootaert, Si1061). thinner preapical av. Tibia with 2 ad, 1 pd. Length of femur, tibia and tarsal segments (in mm): 0.65 : 0.58 : 0.22 : 0.14 : Diagnosis. – A small species without acrostichal bristles. 0.09 : 0.07 : 0.08. Wing without stigma. Third antennal segment elongated, Hind leg. Coxa with a thin, black exterior bristle. Femur with triangular, almost 1.5 times as long as high. Mid femur with a small preapical av, and some of the hairs of the anteroventral 4 to 5 short ventral spine-like bristles. row near tip very slightly lengthened. Tibia about as long as

Figs. 25-27 Teuchophorus spinulosus, new species, male: 25, antenna; 26, mid femur; 27, hypopygium lateral view.

71 Grootaert: Teuchophorus of Singapore femur; with 2-3 short dorsal bristles; 1 very short and weak Paratypes: SINGAPORE: 1 male, Nee Soon (Lower Peirce ventral bristle. Length of femur, tibia and tarsal segments (in Reservoir), 27 Apr.2005 (25094, coll. P. Grootaert, Si789); 1 male, mm): 0.70 : 0.64 : 0.11 : 0.19 : 0.11 : 0.08 : 0.08. Nee Soon, 19 Aug.2005 (25297, coll. P. Grootaert, Si952); 1 male, Wing greyish brown tinged, with dark brown veins. No stigma Nee Soon, 7 Sept.2005 (25323, coll. P. Grootaert, Si1052). and costa between r1 and r2+3 not darkened. r4+5 and m1+2 very parallel. Apical part of cu, 1.5 times as long as tp. Halter Diagnosis. – Third antennal segment nearly 1.5 times as long white. Squama brown, with a black border, and 5 long black as wide: basal part wide, with a fine, elongated tip. No acr. cilia. No stigma. No ventral bristles on femora. Abdomen dorsally dark brown, with a feeble greenish gloss. Sterna brown. Hairs and hindmarginal bristles on terga short, Description. – Male. Body length: 1.75 mm; wing length: black. Hypopygium (Fig. 27) brown, with tips of surstyli 1.5 mm. black. Head. Frons broad, with shining metallic green ground- colour; face concolorous with frons, narrow, near middle as Female unknown. wide an eye facet. Eye facets enlarged in front and below, small above. Palpi small, yellowish-brown, with a black apical Etymology. – The name spinulosus refers to the 4-5 short bristle. Rostrum brown. Occiput shining metallic green. spine-like bristles on the mid femur. Chaetotaxy as usual; a pair of long brown ocellars and verticals, 2 small postocellars. Postocular cilia uniseriate, Discussion. – The short spine-like bristles on the mid femur short, brown. Antenna (Fig. 28) with all segments yellowish- are characteristic. The shape of the third antennal segment is brown; third antennal segment not triangular, but rather onion- also quite unique. shaped with a slender point; almost 1.5 times as long as wide. Thorax. Mesoscutum and scutellum brownish in ground- Distribution and habitat. – Singapore. Riverbanks in rain colour, with a metallic green shine. Pleura with mesopleura forest. yellowish-brown; sternopleura and hypopleura contrastingly pale yellow; pteropleura brown. Long bristles on mesoscutum Teuchophorus acuminatus, new species dark brown. No acr, 5 dc. Scutellum with 2 marginals. (Figs. 28-30) Legs yellow, without distinct bristles. Femora without ventral bristles. No indication of a basal spine, or a hair on a tubercle Material examined. – Holotype male: SINGAPORE: Nee Soon on fore metatarsus. Fore leg: No indication of a basal spinule (Lower Peirce Reservoir), 27 Apr.2005 (25091, coll. P. Grootaert, or hair on tarsomere 1. Length femur, tibia and tarsomeres Si783). (in mm): 0.42 : 0.42 : 0.36 : 0.11 : 0.07 : 0.06 : 0.06.

Figs. 28-30. Teuchophorus acuminatus, new species, male: 28, antenna; 29, hypopygium ventral view; 30, hypopygium lateral view.

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Mid leg: Length femur, tibia and tarsomeres (in mm): 0.53 : Group X. T. pectinatus group 0.59 : 0.39 : 0.27 : 0.084 : 0.07 : 0.07. Hind leg: Length femur, tibia and tarsomeres (in mm): 0.63 This large species group is characterized as follows: spinules : 0.64 : 0.11 : 0.252 : 0.14 : 0.098 : 0.07. at ventral base of fore tarsomere 1 present; Legs usually Abdomen. Tergites brown, sternites pale brown. First tergite poorly bristled; Eyes separated on face, except in T. costalis with short marginals only (not long as in T. temasek and the subgroup; Third antennal segment short, length less than or other species of the T. gratiosus group). Male genitalia large, equal to width, seldom longer than wide; Surstyli separated, elongated. Hypandrium and aedeagus asymmetrical (Fig. 29- but sometimes closely connected, usually long, mostly of 30). about equal length and often finger-shaped. Cerci often sickle- shaped. Hypandrium as a rule not surpassing tips of surstyli. Female unknown. This is a very large group containing 29 species that has been subdivided into 12 subgroups (Meuffels & Grootaert, 2003). Etymology. – The name acuminatus refers to the pointed tip This group is apparently characteristic for the Oriental and of the third antennal segment (acuminatus Lat. for pointed). Australian Regions and radiated here.

Discussion. – The shape of the tip of the third antennal segment suggests that T. acuminatus, new species, might Teuchophorus simplicissimus Meuffels & Grootaert, belong to the T. gratiosus group. In contrast to the T. gratiosus 2003 group, the genitalia are large and there are no long marginal (Figs. 31-34) bristles on the first tergite. Because of the absence of the acrostichals, the rather pale mesoscutum and the asymmetrical Meuffels & Grootaert, 2003: 222, Figs. 192-197. hypandrium, T. acuminatus, new species, is provisionally placed in the T. pauper group. Material examined. – SINGAPORE: 15 males, Sungei Buloh, 6 Oct.2000 (20032, coll. P. Grootaert & N. Evenhuis); 19 males, 22 females, Sungei Buloh, 9 Dec.2002 (22057, coll. P. Grootaert); 7 Distribution and habitat. – Singapore. Riverbanks in rain males, Sungei Buloh, 28 May.2005 (25037, coll. P. Grootaert, forest. Si555); 62 males, Sungei Buloh, 22 Jun.2005 (25121, coll. P.

Figs. 31-34. Teuchophorus simplicissimus Meuffels & Grootaert, male: 31, antenna (Sungei Buloh); 32, hypopygium ventral view (Pulau Tioman); 33, hypopygium ventral view with detail of dorsal surstylus (Sungei Buloh); 34, hypopygium lateral view with detail of aedeagus (Sungei Buloh). ae: aedeagus; del: dorsal epandrial lobe; hy: hypandrium.

73 Grootaert: Teuchophorus of Singapore

Grootaert, Si795); 10 males, Sungei Buloh, 26 Aug.2005, sweep presence of the 5 erect hairs in the second basal quarter netting (25321, coll. P. Grootaert, Si1020); 1 male, Kranji, 27 posteriorly on the hind tibia. Jul.2005 (25254, coll. P. Grootaert, Si926). Distribution and habitat. – Thailand, Malaysia and MALAYSIA: 70 males, 48 females, Pulau Tioman, Monkey Bay, Singapore. It is very common in the upper, less saline parts 14 Jul.2005 (25217, coll. P. Grootaert, Tio30); 14 males, Juara, 20 Jul.2005 (25244, coll. P. Grootaert, Tio76); 67 males, Juara, 20 of mangroves in Singapore. The specimens in Thailand were Jul.2005 (25245, coll. P. Grootaert, Tio86); 33 males, Juara, 20 collected on the sides of pools in a cave system in connection Jul.2005 (25246, coll. P. Grootaert, Tio124). with a mangrove.

Diagnosis. – Third antennal segment hardly as long as wide. Costa not swollen, but darkened between r1 and r2+3. First Teuchophorus limosus, new species tarsomere of fore leg with a basal spinule. Mid femur without (Figs. 35-39) bristles. Mid tibia with a row of ventral spinules in apical third. Hind femur without ventrals. Hind tibia posteriorly in Teuchophorus limosus is referred to as species B in Evenhuis & second basal quarter with about 5 erect hairs. Tip of Grootaert, 2002: 316. hypandrium with a small hook. Material examined. – Holotype male: SINGAPORE: Sungei Buloh, 27 Nov.2003, mangrove (23092, coll. P. Grootaert) in Raffles Discussion. – The specimens in Sungei Buloh are a little Museum. smaller than the ones on Pulau Tioman. Study of the male genitalia (Figs. 33 vs. 32) suggests that they are conspecific. Paratypes: SINGAPORE: 44 males, Sungei Buloh, 27 Nov.2003, It is confirmed by genetic work (Wei Song, Grootaert & mangrove (23092, coll. P. Grootaert); 2 males, 4 females, Sungei Meier, in litt.). For identification one should check the Buloh, 27 Nov.2003 (23091, coll. P. Grootaert); 2 males, Sungei

Figs. 35-39. Teuchophorus limosus, new species male. 35. antenna; 36. mid leg with detail of ventral spinules on apical third of tibia; 37. hind leg; 38. hypopygium ventral view; 39. hypopygium lateral view.

74 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Buloh, 9 Dec.2003 (22057, coll. P. Grootaert); 7 males, Sungei 37) without ventral bristles, with two long av preceding the Buloh, 28 Mar.2005 (25037, coll. P. Grootaert, Si554); 63 males, preapical a. Tibia about as long as femur; 3 dorsal bristles; Sungei Buloh, 22 Jun.2005 (25121, coll. P. Grootaert, Si794); 2 first tarsal segment short; on posterior apical rim a short black males, Sungei Buloh, 11 May.2005 (25159, coll. P. Grootaert, fringe. Length of femur, tibia and tarsal segments (in mm): Si810); 7 males, Sungei Buloh, 6 Jul.2005 (25199, coll. P. Grootaert, 0.52 : 0.56 : 0.11 : 0.14 : 0.1 : 0.07 : 0.07. Si879); 27 males, Sungei Buloh, 26 August.2005, sweep netting (25321, coll. P. Grootaert, Si1019); 4 males, 7 females, Lim Chu Wing hyaline. Costa not thickened between tips of r1 and Kang, 13 Oct.2000 (20051, coll. P. Grootaert & Neal Evenhuis); 3 r2+3, at most darker than the rest of the costa. r4+5 and m1+2 males, Pulau Ubin, Chek Jawa, 19 Nov.2003 (23080, coll. P. apically more or less parallel. Length ratio of basal and apical Grootaert); 3 males, 1 female, Chek Jawa, 2 Dec.2003 (23097, coll. parts of m1+2 about 1:2.5. Halter yellow. Squama brown, P. Grootaert); 1 male, 1 female, Chek Jawa, 11 Dec.2003 (23119, broadly blackened at apex, with dark cilia. coll. P; Grootaert); 3 males, Pandan mangrove, 5 Dec.2003 (23109, Abdomen dorsally dark bronze-brown, subshining. Venter coll. P. Grootaert); 17 males, 10 females, Mandai mangrove, 9 yellowish; sternites brownish at middle. Hairs and marginal Oct.2000 (20038); 3 males, 3 females, Kranji, mangrove, 6 Oct.2000 bristles on terga short, black. Hypopygium (Figs. 38-39) dark (20033, coll. P. Grootaert & N. Evenhuis, species B in Evenhuis & brown. Hypandrium, aedeagus and dorsal epandrial lobes Grootaert, 2002); 2 males, Kranji (beach forest), 27 Jul.2005 (25254, coll. P. Grootaert, Si925). strongly curved and separated from surstyli. Aedeagal complex looks contrastingly black due to the black dorsal Diagnosis. – Antenna brownish black with third antennal epandrial lobes, but hypandrium and aedeagus transparent. segment a little wider than long. No stigma present, but costa between r1 and r2+3 darkened. Acr uniseriate. First tarsomere Female. Body length 1.4-1.7 mm; wing length 1.3-1.45 mm. of fore leg without basal spinule (at most an indistinct hair). In most respects identical to male. Ventral spinules on apical Mid femur with 2 short ventral bristles at base. Mid tibia half of mid tibia absent. ventrally with 4 spinules in apical third. Hind femur without bristles, except for 3 av bristles in apical third. Hind tibia Etymology. – The name limosus (Latin for muddy) refers to with 2 dorsal bristles. Aedeagal complex strong, black, the muddy substrate in the mangrove where the new species separated from surstyli. lives.

Description. – Male. Body length: 1.4-1.6 mm; wing length: Discussion. – The new species looks superficially like T. 1.25-1.4 mm. simplicissimus that is sympatric. It is easily distinguished from Head. Frons subshining metallic green. Face somewhat denser the latter by the strong, black, curved aedeagal complex, the greyish dusted than frons. Palpi small, brown, with a small absence of the posterior erect hairs on the hind tibia and the black bristle. Rostrum brownish yellow. Occiput subshining absence of a basal spinule at the base of the first tarsomere dark metallic green. Chaetotaxy as usual; no postocellars. of the fore leg. Postoculars short, uniseriate, black. Antenna (Fig. 35) short; first and second segments brownish black; third segment dark To identify the species quickly, squeeze the abdomen so that brown, with a lighter coloured apex. Third segment short, the large black aedeagal complex is freed or, look for absence shorter than wide, with a blunt apex. Arista less than 3 times of the erect posterior hairs on the hind tibia to separate the as long as antenna, shortly pubescent; basal segment slightly species from T. simplicissimus. shorter than upper margin of third antennal segment. Thorax. Mesoscutum and scutellum subshining dark metallic Distribution and habitat. – Singapore. Very common in green. Pleurae dark brown, with a metallic green gloss on mangrove, but occurs only in the upper mangrove near pools mesopleura and metapleura; the latter with a black anterior with less saline water or along affluent freshwater streams. border. Acr uniseriate; 5 dc. A minute propleural bristle. Legs, including coxae, yellow. Mid coxa exteriorly with a vague brown longitudinal streak. Teuchophorus tiomanensis, new species Fore leg. Coxa anteriorly with short, sparse, brownish hairs; (Figs. 40-44) towards tip a row of 4-5 rather short brownish bristles. Femur with a very weak preapical pv. Tibia about as long as femur, Material examined. – Holotype male: MALAYSIA: Pulau Tioman, with a short anterodorsal serration, most distinct in apical Paya, 19 Jul.2005 (25237, coll. P. Grootaert, Tio44). half. First tarsal segment without spinules at base, at most an Paratypes: MALAYSIA: 198 males, 118 females, Pulau Tioman, indistinct hair. Length of femur, tibia and tarsal segments (in Paya, 19 Jul.2005 (25237, coll. P. Grootaert, Tio44; 25 males mm): 0.42 : 0.42 : 0.2 : 0.07 : 0.06 : 0.04 : 0.07 selected for DNA sequencing); 10 males, Juara, 20 Jul.2005 (25244, Mid leg. Coxa anteriorly with short brownish hairs, no exterior coll. P. Grootaert, Tio77); 30 males, Juara, 20 Jul.2005 (25245, coll. bristle. Femur (Fig. 36) with a rather long preapical av, and P. Grootaert, Tio87), 16 males, Juara, 20 Jul.2005 (25246, coll. P. a slightly shorter preapical pv; ventrally at base 3 pale bristles Grootaert, Tio126). Females of the Juara samples could not be about half as long as femur is deep; tibia about as long as separated from the females of T. simplicissimus. femur; 2 ad, 1 pd; ventrally on apical two fifths a row of 4- 5 short spinules (Fig. 36); apical bristles short. Length of Diagnosis. – Antenna brownish black with third antennal femur, tibia and tarsal segments (in mm): 0.52 : 0.52 : 0.21 segment as long as wide. No stigma present and costa between : 0.11 : 0.08 : 0.07 : 0.07. r1 and r2+3 only faintly darkened. Acr uniseriate. Fore tibia Hind leg. Coxa with a weak black exterior bristle. Femur (Fig. with dorsal serration. First tarsomere of fore leg with a small

75 Grootaert: Teuchophorus of Singapore basal spinule. Mid tibia ventrally with a row of spinules in mesopleura and metapleura. Acr uniseriate, much shorter than apical half. Hind femur with minute bristles, except for 3 av dc; 5 dc. bristles in apical third. Hind tibia with 3 dorsal bristles; Legs, including coxae, yellow. Mid coxa brownish, exteriorly posteroventrally in apical half a row of hairs about as long as with a vague brown longitudinal streak. tibia is wide. Hypandrium black, pointed. Cercus with long Fore leg. Coxa anteriorly with short, sparse, brownish hairs; apical bristles, surpassing tip of ventral surstylus. Dorsal towards tip a row of 4 brownish bristles. Femur with a very surstylus not developed. weak preapical pv. Tibia about as long as femur, with an anterodorsal serration (Fig. 42). First tarsal segment with a Description. – Male. Body length: 1.6-1.75 mm; wing length: spinule at base. Length of femur, tibia and tarsal segments 1.5-1.6 mm. (in mm): 0.49 : 0.41 : 0.21 : 0.07 : 0.07 : 0.07 : 0.08. Head. Frons subshining metallic green. Face wide, almost as Mid leg. Coxa anteriorly with short brownish hairs, no exterior wide as basal antennal segment. Palpi small, black, with a bristle. Femur with a rather long preapical av, and a slightly small black bristle. Rostrum brown. Occiput subshining dark shorter preapical pv; ventrally with only minute hairs. Tibia metallic green. Chaetotaxy as usual. Postoculars short, about as long as femur; 2 ad, 1 pd; ventrally on apical third uniseriate, black. Antenna (Fig. 40) short; first and second a row of 4-5 short spine-like bristles; apical bristles short. segments brownish black; third segment dark brown, with a Length of femur, tibia and tarsal segments (in mm): 0.59 : lighter coloured apex. Third segment as long as wide, with a 0.59 : 0.25 : 0.17 : 0.09 : 0.08 : 0.07. rounded tip. Arista 3.5 times as long as antenna, shortly Hind leg. Coxa with a weak black exterior bristle. Femur (Fig. pubescent; basal segment longer than upper margin of third 41) without ventral bristles, with two long av preceding the antennal segment. preapical a. Tibia about as long as femur; 3 dorsal bristles; a Thorax. Mesoscutum and scutellum subshining dark metallic weak posterior in basal 2/5; posteroventrally in apical half a green. Pleurae dark brown, with a metallic green gloss on row of hairs about as long as tibia is wide (Fig. 41). Length

Figs. 40-44. Teuchophorus tiomanensis, new species, male: 40, antenna; 41, hind leg anteriorly with detail of tibia in posterior view; 42, fore leg; 43, hypopygium ventral view; 44, hypopygium lateral view.

76 THE RAFFLES BULLETIN OF ZOOLOGY 2006 of femur, tibia and tarsal segments (in mm): 0.59 : 0.63 : Discussion. – T. tiomanensis is related to T. simplicissimus 0.13 : 0.24 : 0.11 : 0.07 : 0.07. but can easily be distinguished from the latter by its cerci Wing hyaline. Costa not thickened between tips of r1 and that are as long as the single surstylus, and that bear a number r2+3, hardly darker than the rest of the costa. r4+5 and m1+2 of long bristles at tip. apically more or less parallel. Apical half of m1+2 about twice as long as tp. Halter white. Squama brown, darker at apex, Distribution and habitat. – Malaysia, Pulau Tioman. with dark cilia. Mangrove. Abdomen dorsally dark bronze-brown, subshining. Sternites brown. Hairs and marginal bristles on terga short, black. Hypopygium (Figs. 43-44) dark brown. Hypandrium broad Teuchophorus bulohensis, new species at base, but tip narrow and elongate. A single short basal (Figs. 45-47) epandrial bristle present. Dorsal surstylus absent. Cercus yellowish, but with a brown tip bearing about 3-4 long bristles. Material examined. – Holotype male: SINGAPORE: Sungei Buloh, 28 Mar.2005 (25037, coll. P. Grootaert, Si553). Female. Body length: 1.4-1.7 mm; wing length: 1.3-1.45 mm. In most respects identical to male. Fore tibia with a weak Diagnosis. – Rather large species with short surstyli; cerci serration. First tarsal segment also with a basal spinule. Mid longer than surstyli. Hind tibia with some short posteroventral tibia without ventral spinules in apical half; a strong av bristle bristles near middle (not erect like in T. simplicissimus). near middle (absent in male), a strong anterior at basal quarter and an anterior just below middle. Hind tibia without a pv Description. – Male. Body length: 1.5 mm; wing length: 1.6 row of hairs. mm. Head. Frons and face shining metallic green. Face broad, Etymology. – The name tiomanensis refers to the type locality about as wide as basal antennal segments. Palpi small, brown, Pulau Tioman, a Malaysian island in the South China Sea. with a small black bristle. Rostrum brownish yellow. Occiput

Figs. 45-47. Teuchophorus bulohensis, new species, male: 45, antenna; 46, hypopygium ventral view; 47, hypopygium lateral view.

77 Grootaert: Teuchophorus of Singapore subshining dark metallic green. Chaetotaxy as usual; minute Wing hyaline. Costa not thickened between tips of r1 and postocellars. Postoculars short, uniseriate, black. Antenna r2+3. r4+5 and m1+2 apically more or less parallel. Length (Fig. 45) short; first and second segments brownish black; ratio of basal and apical parts of m1+2 about 1:2.5. Halter third segment dark brown, with a paler apex. Third segment yellow. Squama brown, broadly blackened at apex, with 6 short, shorter than wide, with a rounded apex. Arista about 3 black cilia. times as long as antenna, shortly pubescent; basal segment Abdomen dorsally dark bronze-brown, subshining. Sternites slightly shorter than upper margin of third antennal segment. brownish. Hairs and marginal bristles on terga short, black. Thorax. Mesoscutum and scutellum subshining dark metallic Hypopygium (Figs. 46-47) dark brown. with short truncate green. Pleurae dark brown, with a metallic green gloss on ventral surstylus. Cercus a little longer than surstyli. mesopleura and metapleura; the latter with a black anterior border. Acr uniseriate; 5 dc. A minute propleural bristle. Female unknown. Legs, including coxae, yellow. Mid coxa exteriorly with a vague brown longitudinal streak. Etymology. – The species is named after the type locality Fore leg. Coxa anteriorly with short, sparse, black hairs; Sungei Buloh, a mangrove nature reserve on the North coast towards tip a row of 4-5 short black bristles. Femur with a of Singapore. preapical pv, without ventral bristles. Tibia about as long as femur, with an anterodorsal serration, most in apical two thirds. First tarsal segment with a small spinule at base. Length Discussion. – The short, truncate ventral surstylus is quite of femur, tibia and tarsal segments (in mm): 0.46 : 0.39 : unique in Teuchophorus. Teuchophorus bulohensis resembles 0.22 : 0.07 : 0.06 : 0.07. T. limosus but is bigger. It differs from T. simplicissimus in Mid leg. Coxa anteriorly with short black hairs, no exterior having the eyes wider separated on face and lacking the erect bristle. Femur with a rather long preapical av (preapical pv bristles on the posterior side of the hind tibia. probably broken); ventrally at base with some short bristles about half as long as femur is deep; tibia about as long as Distribution and habitat. – Singapore. Actually it is only femur, with a strong anterior at middle; 2 ad, 1 pd; ventrally known from the holotype male found in mangrove. on apical half a row of 4-5 black spinules; apical bristles short. Length of femur, tibia and tarsal segments (in mm): 0.59 : 0.59 : 0.29 : 0.17 : 0.98 : 0.98 : 0.07. Unclassified species Hind leg. Coxa with a weak black exterior bristle. Femur without ventral bristles (preapical probably rubbed off). Tibia Teuchophorus krabiensis Meuffels & Grootaert, 2003 about as long as femur; 5 dorsal bristles and some short, but (Figs. 48-49) distinct posteroventral bristles near middle (not erect like in simplicissimus). Length of femur, tibia and tarsal segments Meuffels & Grootaert, 2003: 235, Figs. 229-234. (in mm): 0.57 : 0.59 : 0.14 : 0.21 : 0.14 : ? : ? . Teuchophorus species A - Evenhuis & Grootaert, 2002: 316.

Figs. 48-49. Teuchophorus krabiensis Meuffels & Grootaert, male: 48, antenna; 49, hind leg anterior view with detail of the black bristles in the apical comb in posterior view.

78 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Material examined. – SINGAPORE: 2 males, Labrador Park, 7 Teuchophorus stenostigma Meuffels & Grootaert, 2003 Oct.2000 (20035, coll. P. Grootaert & N. Evenhuis; species A in (Figs. 50-55) Evenhuis & Grootaert, 2002); 2 males, Labrador Park, 31 Jul.2005 (sample no. 25275, coll. P. Grootaert, Si929); 2 males, Semakau Material examined. – SINGAPORE: 3 males, 2 females, Nee Soon, Island, sandy beach near mangrove, 10 Mar.2005 (25009, coll. P. Central Catchment area, acidic swamp forest, 4 Dec.2002 (22054, Grootaert, Si403). coll. P. Grootaert, Si320, Si 444); 1 female, Nee Soon, 9 Mar.2005, (25005, coll. P. Grootaert Si513); 1 male, 1 female, Bukit Timah, MALAYSIA: 2 males, 5 females, Langkawi, Burau Bay, 4 16 Mar.2005 (25021, coll. P. Grootaert, Si514); 3 males, 1 female, Sept.2005, sandy beach (25327, coll. P. Grootaert; all extracted for Sime forest, 14 Mar.2005 (25013, coll. P. Grootaert, Si420); 5 males, DNA sequencing). 3 females, Sime forest, 1 Apr.2005 (25041, coll. P. Grootaert, Si638); 1 male, 2 females, Sime forest, 8 Apr.2005 (25044, coll. P. Grootaert, Diagnosis. – Third antennal segment about 1.5 times as long Si690); 1 male, Sime forest, 2 Jul.2005 (25191, coll. P. Grootaert, as wide (Fig. 48). No stigma: costa darkened, not thickened. Si867). First tarsomere of fore leg with a basal spinule. Mid femur at base with about 5-6 bristles, shorter than femur is wide. Mid MALAYSIA: 12 males, Pulau Tioman, Juara, 20 Jul.2005 (24246, tibia with black spinules ventrally in apical third. Hind femur coll. P. Grootaert, Tio127); 5 males, Johor, Gunung Belumut, 22 without bristles, apart from some anteroventral preapicals. Oct.2005 (25397, coll. P. Grootaert, Si1099). Hind tibia with at least 6 long dorsal bristles. Subapical comb on hind tibia with some dark, thickened bristles (Fig. 49). Diagnosis. – Third antennal segment as long as wide. Wing with a very narrow stigma i.e. costa a little thickened and Discussion. – This species comes close to T. simplicissimus slightly darkened. Acr uniseriate. Fore metatarsus with a and belongs to the group of species with a row of ventral distinct small ventral spinule at base (also in female). Mid spinules on the apical third of the mid tibiae. The black bristles in the posteroventral comb on hind tibia are unique in femur with only minute ventral bristles. Mid tibia on apical Teuchophorus and thus an easy character for identification. third without a ventral row of spinules nor a serration. Hind femur with very short ventral bristles, some longer av Distribution and habitat. – South Thailand, Singapore. Rare; preapicals. Halter dark. Aedeagus not large nor contrastingly on sandy beaches where it prefers wet, shady areas. black like in T. limosus.

Figs. 50-55. Teuchophorus stenostigma, male: 50, antenna; 51, wing; 52, mid leg anteriorly; 53, hind leg anteriorly; 54, hypopygium ventral view; 55, hypopygium lateral view.

79 Grootaert: Teuchophorus of Singapore

Description. – Male. Body length: 1.37-1.55 mm; wing without ventral bristles, at most some hairs ventrally and only length: 1.32-1.5 mm. 3 dorsal bristles. Teuchophorus stenostigma was originally Head. Frons and face with shining dark metallic green ground- described from South Thailand (Trang province) in a similar colour. Face about 3/4 times as wide as width of third antennal habitat as here in Singapore. The original description mentions segment. Palpus small, black, with a small, black apical yellow antenna, yellow halters and largely yellow pleurae. bristle. Rostrum brown. Occiput blackish green. Chaetotaxy The specimens from Singapore are all darker. The shape of as usual; no postocellars. Postoculars uniseriate, short, black. the surstyli is the same, but the bristling on the ventral Antenna (Fig. 50) brown; third segment rounded triangular, surstylus is longer in the Singaporean specimens. At the with downward shifted blunt apex, a little longer than wide. moment we consider them conspecific. Arista about 3 times as long as antenna, shortly pubescent; basal aristal segment slightly more than half as long as third Distribution and habitat. – Thailand, Malaysia, Singapore. antennal segment. River banks in forest. Thorax. Mesoscutum dark brown, paler at sides, but darker toward tip; scutellum blackish. Pleurae largely yellowish brown. Chaetotaxy as usual; acr uniseriate, moderately long. GENERAL DISCUSSION Legs and coxae yellow, but apical 3/4 of hind femora, hind tibiae and tarsi brown. The goal of the present paper is to give a taxonomic account Fore leg. Coxa anteriorly with very short black hairs, that of the genus Teuchophorus occurring in Singapore. Singapore become longer towards apex of coxa. Femur without bristles. is a small island republic at the southern tip of the Malay Tibia slightly shorter than femur, without bristles or serration. Peninsula and further surrounded by Indonesian islands. First tarsal segment ventrally shortly spinulose, and with a Singapore is densely populated, but still possesses some semi- small black spinule at ventral base. Length of femur, tibia natural forests. Thirteen Teuchophorus species are recorded and tarsal segments (in mm): 0.35 : 0.37 : 0.21 : 0.08 : 0.07 here. Nine are new for science while the other four species : 0.06 : 0.07. have been described by Meuffels & Grootaert in 2003. Mid leg. Coxa anteriorly with very short brown hairs. Femur with about 4 ventral bristles at basal third being _ of width Local diversity of femur (Fig. 52); a weak preapical pv, and a yet thinner preapical av. Tibia 2 ad, 1 pd, 1 av, all strong. Length of Table 1 gives a survey of the species distribution per sampled femur, tibia and tarsal segments (in mm): 0.56 : 0.52 : 0.21 site. The data of the terrestrial rain forest and swamp forest : 0.12 : 0.07 : 0.07 : 0.07. are grouped on the left side of the table. As can be seen, Nee Hind leg. Coxa with a thin, black exterior bristle. Femur (Fig. Soon (near Upper Peirce reservoir, a swamp forest, with acidic 53) with a short preapical av, and some of the hairs of the soil) is the most diverse site in Singapore with seven anteroventral row near tip very slightly lengthened. Tibia Teuchophorus species. Sime forest, a rain forest that together about as long as femur; 3 dorsal bristles; 1 very short and with Nee Soon is part of the large Central Catchment, is the weak ventral bristle. Length of tibia and tarsal segments (in second diverse site with six species. The species composition mm): 0.56 : 0.56 : 0.14 : 0.17 : 0.11 : 0.08 : 0.07 : 0.07. however is not the same. Nee Soon has three species of the Wing (Fig. 51) brownish tinged, with dark brown veins. A T. gratiosus group. T. temasek being very abundant in Nee narrow, lanceolate, brownish stigma, occupying less than half Soon, but absent in Sime forest. On the other hand, T. the width of subcostal cell. r4+5 and m1+2 very slightly ornatulus is very abundant in Sime forest, but not present in diverging. Vein Cu black, at least darker than the other veins. Nee Soon. T. spinulosus is also only recorded in Sime forest. Halter somewhat brownish (rather dark in some specimens). Additional sampling in comparable sites will show if a Squama brown, with a blackish border, and 4 long black cilia. correlation can be established between the presence of the Abdomen dorsally dark brown or brown, with a feeble species and the soil type (acidic, soil texture, mud/sand), soil greenish gloss. Sterna brown. Hairs and hindmarginal bristlets humidity and exposure to sunlight. Only two species are on terga short, black. Hypopygium (Fig. 54-55) brown, with recorded from the rain forest in Taban valley at Bukit Timah. tips of surstyli black. Although Taban valley looks like a typical Teuchophorus site i.e. a small stream with large mud and sandy banks, the insect Female. Body length: 1.55 mm; wing length: 1.5 mm fauna there is poor on the whole. A reason is probably that In most respects identical to male. Face hardly broader than the site was heavily disturbed by human activity until ten in male. Wing without stigma, but costa slightly thickened. years ago and vegetation changed considerably during the Oviscapt with a row of short black acanthae. last years from open to closed shrub vegetation underneath high trees. The weekly mosquito fogging in the nearby Discussion. – Teuchophorus stenostigma is the only habitations could also have a negative impact. Singaporean species with a slightly developed stigma, the other species have at most the costa a little darkened. T. The marine sites such as the mangroves at Sungei Buloh, laosensis Olejnicek, 2003, is a similar species, but it has Kranji, Mandai, Pasir Ris, Chek Jawa (Pulau Ubin), and brownish black antennae and yellow halters. The hind tibia Pandan and the sandy beach at Semakau Island and Labrador should bear ventral bristles, but Oleijnicek’s drawing shows Park are grouped at the right side of table 1. Teuchophorus a row of long dorsal bristles. Teuchophorus stenostigma from krabiensis is a species that is only found on sandy beaches Singapore has brown antennae, darkened halters, hind tibiae and here in Singapore it was recorded from the sandy part of

80 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Table 1. Occurrence of Teuchophorus in Singapore.

terrestrial forest marine habitats

Chek Jawa Labrador Park

Mandai

Nee Soon

Padan

Pasir Ris Semakau Island

Seletar

Sungei buloh Bukit Timah

Total specimens

Kranji species name Sime forest Teuchophorus acuminatus 31 4 Teuchophorus antennatus 12 3 Teuchophorus bulohensis 11 Teuchophorus krabiensis 22 4 Teuchophorus limosus 156 2 27 8 3 196 Teuchophorus meieri 2 2 Teuchophorus neesoonensis 5 5 Teuchophorus ornatulus 105 105 Teuchophorus simplicissimus 132 1 1 1 135 Teuchophorus singaporensis 81533 1 1 31 Teuchophorus spinulosus 10 10 Teuchophorus stenostigma 61726 31 Teuchophorus temasek 32 32 Total specimens 57 150 5 9 289 4 28 1 9 3 2 2 559 Number species 7 6 2 2 3 3 2 1 2 1 1 1 13 the beach at Labrador Park and the new, reclaimed sandy characters of these species differ hardly, and no Male beach at Semakau landfill Island. Secondary Sexual Characters are present. Nevertheless, the male genitalia of both species are quite distinct. Dissection, Although a reasonable amount of material was collected (559 preparation of the male genitalia of these small species will specimens), I do not claim that the taxonomic survey of be mandatory to identify Teuchophorus correctly whenever Singapore is complete. In fact, only twelve sites were more the slightest doubts arise on the somatic characters. or less intensively sampled. These sites are locally reputed as quite natural, and undisturbed. But I am sure that there A detailed comparative anatomical study of the male genitalia remain a lot of small, isolated sites all over the island that of Teuchophorus is necessary not only to understand its host isolated populations of Teuchophorus. Only T. function, but also the phylogeny and group relations in bulohensis is a singleton (i.e. a species represented by a single Teuchophorus. The aedeagal complex is composed of the specimen). T. meieri is the only doubleton. Singletons and hypandrium, the aedeagus and a pair of sclerites that I doubletons make up fifteen percent of the recorded fauna. provisionally call here the “dorsal epandrial lobes”. In at least This low figure is an indication that the sampling of the one species, T. pauper, they are broadly fused at their base selected sites was quite representative. Nevertheless for the and bear apical papillae. In many other species the two reason cited above and when looking on a regional basis, more sclerites are separated for most of their length as can be seen species are expected to be present in Singapore. in T. limosus, new species. Hypandrium and the dorsal epandrial lobes form a tube into which the aedeagus can be When looking on a regional scale it is also clear that many moved. The hypandrium is usually symmetrical, but in some more species of Teuchophorus are expected to occur in species it is asymmetrical, raising questions about its function. Southeast Asia. At the moment only seven species are A better understanding of the genital morphology will show recorded from Malaysia, four shared with Singapore. if the 10 species groups tentatively established by Meuffels Singapore has thirteen species. Thailand has nineteen species & Grootaert (2003) will stand. At the same time the relation now, only four shared with Singapore. Indonesia has twenty with the Sympycnus – Chaetogonopteron complex will be species, none shared with Singapore. These observations better understood. Future molecular work will be an show nothing else than the huge under-sampling of the region. independent way to corrobate the data.

Further research in Teuchophorus ACKNOWLEDGEMENTS The problem of cryptic species in Teuchophorus is shown here. Mixed and un-mixed populations of T. pauper and T. The author thanks Prof. Dr. Peter Ng, director of Raffles singaporensis, new species, were found. The somatic Museum for Biodiversity Research, and Prof. Dr. Rudolf

81 Grootaert: Teuchophorus of Singapore

Meier, head of the Evolutionary biology laboratory, for Descriptions of New Species. Raffles Bulletin of Zoology, 50(2): hosting him during his sabbatical stay at NUS. Mr. K. L.Yeo 301-316. assisted in the field. The authorities of National Parks are Grichanov, I., 1995. Olegonegrobovia (Diptera: Dolichopodidae), thanked for issuing the Research permits to collect in the new genus from Uganda. An International Journal of National Parks. The author thanks two anonymous referees Dipterological Research, 6: 125–128. for their useful comments. He finally acknowledges a grant Grichanov, I., 1996. Olegonegrobovia pappi sp.n. from Tanzania in the Action 4 programme of the Belgian Science Policy. (Diptera: Dolichopodidae). An International Journal of Dipterological Research, 7(2): 123–124.

LITERATURE Meuffels, H. & P. Grootaert, 2004. (published in 2003) The genus Teuchophorus in South-East Asia and New Guinea, description of new species, species-groups and their phylogeny (Insecta, Bickel, D. J., 1999. The Oriental genus Mastigomyia Becker (Diptera: Dolichopodidae). Raffles Bulletin of Zoology Diptera, Dolichopodidae). Journal of Natural History 38: 143- (Singapore), 47(1): 287–294. 258. Evenhuis, N. & P. Grootaert, 2002. Annotated Checklist of the Olejnicek, J., 2003. Some Dolichopodidae (Diptera) from Laos. Dolichopodidae (Diptera) of Singapore, with New Records and Biologia, Bratislava, 58: 191-200.

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THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 83-92 Date of Publication: 28 Feb.2006 © National University of Singapore

COPIDOGNATHINES (ACARI: HALACARIDAE) IN MANGROVES OF SINGAPORE. I. DESCRIPTION OF THREE SPECIES

Ilse Bartsch Deutsches Zentrum für Marine Biodiversitätsforschung, Forschungsinstitut Senckenberg, Notkestr. 85, 22607 Hamburg, Germany Email: [email protected]

ABSTRACT: – At least a dozen halacarid species inhabit algal mats on sediments, pneumatophores and stems in mangroves of Singapore. In this paper three of the six most abundant copidognathine species, Acarothrix ampliumeris, new species, A. palustris Bartsch and Copidognathus rhombognathoideus, new species, are diagnosed and the new species described. The genus Acarothrix is thought to be bound to tropical and warm-temperate areas with fluctuating salinity. The generic diagnosis is supplemented. Copidognathus rhombognathoideus is closely related to C. lutarius Bartsch, a species recorded from mangroves in the tropical eastern Indian Ocean.

KEY WORDS. – Singapore, mangrove, Halacaridae, Copidognathinae, new species, descriptions.

INTRODUCTION COLLECTING AREA, MATERIAL AND METHODS

Mangroves are characteristic elements in tropical intertidal Singapore lies in the tropics, at approximately 1°N. The island estuaries and protected, muddy shores. Once 13% of the total city-state is surrounded by the Johor Strait and Strait of land area of Singapore had been covered by mangroves, in Singapore which in turn are connecting the Straits of Malaca the meantime only small patches remain (Liow, 2000). and the South China Sea. Mangrove areas studied on the Mangroves are inhabited by a rich macro- and meiofauna, southern coast are (1) in a small bay of the river Pandan but the knowledge of the meiofauna is inadequate and the (1°18'N 103°45'E) and (2) in a corner of the beach of the aquatic mite fauna almost unknown. Most studies on West Coast Park (1°18'N 103°46'E), on the northern coast mangrove meiofauna deal with that of mudflats and/or leaf (3) at the end of Lim Chu Kang Road (1°27'N 103°42'E) and litter, with nematodes and harpacticoids generally being the (4) near an adjacent camp area (1°26'N 103°42'E). dominant taxa (Sasekumar, 1994; Gee & Somerfield, 1997; Somerfield et al., 1998; Katiresan, 2000; Ólafson et al., 2000; The tide in Singapore is approximately semi-diurnal with Netto & Gallucci, 2003), and, if at all, only scattered records considerable difference in the tidal amplitude (Hopper, 1998/ of mites. Whereas mites are rare on bare mudflats, they are 99.). During the stay in Singapore in September/October abundant amongst the algal cover on pneumatophores, stems 2004, the minimum and maximum tidal range varied from and branches (Gwyther, 2000; Proches et al. 2001, 2004; approximately 10 to 260 cm, respectively. Proches & Marshall, 2002; Bartsch, 2003). Major elements of the mite fauna are the Halacaridae (Prostigmata), the only The samples were collected by the author. The mites were mite family completely adapted to life in the sea. The co- preserved and stored in ethanol and after clearing mounted occurring Oribatida (Cryptostigmata), Astigmata and in glycerine jelly. Holotypes and paratypes are deposited in Mesostigmata are in the majority semiaquatic or terrestrial. Zoological Reference Collection of the Raffles Museum of The pontarachnids (Hydrachnidia, Prostigmata), truly aquatic Biodiversity Research (ZRC), additional material is deposited and allied with fresh water mites, are present but in general in the Senckenberg-Museum, Frankfurt (SMF) and Zoological rare in species. Six of the dozen most abundant halacarid Museum, Hamburg (ZMH). species found in Singapore mangrove areas belong to the subfamily Copidognathinae, three are described below, Abbreviations used in the descriptions are: AD, anterior dorsal descriptions of the others are in preparation. plate; ads, adanal setae; AE, anterior epimeral plate; ds-1 to

83 Bartsch: Copidognathine mites in mangroves. I. ds-5, first to fifth pair of dorsal setae of idiosoma, numbered longer than height of that segment; bipectinate seta of tibia from anterior backward; GA, genitoanal plate; GO, genital III and IV wide. Claws with delicate accessory process but opening; OC, ocular plate(s); P-2 to P-4, second to fourth no pecten. palpal segment; pas, parambulacral seta(e); PD, posterior dorsal plate; PE, posterior epimeral plate(s); pgs, perigenital Etymology. – The anterior portion is unusual wide, hence setae, numbered from anterior backward; sgs, subgenital the specific name ampliumeris, broad shouldered, derived setae. The legs and their segments are numbered I to IV, leg from amplus (Latin), wide, broad and umerus (Latin), segments are trochanter, basifemur, telofemur, genu, tibia, shoulder. and tarsus. Colour. – Idiosoma pale or light brown. Eye spots small, In the descriptions, the position of a seta is given in a decimal black. system, with reference to the length of a plate from its anterior to posterior margin. The length of a leg segment is that along Description. – Female: Idiosoma short and wide, slightly the dorsal margin. The setation formula of the legs starts with flattened; with large interval between insertions of legs II and the trochanter, the parambulacral setae are excluded, the III. Length of idiosoma 294-309 µm, length of holotype 294 solenidia included. The diagnoses of the species are prepared µm, width 175 µm. Median portions of AD and PD delicately on the basis of specimens from Singapore. reticulate. Length of AD 87 µm, width 78 µm; anterior margin arched, posterior margin triangular; pair of gland pores on minute cones at the level of insertion of leg I. OC elongate, SYSTEMATICS but hardly reaching the level of insertion of leg III (Fig. 1), length 104 µm, width 48 µm. With dark eye pigment beneath Genus Acarothrix Bartsch, 1990 anterior portion of plate but no distinctly delimited corneae; gland pore and adjacent pore canaliculus in anterolateral Diagnosis. – Dorsum with five pairs of dorsal idiosomatic margin of OC. Length of PD 172 µm, width 102 µm; anterior setae, pair of adanal setae on anal plate and three to four pairs margin rounded; pair of delicate pores somewhat anterior to of gland pores; posterior pair of pores on cones. Pair of greatest width of PD, pair of gland pores on protuberances in ds-3 in about middle of OC. AE with epimeral pores. Male posterior margin of PD. Pair of ds-1 on AD; ds-2 within genital sclerites with pair of external genital acetabula. Palps striated integument between AD and OC (in holotype left four-segmented, second segment with one seta, third segment ds-2 not seen); ds-3 in middle of OC (at 0.5 relative to length without setae, fourth segment with three setae in basal whorl. of OC); ds-4 and ds-5 on PD near lateral margin. Adanal setae Genua shorter than telofemora and tibiae. Trochanters I to in ventral position, adjacent to anal sclerites. IV with one seta each. Basifemur II with three setae. Tibiae I to IV with two ventral setae each. Tarsus I with large ventral Integument of ventral plates almost smooth. AE wide and seta and one or two small apical setulae; tarsi II to IV without long, extending far beyond insertion of leg II, its length 100 ventral setae. All tarsi with pair of parambulacral setae. µm, width 175 µm; posterior margin very wide, truncate; three Solenidion of both tarsus I and II in dorsolateral position. pairs of ventral setae only about as long as dorsal idiosomatic All tarsi with paired claws, central sclerite with claw-like setae; pair of epimeral pores lateral to apodemes between process. epimera I and II (Fig. 2); diameter of pores 6 µm, pores covered with numerous villi (Fig. 3). Length of PE 182 µm, Type species. – Acarothrix palustris Bartsch, 1990. extending anteriad far beyond insertion of leg III, posteriad only slightly beyond insertion of leg IV; each plate with one dorsal seta and three ventral setae. Length of GA 122 µm, Acarothrix ampliumeris, new species width 87 µm; with three pairs of pgs. GO large, length 72 (Figs. 1-16) µm, width 35 µm, its distance to anterior margin of GA equalling 0.34 of length of GO. Genital sclerites with one Material examined. – Holotype: Female, ZRC.ARA.480, Singapore, pair of sgs. Ovipositor extending slightly beyond GO. northern coast, end of Lim Chu Kang Road, mangrove, Cladophora mat (Chlorophyta) on muddy and sandy sediment, coll. I. Bartsch, Length of gnathosoma 92 µm, width 65 µm; pharyngeal field 7 Oct.2004. panelled, remainder of gnathosoma with almost smooth integument. Rostrum in ventral aspect triangular, almost as Paratypes: One male, ZRC.ARA.481, collecting data as above; one larva, ZRC.ARA.482, collecting data as above; one male, SMF, long as gnathosomal base (Fig. 4); extending almost to about collecting data as above; one female, ZMH, collecting data as above; the level of seta on P-2. Tectum truncate. One pair of one male, ZMH, collecting data as above. maxillary setae on gnathosomal base, the other pair in distal half of rostrum; two pairs of rostral setae at tip of rostrum. Diagnosis. – Length of idiosoma 294-325 µm. OC elongate; P-4 with three basal setae removed from basis by about length corneae lacking. PE more than half length of idiosoma. Pair of P-3. of ds-2 in striated integument. Gland pores between ds-4 and ds-5 small, almost inconspicuous. AE very wide. Male with Legs I and II (Figs. 5 and 6) almost equal-sized and shorter 20-22 pgs. All tibiae with one bipectinate ventromedial and than legs III and IV (Figs. 7 and 8). Telofemora with delicate one long, smooth ventral seta; bipectinate seta on tibia II much villi, though in general integument of legs almost smooth.

84 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Figs. 1-8. Acarothrix ampliumeris, new species, female: 1. idiosoma, dorsal; 2, idiosoma, ventral; 3, left part of AE; 4, gnathosoma, ventral; 5, leg I, medial; 6, basifemur to tarsus II, medial; 7, leg III, medial; 8, leg IV, medial. Scale line = 50 µm.

85 Bartsch: Copidognathine mites in mangroves. I.

Tarsi without fossa membranes. Leg chaetotaxy (solenidia Male: Length of idiosoma 314-325 µm. Dorsal aspect same included, pas excluded): leg I, 1, 2, 5, 4, 6, 6; leg II, 1, 3, 5, as that of female. GA with rounded anterior corners; plate 4, 6, 4; leg III, 1, 2, 3, 3, 5, 4; leg IV, 1, 2, 2-3, 3, 5, 3. somewhat longer than but far from as wide as AE. GO in Telofemur IV with the combinations 2/1 and 2/0 dorsal/ middle of GA, distance both to anterior margin of GA and ventral setae. Ventral seta of genu I very long, slightly longer end of genital sclerites slightly less than length of GO. than that seta on genu II as well as ventral seta of tibia I. Spermatopositor extending beyond GA, its length 95 µm, Ventral setae of genua III and IV slender and short. Tibia I width 72 µm. GA with 10-11 pairs of pgs (Fig. 13). Genital with long ventral and bipectinate ventromedial seta. On tibia sclerites with four pairs of sgs; three pairs of sgs setiform, II that bipectinate ventromedial seta conspicuously long, one pair spur-like. A pair of foveae (external genital acetabula) length about twice height of segment, ventral seta long, immediately posterior to sgs (Figs. 12 and 13), foveae ca 4 smooth. Bipectinate seta of tibia III and IV very wide and µm in diameter and obscured by villi. almost as long as height of segment. Dorsolateral and dorsomedial fossary setae of tarsus I almost equal-sized (Fig. Protonymph: Not seen. 9); solenidion, length 10 µm, immediately adjacent to dorsolateral fossary seta (Fig. 10); famulus represented by Larva: Length of idiosoma 177 µm. OC and PD distinctly minute cap, length 1 µm. Tarsus I with one long ventral seta shorter than in adults. Pair of ds-3 on OC, ds-4 and ds-5 in in basal third and one small eupathid seta near its tip. Basidorsal fossary seta of tarsus II wider than that of tarsus margin of PD (Fig. 14). AE wide; apodemes between epimera I, dorsomedial fossary seta very slender (Fig. 11), dorsolateral I and II extending backward beyond pair of epimeral pores fossary seta close to tip of tarsus, adjacent to solenidion. (Fig. 15); plate with two pairs of ventral setae. PE short, with Tarsus III with four dorsal setae; pair of apical fossary setae single pair of setae. Shape of gnathosoma similar to that of slender. Basidorsal fossary seta of tarsus IV wider than adults. With three pairs of five-segmented legs, basi- and corresponding seta of tarsus III; shape of apical fossary setae telofemora fused. Leg chaetotaxy: leg I, 1, 1+3 (basi- plus similar to setae on tarsi III. Tarsi I and II each with pair of telofemur), 4, 5, 6; leg II, 1, 1+3, 4, 5, 4; leg III, 1, 1+2, 3, doubled pas, tarsus III with eupathid medial and widened but 5, 4. Bipectinate seta of tibia II rather slender, much longer tapering lateral pas, tarsus IV with pair of bristle-like pas. than that seta on tibia I. Both ventral setae on tibia III rather slender, none distinctly bipectinate (Fig. 16). Distance Claws with very delicate accessory process; pecten lacking. between two basal setae of tarsus III equalling height of tarsus. All tarsi with small median claw, that of tarsus I bidentate, on the other tarsi upper tooth almost reduced. Remarks. – Four species are at present known, all recorded

Figs. 9-16. Acarothrix ampliumeris, new species: 9, tarsus I, lateral, female (medial fossary seta and claw in broken line, medial parambulacral setae omitted); 10, tip of tarsus I, lateral, female; 11, tarsus II, lateral, female; 12, genital opening, male; 13, genitoanal plate, male; 14, idiosoma, dorsal, larva; 15, idiosoma, ventral, larva; 16, tibia and tarsus III, medial, larva (lateral setae and claw in broken line). Scale line = 50 µm.

86 THE RAFFLES BULLETIN OF ZOOLOGY 2006 from tropical and warm-temperate shallow water habitats, Complementary description. – Adults: Length of female 294- from muddy areas periodically influenced by fresh water. 320 µm, of male 280 µm. Shape of dorsal plates as illustrated Acarothrix ampliata Bartsch, 2004, was collected amongst (Fig. 17). Costae of PD approximately 20 µm wide, with the vegetation of a small river emptying into Tampa Bay, narrow cerotegumental borders. Pair of epimeral pores at end Florida, USA, A. longiunguis Bartsch, 1997, in soft mud in of apodemes between epimera I and II (Fig. 18); porus a northern Australian mangrove area, A. palustris Bartsch, covered with numerous setiform tines. Male with 11-12 pairs 1990, amongst surface sediment and its cover by green and of pgs; its GO large, distance to anterior margin of GA red algae (Cladophora, Caloglossa) in Hong Kong, China, equalling length of GO (Fig. 19). Basi- and telofemora I and and A. umgenica in Avicennia mangroves in KwaZulu-Natal, II with numerous filaments, trochanters, basi- and telofemora South Africa (Bartsch, 1990, 1997a, 2004; Proches, 2002). III and IV with filaments fused to cerotegumental lamellae. Leg chaetotaxy: leg I, 1, 2, 5, 4, 6, 6; leg II, 1, 3, 5, 4, 6, 4; Distinguishing characters between A. ampliumeris and the leg III, 1, 2, 2-3, 3, 5, 4; leg IV, 1, 2, 2-3, 3, 5, 3. Telofemora others are: the position of ds-2 (in A. ampliumeris within the III and IV in general with 2/1 dorsal/ventral setae, rarely striated integument anterior to the OC - in the other species ventral seta absent. Tibiae I, II and III each with short, wide in the margin or on the OC), the shape of the OC (in A. bipectinate seta, on tibia IV this seta slender and smooth. ampliumeris elongate, its posterior portion wide - in the other Claws almost smooth though vestiges of accessory process species either shorter or more slender), the gland pores on present. Median claw of all tarsi bidentate, that claw of tarsus the PD (in A. ampliumeris one distinct pair on cones near III and IV with long upper and lower tooth and therefore posterior margin - in the other species two distinct pairs of median claws somewhat larger than on tarsi I and II. pores, one pair between the ds-4 and ds-5, the other pair near the posterior margin), and the shape of the pectinate seta of Protonymph: Length of idiosoma 217 µm. Dorsal and ventral tibia II (in A. ampliumeris long - in the other species short plates smaller than in adults. Leg chaetotaxy: leg I, 1, 2, 3, 4, and wide). 5, 6; leg II, 1, 2, 3, 4, 5, 4; leg III, 1, 2, 2, 3, 5, 3; leg IV, 0, 1+2 (basi- plus telofemur), 3, 5, 3. Tibiae I, II and III each Distribution. – Singapore, from algal mats in mangrove areas. with one of the two ventral setae being short, wide and pectinate, the other seta slender and smooth. Tibia IV with two smooth ventral setae. Both tarsi III in the protonymph Acarothrix palustris Bartsch, 1990 examined with three instead of four dorsal setae. (Figs. 17-20) Larva: Length of idiosoma 177 µm. Shape of dorsal and Acarothrix palustris Bartsch, 1990: 205-207, figs. 1-14. ventral plates as illustrated in Bartsch (1990: figs. 8 and 9). PD with pair of cerotegumental ridges. Pair of epimeral pores Material examined. – One female, ZRC.ARA.483, Singapore, at about the level of end of apodemes. Leg chaetotaxy: leg I, northern coast, end of Lim Chu Kang Road, Cladophora mat on 1, 1+3 (basi- plus telofemur), 4, 5, 6; leg II, 1, 1+3, 4, 5, 4; muddy and sandy sediment, coll. I. Bartsch, 7 Oct.2004; one larva, leg III, 1, 1+2, 3, 5, 4. Pectinate seta of tibia I and II wide, ZRC.ARA.484, collecting data as above; one female, SMF, collecting data as above; one female, ZMH, collecting data as above; less than height of each segment. Tibia III with two almost one male, ZMH, collecting data as above; one protonymph, ZMH, smooth ventral setae. Distance between two basal setae of collecting data as above. One larva, author’s collection, Singapore, tarsus III more than height of that segment (Fig. 20). southern coast, river Pandan, sediment, green algae and epibiota on Avicennia pneumatophores in a rockpool, coll. I. Bartsch, 27 Remarks. – The PD and GA of the Singapore specimens Sep.2004. illustrated are slightly more slender than in the holotype and paratype from Hong Kong. The width of the plates proved to Diagnosis. – Length of idiosoma 280-320 µm. PD with pair vary slightly in the material from Singapore. of longitudinal porose costae with narrow cerotegumental ridges, remainder of plate reticulate. OC with two corneae. The adults of the two Acarothrix species (A. ampliumeris and PD with two pairs of gland pores. Pair of ds-2 in margin of A. palustris) present in Singapore mangroves can be OC. Pair of ds-4 and ds-5 in lateral margin of PD, almost distinguished on the basis of the length of OC, presence or equidistant relative to anterior pair of gland pores. Female absence of corneae, shape of AE, length of PE, and shape of and male GA almost ovate. Male GA with 11-12 pairs of ventral setae on tibiae II and IV. Juveniles which have small pgs, interval between anterior margin of GA and that of GO dorsal and ventral plates can be distinguished on the basis of equalling length of GO. Tibiae I and II with six setae each. presence or absence of corneae, position of epimeral pores Tibiae I to III each with one bipectinate and one smooth, relative to length of apodemes, and size of the median claws slender ventral seta; on tibiae IV both ventral setae smooth on the posterior tarsi. In addition, larvae can be distinguished and slender. Paired claws almost smooth. on the basis of presence or absence of cerotegumental ridges on the PD and position of the basal seta on tarsus III. Colour. – Gut content pale or slightly brown; the male examined darker than females. Dorsum with three small black Distribution. – Acarothrix palustris inhabits mangroves in eye spots. Hong Kong and Singapore (Bartsch, 1990, and present record).

87 Bartsch: Copidognathine mites in mangroves. I.

Genus Copidognathus Trouessart, 1888 one male, ZRC.ARA.487, collecting data as above; one protonymph, ZRC.ARA.488, collecting data as above; one female, SMF, Diagnosis. – Dorsum with five pairs of dorsal idiosomatic collecting data as above; one female, ZMH, collecting data as above; setae, pair of adanal setae on anal plate and four pairs of gland one protonymph, one larva, ZMH, collecting data as above. pores, the latter may be almost inconspicuous or large and Diagnosis. – Length of idiosoma 353-414 µm. Dorsal plates opening on cones. Pair of ds-3 in striated integument or on with delicate porosity in large hardly raised areolae. OC about PD. AE with pair of epimeral pores. Pair of genital acetabula twice as long as wide, with two corneae. PD with two pairs internal. Palps four-segmented; P-2 with one seta; no seta on of wide porose costae. Epimeral pores large, ovate, their P-3 (there may be a small spiniform cuticular process); P-4 margins smooth. Male GA with seven to eight pairs of pgs. with three setae in basal whorl. Genua of legs shorter than Gnathosoma short, length of rostrum half that of gnathosomal telofemora and tibiae. Trochanters I to IV with 1, 1, 1, 0 setae. base. Tibiae I to IV with 2, 2, 1, 1 bipectinate setae. Tarsi III Basifemur II with two setae. Tibiae I and II with three ventral and IV with lateral but no medial pas. setae each (rarely two setae), tibiae III and IV with two ventral setae each. Tarsus I ventrally with one seta in about middle Etymology. – The species is characterized by a dark-green of segment and two small eupathid apical setae; tarsus II gut content and a gnathosoma with a short rostrum, hence, at without ventral setae; both tarsus I and II with pair of low magnification (20-25x), it may easily be mistaken for a parambulacral setae. Tarsi III and IV in general with pair of representative of the genus Rhombognathus. The specific parambulacral setae but no ventral setae, rarely one of name with the suffix -ides (Greek), Latinized -ideus, refers parambulacral setae moved to a ventral position or lacking. to this similarity. Solenidion of both tarsus I and II in dorsolateral position. All tarsi with paired claws, central sclerite with claw-like Colour. – Content of idiosoma dark, green or grey-green. process. Idiosoma and legs generally fouled. Type species. – Copidognathus glyptoderma Trouessart, Description. – Female: Length of idiosoma 385-414 µm, that 1888. of holotype 414 µm, width 264 µm. Porose areolae of dorsal plates hardly raised; areolae with delicate porosity. Length of AD 117 µm, width 135 µm. Posterior margin almost Copidognathus rhombognathoideus, new species truncate. AD with three wide almost fused areolae (Fig. 21); (Figs. 21-41) anterior third of AD, the porose areola included, as well as paired areolae with delicate porosity. Length of OC 132 m, Material examined. – Holotype. Female, ZRC.ARA.485, Singapore, µ northern coast, end of Lim Chu Kang Road, coll. I. Bartsch, 7 width 67 µm. Plate around corneae slightly raised; integument Oct.2004. within raised area and along lateral margin of OC porose. No distinct gland pore but small pore canaliculus in lateral Paratypes. One female, ZRC.ARA.486, collecting data as above; margin. Triangular posterior portion of OC extending to the

Figs. 17-20. Acarothrix palustris Bartsch: 17, idiosoma, dorsal, female: 18, idiosoma, ventral, female; 19, genitoanal plate, male; 20, tibia and tarsus III, medial, larva. Scale line = 50 µm.

88 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Figs. 21-28. Copidognathus rhombognathoideus, new species, female: 21, idiosoma, dorsal; 22, idiosoma, ventral; 23, left part of AE; 24, gnathosoma, ventral; 25, basifemur to tarsus I, medial; 26, basifemur to tarsus II, medial (in holotype one of ventral setae of that telofemur lacking); 27, leg III, medial; 28, leg IV, medial (ambulacrum with claws broken). Scale line = 50 µm.

89 Bartsch: Copidognathine mites in mangroves. I.

Figs. 29-41. Copidognathus rhombognathoideus, new species: 29, basifemur to tarsus I with porose areolae, lateral; 30, tip of tarsus I, lateral, female (medial setae and claw omitted); 31, tarsus II, lateral, female (medial setae and claw omitted); 32, tarsus III, lateral, female (medial fossary seta and claw omitted); 33, genitoanal plate, male; 34, genital opening, male; 35, idiosoma, dorsal, protonymph; 36, idiosoma, ventral, protonymph; 37, tibia and tarsus III, medial, protonymph; 38, idiosoma, dorsal, larva; 39, idiosoma, ventral, larva; 40, tarsus III, lateral, larva (lateral seta and claw in broken line), 41, idiosoma, dorsal, female. Scale line = 50 µm.

90 THE RAFFLES BULLETIN OF ZOOLOGY 2006 level of insertion of leg III. Length of PD 274 µm, width 197 Male: Length of idiosoma 353 µm. Dorsal aspect similar to µm; anteriad extending far between OC. PD with two pairs that of female. Anterior margin of GA wide, truncate, length of porose costae, 20-25 µm wide, and five panels with small of plate 179 µm, width 150 µm. Length of GO 62 µm. foveae. Distinct gland pores lacking. Dorsal idiosomatic setae Distance to anterior margin of GA 1.3 times length of GO. short; ds-1 on AD, ds-2 within anteromedial margin of OC, Seven to eight pairs of pgs arranged scatteredly around GO ds-3 to ds-5 on PD within lateral margin of medial porose (Fig. 33). Spermatopositor large, length 115 µm, width 86 costae, ds-5 immediately anterior to the level of insertion of µm, extending to anterior margin of GA. Genital sclerites leg IV; ds-3 to ds-5 situated almost equidistant. Adanal setae with four pairs of sgs, three of them setiform, one spiniform close to anal cone. (Fig. 34). With single pair of internal genital acetabula.

Integument of ventral plates uniformly porose. Length of AE Protonymph: Length of idiosoma 329-334 µm. OC and PD 127 µm, width 238 µm. Camerostome with lamella. Pair of distinctly shorter than in adults (Fig. 35). AD with three small, lamellar epimeral processes in medial edge of insertion porose areolae. PD with pair of porose costae, about 25 µm of leg I. Ventral setae longer than dorsal idiosomatic setae. wide, and three longitudinal panels with small foveae. Pair AE with three pairs of setae; pair of ovate epimeral pores of ds-2 within striated integument. Pairs of ds-3 to ds-5 on about 10 µm in length (Fig. 22). Wall of epimeral porus PD, ds-3 and ds-4 close together. AE with pair of ovate slightly raised, smooth, without tines (Fig. 23). Length of PE epimeral pores. PE shorter than in adults, with one dorsal 224 µm, extending distinctly beyond insertion of leg IV. and two ventral setae. Genital plate separated from anal plate Length of GA 213 µm, width 157 µm; anterior margin (Fig. 36). Primordial genital slit with numerous papillae. truncate; with three pairs of pgs inserted as illustrated. Length Shape and setation of gnathosoma as in adults. Leg of GO 90 µm, width 60 µm; ovipositor extending beyond chaetotaxy: leg I, 1, 2, 3, 4, 5, 7; leg II, 1, 2, 3, 4, 5, 4; leg III, GO to the level of anterior pair of pgs. Interval between 1, 1, 2, 3, 5, 4; leg IV, 0, 1+2, 3, 5, 3. Ventromedial seta of anterior margin of GA and GO equalling 0.8 times length of tibia III short and slightly bipectinate (Fig. 37). Basalmost GO. seta of tarsus III situated about halfway between segment’s base and fossary seta. Gnathosoma, rostrum and palps short; length of gnathosoma 112 µm, width 87 µm; its length 0.27 times that of idiosoma. Larva: Length of idiosoma 224 µm. Shape of dorsal plates Rostrum triangular in ventral aspect, its length half that of similar to those of protonymphs (Fig. 38). AE with pair of gnathosomal base (Fig. 24). Palps slender, short, but ovate epimeral pores and two pairs of ventral setae. PE with extending beyond tip of rostrum. P-4 with three basal setae; single seta (Fig. 39). Chaetotaxy of the three pairs of five- apically with setula and two spurs. Tectum truncate. segmented legs: leg I, 1, 1+3, 4, 5, 7; leg II, 1, 1+3, 4, 5, 4; leg III, 1, 1+2, 3, 5, 4. Ventromedial seta of tibiae I and II Legs I and II almost equal-sized (Figs. 25 and 26). Telofemora large and distinctly bipectinate, that seta of tibia III short, I and II somewhat longer than these legs’ tibiae; telofemora slender, though bipectinate. Basalmost seta of tarsus III close III and IV shorter than tibiae (Figs. 27 and 28). Length:height to base of segment (Fig. 40). ratio of telofemora I and II 2.2, that of telofemora III and IV 2.3-2.4. Tarsi without fossa membranes. Integument of lateral Variations. – The anterior part of the PD is rather wide in flanks of basifemora, telofemora and tibiae I and II and those the holotype female but less wide in a paratype female (Fig. of trochanters, telofemora and tibiae III and IV with porose 41). Telofemur II mostly bears 3/2 dorsal/ventral setae; in areolae (Fig. 29). Leg chaetotaxy: leg I, 1, 2, 5, 4, 7, 7; leg the holotype one ventral seta is lacking. II, 1, 2, 4-5, 4, 7, 4; leg III, 1, 2, 2, 3, 5, 4; leg IV, 0, 2, 2, 3, 5, 3. Telofemur I unilaterally with either 3/2 or 3/1 dorsal/ Remarks. – Copidognathus rhombognathoideus is most ventral setae. Tibiae I and II each with two wide, similar to C. lutarius Bartsch, 2003. They share characters conspicuously bipectinate ventromedial setae and one slender, such as the ornamentation of the dorsal plates, ovate epimeral smooth ventral seta; tibiae III and IV each with one bipectinate pores, very short triangular rostrum and short palps, wide ventromedial and one long, slender ventral seta. Dorsal and bipectinate setae on the tibiae, single pas on the tarsi III and dorsomedial setae of tarsi I and II at almost the same level, IV, and dark, green gut content. Distinguishing characters dorsolateral seta moved towards tip of tarsus, close to are: the size of the porose areolae on AD (three large, fused solenidion. Solenidia short, baculiform (Figs. 30 and 31), that areolae in C. rhombognathoideus, three small areolae in C. of tarsus I 5 µm in length, of tarsus II 7 µm. Tarsus I with lutarius), the shape of the PD (anterior margin truncate in C. short lamelliform famulus. Tarsi III and IV with four (Fig. rhombognathoideus but ovate in C. lutarius), the position of 32) and three dorsal setae, respectively. Tarsus I with three the ventral seta on tarsus I (in basal half in C. ventral setae, one level with dorsal seta and two eupathid setae rhombognathoideus but moved to distal half in C. lutarius). near tip of tarsus, close to pair of doubled pas. Tarsus II with pair of pas singlets, tarsi III and IV with slender lateral pas; Copidognathus balakrishnani Chatterjee, 2000, a species medial pas absent. found amongst green algae in Cochin backwater, Kerala, India (Chatterjee, 2000), and C. caloglossae Proches, 2002, from Paired claws slender, accessory process vestigial. Pecten Bayhead Lagoon, Durban Harbour and mangrove forests, lacking. Median claw bidentate. KwaZulu-Natal and Inhambane, South Africa and

91 Bartsch: Copidognathine mites in mangroves. I.

Mozambique (Proches, 2002), have similar short rostra and Bartsch, I., 1997a. Copidognathinae (Halacaridae, Acari) from palps as the two above mentioned species, a PD with four Northern Australia; description of four new species. In: Hanley, wide porose costae, legs with porose integument, and J. R., G. Caswell, D. Meegirian & H. K. Larson (eds.), conspicuously stout bipectinate setae on tibiae I and II. Proceeding of the Sixth International Marine Biological Workshop. The marine flora and fauna of Darwin Harbour, Copidognathus balakrishnani differs from C. lutarius and C. Northern Territory, Australia. Museum & Art Gallery of the rhombognathoideus in that the PD outside the porose costae Northern Territory, Darwin. Pp. 231-243. is reticulate (with delicate foveae in C. lutarius and C. Bartsch, I., 1997b. A new species of the Copidognathus tricorneatus rhombognathoideus), there are three panels outside the porose group (Acari: Halacaridae) from Western Australia with a review costae (five in C. lutarius and C. rhombognathoideus), and of this species-group. Species Diversity, 2: 155-166. tarsi III and IV bear two pas (a single one in C. lutarius and Bartsch, I., 2003. Mangrove halacarid fauna (Halacaridae, Acari) C. rhombognathoideus). The anterior margin of the PD of C. of the Dampier region, Western Australia, with description of caloglossae is much wider than in the other species, the five new species. Journal of Natural History, 37: 1855-1877. ds-2 is near the medial corner of the OC instead of in the Bartsch, I., 2004. Acarothrix ampliata (Arachnida: Acari: anteromedial margin as in the other species, and the GA in Halacaridae), a new halacarid mite from Florida, with notes on both female and male is very long compared to the AE. external genital acetabula. Species Diversity, 9: 259-267. Chatterjee, T., 2000. Two new species of Copidognathus Copidognathus lutarius and C. rhombognathoideus differ (Halacaridae: Acari) from Kerala. Journal of the Bombay from Copidognathus congeners by the combination of (1) Natural History Society, 97: 253-259. dark, green or grey-green gut content, (2) very short, triangular Gee, J.M. & P. J. Somerfield 1997. Do mangrove diversity and leaf rostrum and small palps, (3) large ovate epimeral pores with litter decay promote meiofaunal diversity? Journal of smooth margins, (4) short solenidia, and (5) absence of a Experimental Marine Biology and Ecology, 218: 13-33. medial parambulacral seta on tarsi III and IV. The habitus as Gwyther, J., 2000. Meiofauna in phytal-based and sedimentary well as the collecting data of C. balakrishnani and C. habitats of a temperate mangrove ecosystem – a preliminary caloglossae demonstrate relationship to C. lutarius and C. survey. Proceedings of the Royal Society of Victoria, 112: 137- rhombognathoideus, but the remarkable colour of the gut 151. content is not mentioned in the descriptions of C. Hopper, M., 1998/99. WXTide 32. (freeware). balakrishnani and C. caloglossae (Chatterjee, 2000; Proches, Katiresan, K., 2000. A review of studies on Pichavaram mangrove, 2002), the epimeral pores are not mentioned, in neither the southeast India. Hydrobiologia, 430: 185-205. text nor the illustrations (Chatterjee, 2000; Proches, 2002), Liow, L. H., 2000. Mangrove conservation in Singapore: A physical and according to Chatterjee (2000), tarsi III and IV of C. or a psychological impossibility? Biodiversity and Conversation, balakrishnani bear both medial and lateral pas, and, according 9: 309-332. to Proches (2002: fig. 3G), at least tarsus III of C. caloglossae Netto, S. A. & F. Gallucci, 2003. Meiofauna and macrofauna has both a lateral and medial pas. The similarity in the general communities in a mangrove from the Island of Santa Catarina, shape may be purely accidental, comparable with the overall South Brazil. Hydrobiologia, 505: 159-170. similarity demonstrated by some species of the Ólafson, E.; S. Carlström & S. G. M. Ndaro, 2000. Meiobenthos of Copidognathus tricorneatus-group and Arhodeoporus hypersaline tropical mangrove sediment in relation to spring eclogarius-group (Bartsch, 1997b), but the four species may inundation. Hydrobiologia, 426: 57-64. also be closely related and form a species group and the Proches, S., 2002. New species of Copidognathinae (Acari: postulated differences the result of the somewhat cursory Halacaridae) from southern Africa. Journal of Natural History, descriptions of C. balakrishnani and C. caloglossae. 36: 999-1007. Proches, S., D. J. Marshall, K. Ugrasen & A. Ramcharan, 2001. Distribution. – Singapore, amongst muddy sediment and algal Mangrove pneumatophore assemblages and temporal turf on pneumatophores. patterns. Journal of the Marine Biological Association of the United Kingdom, 81: 545-552. Proches, S. & D. J. Marshall, 2002. Epiphytic algal cover and ACKNOWLEDGEMENTS sediment deposition as determinants of arthropod distribution and abundance on mangrove pneumatophores. Journal of the The study of the halacarid fauna of Singapore was financed Marine Biological Association of the United Kingdom, 82: 937- by a grant of the National University of Singapore which is 942. gratefully acknowledged. A special thank is due to Dr. Darren Sasekumar, A., 1994. Meiofauna of a mangrove shore on the west C. J. Yeo, Raffles Museum of Biodiversity Research, and coast of peninsular Malaysia. Raffles Bulletin of Zoology, 42(4): the staff of the Museum, who helped in many ways to make 901-915. my stay a success. Somerfield, P. J., J. M. Gee & C. Aryuthaka, 1998. Meiofaunal communities in a Malaysian mangrove forest. Journal of the Marine Biological Association of the United Kingdom, 78: 717- LITERATURE CITED 732. Trouessart, E., 1888. Note sur les acariens marins recueillis par M. Bartsch, I., 1990. Acarothrix palustris gen. et spec. nov. Giard au laboratoire maritime de Wimereux. Compte rendu (Halacaroidea, Acari), ein Bewohner der Salzwiesen Südchinas. hebdomadaire des séances de l’Académie des Sciences, Paris, Zoologischer Anzeiger, 224: 204-210. 107: 753-755.

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THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 93-101 Date of Publication: 28 Feb.2006 © National University of Singapore

TWO NEW SPECIES OF CYDRELA THORELL (ARANEAE: ZODARIIDAE) FROM THAILAND

Pakawin Dankittipakul Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand and The University of Auckland, Private Bag 92019, Auckland, New Zealand E-mail: [email protected]

Rudy Jocqué Section of Invertebrates, Musée Royal de l’Afrique Centrale, B-3080 Tervuren, Belgium. E-mail: [email protected]

ABSTRACT. – Two new species of the spider genus Cydrela Thorell, 1873 (Zodariidae: Cydrelinae) are described from Thailand. Cydrela pristina, new species (male + female), was collected in primary montane cloud forests near the summit of Doi Inthanon, the highest mountain culminating at 2,563 m asl. Cydrela decidua, new species (female), was collected from a mixed deciduous forest of Phu Phan, a series of mountainous ranges situated in the northeastern part of the kingdom. The species previously described from Tibet, C. linzhiensis (Hu, 2001), new combination, is here transferred from Storena Walckenaer, 1805.

KEY WORDS – Cydrelinae, Storena, new combination, Tibet, Oriental Region, Doi Inthanon, Phu Phan.

INTRODUCTION Region, especially in entirely contrasting type of habitat, is remarkable and throws a new light on the ecological range During recent fieldwork in Thailand several interesting spider of the spider subfamily Cydrelinae. The present study expands specimens were obtained from the summit of Doi Inthanon- the known distribution of the Cydrelinae towards the east and the highest mountain in the country, culminating at 2,563 m. provides a better understanding of the actual geographical The mountain consists of a number of remnant patches of range of this subfamily. primary tropical montane cloud forests. Most of the specimens examined are members of the spider family Amaurobiidae, At present six zodariid genera (Asceua Thorell, Cydrela, which is fairly common and can be considered as an abundant Euryeidon Dankittipakul & Jocqué, Heradion Dankittipakul group of spiders inhabiting evergreen hill forests of northern & Jocqué, Mallinella Strand, and Storenomorpha Simon) Thailand (Dankittipakul et al., 2006; Dankittipakul & Wang, have been reported from Thailand. The present description 2003, 2004). These forests also harbour many remarkable of new Cydrela species once more indicates that the forest species of the Zodariidae that are well adapted for living in floor fauna of Southeast Asia is still very superficially known. constantly moist forests of Southeast Asia. As a result, two new genera of the subfamily Zodariinae Simon were consequently described (Dankittipakul & Jocqué, 2004). MATERIAL AND METHODS Surprisingly, the expedition also led to a discovery of members of the subfamily Cydrelinae Simon, specifically the All illustrations were made with a Nikon SMZ 800 two Cydrela species described herein. stereomicroscope equipped with a drawing tube. Body measurements are in millimetre. Measurements of leg The Cydrelinae consists of several zodariid genera with segments were taken from the dorsal side. Epigynes were species that live in savannas, semideserts or other arid areas drawn in natural and cleared state (after immersing in lactic of the African continent. Only a few species of the genus acid for 10–20 minutes). Male palps were drawn in lateral Capheris Simon, C. escheri Reimoser and C. nitidiceps and ventral view. Structures examined with the scanning Simon, were known from India, the easternmost point in the electron microscope (JEOL LV–5400) were dried after range of the subfamily. Therefore, Cydrela Thorell was immersion in hexamethyldisilazane (Brown, 1993), stud- previously considered an African genus with fourteen nominal mounted and sputter-coated for observation and photography. species. The presence of Cydrela species in the Oriental

93 Dankittipakul & Jocqué: Two new species of Cydrela

The specimens examined are deposited in the collections of III, narrowed in front. Profile with raised cephalic area and the Muséum d’histoire naturelle, Genève (MHNG), slightly dip near fovea (Fig. 16); highest point between PME Koninklijk Belgisch Instituut voor Natuurwetenschappen, and deep fovea. Tegument smooth. Carapace light brown, Brussels (KBIN) and in the collection of the first author (PDC) silvery hairs present on cervical groove and ocular area. Legs which will be deposited in the MHNG later. dark green, posterior legs with numerous strong spines.

Abbreviations used in the text and in the figures are as follow: Eyes small, in three rows (2–2–4) (Figs. 11, 15); AER strongly ALE, anterior lateral eyes; ALS, anterior lateral spinnerets; procurved, ALE located in front of AME; PER slightly AME, anterior median eyes; LE, lateral eyes; MAp, Major recurved, much wider than anterior one; all eyes circular; ampullate gland spigot; ME, median eyes; MOQ, median AME dark, remainder pale; AME the smallest, hardly larger ocular quadrangle; MS, median septum of epigyne; PER, than other six, which are subequal; AME their diameter apart, posterior eyes row; Pir, piriform gland spigot; PLE, posterior about their diameter from ALE; PME more than their diameter lateral eyes; PLS, posterior lateral spinnerets; PME, posterior apart, at about 4–5 times that distance from PLE. MOQ longer median eyes; PMS, posterior median spinnerets; RTA, than wide, about 1.0-1.4 times anterior width, and much wider retrolateral tibial apophysis. Spination: d, dorsal; pl, behind than in front. Clypeus high, 5.2 to 5.8 times diameter prolateral; r, retrolateral; v, ventral. of ALE, vertical. Eye sizes and interdistances: AME 0.04, ALE 0.08, PME 0.09, PLE 0.08; AME–AME 0.06, AME– ALE 0.06, PME–PME 0.07, PME–PLE 0.20, ALE–PLE 0.30; TAXONOMY MOQ 0.34 long, front width 0.14, back width 0.30. Clypeus 0.46 high. FAMILY ZODARIIDAE THORELL, 1881 Chilum poorly developed, represented by lightly chitinized Cydrela Thorell, 1873 area, without hairs; double, poorly delimited. Chelicerae (Figs. 15, 16) strong, rather long; with relatively large lateral Cydrela pristina, new species condyle; anterior surface densely haired, with pale promesal (Figs. 1–12, 14–22) patch and retromesal gland (Fig. 7); no teeth present; fangs relatively long with thick base. Maxillae (Fig. 14) slightly Type material. – HOLOTYPE: Male (MHNG) THAILAND, Chiang swollen at base; rather broad, with anteromesal scopulae. Mai Province, Chomthong District, Doi [northern Thai, Doi = Mt.] Labium (Fig. 14) roughly triangular, with narrowed base. Inthanon National Park, Doi Inthanon, Kew Mae Pan, 2,000–2,100 Sternum (Fig. 14) roughly triangular, almost straight anterior m, pitfall trapping, coll. S. Sonthichai & P. Dankittipakul, 25 Mar. margin with small indentation on either side of center; slightly –22 Apr.2000. longer than wide. Precoxal sclerite small, lightly sclerotized. PARATYPES: 1 female (MHNG), 2,450 m, near the summit of Doi Inthanon, pitfall trapping, coll. S. Sonthichai & P. Dankittipakul, Leg formula 4123 (8.77, 7.70, 6.87, 6.71). Leg measurements: 25 Dec.1999 – 23 Jan.2000; 1 female (MHNG), 2,250 m, leaf litter femora: I 2.05, II 1.95, III 1.88, IV 2.30; patellae: I 0.91, II sifting, coll. S. Sonthichai & P. Dankittipakul, 15 Jan.2000; 5 females 0.94, III 0.90, IV 1.06; tibiae: I 1.80, II 1.40, III 1.20, IV (MHNG, PDC ZO–00159/60), 2,540 m, leaf litter sifting, coll. P. 1.60; metatarsi: I 1.56, II 1.48, III 1.74, IV 2.40; tarsi: I 1.38, Dankittipakul, 4–7 Jul.2002; 1 male (MHNG TH–00/03), 2,500 m, II 1.10, III 0.99, IV 1.41. Spination: spines usually few and leaf litter sifting, coll. P. J. Schwendinger, 28 Oct.2000; 1 female short on legs I and II, more numerous long, slender spines on (KBIN), 1,630 m, coll. P. J. Schwendinger, 25 Feb.1986; 3 females legs III and IV. Hinged hairs present. Metatarsal preening (MHNG), 1,650–1,680 m, leaf litter sifting, coll. T. Chami-Kranon bush present on leg III. Trichobothria in 2 rows on tibiae, in & A. Hasalem, 12 Jan.2005. 1 row on metatarsi and tarsi. Scopulae rarely present, mostly Diagnosis. – The male of C. pristina, new species can be absent or spiniform. Pair tarsal claws with 8 teeth. distinguished from other members of the genus Cydrela by the bifid tibial apophysis (Figs. 9, 20), the lightly sclerotized Spination: femora I d1–1–1 pl1 II d1–1–1 pl1 rl1 III d1–1– conductor (Figs. 17, 20) on the retrolateral side of the 1 pl2 rl2 IV d1–1–1 rl1; patellae II pl1 III d1 pl2 rl1 IV d1 membranous tegulum (Fig. 17) and the elongate embolus pl1 rl1; tibiae I pl1–1–1 v2–1–2 II pl1–1–1 v2–2–1 III d1–1– (Figs. 17, 19). Females can be recognized by the presence of 1 pl1–1–1 rl1–1 v2–2–2 IV d1–1–1 pl1–1–1 rl1–1 v2–1–1– a median septum and a hood on the epigyne (Figs. 10, 21), 2; metatarsi I v3–2–2 II pl1 v3–2–2 III d1 pl1–1 rl1–1 v3–2– the small, anteriorly located copulatory orifices (Figs. 10, 21) 2 IV d2–1 pl1–1–1 rl1–1 v2–1–1–1–1–3 tarsi II v1–1–1 III and the strongly sclerotized spermathecae (Fig. 22). v1–1–1–1–1–1 IV v2–1–1–1–1.

Etymology. – The species name is a Latin adjective meaning Abdomen (Fig. 11) oval, with two dorsal sigilla; sparsely “of former times” referring to the pristine forests the clothed with white hairs. Dorsum gray, provided with simple specimens were found in. pattern consisting of two longitudinal bands, followed by pairs of pale patches, fused just in front of the spinnerets. Tracheal Description. – Male (holotype). Total length 6.97. Carapace spiracle narrowed, lightly sclerotized. Six spinnerets (Figs. 3.61 long, 2.32 wide. Abdomen 3.35 long. 1, 3-5): Anterior pair relatively long, cylindrical or slightly conical, clearly bisegmented; PLS slightly longer than half Prosoma (Fig. 11) longly oval, widest between coxae II and length of ALS, cylindrical, faintly bifurcated; PMS conical,

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Figs. 1-6. Cydrela pristina, new species: 1, spinnerets of male; 2, spinnerets of female; 3, anterior lateral spinnerets; 4, anterior lateral spinnerets, different view; 5, posterior lateral spinneret with minor ampullate gland spigot; 6, female palpal tarsi. Abbreviations: Pir, piriform gland spigot; MAp, Major ampullate gland spigot.

95 Dankittipakul & Jocqué: Two new species of Cydrela tiny, provided with few spigots. Colulus represented by two from fovea to PME; chelicerae densely clothed with strong groups of hairs. Epiandrum well developed, provided with hairs. Spinnerets as in Fig. 2. Female palp (Fig. 6) swollen; two sclerotized depressions. triangular tarsi ventrally flattened; provided with large terminal claw and pairs of spines on prolateral side (Fig. 6). Male palp (Figs. 9, 17–20): Tibia with a large and protruding retrolateral tibial apophysis (Fig. 17), slightly bifid as seen Eye sizes and interdistances: AME 0.06, ALE 0.10, PME from in front (Figs. 9, 20), dorsally with small triangular 0.10, PLE 0.10; AME–AME 0.07, AME–ALE 0.06, PME– appendage curved inward (Fig. 18). Cymbium with few spines PME 0.10, PME–PLE 0.28, ALE–PLE 0.44; MOQ 0.26 long, on distal part; retrolateral side protruding (Fig. 20), front width 0.18, back width 0.30. Clypeus 0.61 high. accommodating the conductor; cymbial fold absent. Subtegulum well-developed. Tegulum (Fig. 17) simple, Leg formula 4132 (8.95, 7.15, 6.90, 5.28). Leg measurements: widened at extremity, posterior portion lightly sclerotized; femora: I 2.13, II 1.95, III 2.18, IV 2.37; patellae: I 1.00, II anterior part forming a groove, accommodating elongate 1.00, III 1.02, IV 1.08; tibiae: I 1.68, II 1.40, III 1.24, IV embolus, originating on lateral base of tegulum. 1.72; metatarsi: I 1.32, II 1.40, III 1.60, IV 2.50; tarsi: I 1.02, II 0.93, III 0.86, IV1.28. Female (paratype). Total length 8.61. Carapace 4.0 long, 2.50 wide. Abdomen 4.61 long. Spination: femora I d1–1–1 pl1 II d1–1–1 pl1 III d1–1–1 pl1 rl1 IV d1–1–1 rl1; patellae III d1–1 pl1–1 rl1 IV d1–1 pl1– Coloration and pattern similar to male but usually with darker 1 rl1; tibiae I pl1 v1–1 II pl1 v1–1 III d1–1–1 pl1–1–1 rl1– (Fig. 12), smaller females usually pale, mostly orange; legs 1 v2–1–1–1–1 IV d1–1 pl1–1–1 rl1–1 v2–1–1–1–1; metatarsi of normal length; carapace slightly wider in front; carapace I v2–1 II v2–2–3 III d1 pl1–1–1 rl1–1–1 v2–2–3 IV d1 pl1– and opisthosoma without silvery hairs; a row of hairs running 1–1–1 rl1–1–1 v1–1–1–1–3 tarsi III v2–2–2 IV v2–2.

Figs 7-10. Cydrela pristina, new species: 7, cheliceral gland; 8, tarsal trichobothrium; 9, right male palp, retrolateral view; 10, female epigyne, ventral view.

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Epigyne (Figs. 10, 21, 22): Simple, strongly sclerotized plate, It is interesting to note that only four specimens examined provided with median septum and hood (Figs. 10, 21). Small, were collected from evergreen hill forest at about 1,630–1,680 copulatory orifices anteriorly situated (Fig. 21). Copulatory m asl. These specimens were obtained by leaf litter sifting in ducts short, strongly sclerotized, leading to round, posteriorly a damp swamp between steep ridges in evergreen hill forest. located spermathecae (Fig. 22). Fertilization ducts simple. At this altitude forest litter of only three square metres collected near the swamp contained three females and two Natural history.- Most of the specimens of C. pristina, new juveniles whilst the remaining 47 square metres of species examined were collected from primary tropical decomposing leaf litter layer nearby were inhabited by other montane cloud forests (2,000–2,540 m) near the summit of zodariid genera, i.e. Asceua, Euryeidon and Mallinella. Doi Inthanon. The forest floor was covered with a thick layer (9–22 cm depth) of decomposing organic matter. The annual Distribution.- Known only from the type locality and mean temperature of these forests is 9 and 12 ºC at the summit surroundings. and Kew Mae Pan, respectively. The constantly low temperature is responsible for the formation of the daily cloud cover above 1,800 m of the mountain. The annual rainfall is Cydrela decidua, new species over 2,300 mm where the spiders were collected. During the (Figs. 13, 23, 24) rainy season (July to late October) relative humidity of the air reaches 100 % and the soil becomes saturated with water. Type material. – HOLOTYPE: Female (MHNG), northeastern Additional physical factors of the type and other localities THAILAND, border between Kalasin and Sakon Nakhon Province, were also reported by Dankittipakul & Sonthichai (2002). Phu [= Mt.] Phan, 524 m, partially dry stream bed in a mixed

Figs. 11-13. Cydrela pristina, new species: 11, male holotype, habitus, dorsal view; 12, female paratype, dorsal view of abdomen. Cydrela deciduas, new species. 13, female holotype, habitus, dorsal view. Scale bar = 1.0 mm.

97 Dankittipakul & Jocqué: Two new species of Cydrela evergreen deciduous gallery forest ca. 1 km of the main road near and slightly dip near fovea; highest point half way between the summit, coll. P. J. Schwendinger, 7-8 Dec.1995. PME and deep fovea. Tegument smooth. Carapace orange in colour, with a few long hairs on ocular area. Legs yellow, Diagnosis. – Somatic features of the female C. deciduas, new anterior legs provided with a few weak spines, posterior legs species resemble that of C. pristina, new species but can be with numerous short ones. easily distinguished from the latter species by its smaller size and an epigyne with large genital orifices (Fig. 23), the Eyes small, in three rows (2–2–4); AER strongly procurved, internal structure of the epigyne with strongly convoluted ALE located just in front of AME; PER slightly recurved, entrance ducts and the kidney-shaped spermathecae (Fig. 24). much wider than anterior one, PER clearly separated from AER; all eyes circular; AME dark, remainder pale; AME the Etymology. – The species name is a Latin adjective meaning smallest, hardly larger than other six, which are subequal; “fallen” and refers to the semi-deciduous woodland the AME their diameter apart, about their diameter from ALE; specimen was found in. PME more than their diameter apart, at about 3 times that distance from PLE. MOQ longer than wide and always much Description. – Female (holotype). Total length 6.05. Carapace wider behind than in front. Eye sizes and interdistances: AME 2.46 long, 1.51 wide. Abdomen 3.14 long. 0.06, ALE 0.08, PME 0.06, PLE 0.06; AME–AME 0.04, AME–ALE 0.08, PME–PME 0.08, PME–PLE 0.20, ALE– Prosoma longly oval, widest between coxae II and III, PLE 0.30; MOQ 0.30 long, front width 0.14, back width 0.20. strongly narrowed in front. Profile with raised cephalic area Clypeus 0.26 high, convex, strongly retreating.

Figs. 14-16. Cydrela pristina, new species. Sternum (14); carapace, lateral (15) and frontal (16) views. Scale bar = 1.0 mm.

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Figs. 17-22. Cydrela pristina, new species. Male holotype (17-20) and female paratype (21, 22). Male palp: ventral (17), dorsal (18), prolateral (19) and retrolateral (20) views. Female epigyne (21) and its internal structure (22). Scale bars = 0.5 mm.

Figs. 23, 24. Cydrela decidua, new species. Female holotype (23, 24). Epigyne (23), and its internal structure (24). Scale bars = 0.5 mm.

99 Dankittipakul & Jocqué: Two new species of Cydrela

Chilum indistinct. Chelicerae strong, rather broad; anterior Remarks. – Cydrela linzhiensis (Hu, 2001), new combination surface densely clothed with a patch of very thick hairs; fangs is considered a member of the Cydrelinae due to its peculiar relatively long with thick base, no teeth present on cheliceral eye arrangement (2–2–4), an elevated pars cephalica of the groove. Maxillae slightly swollen at base; rather broad, with carapace, narrowing in front and the presence of a basolateral anteromesal scopulae. Labium roughly triangular, with extension on the maxillae. Cydrela linzhiensis, new narrowed base. Sternum roughly triangular, almost straight combination can be recognized by its very short, tiny, anterior anterior margin with small indentation on either side of center; pin-shaped embolus situated near the apex of the tegulum slightly longer than wide. Precoxal sclerite absent. and a dorsally concave retrolateral tibial apophysis. Posterior spinnerets divided into two segments, not in one piece as Leg formula 4123 (5.60, 4.84, 3.84, 3.73). Leg measurements: illustrated in Figs.14.1 and 14.4 given by Hu (2001). femora: I 1.42, II 1.21, III 1.02, IV 1.56; patellae: I 0.56, II 0.56, III 0.48, IV 0.56; tibiae: I 1.50, II 0.83, III 0.75, IV 1.05; metatarsi: I 0.65, II 0.65, III 0.90, IV 1.52; tarsi: I 0.71, ACKNOWLEDGMENTS II 0.62, III 0.58, IV 0.91.Spination: Spines usually few and thin on legs I and II, more numerous short, stout spines on We are grateful to Dr Peter J. Schwendinger for providing legs III and IV. Metatarsal preening bush present on leg III material from his personal collection and for the loan of absent. Scopulae absent. Pair tarsal claws with tiny teeth, specimens from the MHNG. Dr Xin-Ping Wang (California generally more than 10. Academy of Sciences) and Professor Shuqiang Li (Institute of Zoology, Chinese Academy of Sciences) kindly provided Spination: femora I d1–1 II d1–1 III d1 IV d1; patellae III literature and shared their expertise on Chinese spiders. Jim pl1–1–1 d1 rl 1 ap 2 IV pl1–1–1 d1 rl1 ap 2; tibiae I v1 II pl1 Xu (The University of Auckland, UOA) translated the v1–1–1 III pl1–1–1 d3 rl 2 v2 IV pl1–1–1 d3 rl2 v1–1–1; Chinese text. Alan D. Newson (UOA) kindly improved the metatarsi I v2–1 II v4–2–1 III pl1–1–1 d1 rl2 v4–1–1 IV pl1– English text. Work on this paper was carried out while the 1–1–1 d1 rl3 v4–1–1–1. first author was based at Chiang Mai University (CMU). He is grateful to people who provided support, in particular to Abdomen oval. Dorsum gray, provided with a simple pattern Professor Saowapa Sonthichai (CMU), Dr Angoon consisting of two pairs of spots, anterior one largest, followed Lewvanich (The Royal Academy of Thailand, Bangkok), Dr by a pale folium. Six spinnerets. Anterior pair relatively long, Wipada Vungsilabutr (Department of Entomology and cylindrical or slightly conical, bisegmented; posterior Zoology, Ministry of Agriculture, Bangkok) and staffs of the spinnerets much shorter than AS. Colulus represented by two Doi Inthanon National Park. groups of hairs. The Royal Forest Department gave permission to collect Epigyne (Figs. 23, 24): Simple sclerotized plate with two large spiders in protected areas. Thailand Research Fund TRF/ posterior genital orifices (Fig. 23) connected to a pair of BIOTEC Special Program for Biodiversity Research and receptacles (Fig. 24) then forming thick copulatory ducts Training (project number BRT_T 542094 and BRT_R which arise posteriorly; spermathecae kidney-shaped (Fig. 145002) supported the first author at the beginning of this 24). project.

Natural history. – The female holotype was collected from a 2–3 cm long burrow with a flap-shaped door (not a real LITERATURE CITED trap door) that closed the entrance of the burrow. The burrow was located on a vertical road bank of a main road in a dry Brown, D., 1993. A further chemical alternative to critical-point deciduous forest. Hewitt (1914) first observed that an African drying for preparing small (or large) flies. Fly Times, 11: 10. species, C. friedlanderae Hewitt, 1914, live in burrows with Dankittipakul, P. & R. Jocqué, 2004. Two new genera of the trap doors but the species was believed to be a member of Zodariidae (Araneae) from Southeast Asia. Revue suisse de Capheris instead of Cydrela (Jocqué, 1991). Several members Zoologie, 111(4): 749–784. of the genus Psammorygma Jocqué, another genus of the Dankittipakul, P., T. Chami-Kranon & X.-P. Wang, 2005. New and Cydrelinae, have been recorded (unpublished) to live in poorly known species of coelotine spiders (Araneae, burrows with trap doors. We document here the evidence of Amaurobiidae) from Thailand. Zootaxa, 970: 1–11. burrow construction of the genus Cydrela. Dankittipakul, P. & S. Sonthichai, 2002. Spiders of Doi Inthanon National Park. In: Baimai, V. (ed.), BRT Research Reports 2002. Distribution. – Known only from the type locality. Biodiversity Research and Training Program, Bangkok. Pp. 140– 147. Dankittipakul, P., S. Sonthichai & X.-P. Wang, 2006. Ten new Cydrela linzhiensis (Hu, 2001), new combination species of coelotine spiders from Thailand. Revue suisse de Zoologie, 113(1): in press. Storena linzhiensis Hu, 2001: 92, Figs. 14.1–6 (description male). Dankittipakul, P. & X.-P. Wang, 2003. New species of coelotine spiders from northern Thailand I. Revue suisse de Zoologie, 110 Type material. – Holotype: Male, Tibet, Linzhi Province, 3,000 m, (4): 723–737. 13 May.1987 (not examined).

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Dankittipakul, P. & X.-P. Wang, 2004. New species of coelotine Hu, J. L., 2001. Spiders in Qinghai-Tibet Plateau of China. Henan spiders from northern Thailand II. Revue suisse de Zoologie, Science and Technology Publishing House. 658 pp. 111(3): 539–550. Jocqué, R., 1991. A generic revision of the spider family Zodariidae Hewitt, J., 1914. Descriptions of new Arachnida from South Africa. (Araneae). Bulletin of the American Museum of Natural History, Records of Albany Museum Grahamstown, 3: 1–37. 201: 1–160.

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THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 103-117 Date of Publication: 28 Feb.2006 © National University of Singapore

THREE NEW AND FOUR NEWLY RECORDED SPECIES OF LINYPHIINAE AND MICRONETINAE SPIDERS (ARANEAE: LINYPHIIDAE) FROM NORTHERN VIETNAM

Lihong Tu1,2 & Shuqiang Li1,* 1Institute of Zoology, Chinese Academy of Sciences, Beijing100080, P. R. China 2College of Life Science, Capital Normal University, Beijing100037, P. R. China E-mail: [email protected] (*Corresponding author)

ABSTRACT. – Seven species belonging to six linyphiine and micronetine genera collected from northern Vietnam are diagnosed, described and illustrated. Among them, three species are new to science: Bathyphantes floralis, new species, Neriene oxycera, new species, and Nesioneta ellipsoidalis, new species. The species Prosoponoides sinensis (Chen, 1991), new combination, Microbathyphantes aokii (Saito, 1982), new combination, Parameioneta bilobata Li & Zhu, 1993, and Neriene cavaleriei (Schenkel, 1963) are newly recorded from Vietnam. Microbathyphantes aokii is also considered here as a senior of Bathyphante dipetalus Chen & Yin, 2000, new synonymy.

KEY WORDS. – Linyphiidae, Linyphiinae, Micronetinae, taxonomy, new species, Vietnam.

INTRODUCTION illustrations were made using a drawing tube. Male palps and female epigyna were examined and illustrated after they were This is the fifth publication on taxonomic study of spiders dissected from the spider’s bodies. Vulvae were cleared in and small crustaceans from northern Vietnam based on the boiling KOH solution to dissolve nonchitinous tissues, and material collected by the second author in December 2000 the embolic divisions of male palps were excised by breaking (Hou & Li 2003; Peng & Li 2003; Tu & Li 2004; Li & Liang the column (the membranous connection between the 2002). This survey began at 4 December 2000, and conducted suprategulum and the radix). For examination of the genital from Ha Jiang to Cao Bang Provinces, then along Lang Son structures under a compound microscope, male palps and Province to Hanoi. After 22 December, the field collection epigyna were immersed in 75% alcohol solution, while was made in a research station of Hanoi University in Son embolic divisions and vulvae were mounted in Hoyer’s Toy Province, which located about 70km north of Hanoi. This Solution. survey finished in 24 December 2000. For each species, only the name that appeared in the original Linyphiinae and Micronetinae spiders are one of the largest description, new synonyms and new combinations are listed. groups of small spiders within the araneoid family Synonyms listed in Platnick’s spider catalogue (Platnick, Linyphiidae. It is recorded that there have been about 819 2005) are not repeated here. Information about the distribution linyphiines and 1114 micronetines species from the world of each species in Vietnam is provided at the provincial level. (Tanasevitch, 2004), but none has been reported from If not otherwise stated the material examined for this paper Vietnam before. As Vietnam is known as very diverse area is deposited in the Institute of Zoology, Chinese Academy of in the temperate regions of the northern hemisphere, several Sciences in Beijing, China (IZCAS); abbreviations for other hundreds species of linyphiine and micronetine spiders are depositaries are: Muséum d’histoire naturelle (MHNG), expected to be discovered from there. More survey and further Genève, Switzerland; Zoological Museum of Turku study on linyphiid spiders of Vietnam are expected in the University, Finland (MZT) and Hunan Normal University future. (HNU), Changsha, China.

The leg measurements are given in the following sequence: MATERIAL & METHODS total (femur, patella+tibia, metatarsus, tarsus). All measurements are in millimeters. Due to without a uniform Specimens were examined and measured under an SZ11- terminology for homologous structures of Linyphiinae and Olympus stereomicroscope. Further details were studied Micronetinae, terminology for the somatic morphology and under an Olympus BX40 compound microscope. All genital structures of Linyphiinae is after van Helsdingen

103 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

(1969), and those of Micronetinae is after Saaristo & (Fig. 1A). Sternum slightly darker than carapace. Abdomen Tanasevitch (1996). Abbreviations used as followed: gray, with white and black spots. Lengths of legs: I 3.80 (0.93+1.17+1.00+0.70), II 3.50 (0.93+1.07+0.90+0.60), III Somatic characters: AER- anterior eye row; ALE- anterior 2.54 (0.67+0.77+0.60+0.50), length of leg IV unknown. All lateral eye; AME- anterior median eye; AME-ALE- distance tibia with two dorsal spines, Ti I additionally with one between AME and ALE; AME-AME- distance between prolateral and one retrolateral spine, Ti II with one retrolateral AMEs; AMEd- diameter of AME; AMEr- radius of AME; spine. Fe I and II with small spine dorsally. Tm I: 0.28. Tm PER- posterior eye row; PLE- posterior lateral eye; PME- IV unknown. posterior median eye; PMEd- diameter of PME; PME-PLE- distance between PME and PLE; PME-PME- distance Palp (Figs. 1B-C). Patella with long dorsal bristle. Tibia between PMEs. unmodified with one prodorsal, two retrodorsal trichobothria. Paracymbium J-shaped, distally narrowing with blunt apex. Male palp: ALP- anterior projection of LC; ATA- anterior Sperm duct in tegulum with three turnings in retrolateral view. terminal apophysis; DEA- dorsal embolic apophysis; DLP- Pit-hook on distal part of suprategulum bifid, one point, one dorsal projection of LC; DTA- dorsal tibia apophysis; E- stout, both strongly sclerotized (Fig. 1D). Embolic division embolus; EM- embolic membrane; EP- embolus proper; LC- (Fig. 1D): radix small and simple, somewhat S-shaped; lamella characteristica; LLP- lateral projection of LC; MTA- embolus spiraling forwards, spiral axis parallel with that of median terminal apophysis; P- paracymbium; PCA- proximal palp; embolic membrane extending out through the spiraling cymbial apophysis; PEA- posterior embolic apophysis; PH- embolus, flower-shaped, with rotation at stem portion and pit-hook on distal part of suprategulum; PLP- posterior thread-like projections on prolateral surface of apical portion projection of LC; PTA- posterior terminal apophysis; R- (Figs. 1F-G); two terminal apophyses in ventral view, the radix; SPT- suprategulum; TA- terminal apophysis; TH- outer one membranous, quadrate and slightly convex, its inner thumb of embolus; TRS- transversal sclerite; VEA- ventral side molar-shaped with many dentils, the inner one strongly embolic apophysis. sclerotized; lamella characteristica (Figs. 1C, 1E) large triangular plate-shaped, dorsal projection pointed, anterior Epigynum: DPS- distal portion of scape; EG- entrance groove; projection extending, turning with hooked apex; long thread- FG- fertilization groove; LD- depression of lateral plate; like membrane arising from inner surface of lamella LWS- lateral wing of scape; PMP- posterior median plate; characteristica. MPS- median portion of scape; PPS- proximal portion of scape; RMP- ridge of median plate; S- spermatheca; SC- Description of female. – Total length: 2.17. Carapace: 0.85 scape; TP- turning point. long, 0.57 wide. Abdomen: 1.33 long, 0.90 wide. Carapace grayish brown, with black margin. Abdomen gray, without spots. Lengths of legs: I 3.50 (0.90+1.03+0.97+0.60), II 3.26 TAXONOMY (0.83+1.03+0.80+0.60), III 2.47 (0.67+0.80+0.60+0.40), IV 3.25 (0.90+1.07+0.73+0.55). Tm I: 0.17. Tm IV absent. Other Bathyphantes floralis, new species somatic characters same as male. (Fig. 1) Epigynum (Figs. 1H-J). Ventral wall simple and convex, Type specimens. – Holotype: male, Van Hoa Village, Son Tay folding at middle. Long, spiraling entrance grooves partly Province, Bavi District, Vietnam (131), 23 Dec.2000. Paratypes: 1 visible through it. Fingerlike scape arising from dorsal wall, male and 1 female, same data as for holotype; 1 female, Van Hoa visible in ventral view, furnished with pit at distal end. Village, Son Tay Province, Bavi District, Vietnam (142), 23 Dec.2000; 1 female, Gao Bao Village, Ha Jiang Province, Vietnam Etymology. – The specific name comes from Latin floralis (017), 9 Dec.2000. (flower-like) and refers to the shape of embolic membrane. Diagnosis. – The male of Bathyphantes floralis, new species, can be easily distinguished from other Bathyphantes by the Distribution. – Vietnam (Son Tay, Ha Jiang). shape of the lamella characteristica (Fig. 1E), the bifurcate pit-hook on the distal part of suprategulum (Fig. 1D) and by the flower-shaped embolic membrane (Figs. 1F-G). The Microbathyphantes aokii (Saito, 1982), female can be identified by the long, conspicuously diverging new combination and spiraling entrance grooves (Fig. 1J). (Fig. 2)

Description of male. – Total length 2.17. Carapace: 0.87 long, Bathyphantes aokii Saito, 1982: 34, Figs. 3-5, 10. Bathyphantes dipetalus Chen & Yin, 2000: 89, Figs. 21-27 (new 0.67 wide. Abdomen: 1.30 long, 0.57 wide. Carapace pale synonymy). gray. Eyes with black surroundings, AMEs smallest, others subequal; AER recurved, AME-AME shorter than AMEd, Material examined. – 1 male and 6 females, Viet Lann Village, Ha AME-ALE longer; PER straight, PME-PME shorter than Jiang Province, Vietnam (042), 10 Dec.2000. PMEd, PME-PLE longer; ALE and PLE juxtaposed. Chelicerae whitish brown, stridulatory ridges absent, fang Additional type material examined. – 1 male (HNU), holotype of groove with three promarginal and four retromarginal teeth Bahtypnantes dipetalus Chen & Yin, 2000, Daoxian County, Hunan

104 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Fig. 1. Bathyphantes floralis, new species: A, left chelicera, frontal view; B, left male palp, retrolateral view; C, ditto, prolateral view; D, embolic division, ventral view; E, lamella characteristca, prolateral view; F, embolic membrane, retrolateral view; G, ditto, prolateral view; H, epigynum, ventral view; I, ditto, lateral view; J, vulva, dorsal view. Scale bars = 0.05 mm.

105 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

Fig. 2. Microbathyphantes aokii (Saito, 1982): A, left chelicera, frontal view; B, left male palp, retrolateral view; C, ditto, prolateral view; D, embolic division, ventral view; E, epigynum, ventral view; F, ditto, lateral view; G, vulva, dorsal view. Scale bars = 0.1 mm.

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Province, China, 19 Aug.1987; 1 female (HNU), paratype of Microbathyphantes aokii (Saito, 1982), new combination, can Bahtypnantes dipetalus Chen & Yin, 2000, same data as holotype. be distinguished from M. asiaticus and M. spedani by the rolling and pointed apex of anterior projection of lamella Diagnosis. – The male of M. aokii can be distinguished from characteristica, which was not bifurcated as both in M. other Microbathyphantes by the anterior projection of lamella asiaticus and M. spedani according to the illustrations by van characteristica which was ribbon-shaped with rolling and Helsdingen. Furthermore, the study on types of Bathyphantes pointed apex (Figs. 2B-D). dipetalus Chen & Yin, 2000, shows that Microbathyphantes aokii (Saito, 1982) is a senior synonym of Bathyphante Description of male. – Total length: 1.90. Carapace: 0.90 dipetalus Chen & Yin, 2000. long, 0.68 wide. Abdomen: 1.00 long, 0.73 wide. Carapace pale brown. Eyes with black surroundings; AME smaller, Distribution. – Vietnam (Ha Jiang), China, Japan. others subequale; AER recurved, PER straight, eyes separated by about AMEd, ALE and PLE juxtaposed. Chelicerae brown; stridulatory ridges absent; fang groove with three promarginal Neriene cavaleriei (Schenkel, 1963) and three retromarginal teeth (Fig. 2A). Lengths of legs: I (Fig. 3) 3.85 (1.10+1.30+0.75+0.70), II 3.73 (1.07+1.23+0.83+0.60), III 2.94 (0.87+0.90+0.67+0.50), IV 3.80 Linyphia cavaleriei Schenkel, 1963: 119, Fig. 71. (1.00+1.17+0.90+0.73). Only femur I with short dorsal spine. Neriene cavaleriei van Helsdingen, 1969: 153, Figs. 200-204. All tibia with two dorsal spines; tibia I with additional one prolateral and one retrolateral spine; tibia II with one Material examined. – 1 male and 1 female, Sac Ha Village, Cao retrolateral spine. Tm I: 0.27. Tm IV absent. Abdomen gray, Bang Province, Vietnam (075), 17 Dec.2000; 1 female, Sac Ha without spots. Village, Cao Bang Province, Vietnam (069), 17 Dec.2000; 1 female, Van Hoa Village, Sac Tay Province, Bavi Distr., Vietnam (128), 24 Dec.2000. Male palp (Figs. 2B-C). Tibia with one prodorsal, two retrodorsal trichobothria. Cymbium unmodified. Diagnosis. – The male N. cavaleriei can be recognized by Paracymbium J-shaped with short hairs on basal part, slightly the shape of paracymbium (Fig. 3C) and by the longer tapering off distally. Tegulum with membranous extension terminal apophysis that with more than five coils (Fig. 3I). covering apical part of embolus. Embolic division (Fig. 2D) The female can be distinguished by the bell-shaped epigynum characters by well-developed lamella characteristica with four with two rows of translucent areas (Fig. 3J) and vulva with projections: lateral one sharp triangular, anterior margin five parallel spiral grooves (Fig. 3K). roughened by small warts; dorsal one blunt and turning inwards; posterior one translucent and hooked; anterior one Description of male. – Total length: 5.04. Carapace: 2.02 longest, ribbon like and extending forwards with apex point long, 1.51 wide. Abdomen: 3.15 long, 1.01 wide. Carapace and slightly rolling. Embolus smooth coil more than one brown; cephalic portion slightly elevated. PME biggest, circle, then extending forwards, companied by the anterior situated on black tubercles, others subequal. AER recurved, extension of lamella characteristca. AME-AME about AMEd, AME-ALE longer; PER slightly procurved, PME-PLE about PMEd, PME-PME longer; ALE Description of female. – Total length: 1.80. Carapace: 0.77 and PLE juxtaposed. Chelicerae brown, stridulatory ridges long, 0.80 wide. Abdomen: 1.13 long, 0.87 wide. Lengths of absent, promargin of fang groove with six teeth, but the legs: I 3.30 (0.90+1.00+0.83+0.57), II 3.14 biggest one not on the same line with the other five small (0.83+0.97+0.77+0.57), III 2.44 (0.67+0.70+0.57+0.50), IV ones, retromargin with six teeth (Fig. 3B). Lengths of legs: 3.23 (0.90+1.00+0.80+0.53). Tm I: 0.24. Tm IV absent. Other I 12.10 (2.96+3.53+3.72+1.89), II 9.83 somatic characters same as in male. (2.58+2.84+2.96+1.45), III 6.74 (1.89+1.95+1.95+0.95), IV 9.51 (2.77+2.58+2.90+1.26). Each patella and tibia with two Epigynum (Figs. 2E-F). Doorframe-shaped in ventral view, dorsal spines, Ti I-II additionally with one prolateral and one posterior margin elevated and incised beside posterior median retrolateral spine, Ti III-IV with one prolateral spine. Fe I-IV plate. Posterior median plate concave centrally and two ridges with small dorsal spine. Tm I: 0.18. Tm IV absent. Abdomen arising from lateral side towards center. Entrance grooves cylindriform with conspicuous posterodorsal tubercle above (Fig. 2G) start before the middle ridges, running either way spinnerets; dark gray with whitish spots laterally and dorsally, in a half circle up to the conspicuous turning points, then ventral surface uniformly black. another half circle reversely and entering spermatheca anterally. Fertilization grooves running along both lateral Male palp (Figs. 3C-E). Patella short, with long spine dorsally. sides and ended behind the middle ridges. Tibia wider than long with several long spines and one prodorsal, two retrodorsal trichobothria. Cymbium Remarks. – Compared with the illustrations of unmodified. Paracymbium somewhat J-shaped with short Microbathyphantes asiaticus van Helsdingen, 1985, and M. hairs on baslal part, distal arm well-developed with two free spedani (Locket, 1968) provided by van Helsdingen (1985, ends, one triangular pointing backwards, one more sharper, Figs. 11-17), the genital structures of the current species show pointing forwards. Pit-hook on distal part of suprategulum striking resemblance to that of Microbathyphantes, and as normal shape of Neriene hammeni group (van Helsdingen, herewith transfer it to genus Microbathyphantes. 1969), L-shapped with hooked apex (Fig. 3F). Embolic

107 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

Fig. 3. Neriene cavaleriei (Schenkel, 1963): A, female left chelicera, frontal view; B, male left chelicera, frontal view; C, male left palp, retrolateral view; D, ditto, ventral view; E, ditto, prolateral view; F, distal part of suprategulum; G, embolic division, ventral view; H, embolus, dorsal view; I, terminal apophysis, lateral view; J, epigynum, ventral view; K, vulva, dorsal view. Scale bars = 0.1 mm.

108 THE RAFFLES BULLETIN OF ZOOLOGY 2006 division (Fig. 3G) prominently characters by large lamella Tibia almost as long as cymbium, with several long spines characteristica in prolateral view, with slender and sharp on lateral and ventral surfaces, and one prodorsal, two posterior tip, blunt anterior tip, point tip at end of lateral retrodorsal trichobothria. Paracymbium J-shaped, basal part projection and additional tooth-like projection on dorsal plate-shaped with short hairs, distal arm narrow, curved, margin. Terminal apophysis conspicuously long with more pointing forwards with cluster of thin hairs at tip. Pit-hook than five coils (Fig. 3I), basal coil saucer-shaped, others with on distal part of suprategulum long, extending out of thin transversal grooves. Embolus visible at base of terminal cymbium, bending laterally. Embolic division (Fig. 4F) more apophysis covered by embolic membrane, with falciform simple than that of most Neriene: Lamella characteristica with appendage distally which sclerotized and serrated at basal long, slender lateral projection and posterior projection part of anterior margin (Fig. 3H). Transversal sclerite pointing forward and backward respectively; short, blunt somewhat rectangular, lying between lateral projection of anterior projection and dorsal projection. Terminal apophysis lamella characteristica and base of terminal apophysis. short, twisted element with no more than half coil. Embolus covered by embolic membrane with thumb-like appendage Description of female. – Total length: 5.29. Carapace: 2.02 apically, embolus proper blunt. long, 1.51 wide. Abdomen: 3.15 long, 1.01 wide. Lengths of legs: I 11.03 (2.90+3.34+3.15+1.64), II 8.95 Female unknown. (2.46+2.65+2.52+1.32), III 6.04 (1.76+1.76+1.70+0.82), IV 8.69 (2.58+2.46+2.52+1.13). Tm I: 0.20. Tm IV absent. Other Etymology. – The specific name comes from Latin oxycerus somatic characters same as male. (sharp) and refers to the sharp lateral and posterior apophyses of lamella characteristica. Epigynum (Fig. 3J). In ventral view, ventral plate bell-shaped, with two rows of translucent areas. Posterior margin deeply Remarks. – Although the embolic division of this new species concave in median and convex laterally. Scape arising from is relatively simple, the pattern is reminiscent of the a Neriene dorsal wall, turning up ventrally, with pit at tip. Vulva (Fig. male palp. The general morphology of male palp, particularly 3K) as long as wide, fertilization grooves started from ventral the shape of embolus, which is similar to that of N. emphana, wall, with five parallel coils, turning points laterally of apices, may show some relationship between them. However, neither pointing inwards. terminal sclerite nor transversal sclerite exist in this new species that suggest that it does not belong in the N. emphana Distribution. – Vietnam (Cao Bang, Sac Tay), China. group.

Distribution. – Known only from the type locality. Neriene oxycera, new species (Fig. 4) Nesioneta ellipsoidalis, new species Type specimens. – Holotype: male, Van Hoa Village, Son Tay (Figs. 5, 6) Province, Bavi District, Vietnam (142), 23 Dec.2000. Type specimens. – Holotype: male, Tan Linh Village, Son Tay Diagnosis. – Male of Neriene oxycera, new species, can be Province, Bavi District, Vietnam (138), 24 Dec.2000. Paratypes: 6 distinguished by the shape of lamella characteristica and the males and 9 females, Tan Linh Village, Son Tay Province, Bavi simple terminal apophysis (Figs. 4C-F). District, Vietnam (138), 24 Dec.2000; 1 female, Ha Jiang Town, Ha Jiang Province, Vietnam (006), 8 Dec.2000; 3 females, Viet Lann Description of male. – Total length: 2.90. Carapace: 1.37 Village, Ha Jiang Province, Vietnam (022), 10 Dec.2000; 5 females, long, 1.00 wide. Abdomen: 1.60 long, 0.90 wide. Carapace Viet Lann Village, Ha Jiang Province, Vietnam (023), 10 Dec.2000; 1 female, Viet Lann Village, Ha Jiang Province, Vietnam (026), 10 grayish brown, unmodified. Eyes (Fig. 4A) subequal. AER Dec.2000; 1 male and 1 female, Viet Lann Village, Ha Jiang recurved, intervals between eyes of AER shorter than AMEd; Province, Vietnam (032), 10 Dec.2000; 6 females, Viet Lann Village, PER straight, PME-PME about PMEd, PME-PLE longer; Ha Jiang Province, Vietnam (033), 10 Dec.2000. ALE and PLE juxtaposed. Chelicerae (Fig. 4B) brown, furnished with small warty granulations anterolaterally, Additional compared material examined. – 1 female of Nesioneta stridulatory ridges absent, promargin of fang groove with two benoiti (van Helsdingen, 1978) (MZT AA0.584), Seychelles, La teeth, first much bigger than second, retromargin with two Passe Gesthause on yard, coll. M. Saaristo, 11 Jan.1999; 1 male of small teeth. Lengths of legs: I 8.90 (2.33+2.67+2.50+1.40), Nesioneta benoiti (van Helsdingen, 1978) (MZT AA0.585), II 6.86 (1.73+2.03+2.00+1.10), III 4.00 Seychelles, Silhouette Anse Lacars-Anse Parates sifting leaves of Dendrobium umbellatum, coll. M. Saaristo, 7 Jan.1999. (1.03+1.07+1.23+0.67), IV 5.66 (1.50+1.60+1.73+0.83). Each patella and tibia with two dorsal spines, Ti I-II Diagnosis. – The male of this new species can be easily additionally with one prolateral spine. Fe I-III with small distinguished by the special shape of male palpal tibia and dorsal spine, Fe IV without. Tm I: 0.14. Tm IV absent. the posterior tooth of cymbium (Figs. 5A-C), as well as the Abdomen cylindriform, without tubercle, gray with whitish combined characters of embolic division (Figs. 5D-E); female spots laterally and dorsally. can be identified by the round and protruding epigynum with vase-shaped proximal portion of scape and large oval Male palp (Figs. 4C-E). Patella short, with long spine dorsally. depression of lateral plates on each side (Fig. 5F).

109 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

Fig. 4. Neriene oxycera, new species: A, ocular area of male, dorsal view; B, male left chelicera, frontal view; C, left male palp, retrolateral view; D, ditto, ventral view; E, ditto, prolateral view; F, embolic division, ventral view. Scale bars = 0.1 mm.

110 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Fig. 5. Nesioneta ellipsoidalis, new species: A, left male palp, retrolateral view; B, ditto, prolateral view; C, ditto, ventral view; D, embolic division, ventral view; E, ditto, dorsal view; F, epigynum, ventral view; G, ditto, dorsal view; H, ditto lateral view; I, vulva, dorsal view. Scale bars = 0.1 mm.

111 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

Description of male. – Total length: 1.80. Carapace: 0.80 anterior-ventrally, also carrying hair. Tibia with two long, 0.60 wide. Abdomen: 1.10 long, 0.60 wide. Carapace retrolateral and one prolateral trichobothria. Cymbium normal grayish brown with black margin. Eyes subequal, with black shape, with tooth-like apophysis posteriorly. Paracymbium surroundings; AER recurved, PER straight, eyes separated as usual shaped for Nesioneta, apical branch ending in flat, by AMEr, ALE and PLE juxtaposed. Chelicerae brown, translucent blade-shaped apex, strong tooth on outer margin stridulatory ridges present on ectal side, fang groove with of bottom. Pit-hook on distal part of suprategulum bifurcate, three promarginal teeth, second of which biggest, and three one point, slightly hooked, one truncate. Embolic division small retromarginal teeth. Sternum darker than carapace. (Figs. 5D-E): radix large boat-shaped with dorsal bulge and Abdomen dark gray, with a large white chevron. Lengths of anterior hook as usual, all other elements in planted on basal legs: I 2.80 (0.73+0.90+0.70+0.47), II 2.46 and curved middle part. Lamella characteristica long and (0.63+0.80+0.60+0.43), III 1.97 (0.57+0.60+0.47+0.33), IV sharp. Terminal apophysis consisted of three parts, anterior 2.61 (0.80+0.77+0.67+0.37). Tibia dorsal spines: 2-2-2-2. Tm one large with many dentate-like projections and point tip, I: 0.28. Tm IV absent. middle one short, plate-shaped and strongly sclerotized with scaled surface, posterior one slightly sclerotized with serrated Male palp (Figs. 5A-C). Patella with long bristle dorsally. apex. Embolus (Figs 5E, 6A) connected with radix by Tibia widely depressed dorsal-posteriorly, two tips of cylindrical membrane, its main body oval plate-shaped with retrolateral margin and one dorsal tip of prolateral margin two small apophyses posteriorly (Fig. 6A) and two large rising into lobes, each carrying hair; another process exists apophyses anteriorly, dorsal one long and strongly sclerotized

Fig. 6. Nesioneta ellipsoidalis, new species: A, embolus, dorsal view. Nesioneta benoiti (van Helsdingen, 1978): B, embolus, dorsal view; C, embolic division, ventral view; D, ditto, dorsal view. Scale bars: A-B = 0.05 mm; C-D = 0.1 mm.

112 THE RAFFLES BULLETIN OF ZOOLOGY 2006 extending forward, ventral one slightly sclerotized with sharp Additional compared material examined. – 1 female and male of process in the middle and its tip rolling back. Seminal duct Parameioneta spicata Locket, 1982, Phetchabun Province, Lom Sak, traversing in embolus plate and retracing back, embolus church yard, Thailand, coll. E. Huitula & P. Lehtinen, 27 Oct.–18 Nov.1976. proper stout, near base, covered ventrally by sclerotic extension of cylindrical membrane. Diagnosis. – The male P. bilobata can be easily distinguished by the large hooked dorsal apophysis of tibia, triangular tooth Description of female. – Total length: 1.83. Carapace: 0.70 on outer margin of paracymbium (Fig. 7A) and the characters long, 0.55 wide. Abdomen: 1.50 long, 0.73 wide. Lengths of of embolic division (Fig. 8); the female by the shape of legs: I 2.50 (0.70+0.77+0.60+0.43), II 2.30 epigynum in ventral view and the fork-shaped distal portion (0.67+0.70+0.53+0.40), III 1.90 (0.67+0.43+0.47+0.33), IV of scape (Figs. 7F-I) in dorsal view. 2.44 (0.77+0.73+0.57+0.37). Tibia dorsal spines: 2-2-2-2. Tm I: 0.29. Tm IV absent. Other somatic characters same as male. Description of male. – Total length: 1.50. Carapace: 0.72 long, 0.50 wide. Abdomen: 0.78 long, 0.44 wide. Carapace Epigynum (Figs. 5F-I). Half round and convex in ventral view, yellowish brown, with black margin. Eyes subequal, with enlarged lateral plates forming large oval depressions on each black surroundings; AER recurved, PER straight, eyes side. Scape “S” shaped folding. Margin of proximal portion separated by AMEd, ALE and PLE juxtaposed. Chelicerae parallel originally and widened to broadly rounded posterior brown, fang groove with four promarginal, four retromarginal margin. Inner scape invisible in ventral view. Distal portion teeth. Sternum darker than carapace. Abdomen dark gray, with two wings, translucent, hardly discernable in dorsal view, with white spots. Lengths of legs: I 2.63 tip furnished with pit extending outside posterior margin (0.66+0.86+0.61+0.50), II 2.12 (0.59+0.69+0.50+0.34), III visible in ventral view. 1.62 (0.50+0.50+0.34+0.28), IV 2.25 (0.66+0.69+0.56+0.34). Tibia dorsal spines: 2-2-2-2. Tm I: 0.27. Tm IV absent. Etymology. – The specific name comes from Latin Male palp (Figs. 7A-C). Patella with long spine dorsally. Tibia ellipsoidalis (ellipse) in reference to the shape of the embolus dorsal apophysis hook-shaped with multiple teeth in planted of the male palp. in concave area (Fig. 7E). Paracymbium U-shaped with black triangular tooth on outer margin. Pit-hook on distal part of Remarks. – The new species is similar to Nesioneta benoiti suprategulum with long stick and hooked apex. Embolic (van Helsdingen, 1978), which was transferred from genus division (Figs. 7D, 8A): lamella characteristic with large Meioneta by Saaristo (1995). Comparison of the two species sclerotic at base, two long narrow branches out extending, revealed that the differences between them mainly one knife-shaped, strongly sclerotized, the other less concentrated on embolic division (Figs. 6A-B): 1) the sclerotized with fringy apex. Terminal apophysis with two different shapes of their dorsal embolic apophyses; 2) the free ends, anterior one wide, petal-shaped, posterior one long middle process of the ventral embolic apophysis in the new with tapering tip. Embolus proper stout (Figs. 7D, 8B-C). species is long and sharp, while it is short in N. benoiti; 3) Thumb spike-like, with wide base and some thread-like the oval plate-shaped main body of embolus of this new projections along its outer margin. Embolic membrane tail species has two small posterior embolic apophyses, while N. shaped with many papillae on it. benoiti has only one. Additionally, in this new species, the dentate-like projections on the dorsal surface of anterior Description of female. – Total length: 1.72. Carapace: 0.59 terminal apophysis in N. benoiti seem much smaller in central long, 0.40 wide. Abdomen: 1.13 long, 0.72 wide. Lengths of than those along the margin (Fig. 6C). legs: I 2.47 (0.69+0.78+0.56+0.44), II 2.06 (0.59+0.69+0.44+0.34), III 1.56 (0.50+0.44+0.34+0.28), IV Distribution. – Known only from the type locality. 2.19 (0.63+0.72+0.50+0.34). Tibia dorsal spines: 2-2-2-2. Tm I: 0.25. Tm IV absent. Other somatic characters same as male.

Epigynum (Figs. 7F-H). Protruding, scape “S” shape Parameioneta bilobata Li & Zhu, 1993 puckered, proximal portion triangular, middle portion and (Figs. 7, 8) distal portion folded under it, pressed tightly and invisible in lateral view. Distal portion forkshaped, very translucent, with Parameioneta bilobata Li & Zhu, in Song, Zhu & Li, 1993: 867, pit in central and bifid apex. Figs. 28A-M; Li & Zhu, 1995: 45, Figs. 5a-m.

Material examined. – 4 males and 2 females, Gao Bao Village, Ha Distribution. – China, Vietnam (Ha Jiang Province). Jiang Province, Vietnam (017), 9 Dec.2000; 4 females, Gao Bao Village, Ha Jiang Province, Vietnam (015), 9 Dec.2000; 4 females, Viet Lann Village, Ha Jiang Province, Vietnam (022), 10 Dec.2000; Prosoponoides sinensis (Chen, 1991), new combination 4 females, Viet Lann Village, Ha Jiang Province, Vietnam (023), (Fig. 9) 10 Dec.2000; 3 females, Viet Lann Village, Ha Jiang Province, Vietnam (032), 10 Dec.2000; 1 female, Viet Lann Village, Ha Jiang Neriene sinensis Chen, 1991: 164, Figs. 2A-D. Province, Vietnam (032), 10 Dec.2000; 3 females, Viet Lann Village, Ha Jiang Province, Vietnam (033), 10 Dec.2000; 1 female, Viet Material examined. – 1 male and 1 female, Tan Linh Village, Son Lann Village, Ha Jiang Province, Vietnam (042), 10 Dec.2000. Tay Province, Vietnam (131), 23 Dec. 2000.

113 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

Fig. 7. Parameioneta bilobata Li & Zhu, 1993: A, left male palp, retrolateral view; B, ditto, prolateral view; C, ditto, ventral view; D, embolic division, dorsal view; E, palpal tibia, dorsal view; F, epigynum, ventral view; G, ditto, lateral view; H, ditto dorsal view; I, vulva, dorsal view. Scale bars = 0.1 mm.

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Additional Material examined. – 1 male (MHNG), holotype of Male palp (Figs. 9B-D). Patella short. Tibia wider than long, Prosoponoides hamatus Millidge & Russell-Smith, 1992, Ketambe, with one prodorsal, one retrodorsal trichobothrium. Cymbium Gunung Leuser, northern Sumatro, coll. C. Deeleman, 3 May.1976; unmodified. Paracymbium polliwog-shaped, main body stout, 1 female (MHNG), holotype of Prosoponoides kaharianum Millidge strongly sclerotized with short hairs, distal “tail” part narrow & Russell-Smith, 1992, Swampy primary forest, kaharian, Central and thin, hardly visible. Large pit-hook on distal part of , coll. C. Deeleman, 2 Sep.1985; 1 male (MHNG), holotype of Prosoponoides simile Millidge & Russell-Smith, 1992, suprategulum well developed, with tip bifurcated in ventral Khao Yao National Park, Thailand, coll. C. Deeleman, 22 Dec.1988. view. Embolic division (Fig. 9E) characters by well- developed lamella characteristica with lateral projection wide Diagnosis. – The male of Prosoponoides sinensis can be and flat, bending dorsally, sharp apex pointing forwards, recognized by the large lateral projection of lamella anterior and two dorsal projections blunt, triangular posterior characteristica (Figs. 9B-E); the female by the epigynum with projection pointing backwards. Terminal apophysis with two a septum that divides atrium into two compartments (Fig. free ends, anterior one long, narrow, ribbon-like, posterior 9F). one sclerotized. Embolus long, more than half circle, arising from inside of radix, tracing out of cymbium through rolled Description of male. – Total length: 2.70. Carapace: 1.50 embolic membrane. long, 1.00 wide. Abdomen: 1.53 long, 1.00 wide. Carapace yellowish brown, unmodified. Eyes with black surroundings; Description of female. – Total length: 3.20. Carapace: 1.30 AME smallest, others subequal; AER recurved, AME-ALE long, 1.10 wide. Abdomen: 2.0 long, 1.50 wide. Lengths of about AMEd, AME-AME shorter; PER straight, eyes legs: I 5.90 (1.60+1.83+1.60+0.87), II 5.17 separated by about PMEd; ALE and PLE juxtaposed. (1.50+1.67+1.40+0.60), III 3.23 (0.90+1.00+83+0.50), IV Chelicerae brown; stridulatory ridges absent; fang groove with 4.74 (1.27+1.40+1.37+0.70). Tm I: 0.26. Tm IV absent. Other three promarginal, two retromarginal teeth (Fig. 9A). Lengths somatic characters same as male. of legs: I 5.90 (1.67+1.80+1.60+0.83), II 5.23 (1.50+1.60+1.40+0.73), III 3.03 (0.83+0.90+0.80+0.50), IV Epigynum (Fig. 9F). Two openings of entrance grooves with 3.94 (1.20+1.27+1.20+0.67). Each patella and tibia with two curved, well sclerotized margins. Atrium formed by enlarged dorsal spines; Ti I-II additionally with one prolateral, one openings with septum, which divides atrium into two. retrolateral spine. Fe I-IV with one small spine dorsally. Tm Triangular scape extending from dorsal wall of atrium, I: 0.21. Tm IV absent. Abdomen oval shape, dorsal surface strongly sclerotized with big pit, slightly turning upwards in pale white with black longitudinal spot on median line from lateral view. Vulva (Figs. 9G-H) as long as wide, entrance which send out several short transversal spots, last one third grooves started from ventral wall, with two and half coils, part black, ventral surface uniformly dark gray. turning points laterally of apices, pointing inwards.

Fig. 8. Parameioneta bilobata Li & Zhu, 1993: A, embolic division (removing embolus), ventral view; B, embolus, ventral view; C, ditto, ventrolateral view. Scale bars = 0.1 mm.

115 Tu & Li: Linyphiinae and Micronetinae spiders from Vietnam

Fig. 9. Prosoponoides sinensis (Chen, 1991): A, left chelicera, frontal view; B, left male palp, retrolateral view; C, ditto, ventral view; D, ditto, prolateral view; E, embolic division, ventral view; F, epigynum, ventral view; G, vulva, dorsal view; H, ditto, ventral view. Scale bars = 0.1 mm.

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Remarks. – Compared with the illustrations of Neriene Helsdingen, P. J. van., 1978. Contributions à l’étude de la faune clathrata (Sundevall, 1830) provided by van Helsdingen terrestre des îles granitiques de l’archipel des Séchelles (Mission (1969: Figs. 83-91), the genital structures of present species P. L. G. Benoit - J. J. Van Mol 1972). Linyphiidae (Araneae). show several differences: 1) without the coil-shaped terminal Revue de Zoologie Africaine, 92: 889-898. apophysis which is typical character of male palp of Neriene; Helsdingen, P. J. van., 1985. Araneae: Linyphiidae of Sri Lanka, 2) embolus long, thread like; 3) epigynum with posterior with a note on Erigonidae. Entomologica scandinavica (Suppl.), margin of ventral wall extending in the middle to form a 30: 13-30. septum which divides atrium into two. These characters do Hou, Z. E. & S. Q. Li, 2003. Terrestrial talitrid amphipods not corresponding well with the genital type of Neriene, but (Crustacea: Amphipoda) from China and Vietnam: studies on show striking resemblance to that of Prosoponoides hamatum the collection of IZCAS. Journal of Natural History, 37(20): 2441-2460. Millidge & Russell-Smith (1992: Figs. 1-4, 8-11). As a result, N. sinensis was transferred to Prosoponoides here in this Li, S. Q. & X. Q. Liang, 2002. Caridean prawns of northern Vietnam study. (Decapoda: Atyidae: Palaemonidae). Acta Zootaxonomic Sinica, 27(4): 707-716. Distribution. – Vietnam (Cao Bang, Sac Tay), China. Li, S. Q. & C. D. Zhu, 1995. Five new species of linyphiid spiders from China (Araneae: Linyphiidae). Acta zootaxonomica sinica, 20: 39-48. ACKNOWLEDGMENTS Locket, G. H., 1968. Spiders of the family Linyphiidae from Angola. Publicações culturais da Companhia de Diamantes de Angola, 71: 61-144. The authors are indebted to Dr Michael I. Saaristo (Centre for Biodiversity, University of Turku, Finland) and Dr Marples, B. J., 1955. Spiders from western Samoa. Journal of the Xinping Wang (University of Florida, USA) for comments Linnean Society of London, 42: 453-504. on this paper. Thanks are given Dr Peter Schwendinger Peng, X. J. & S. Q. Li, 2003. New localities and one new species (Muséum d’histoire naturelle, Genève, Switzerland) for of jumping spiders (Araneae: Salticidae) from Northern providing compared material studied. This study was Vietnam. Raffles Bulletin of Zoology, 51(1):21-24. supported by the National Natural Sciences Foundation of Platnick, N. I., 2005. The world spider catalog, version 5.5. China (NSFC-30270183, 30370263, 30310464, 30470213, American Museum of Natural History, online at http:// 30499341), by the National Science Fund for Fostering research.amnh.org/entomology/spiders/catalog81-87/ index.html. Talents in Basic Research (NSFC-J0030092), by the Beijing Natural Science Foundation (6052017) and partly also by the Saaristo, M. I. & A. V. Tanasevitch, 1996. Redelimitation of the Kadoorie Farm and Botanic Garden, Hong Kong Special subfamily Micronetinae Hull, 1920 and the genus Lepthyphantes Administrative Region, China. Menge, 1866 with descriptions of some new genera and species (Aranei, Linyphiidae). Berichte des Naturwissenschaftlich- Medizinischen Vereins in Innsbruck (Innsbruck), 83:163-186. LITERATURE CITED Saaristo, M. I., 1995. Linyphiid spiders of the granitic islands of Seychelles (Araneae, Linyphiidae). Phelsuma, 3: 41-52.

Chen, J. & C. M. Yin, 2000. On five species of linyphiid spiders Saito, H., 1982. Soil dwelling linyphiine and erigonine spiders from from Hunan, China (Araneae: Linyphiidae). Acta Arachnologica Ogasawara Islands, Japan. Edaphologia, 25-26: 33-39. Sinica, 9: 86-93. Schenkel, E., 1963. Ostasiatische Spinnen aus dem Muséum Chen, Y. F., 1991. Two new species and two new records of linyphiid d’Histoire naturelle de Paris. Mémoires Muséum national spiders from China (Arneae [sic]: Linyphiidae). Acta d’histoire naturelle, Paris (A, Zoologie), 25: 1-481. Zootaxonomica Sinica, 16: 163-168. Song, D. X., M. S. Zhu & S. Q. Li, 1993. Arachnida: Araneae. Helsdingen, P. J. van., 1969. A reclassification of the species of Animals of Longqi Mountain, 1993: 852-890. Linyphia Latreille based on the functioning of the genitalia Tu, L. H. & S. Q. Li, 2004. A preliminary study of erigonine spiders (Araneida, Linyphiidae), I. Zoologische Verhandelingen, Leiden, (Linyphiidae: Erigoninae) from Vietnam. Raffles Bulletin of 105: 1-303. Zoology, 52(2): 419-433.

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THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 119-124 Date of Publication: 28 Feb.2006 © National University of Singapore

RARE SPIDERS OF THE GENUS CYCLOCOSMIA (ARACHNIDA: ARANEAE: CTENIZIDAE) FROM TROPICAL AND SUBTROPICAL CHINA

M. S. Zhu*, J. X. Zhang and F. Zhang College of Life Sciences, Hebei University, Baoding, Hebei 071002, P. R. China Email: [email protected] (*Corresponding author)

ABSTRACT. – Two species of the ctenizid spider genus Cyclocosmia from tropical and subtropical China are diagnosed, described and illustrated. One of them, Cyclocosmia latusicosta, is found to be new to science. The relationships between all known species of the genus Cyclocosmia are discussed.

KEY WORDS. – Araneae, Ctenizidae, Cyclocosmia, new species, China

INTRODUCTION This study is a part of our research on the taxonomy and biogeography of Chinese mygalomorph spiders funded by Spiders of the genus Cyclocosmia Ausserer, 1871, are the NSFC, in which mygalomorphs are considered as fascinating because their abdomen is abruptly truncated and important objects for studies on biogeography and phylogeny. ends in a hard, heavily sclerotized disc strengthened by series Spiders of the genus Cyclocosmia are interesting of raised ribs separated by narrow grooves (Gertsch & mygalomorphs with unique morphology and biology. In this Platnick, 1975). The spiders usually live at steeply sloping study, two species of Cyclocosmia from tropical and banks of sandy clay, covered with moist leaf litter, where subtropical China are reported and one of them is considered they dig 7-15 cm deep vertical burrows with a hinged door to be new to science. attached to the entrance. Like all other trapdoor spiders, Cyclocosmia leads an extremely sedentary existence. It is distinct in having developed a morphological defense to cope MATERIALS AND METHODS with intruders. When the spider retreats headfirst to the bottom of its burrow, the abdominal disc fits tightly against the round All measurements in this paper are in millimeters. Leg walls of the burrow and forms an impenetrable false bottom measurements are given as: total length (femur, patella, tibia, (Gertsch & Wallace, 1936; Gertsch & Platnick, 1975). Due metatarsus, tarsus). Dissected epigynes were cleared in 10% to the successful camouflage of the trapdoor, Cyclocosmia is KOH for approximately 15 minutes at room temperature of very difficult to find and collect, and therefore it is often 18-25°C and then rinsed in distilled water. Abdominal rib considered as one of the rarest spiders. counts follow Gertch & Platnick (1975). SEM micrographs were taken on a KYKY 2800 scanning electron microscope. Being a small genus of the family Ctenizidae, Cyclocosmia All specimens are deposited in the Museum of Hebei only comprises six named species worldwide (Platnick, 2005; University, Hebei, China (MHBU). The abbreviations used Schwendinger, 2005): C. truncata (Hentz, 1841) and C. are: ALE – anterior lateral eyes, AME – anterior median eyes, torreya Gertsch & Platnick, 1975, in the USA, C. loricata PLE – posterior lateral eyes, PME – posterior median eyes, (C. L. Koch, 1842) in Mexico and Guatemala, C. ricketti MOA – median ocular area. (Pocock, 1901) in China, C. siamensis Schwendinger, 2005, and C. lannaensis Schwendinger, 2005, in Thailand. Gertsch & Platnick (1975) revised the then known species of this TAXONOMY genus and pointed out that the ribs and grooves of the abdominal disc, the number of setae on the seam of the disc, Cyclocosmia Ausserer, 1871 and the shape of the spermathecae are valuable characters for species discrimination. However, they found no significant Cyclocosmia Ausserer, 1871: 144 (Type species by monotypy: differences in the male palps among three American species, Mygale truncata Hentz, 1841); Gertsch & Platnick, 1975: 5. but the male palps in the species from Thailand are distinct Chorizops Ausserer, 1871: 144 (Type species by monotypy: from each other (Schwendinger, 2005). Actinopus loricatus C. L. Koch, 1842); Roewer, 1942: 146;

119 Zhu et al.: Chinese Cyclocosmia

Bonnet, 1956: 1078. First synonymized by Banks (1910), but genera, Galeosoma and Idiosoma of the family Idiopidae, removed from synonymy by Petrunkevitch (1911) and all have similar abdominal form, they are not closely related to subsequent authors until resynonymized by Gertsch & Platnick Cyclocosmia. These idiopid spiders possess eyes arranged in (1975). three rows rather than two, which is characteristic for Halonoproctus Pocock, 1901: 209 (Type species by monotypy: ctenizids. In addition, the abdomen of the genus Galeosoma Halonoproctus ricketti Pocock, 1901). First synonymized by Simon, 1903. is abruptly truncate but bears no ribs or grooves, whereas that of the genus Idiosoma bears ribs or grooves but is only Diagnosis. – Cyclocosmia can be easily distinguished from moderately truncate (Gertsch & Platnick, 1975). all other ctenizids by its distinctly truncate abdomen forming a caudal disc bearing ribs and grooves. Although two other Description. – See Gertsch & Platnick (1975).

KEY TO CHINESE SPECIES OF CYCLOCOSMIA

1. Abdominal disc with 23-33 ribs on each side; an elevated central zone inside each of the upper muscle impressions absent; length of spermathecae less than one and a half times their width ...... Cyclocosmia ricketti – Abdominal disc with 20-23 ribs on each side; an elevated central zone inside each of the upper muscle impressions present; length of spermathecae two times their width...... Cyclocosmia latusicosta

Cyclocosmia ricketti (Pocock, 1901) (Figs. 1-2)

Halonoproctus ricketti Pocock, 1901: 209, pl. 21, fig. 1. Cyclocosmia ricketti: Simon, 1903: 885, Figs. 1044-1047; Gertsch Fig. 1. Abdomen of Cyclocosmia ricketti (Pocock, 1901), caudal & Platnick, 1975: 18-19, Figs. 28, 29, 32, 36 (part); Chen & view. Zhang, 1991: 32, Fig. 25; Song, Zhu & Chen, 1999: 36, Figs. 16H, K-L (part); Schwendinger, 2005: 227, Figs. 2-8, pl. 1D.

Fig. 2. Cyclocosmia ricketti (Pocock, 1901): A, habitus of female; B, ocular area, dorsal view; C, abdomen, caudal view; D, same, lateral view; E, spermathecae, dorsal view. Scale bars: A, B, E = 1.0 mm, C-D = 2.0 mm.

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Material examined. – 1 female, Taishun, Zhejiang, China, coll. Z. formula: 3214. Tibia I 1.98 wide, tibia II 1.98 wide, tibia III F. Chen, Dec.1989. 2.25 wide, tibia IV 2.25 wide. Abdomen (Fig. 2C-D) funnel-shaped and dark yellow-brown. Diagnosis. – Cyclocosmia ricketti differs from the American species Caudal disc slightly concave, 13.95 in transversal diameter of Cyclocosmia in the abdominal disc with 23-33 ribs on each side and 15.30 in longitudinal diameter, with two ribs running (Fig. 2C) and the parallel-sided spermathecae (Fig. 2E). It is similar to Cyclocosmia latusicosta, new species, in the shape of dorsoventrally and 32/33 radiating ribs on each side; each spermathecae, but can be distinguished from the latter by the length rib angle with 25 or more bristles. Abdominal disc with six of spermathecae being less than one and a half times its width (Fig. well-marked muscle impressions. Four spinnerets, with inner 2E), abdominal disc with 23-33 ribs on each side, and an elevated pair small and one-segmented, and outer pair much longer central zone absent inside the upper pair of muscle impressions (Fig. and three-segmented. Two spermathecae (Fig. 2E), sack-like, 2C). It is also similar to Cyclocosmia siamensis Schwendinger, 2005 parallel-sided, each one with a length less than one and a (Schwendinger, 2005: 231, Figs. 9-30, pl. 1C) in the shape of half times its width. spermathecae, but differs from the latter in the upper and median pair of muscle impressions on opisthosomal disc separated by one Male. Unknown. transversal rib, all ribs lacking hairs with dark, short, cylindrical, upright proximal portion and light, long, flat, reclining distal portion, rib angles lacking bottlebrush-like bristles bent toward the centre Distribution. – China (Fujian, Hunan, Zhejiang, Sichuan). (Fig. 2C). Remarks. – Song et al. (1999: 36, Figs. 16H, K-L, pl. 1A-B) Description of new female. Total length, including chelicerae, provided the illustrations of Cyclocosmia ricketti basing on 25.83; chelicerae 4.32 long; carapace 11.07 long, 8.91 wide; a female specimen collected from Zhangjiajie, Hunan abdomen 15.57 long, 15.30 wide. Province, China (drawn by Mr. J. F. Wang). However, the Carapace (Fig. 2A) red-brown and smooth, with a few drawing of the disc (Song et al., 1999: Fig. 16L) was copied marginal hairs and a long bristle in front of ocular area, four from Gertsch & Platnick, 1975: Fig. 29, and it showed to be long bent bristles in longitudinal row running through ocular C. siamensis. area. Carapace curved in front and straight behind, widest at coxae II. Ocular area black, with a black band in front of fovea and beside ocular area respectively. Cervical groove Cyclocosmia latusicosta, new species and radial furrows distinct. Fovea deep and procurved, U- (Figs. 3-6) shaped, its greatest width occupying one fourth of carapace width at that point. Holotype. – Female, Aidian Village (21°49'N 107°18'E), Ningming Eyes (Fig. 2B) set on low tubercle, ocular area 1.00 long, County, Guangxi, China, coll. M. S. Zhu, J. X. Zhang and F. Zhang, 25 Aug.2004. 2.43 wide anteriorly, 2.43 wide posteriorly, occupying two fifths of carapace width at that point. Clypeus height 1.46. Paratypes. – 4 females, same data as for the holotype. Anterior eye row straight and posterior eye row recurved, both rows almost equal in length. Ratio of eyes, ALE: AME: Diagnosis. – The new species is similar to Cyclocosmia PLE: PME (0.53: 0.45: 0.40: 0.25). ALE-AME 0.30, AME- ricketti, especially in the shape of its spermathecae, but differs AME 0.20, PLE-PME 0.08, PME-PME 1.18. MOA 0.91 long, from the latter by the spermathecae being two times longer 1.10 wide in front, 1.68 wide at back. than wide (Figs. 6F-J), abdominal disc with 20/23 ribs on Chelicerae red-brown, inner margin with 11 teeth, outer each side, and each upper muscle impression with an elevated margin with nine teeth and 15 denticles between them. central zone (Fig. 6B). It is also similar to Cyclocosmia Rastellum raised on prominent angled projection and siamensis Schwendinger, 2005 (Schwendinger, 2005: 231, consisting of many short black teeth. Labium yellow-brown, figs. 9-30, pl. 1C) in the shape of spermathecae, but differs 1.56 long, 2.28 wide, with three black cuspules anteriorly. Maxilla yellow-brown, 4.50 long, 2.61 wide, armed with 15 black cuspules at base. Sternum yellow-brown, 6.75 long, 6.30 wide, with numerous long black setae, two pairs of small round sigilla near margin opposite coxae I and II, and a pair of large, irregularly outlined posterior sigilla. Palpal patella with one proventral distal spine. Legs yellow-brown, with tibiae, metatarsi and tarsi darker in color, short and stout, with erect setae ventrally. Tibiae and tarsi of pedipalps, and tibiae, metatarsi and tarsi of legs I and II with numerous horn-like spines, metatarsus III with a few short dorsal and two ventral spines, tarsus III with a few dark spines prolaterally and ventrally, metatarsus IV with four short spines. Paired claws with a single large tooth, unpaired claw lacking tooth. Pedipalp with single claw bearing one tooth. Measurements of legs: I 21.15 (7.20, 4.14, 4.23, 3.60, 1.98), II 18.09 (5.85, 4.05, 3.15, 3.06, 1.98), III 17.46 (5.58, 4.23, Fig. 3. Burrow entrance and trapdoor of Cyclocosmia latusicosta, 2.52, 2.97, 2.16), IV 22.32 (6.21, 5.40, 4.23, 4.14, 2.43). Leg new species.

121 Zhu et al.: Chinese Cyclocosmia from the latter in the upper and median pair of muscle Legs black-brown, short and stout, with erect setae and hairs impressions on opisthosomal disc separated by one transversal ventrally. Legs III and IV distinctly thicker and stronger than rib, all ribs lacking hairs with dark, short, cylindrical, upright legs I and II. Tibiae and tarsi of pedipalps and tibiae, metatarsi proximal portion and light, long, flat, reclining distal portion, and tarsi of legs I and II with numerous horn-like spines, abdominal disc with 20-23 ribs on each side, and each upper metatarsus and tarsus III with a few scattered spines dorsally muscle impression with an elevated central zone (Fig. 6B). and ventrally, metatarsus IV with several small spines at tip. Paired claws with a single large tooth, occasionally Description. – Female. Holotype total length, including chelicerae, 33.56; chelicerae 4.50 long; carapace 12.78 long, accompanied by a denticle, unpaired claw lacking tooth. 11.25 wide; abdomen 17.20 long, 16.38 wide. Pedipalp with single claw bearing one tooth. Measurements Carapace black-brown and smooth, with a few hairs and four of legs:I 22.95 (8.55, 4.95, 4.14, 3.60, 1.71), II 21.60 (7.65, long bent bristles in longitudinal row running through ocular 4.50, 3.51, 3.69, 2.25), III 20.88 (6.03, 4.95, 3.42, 3.78, 2.70), area. Carapace curved in front and straight behind, widest at IV 27.27 (7.29, 5.58, 4.50, 6.93, 2.97). Leg formula: 3214. coxae III. Ocular area darker in color. Cervical groove and Tibia I 2.52 wide, tibia II 2.25 wide, tibia III 2.97 wide, tibia radial furrows distinct. Fovea deep and procurved, U-shaped, IV 2.88 wide. its greatest width occupying one fifth of carapace width at Abdomen (Figs. 6B-D) dark yellow-brown, with crescent- that point. shaped sclerite posterior to spinnerets and anus not connected Eyes (Fig. 6A) set on low tubercle, ocular area 1.25 long, to ventral median rib angle. Caudal disc convex, with two 3.06 wide anteriorly, 3.09 wide posteriorly, occupying one ribs running dorsoventrally and 20/23 radiating ribs on each third of carapace width at that point. Clypeus height 1.80. side; ribs wide and grooves deep; each rib angle distinctly Anterior eye row straight and posterior eye row slightly recurved, both rows subequal in width. Ratio of eyes, ALE: elevated carrying 46-52 long and smooth bristles and a few AME: PLE: PME (0.75: 0.51: 0.54: 0.37). ALE-AME 0.40, bottlebrush-like bristles rising from inner surface of rib angle AME-AME 0.27, PLE-PME 0.08, PME-PME 1.39. MOA and bent towards disc center. All ribs with many granular 1.05 long, 1.29 wide at front, 2.19 wide at back. structures in different size (Fig. 4). Abdominal disc with six Chelicerae (Fig. 6E) black-brown, inner margin with 10-12 well-marked muscle impressions, the upper pair with an teeth of different size, outer margin with eight or nine large elevated central zone connected to the outer rim of each of teeth and 23 denticles between them. Rastellum raised on these muscle impressions, and separated from median pair prominent angled projection and consisting of many short by only one transversal rib, second (lower) transversal rib black teeth. Labium dark red-brown, 2.16 long, 2.52 wide, wide, running into upper portion of ring around median with three black cuspules anteriorly. Maxilla dark red-brown, muscle impression. Disc surface with only three pairs of 5.40 long, 3.15 wide, armed with 12-17 cuspules at base and bristles on the rims of muscle impressions. Four spinnerets, many small cuspules in anterior part. Sternum red-brown, with inner pair small and one-segmented, and outer pair much 7.47 long, 7.56 wide, with numerous long black setae, two pairs of small round sigilla near margin opposite coxae I and longer and three-segmented. Two sack-like, parallel-sided II, and a pair of large indistinctly outlined posterior sigilla. spermathecae (Fig. 6F), each one almost two times longer Palpal patella with one proventral distal spine. than wide.

Male. Unknown.

Etymology. – Latin: latus = wide; Latin: costa = rib.

Variation. Measurements and rib counts (females, n=5): Body length: 33.03-33.56, carapace length 12.78-13.50, width 11.25-12.24; rib counts 20-23 (20/21, 21/20, 21/21, 22/21, 20/23). Spination of patellae: One female possesses one proventral spine on its right palpal patella, two proventral spines on the other palpal patella. Another female has one proventral spine on its right palpal patella, and none on the other palpal patella. Other specimens are the same as holotype. Labium with 2-3 cuspules.

Burrow. Burrows found at a sloping earth bank with moist leaf litter were about 16 cm in depth and 22 mm in diameter, narrowed near the bottom. The hinged entrance door was 32 mm in diameter, made of moss, earth and silks. The inside of the burrow was lined with a thin silk layer (Fig. 3). Fig. 4. Abdomen of Cyclocosmia latusicosta, new species, paratype, caudal view.

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Distribution. – Known only from the type locality in with many knob-like tubercles. Although the shape of southeastern China, near the border to Vietnam. spermathecae of C. lannaensis is similar to the American species C. loricata, it seems to be more closely related to the other Asian species than the American species, because its DISCUSSION apices of spermathecae are round but not “T”-shaped as C. loricata. Among the American species, C. truncata and C. According to the shape of spermathecae, the two Chinese torreya from the USA seem to be more closely related to species of the genus Cyclocosmia (C. ricketti and C. each other. As Gertsch & Platnick (1975) pointed out, their latusicosta, new species) and one of the Thailand species (C. spermathecae are so similar that it is very difficult to siamensis) seem to be more primitive than the other Thailand distinguish them on the basis of genitalic characters alone. species (C. lannaensis) and the three American species (C. The distribution and relationship of these seven species truncata, C. torreya and C. loricata) because their suggest that this genus already existed before the splitting of spermathecae are simple and parallel-sided, whereas the Laurasia, and that the Asian and American species were later spermathecae of the latter are more complex. Among the four geographically separated. Asian species, C. ricketti and C. latusicosta, new species, from China are more closely related because their ribs of disc are smooth and lack setae. C. lannaensis is more derived than ACKNOWLEDGEMENTS the other three Asian species due to its medially constricted spermathecae with knob-shaped apices, and its ribs of disc This study was supported by grants from the NSFC

Fig. 5. Cyclocosmia latusicosta, new species, paratype (SEM micrographs). A, detail of ribs; B, detail of inner side of rib angles showing two kinds of bristles; C, detail of long and smooth bristles; D, detail of bottlebrush-like bristles.

123 Zhu et al.: Chinese Cyclocosmia

(30170118) to M. S. Zhu and in part by the NSFC (30130040) Hentz, N. M., 1841. Species of Mygale of the United States. to D. X. Song. We thank Tinghui Li (Southern Spider Proceedings of the Boston Society of Natural History, 1: 41-42. Research and Breeding Institute of Nanning, Guangxi, China) Koch, C. L., 1842. Die Arachniden. Nürnberg, vol. 9, 108 pp. for his assistance during the expedition to Guangxi Province Petrunkevitch, A., 1911. A synonymic index-catalogue of spiders in August 2004. Many thanks are also due to Dr. P. J. of North, Central and South America with all adjacent islands, Schwendinger for providing the valuable reference. Greeland, Bermuda, West Indies, Terra del Fuego, Galapagos, etc. Bulletin of the American Museum of Natural History, 29: 1-791. LITERATURE CITED Platnick, N. I., 2005. The World Spider Catalog, version 5.5. American Museum of Natural History, online at http:// research.amnh.org/entomology/spider/catalog81-87/index.html Ausserer, A., 1871. Beiträge zur Kenntniss der Arachniden-Familie der Territelariae Thorell (Mygalidae Autor). Verhandlungen der Pocock, R. I., 1901. On some new trap-door spider from China. Zoologisch-Botanischen Gesellschaft in Wien, 21: 117-224. Proceedings of the Zoological Society of London, 1901: 207- 215. Bonnet, P., 1956. Bibliographia araneorum. Toulouse, 2 (2): 919- Roewer, C. F., 1942. Katalog der Araneae von 1758 bis 1940. 1926. Bremen, 1. Band, 1040 pp. Chen, Z. F. & Z. H. Zhang., 1991. Fauna of Zhejiang: Araneida. Schwendinger, P. J., 2005. Two new Cyclocosmia (Araneae: Zhejiang Science and Technology Publishing House, 356 pp. Ctenizidae) from Thailand. Revue suisse de Zoologie, 112 (1): Gertsch, W. J. & N. I. Platnick., 1975. A revision of the trapdoor 225-252. spider genus Cyclocosma (Araneae, Ctenizidae). American Simon, E., 1903. Histoire naturelle des Araignees (Araneides). Paris, Museum Novitates, 2580: 1-20. vol.2, pt. 4, pp. 669-1080. Gertsch, W. J. & H. K. Wallace., 1936. Notes on new and rare Song, D. X., M.. S. Zhu & J. Chen., 1999. The spiders of China. American mygalomorph spiders. American Museum Novitates, Hebei Science and Technology Publishing House, Shijiazhuang, 884: 1-25. 640 pp.

Fig. 6. Cyclocosmia latusicosta, new species. A, ocular area, dorsal view, holotype; B, abdomen, caudal view, holotype; C, same, lateral view, holotype; D, posterior portion of opisthosoma, ventral view, holotype; E, left chelicera, prolateral view, holotype; F-J, spermathecae, dorsal view: F, holotype; G-J, paratypes. Scale bars: A, F-J = 1.0 mm, B-E = 2.0 mm.

124 THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 125-127 Date of Publication: 28 Feb.2006 © National University of Singapore

OOPHANA TIOMANENSIS, A NEW STREPTAXID (GASTROPODA: PULMONATA: STREPTAXIDAE) FROM PULAU TIOMAN, PENINSULAR MALAYSIA

Reuben Clements Department of Biological Sciences, National University of Singapore, Science Drive 4, Singapore 117543, Republic of Singapore.

ABSTRACT. – A new species of terrestrial gastropod belonging to the family Streptaxidae Gray, 1860, is described from Pulau Tioman, Peninsular Malaysia. Oophana tiomanensis, new species, is compared with two congeners that are nearest geographically (i.e., Thailand) and similarly possess three teeth on the peristome. Oophana elisa Gould, 1856, differs from this new species by having the penultimate whorl carinate to the left, while O. bulbulus (Morelet, 1862) differs by being larger (height - 15 mm) and distinctly less oblique. The new species being described here is also the first from the genus Oophana Ancey, 1884, to be recorded from Peninsular Malaysia.

KEY WORDS. – Gastropod, systematics, Oophana, new species, Pulau Tioman, Malaysia.

INTRODUCTION congeners thus far (J. J. Vermeulen, pers. comm.). Anatomical characteristics (e.g., the genital system; Stoliczka, 1871; The pulmonate gastropod family Streptaxidae Gray, 1860, Berry, 1963) could also be used to distinguish species within has a circumtropical distribution that includes South America, this family, but such information remains too scarce for Africa, southern Asia and several Indo-Pacific islands worthwhile comparisons to be made. Until comparative (Richardson, 1988). From the paucity of biological and studies on the anatomy of its congeners are made available, ecological information available on streptaxids, we know that a description of this new species based on shell characteristics they are carnivorous, oviparous or viviparous, and usually should suffice. Specimens examined are deposited in the occur in leaf litter and decaying wood habitats in limestone Zoological Reference Collection (ZRC), Raffles Museum of areas (Berry, 1963; Van Bentham Jutting, 1954). Biodiversity Research, National University of Singapore.

In Peninsular Malaysia, taxonomic work on streptaxids was chiefly pioneered by Van Bentham Jutting (1954, 1961a, b). TAXONOMY Four streptaxid genera (Discartemon Pfeiffer, 1856; Gulella Pfeiffer, 1856; Haploptychius Möllendorf, 1905; and Oophana Ancey, 1884 Sinoennea Kobelt, 1904) are known to occur within the region. Since 1961, however, no new streptaxids have been Oophana Ancey, 1884: 508; Tryon, 1885: 58; Kobelt, 1906: 101; added to the list of 41 species already described from Thiele, 1931: 730; Bentham Jutting, 1954: 95; Zilch, 1960: 562. Peninsular Malaysia (Van Bruggen, 1972; Maassen, 2001). Type species. – Oophana bulbulus (Morelet, 1862). Four decades later, an undescribed streptaxid was found on Pulau Tioman, Peninsular Malaysia. The discovery of this Remarks. – Richardson’s (1988) keys for Streptaxidae, in new species is not surprising because a malacological survey which the genus Oophana is distinguished from other genera of the island has never been performed. In this paper, I report by having two (sometimes only one) parietal lamellae and a new streptaxid belonging to a fifth genus Oophana Ancey, three or more other teeth, are followed here. In Peninsular 1884, which is considered a new record for Peninsular Malaysia, Van Bentham Jutting (1954, 1961a) described seven Malaysia. species of Oophana under the subgenus Haploptychius, which differs from Oophana sensu stricto by having only one parietal lamella present. However, Haploptychius has since been MATERIAL AND METHODS treated as a full genus (see Van Bruggen, 1972; Richardson, 1988). Therefore, it can be said that prior to this report, no Descriptions of Oophana tiomanensis, new species, are based representatives of Oophana occured in Peninsular Malaysia. on shell characters, which have been used for most of its

125 Clements: A new streptaxid from Pulau Tioman

Oophana tiomanensis, new species differs from this new species by having the penultimate whorl (Figs. 1-2) carinate to the left, while O. bulbulus (Morelet, 1862) differs by being larger (height - 15 mm) and distinctly less oblique. Materials examined. – Holotype - 1 ex. (Height 10.4 mm; Width 11.4 mm; Height of aperture 6.0 mm) (ZRC.MOL.90), beneath leaf Distribution. – Gunung Kajang, Pulau Tioman, Peninsular litter on soil, 50 m elevation, Gunung Kajang (2º 46'N 104º 9'E), Malaysia (present study). Pulau Tioman, Peninsular Malaysia, coll. R. Clements, 18 Aug.2003.

Paratypes – 2 ex. (11.8 mm; 11.4 mm; 6.9 mm) (ZRC.MOL.91) and (10.9 mm; 11.4 mm; 5.4 mm) (ZRC.MOL.92), same locality ACKNOWLEDGEMENTS data as holotype. I am very grateful to J. J. Vermeulen for authenticating this Diagnosis. – Shell obliquely ovoid, white or cream coloured, new streptaxid and for his valuable comments, W. J. M. about opaque, with 6-7 whorls. Spire moderately rounded; Maassen for help with the bibliography, S. Y. Chan and S. L. penultimate whorl more convex on left side; last whorl Loke for initially drawing my attention to the distinctiveness protruding to the right side, moderately rounded, slightly 1 angular just below the periphery. First 2 /2 whorls smooth; other whorls sculptured with low, rounded collabral ribs; those on early whorls closely packed, about 8-12 to the mm; those on last whorl furthest apart, about 4 to the mm, continuing into the umbilicus. Suture distinct but not deep. Apex rounded. Base relatively inflated. Umbilicus about 1 mm wide. Aperture oblique, semi-elliptic. Peristome continuous, reflected, thickened, glossy, whitish or yellowish-brown. Parietal side with distinct, slightly undulating lamella continuing 4-5 mm into the aperture. Palatal, basal and columellar side each with single, distinct, knob-shaped tooth. All teeth arranged in cross-shaped manner. Animal with orange-reddish body and tentacles. Foot yellowish.

Etymology. – The species name is derived from the type locality, Pulau [=Island] Tioman.

Remarks. – Outside of Peninsular Malaysia, the nearest congeners of O. tiomanensis, new species, are located in Thailand and other parts of Indo-China. After examining descriptions of eight Oophana species occurring in neighbouring Thailand and Indo-China (see Van Bentham- Jutting, 1954), only two were suitable for comparison with O. tiomanensis, new species, because they similarly possess three teeth on the peristome. Oophana elisa Gould, 1856,

Fig. 1. Live animal of Oophana tiomanensis, new species, from Pulau Tioman, Peninsular Malaysia. The body and tentacles are Fig. 2. Shell of Oophana tiomanensis, new species. Holotype orange-red while the foot is yellow. Photo: H. H. Tan. (ZRC.MOL.83) from side, top and bottom.

126 THE RAFFLES BULLETIN OF ZOOLOGY 2006 of this particular species and H. H. Tan for the photograph of Richardson, C. L. 1988. Streptaxacea: Catalog of species Part I the live snail. Streptaxidae. Tryonia, 16: 1-326. Stoliczka, F. 1871. Notes on the terrestrial Mollusca from the neighbourhood of Moulmein, with descriptions of new species. LITERATURE CITED The Journal of the Asiatic Society of Bengal, 40: 143-177. Thiele, J. 1931. Handbuch der Systematischen Weichtierkunde Tiel Ancey, M. C.-F. 1884. Sur les divisions proposes dans le genre 1. Amsterdam. 735 pp. Streptaxis. Le Naturaliste, 6: 399. Tryon, G.W., Jr. 1885. Manual of Conchology: Testacellidae, Berry, A. J. 1963. The anatomy of two Malayan limestone hill Oleacinidae, Streptaxidae, Helicoidea, Vitrinidae, Limacidae, Streptaxidae, Sinoennea kanchingensis Tomlin and Oophana Arionidae Volume I. Philadelphia. 363 pp. diaphanopepla van Bentham Jutting, with special reference to Van Bentham Jutting, W. S. S. 1954. The Malayan Streptaxidae of the genital system. Proceedings of the Malacological Society of the genera Discartemon and Oophana. Bulletin of the Raffles London, 35: 139-150. Museum, 25: 71-106. Gould, A. A. 1856. Descriptions of new species of shells. Van Bentham Jutting, W. S. S. 1961a. Additional new species and Proceedings of the Boston Society of Natural History, 6: 11-16. new localities of the family Vertiginidae and the genera Oophana Gray, J. E. 1860. On the arrangement of the land pulmoniferous and Opisthostoma from Malaya. Bulletin of the Raffles Museum, Mollusca into families. The Annals and Magazine of Natural 26: 34-48. History, 6: 267-269 Van Bentham Jutting, W. S. S. 1961b. The Malayan Streptaxidae, Kobelt, W. 1904. Nachrichtsblatt der Deutschen genera Huttonella and Sinoennea. Bulletin of the Raffles Malakozoologischen Gesellschaft, 36: 27-29. Museum, 26: 5-33. Kobelt, W. 1906. Die gattung Paludina L.A.M. Neue Folge in Martini Van Bruggen, A. C. 1972. On a new Streptaxid (Mollusca, and Chemnitz System Conchylien Cabinet 1, 12b: 101. Gastropoda, Pulmonata) from a Sangihe Island, Malay Maassen, W. J. M. 2001. A preliminary checklist of the non-marine Archipelago, with notes on the distribution of Streptaxids in molluscs of West-Malaysia. “A handlist”. De Kreukel, Southeast Asia. Proceedings of the Koninklÿke Nederlandse Amsterdam. 155 pp. Academie van Wetenschappen, Amsterdam (C), 75: 391-401. Morelet, P. M. A. 1862. In: Guérin-Méneville, F. E., Revue et Zilch, A. 1960. Gastropoda: Euthyneura. In: Wenz, W. (ed.), Magasin de Zoologie, 14: 477. Handbuch der Paläozoologie. Berlin. 834 pp. Pfeiffer, L. 1856. Versuch einer Anordnung der Heliceen nach natürlichen Gruppen. Malakozoologische Blätter, 2: 112-183.

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THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 129-155 Date of Publication: 28 Feb.2006 © National University of Singapore

A COLLECTION OF AMPHIBIANS AND FROM HILLY EASTERN CAMBODIA

Bryan L. Stuart The Field Museum, Department of Zoology, Division of Amphibians & Reptiles, 1400 S. Lake Shore Drive, Chicago, Illinois 60605-2496 U.S.A. The University of Illinois at Chicago, Department of Biological Sciences, 845 W. Taylor, Chicago, Illinois 60607 U.S.A. Email: [email protected] (Corresponding author)

Ko Sok Wildlife Protection Office, Ministry of Agriculture, Forestry and Fisheries, Phnom Penh, Cambodia

Thy Neang Protected Area Office, Department of Nature Conservation and Protection, Ministry of Environment, Phnom Penh, Cambodia

ABSTRACT. – The first amphibian and collection is reported from hilly eastern Cambodia since Henri Mouhot’s work in 1859. The collection contains 30 species of amphibians and 42 species of reptiles. Leptobrachium mouhoti, new species, and Ophryophryne synoria, new species, are described. Of the remaining collection, 11 species of amphibians (39.3%) and seven species of (33.3%) are reported from Cambodia for the first time. Comparisons of the show strong faunal overlap with that of mountainous central Vietnam, but little faunal overlap with the Cardamom Mountains of southwestern Cambodia.

KEY WORDS. – Cambodia, amphibians, reptiles, new species.

INTRODUCTION homalopsine watersnakes at Tonle Sap Great Lake (Stuart et al., 2000; Murphy et al., 2002) and on the trade and The amphibian and reptile fauna of Cambodia has received distribution of turtles (Lehr & Holloway, 2000, 2002; Touch very limited attention. Bourret’s monographs on the et al., 2000; Stuart et al., 2002; Holloway, 2003; Stuart and amphibians (1942), turtles (1941a), and snakes (1936) of Platt, 2004). French Indochina (= Cambodia, Laos, and Vietnam), published during the height of French colonial power in the Three upland areas in Cambodia contain sufficient topography region, remain the major works on Cambodia, supplemented to harbor swift, rocky streams: the Cardamom and Elephant only by Saint Girons’ (1972a) monograph on Cambodian Mountains in the southwest, a cluster of low hills in the north- snakes. Consequently, the snakes of Cambodia are the best center and along the northern border, and the hills and known component of the herpetofauna, and the the mountains in the extreme east. These eastern uplands in least known. Since Saint Girons, years of civil conflict and Mondolkiri, Ratanakiri, and Stung Treng Provinces are part security concerns in Cambodia have hampered new of the Annamite Mountains that separate Vietnam from Laos investigations by foreign and national workers. A recent, and Cambodia. More specifically, the uplands in eastern improved political situation has made new herpetological Mondolkiri Province (maximum elevation 1,078 m) are the fieldwork possible, and that attention has largely focused on lower slopes of the Langbian (=Da Lat) Plateau in southern the Cardamom Mountains of southwestern Cambodia, Vietnam, and the uplands in eastern Ratanakiri Province including their low elevation river valleys. This new fieldwork (maximum elevation over 1,500 m) are the lower slopes of in the Cardamoms has resulted in publications on the the Kontum Plateau in central Vietnam. amphibian fauna, including descriptions of three new species (Ohler et al., 2002), a new species (Daltry & Only one herpetological collection has been made in the Wüster, 2002), and the status of two critically endangered, eastern uplands of Cambodia. Henri Mouhot (1826-1861), large reptiles, Crocodylus siamensis and Batagur baska French explorer and naturalist, made natural history (Daltry & Chheang, 2000; Platt et al., 2003a, b; Holloway & collections in Thailand, Cambodia, and Laos from 1858-1861 Heng, 2004). Elsewhere in Cambodia, new work has been under the sponsorship of the Royal Geographical and conducted on the trade and reproductive biology of Zoological Societies of London. Although Mouhot was

129 Stuart et al.: Amphibians and reptiles from Cambodia primarily interested in ornithology and conchology, he did with Vietnam. Phnom Nam Lyr Wildlife Sanctuary lies collect some specimens of amphibians and reptiles during slightly northeast of Seima Biodiversity Conservation Area, his voyage (Mouhot, 1864). His herpetological collections and a continuous strip of forest along the Cambodian border from Cambodia were sent to the British Museum, where they connects both areas. The sanctuary is 47,500 ha in size, and were described by John Edward Gray and Albert Günther ranges from 300 m elevation to a maximum of 1,078 m (Gray, 1861a, b; Günther, 1861, 1864). During his expedition, elevation on the summit of Phnom Nam Lyr Mountain. Lower Mouhot made a single trip to eastern Cambodia. In 1859 he elevations in the sanctuary are dominated by grasslands with traveled by boat from Phnom Penh up the Mekong River to deciduous dipterocarp forest, gallery evergreen forest, and “Pemptiélan” (probably Kampong Cham today), then “150 bamboo forest, and Phnom Nam Lyr Mountain is covered by miles” eastward overland to a village called “Brelum,” where evergreen forest, bamboo forest, and a mixture of the two. he lived from 16 August-29 November among the Stiên (= Phnom Nam Lyr Wildlife Sanctuary was established on 01 Stieng) hill tribe (Mouhot, 1864). Ashburton in Mouhot November 1993 under the Royal Decree on Creation and (1864) gave coordinates for Brelum as “lat. 11o58’, long. Designation of Protected Areas in Cambodia, signed by His 107o12,” and if taken to be precise, place Brelum inside Majesty the King Norodom Sihanouk. The senior author, An present-day Vietnam approximately 25 km from the Dara, and Suon Phalla collected specimens of amphibians Mondolkiri Province, Cambodia border. Brelum was and reptiles at Phnom Nam Lyr Wildlife Sanctuary from 14- considered within the confines of Cambodia at the time of 21 June 2000 (Fig. 1). Mouhot’s visit, and the border has probably been redefined since. In any case, Mouhot certainly traversed through, and Virachey National Park, formerly known as Virachey worked in or very close to, present-day hilly Mondolkiri Protected Area, is situated in Ta Veng, Von Sai, and Andong Province, Cambodia, during his time in Brelum. Bufo Meas Districts in Ratanakiri Province, and in Siem Pang galeatus, described by Günther from Mouhot’s Cambodian District in Stung Treng Province. The national park is 332,500 collection, is at least one example of a species that Mouhot ha in size, and is bordered on the north by Laos and on the must have obtained during the Brelum expedition; this upland east by Vietnam. Hills and low mountains dominate the species of toad is known to occur in eastern Cambodia (see topography of the park, with most areas above 400 m below) and central and southern Vietnam (Smith, 1921; Inger elevation, and with maximum elevations above 1,500 m et al., 1999; Liu et al., 2000), but not southwestern Cambodia elevation near the Laos border. Most of the park is covered (Ohler et al., 2002; B. L. Stuart and D. A. Emmett, by bamboo forest and evergreen forest, usually in varying unpublished data). degrees of mixture. Virachey National Park was established on 01 November 1993 under the Royal Decree on Creation Here we report on new herpetological collections made in 2000 and 2003 in the uplands of eastern Mondolkiri, northeastern Ratanakiri, and northeastern Stung Treng Provinces, Cambodia.

STUDY AREAS

The Seima Biodiversity Conservation Area, part of what was formerly the Samling Forest Concession, is situated in Keo Seima and O’Rang Districts, Mondolkiri Province, at the border with Vietnam. The Seima Biodiversity Conservation Area is 305,400 ha in size, and ranges from 109 m elevation in Keo Seima to 800 m elevation in O’Rang. Higher elevations are predominantly covered by evergreen and evergreen mixed with deciduous forests, and lower elevations are dominated by grasslands with deciduous dipterocarp forest, gallery evergreen forest, and bamboo forest. The Cambodian Forestry Administration and Wildlife Conservation Society (WCS) have been working together to promote biodiversity conservation in the Seima Biodiversity Conservation Area since March 2000. The authors collected specimens of amphibians and reptiles at the Seima Biodiversity Conservation Area from 30 October-10 November and 09- 12 December 2003, with a few specimens collected at other times by colleagues (names provided in Species Accounts) that were donated to us for the purpose of this work (Fig. 1).

Phnom Nam Lyr Wildlife Sanctuary is situated in O’Rang and Pichrada Districts, Mondolkiri Province, at the border Fig. 1. Map showing the collecting localities in this study.

130 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Table 1. Complete locality information for district names provided in Species Accounts.

District Protected Area Province Keo Seima Seima Biodiversity Conservation Area Mondolkiri O’Rang Seima Biodiversity Conservation Area Mondolkiri Pichrada Phnom Nam Lyr Wildlife Sanctuary Mondolkiri Ta Veng Virachey National Park Ratanakiri Von Sai Virachey National Park Ratanakiri Siem Pang Virachey National Park Stung Treng

and Designation of Protected Areas in Cambodia, signed by SPECIES ACCOUNTS His Majesty the King Norodom Sihanouk. The authors collected specimens of amphibians and reptiles at Virachey Nineteen species characteristic of anthropogenically-modified National Park from 19 November-05 December 2003 (Fig. environments were collected (Table 2) near villages, 1). agricultural lands, and roads, with some also taken in open grassland or disturbed forest. All are currently recognized to have wide ranges in mainland Southeast Asia. METHODS

Specimens were caught in the field by hand, preserved in Megophryidae 10% buffered formalin, and later transferred to 70% ethanol. Tissue samples were taken by preserving pieces of liver in Leptobrachium mouhoti, new species 95% ethanol or 20% DMSO/EDTA-salt saturated storage (Figs. 2-4) buffer before the specimens were fixed in formalin. Specimens and tissues were deposited and comparative Material examined. – Holotype: FMNH 262756, adult male, in material was examined in The Field Museum, Chicago shallow water among tree roots at bank of O Doeung Por Stream in (FMNH). Comparative material was also examined from the hilly evergreen forest, near 12°18'08.4"N 107°03'08.1"E, 500 m holdings of the Muséum national d’Histoire naturelle, Paris elev., Seima Biodiversity Conservation Area, O’Rang District, Mondolkiri Province, Cambodia; coll. B. L. Stuart, K. Sok, and T. (MNHN). Neang, 2 Nov.2003.

Measurements were made with dial calipers to the nearest Paratypes: FMNH 261757-60, 261762-63 (six adult males), FMNH 0.1 mm. Measurement abbreviations used are: SVL = snout- 261761 (one adult female), evergreen mixed with deciduous and vent length; HDL = head length from tip of snout to rear of bamboo forest, near 12°18'06.4"N 107°03'06.1"E, 600-700 m elev., the jaws; HDW = head width at the commisure of the jaws; Phnom Nam Lyr Mountain, Phnom Nam Lyr Wildlife Sanctuary, SNT = snout length from tip of snout to the anterior corner O’Rang District, Mondolkiri Province, Cambodia, coll. B. L. Stuart, of the eye; EYE = diameter of the exposed portion of the D. An, and P. Suon, 16-17 Jun.2000. FMNH 261764 (one adult eyeball; IOD = interorbital distance; TMP = horizontal male), gallery evergreen forest along stream, near 12°29'49"N 107 29'33"E, 700 m elev., Phnom Nam Lyr Mountain, Phnom Nam diameter of tympanum; TEY = tympanum-eye distance from ° Lyr Wildlife Sanctuary, O’Rang District, Mondolkiri Province, anterior edge of tympanum to posterior corner of the eye; TIB = tibia length; TAL = tail length; TTL = total length; BDW = maximum body width; FOT = foot length from proximal edge of inner metatarsal tubercle to tip of fourth toe; IML = length of inner metatarsal tubercle; IMW = width of inner metatarsal tubercle; CPL = maximum straight carapace length, including shell projections; CPW = maximum carapace width.

Only species with preserved voucher specimens are reported here. Specimens with affinities to Limnonectes hascheanus, Philautus banaensis and Cyrtodactylus irregularis were collected during this fieldwork, but will be treated elsewhere.

For the sake of brevity, localities are given with district name as the largest political unit (Table 1).

Fig. 2. Eye of Leptobrachium mouhoti, new species, holotype male (FMNH 262756), in life.

131 Stuart et al.: Amphibians and reptiles from Cambodia

Table 2. Voucher numbers and district localities of species in the collection that are characteristic of anthropogenically-modified environments.

Taxon FMNH number Locality Bufonidae Bufo melanostictus Schneider 261880, 262751-52 Keo Seima 262750 Siem Pang 262749 Ta Veng Microhylidae Microhyla heymonsi Vogt 262058-59 Pichrada Microhyla ornata (Duméril and Bibron) 262699-700, 262703 Pichrada Microhyla pulchra (Hallowell) 262785 Keo Seima 262677 Pichrada Ranidae Hoplobatrachus rugulosus (Wiegmann) 262748, 262590-91 Keo Seima 262596-97 Pichrada Fejervarya limnocharis (Gravenhorst) 262770-74, 262932 Keo Seima 262769 O’Rang 261990-96 Pichrada Occidozyga lima (Gravenhorst) 261806, 262780-82 Keo Seima 261785-86 Pichrada Occidozyga martensii (Peters) 261807-08, 262825-31 Keo Seima 262823-24 O’Rang 261813-22 Pichrada 262822 Siem Pang Rhacophoridae Polypedates leucomystax (Gravenhorst) 262810-13, 263130-35 Keo Seima 263152-71 Pichrada 262808-09 Siem Pang 262805-07 Von Sai Calotes versicolor (Daudin) 262046-47 Pichrada Scincidae Mabuya macularia (Blyth) 262978-79 Keo Seima 262977 O’Rang 261834 Pichrada Mabuya multifasciata (Kuhl) 262953 Siem Pang 262952 Ta Veng Gekkonidae Cosymbotus platyurus (Schneider) 261831 Keo Seima Gehyra mutilata (Wiegmann) 262976 O’Rang Gekko gecko (Linnaeus) 261848-49 Pichrada Enhydris plumbea (Boie) 259222 Pichrada Ptyas korros (Schlegel) 263017 O’Rang Rhabdophis subminiatus (Schlegel) 263010 Siem Pang 259207-09 Pichrada Bungarus fasciatus (Schneider) 263019 Ta Veng

Cambodia, coll. B. L. Stuart, D. An, and P. Suon, 22 Jun.2000. Comparative material examined: Leptobrachium banae FMNH FMNH 261765 (one adult male), gallery evergreen forest along 258079, Xe Kong Province, Laos. Leptobrachium buchardi FMNH stream, near 12°29'41"N 107°31'09"E, 700 m elev., Phnom Nam 258080-90, Champasak Province, Laos. Leptobrachium chapaense Lyr Mountain, Phnom Nam Lyr Wildlife Sanctuary, O’Rang District, FMNH 258065, 258067-71, Bolikhamxay Province, Laos. Mondolkiri Province, Cambodia, coll. B. L. Stuart, D. An, and P. Leptobrachium hendricksoni FMNH 178260-61, paratypes, Pahang, Suon, 23 Jun.2000. FMNH 262753-55 (three adult males), hilly Malaysia. Leptobrachium pullum FMNH 196015, syntype, Annam, Vietnam. Leptobrachium smithi FMNH 172661-62, 173973-75, evergreen forest, waterfall on O Ngeugn Stream, 12°18'50.5"N Chiang Mai, Thailand; FMNH 263263, Vientiane Province, Laos; 107°06'03.4"E, 500 m elev., Seima Biodiversity Conservation Area, FMNH 258091-92, 258097, 258101-06, Sayaboury Province, Laos. O’Rang District, Mondolkiri Province, Cambodia, coll. B. L. Stuart, Leptobrachium xanthospilum FMNH 252914, 252920, Gia Lai K. Sok, and T. Neang, 11 Dec.2003; FMNH 262757 (one adult Province, Vietnam. Additional data on L. banae and L. xanthospilum male), same data as holotype except date of collection on 6 taken from original description (Lathrop et al., 1998), on L. buchardi Nov.2003. taken from original description (Ohler et al., 2004), on L. chapaense

132 THE RAFFLES BULLETIN OF ZOOLOGY 2006 taken from Lathrop et al. (1998), on L. hendricksoni taken from subarticular tubercle and tip continuing as a fringe to base of original description (Taylor, 1962), on L. pullum taken from original tip, on preaxial and postaxial sides of toe IV to level of tip of description (Smith, 1921) and Matsui et al. (1999), and on L. smithi toe V continuing as a fringe to base of tip, and on toe V to taken from original description (Matsui et al., 1999). Complete data midway between base and tip; a distinct, oval inner metatarsal on L. hasseltii taken from Dubois and Ohler (1998), Iskandar (1998), tubercle, no outer metatarsal tubercle; heels non-overlapping and Matsui et al. (1999). when legs are held at right angles to body. Etymology. – The specific epithet is a patronym honoring Henri Mouhot (1826-1861), French explorer, naturalist, and Skin on dorsum and dorsal surface of forelimbs smooth with earliest scientific collector of amphibian and reptile specimens fine network of ridges; skin on dorsal surface of hindlimbs in hilly eastern Cambodia. smooth; skin on flank, chin, belly, and ventral surface of thigh with dense, small glandular warts; skin on ventral surface of Diagnosis. – A Leptobrachium having males with SVL 51.6- forelimbs and tibiotarsus smooth; large oval axillary gland 64.7 and female 70.2; eye black with a narrow, orange-red on ventrolateral surface slightly posterior to insertion of crescent on outer margin from the anterior corner over the forelimb with body; distinct, whitish, round femoral gland eye to lower rear corner; uniform dark gray dorsum; flank on posteroventral surface of thigh, closer to knee than vent. gray with small light spots; venter gray or brown with small light spots. Colour of holotype in life. – Dorsum uniform dark gray-black, with three scattered round gray spots on back near groin; flank Description of holotype. – Habitus moderately stocky; head dark gray with small, scattered white spots, usually on warts; wider than long. Snout rounded in dorsal view, slightly dorsal surface of forelimb and tibiotarsus with dark gray projecting beyond lower jaw in profile; nostril closer to tip bands; anterior and dorsal surface of thigh with silver-copper of snout than to eye; canthus rostralis distinct, constricted bands; eye black with a narrow, orange-red crescent on outer behind nostrils; lores oblique, slightly concave; diameter of margin from the anterior corner over the eye to lower rear eye less than length of snout; interorbital distance much corner (orange-red colouration visible only in posterior corner greater than width of upper eyelid; tympanum round, weakly of eye in relaxed living animal, remainder visible when upper visible, slightly elevated relative to skin of temporal region, eyelid is maximally retracted); black spot at anterior and diameter smaller than that of the eye, diameter greater than posterior corner of eye and on tympanum; chin, chest, belly the distance between the tympanum and eye; no pineal and ventral surface of thigh gray with small light gray spots ocellus; large, slit-like vocal sac openings on floor of mouth; on warts, gray darkening posteriorly; ventral surface of no nuptial pad. forelimb and tibiotarsus dark gray with distinct light gray marbling. Tips of all fingers blunt, not swollen; relative finger lengths II = IV < I < III; two oval palmar tubercles in contact, low Colour of holotype in preservative. – Dorsum and dorsal callous bumps on ventral surface of fingers; no finger surface of limbs uniform dark gray, with three scattered round webbing. Tips of toes blunt, slightly swollen; relative toe gray spots on back near groin; flank dark gray with small, lengths I < II < V < III < IV; webbing on toe I to base of tip, scattered white spots, usually on warts; grayish-brown on preaxial side of toe II to toe articulation, on postaxial side crossbars on anterior surface of thigh and anterior and of toe II to base of tip, on preaxial side of toe III to level of posterior surface of tibia; eye gray-blue with a narrow, light proximal subarticular tubercle continuing as a fringe to base orange crescent bordered with black on outer margin from of tip, on postaxial side of toe III to midway between proximal the anterior corner over the eye to lower rear corner (visible

Fig. 3. Dorsal view of Leptobrachium mouhoti, new species, Fig. 4. Ventral view of Leptobrachium mouhoti, new species, holotype male (FMNH 262756), in preservative. holotype male (FMNH 262756), in preservative.

133 Stuart et al.: Amphibians and reptiles from Cambodia

Table 3. Measurements (mm) of adult Leptobrachium mouhoti, new species. Abbreviations defined in the text.

Measurement Males Female Holotype and paratypes Paratype Range; Mean ± S.D. (N = 13) (N = 1) SVL 51.6-64.7; 58.3 ± 3.5 70.2 HDL 20.5-24.7; 22.7 ± 1.2 25.0 HDW 23.9-29.0; 26.5 ± 1.4 29.8 SNT 8.6-11.8; 10.1 ± 1.0 11.7 EYE 6.6-7.6; 7.1 ± 0.3 8.1 IOD 6.7-9.5; 8.1 ± 0.9 8.6 TIB 17.3-23.0; 20.2 ± 1.9 22.6 IML 3.1-3.9; 3.6 ± 0.2 4.3 IMW 1.1-1.9; 1.5 ± 0.2 1.5 Range; Median (N = 13) (N = 1) HDL:HDW 0.82-0.88; 0.86 0.84 HDL:SVL 0.37-0.41; 0.39 0.36 TIB:SVL 0.32-0.38; 0.35 0.32 IML:IMW 1.79-3.18; 2.40 2.87 IML:SVL 0.06-0.07; 0.06 0.06

only by retracting upper eyelid); black spot at anterior and red eye colouration restricted to a narrow crescent on the posterior corner of eye and on tympanum; ventral surface anterior, dorsal, and posterior outer margins of eye (upper grayish-brown with small creamy-white spots on warts, half of iris scarlet in L. pullum). Leptobrachium mouhoti concentrated anteriorly, some larger creamy-white spots on differs from L. smithi by having a uniform dark gray dorsum chest; grayish-brown marbling on ventral surface of forelimb (large dark dorsal markings in L. smithi), having the flank and tibia; axillary and femoral glands creamy-white. dark gray with small, scattered white spots (flank with distinct dark spots in L. smithi), and having orange-red eye colouration Measurements. – Holotype: SVL 64.0, HDL 24.7, HDW restricted to a narrow crescent on the anterior, dorsal, and 28.1, SNT 11.8, EYE 7.1, IOD 9.5, TIB 23.0, IML 3.6, IMW posterior outer margins of eye (upper half of iris scarlet or 1.5. yellow in L. smithi).

Variation. – Black spots or streak below edge of canthus in An additional four species of Leptobrachium occur in parts some paratypes. The scattered round gray spots on back near of Laos and Vietnam near to eastern Cambodia: L. buchardi groin are present only in two other specimens, but in different Ohler, Teynié & David, 2004, L. banae Lathrop, Murphy, locations on the back. Ventral pigmentation is more brown Orlov & Ho, 1998, L. chapaense (Bourret, 1937), and L. than gray in some paratypes. Measurements of paratypes xanthospilum Lathrop, Murphy, Orlov & Ho, 1998. summarized in Table 3. Leptobrachium mouhoti differs from L. buchardi by having a narrow orange-red crescent on the anterior, dorsal, and Comparisons. – Four other species of Leptobrachium have posterior outer margins of eye (upper part of iris pale green reddish (orange, red, or scarlet) eye colouration: L. hasseltii in L. buchardi), and having the female with SVL 70.2 (female Tschudi, 1838, L. hendricksoni Taylor, 1962, L. pullum L. buchardi 49.5). Leptobrachium mouhoti differs from L. (Smith, 1921), and L. smithi Matsui, Nabhitabhata & Panha, banae by having gray and silver-copper bands on limbs 1999. Leptobrachium mouhoti differs from L. hasseltii by (orange-red or red bands on limbs in L. banae), and having having a uniform dark gray dorsum (discrete dark rounded a narrow orange-red crescent on the anterior, dorsal, and spots on dorsum in L. hasseltii) and lacking longitudinal rows posterior outer margins of eye (dorsal half of iris and outer of ridges on hindlimb (present in L. hasseltii). Leptobrachium margin of eye white in L. banae). Leptobrachium mouhoti mouhoti differs from L. hendricksoni by having males with differs from L. chapaense by having a narrow orange-red SVL 51.6-64.7 (male L. hendricksoni 45-46), having a dark crescent on the anterior, dorsal, and posterior outer margins gray dorsum (lavender-brown dorsum in L. hendricksoni), of eye (upper half of iris black, white, or sky blue in L. and having the venter gray or brown with small light spots chapaense). Leptobrachium mouhoti differs from L. (venter white with small black spots in L. hendricksoni). xanthospilum by having males with SVL 51.6-64.7 and Leptobrachium mouhoti differs from L. pullum by having female 70.2 (male L. xanthospilum 62.8-73.4, female L. males with SVL 51.6-64.7 and female 70.2 (male L. pullum xanthospilum 83.2-84.8), having the flank dark gray with 44.4-47.0, female L. pullum 49.4-52.8) and having orange- small, scattered white spots (flank with distinct large yellow

134 THE RAFFLES BULLETIN OF ZOOLOGY 2006 spots in L. xanthospilum), and having a narrow orange-red crescent on the anterior, dorsal, and posterior outer margins Comparative material examined: Ophryophryne gerti FMNH of eye (upper half of iris white in L. xanthospilum). 252899, 252901, Gia Lai Province, Vietnam. Ophryophryne hansi FMNH 252880, holotype, Gia Lai Province, Vietnam; FMNH 252873, 252875, 252878-79, 252882, 252884, 252892-93, Ecology. – Specimens were collected at night on leaf litter paratypes, Gia Lai Province, Vietnam. Ophryophryne microstoma of the forest floor, often near streams. The species was FMNH 254250-51, Vinh Phu Province, Vietnam. Additional data frequently heard calling at night in June, November, and on O. gerti and O. microstoma taken from original descriptions December, suggesting it calls year-round. FMNH 262754 was (Boulenger, 1903; Ohler, 2003). Complete data on Ophryophryne calling from a muddy depression under a 25 cm diameter pachyproctus taken from original description (Kou, 1985) and Ohler rock 2.5 m from a wide, swift stream, and FMNH 261762 (2003). was calling from a depression under a log 4 m from a stream. FMNH 261760-61 were in amplexus at night while floating Diagnosis. – An Ophryophryne having males with SVL 45.8- in a shallow, 3 m wide stream pool with sandy substrate. 47.3; tympanum diameter 80% of the eye diameter; second finger shorter than fourth finger; small tubercles containing black spinules on maxilla, mandible, and rear of back; fine, Ophryophryne hansi Ohler colourless spinules on dorsal surface of tibia and ventral surface of tarsus; strong dorsolateral glandular ridge from Material examined. – Siem Pang: FMNH 262777, bamboo mixed above shoulder to level of 4/5 of distance between axilla and with evergreen forest, O Chay Stream, 14°17'38.7"N 106°36'44.8"E, groin; large tubercles on rear of back; large warts on flanks; 370 m elev., 2 Dec.2003. black nuptial pad covering most of dorsal surface of first phalange of second finger. Remarks. – A single male (SVL 45.6) closely resembles the holotype (FMNH 252880) and male paratypes (FMNH Description of holotype. – Habitus moderately stocky; head 252873, 252875, 252878-79, 252884, 252892-93), which we slightly wider than long. Snout truncate in dorsal view, have examined, from Gia Lai Province, Vietnam. The obliquely projecting beyond lower jaw in profile; nostril specimen has small tubercles on the side of the head; larger closer to tip of snout than to eye; canthus rostralis constricted, tubercles on the top of the head, upper eyelid, and dorsum; rounded; lores vertical; diameter of eye slightly greater than warts on the flank and groin; a distinct supratympanic fold length of snout; interorbital distance slightly smaller than from eye to behind axilla; no skin folds or ridges on the dorsal width of upper eyelid; single, distinct, pointed supraorbital surface of the body; a dark brown oval nuptial pad on the horn projecting from upper eyelid, about equal in length to dorsal and medial surface of the first finger from its base to distance from nostril to tip of snout; tympanum round, visible, the articulation and in a small round patch on the dorsal slightly elevated relative to skin of temporal region, diameter surface of the base of the second finger; the second finger smaller than that of the eye, diameter greater than the distance slightly shorter than the fourth finger; the tympanum to eye between the tympanum and eye; no pineal ocellus. distance greater than the horizontal diameter of the tympanum; and dark gray colouration in preservative. The Tips of all fingers blunt, slightly swollen near tips; relative Cambodian specimen disagrees with Ohler’s (2003) original finger lengths I < II < IV < III; palmar tubercle indistinct, no description by having the tympanum diameter larger than half subarticular or supernumerary tubercles; no finger webbing. (62.5%) of the eye diameter, and is slightly larger than males Tips of toes like fingers; relative toe lengths I < II < V < III in the type series (SVL 35.3 – 43.0 mean ± SD 38.8 ± 2.5, N = 7). The specimen illustrated in Figure 2 of Ohler (2003) is not the holotype of O. hansi (FMNH 252880), nor is it referable to this species, but the text description of the holotype matches FMNH 252880.

The specimen was taken at night (1918 hrs.) on a 5 cm diameter rock midstream in a 7 m wide shallow, swift rocky stream.

This is the first report of the genus in Cambodia.

Ophryophryne synoria, new species (Fig. 5)

Material examined. – Holotype: FMNH 262779, adult male, O Chung Chry Stream, near 12°17'30"N 107°03'06"E, 500 m elev., Seima Biodiversity Conservation Area, O’Rang District, Mondolkiri Province, Cambodia, coll. B. L. Stuart, K. Sok, and T. Neang, 4 Nov.2003. Fig. 5. Ophryophryne synoria, new species, holotype male (FMNH 262779), in life. Paratype: FMNH 262778, adult male, collected with the holotype. 135 Stuart et al.: Amphibians and reptiles from Cambodia

< IV; no webbing between toes I and II, webbing confined to the second finger shorter than the fourth finger (equal in O. base between other toes; an oval, indistinct inner metatarsal gerti). Ophyrophryne synoria differs from O. hansi Ohler, tubercle, no outer metatarsal tubercle. 2003, by having a strong dorsolateral glandular ridge from above shoulder to level of 4/5 of distance between axilla and Skin on dorsum shagreened, small tubercles on tympanum, groin (absent in O. hansi) and having a black nuptial pad temporal region, snout, and from posterior half of upper eyelid covering most of the dorsal surface of the first phalange of to area above shoulder, larger tubercles on rear of back; small the second finger (black nuptial pad on the second finger tubercles containing black spinules on maxilla, mandible, and small, round, and weakly visible in O. hansi). Ophyrophryne rear of back; fine, short, colourless spinules on dorsal surface synoria differs from O. microstoma Boulenger, 1903, by of tibia and ventral surface of tarsus; distinct glandular having a strong dorsolateral glandular ridge from above supratympanic fold from eye to behind axilla; strong shoulder to level of 4/5 of distance between axilla and groin dorsolateral glandular ridge from above shoulder to level of (absent in O. microstoma), having large tubercles on rear of 4/5 of distance between axilla and groin; a delicate glandular back (absent in O. microstoma), having large warts on the V-shaped ridge on dorsum, with tips at level of tympanum flanks (granules in O. microstoma), and having the second and apex at level slightly posterior to shoulder; a delicate finger shorter than the fourth finger (equal in O. microstoma; glandular inverted V-shaped ridge on dorsum, with tips at Ohler, 2003). Ophyrophryne synoria differs from O. level of 2/3 of distance between axilla and groin and apex at pachyproctus Kou, 1985 by having males with SVL 45.8- level of 1/2 of distance between axilla and groin; a delicate, 47.3 (male O. pachyproctus SVL 28-30) and lacking the anus glandular, medial longitudinal ridge connecting these apices; terminal on a dermal protuberance (present in O. one to three delicate oblique ridges on dorsal surface of tibia; pachyproctus). two small transverse folds on posterior edge of upper eyelid; large warts on sides intermixed with smaller warts; two small Ecology. – The holotype and paratype were collected together pointed, projecting dermal flaps above vent; skin on venter at night in a swift, 30 cm wide stream flowing over solid shagreened; black nuptial pad on dorsal and medial surface rock substrate in hilly evergreen forest. The paratype was of finger I from its base to near distal joint and on most of taken at 1935 hrs. on a boulder under a fallen tree at the edge dorsal surface of the first phalange of finger II. of the water. The holotype was taken at 1940 hrs. under a rock ledge 15 cm from the water. Colour of holotype in life. – Dorsum olive-brown with large, yellowish-green spots; dorsal surface of head yellowish-green from tip of snout to eyes, olive-brown V-shaped marking on Bufonidae crown between supraorbital horns with apex pointing posteriorly; temporal region olive-brown; pupil black, Bufo galeatus Günther outlined in orange, diamond-shaped in horizontal orientation; (Fig. 6) iris copper with black streaks radiating from pupil; upper surface of limbs with dark gray and black bands; black spots Material examined. – Pichrada: FMNH 261766-67, evergreen on outer surface of forelimbs and tibia; sides purplish-gray mixed with deciduous and bamboo forest, Phnom Nam Lyr with grayish-white flecking and large black spots; throat Mountain, along O Nam Lyr Stream, near 12°32'16"N 107°32'00"E, uniform dark purplish-gray; chest and anterior half of belly 600 m elev., 17 Jun.2000; FMNH 261768-69, evergreen mixed with deciduous and bamboo forest, Phnom Nam Lyr Mountain, near purplish-gray; posterior half of belly and lower surface of 12°32'16"N 107°32'00"E, 600-700 m elev., 18-21 Jun.2000. limbs purplish-gray with grayish-white flecking; vent and posterior surface of thigh near vent black, posterior surface O’Rang: FMNH 262759, hilly evergreen forest, O Rokhlong Stream, of thigh near tibio-tarsal articulation dark gray with black near 12°18'35.3"N 107°04'28.0"E, 500 m elev., 12 Dec.2003; FMNH and white spots; dorsal surface of foot and tarsus white with 262760, hilly evergreen forest, O Ronas Stream, 12°17'30.7"N black flecking; ventral surface of foot and tarsus black. 107°03'06.0"E, 450 m elev., 3 Nov.2003; FMNH 262761-62, hilly

Measurements. – Holotype: SVL 47.3, HDL 13.8, HDW 14.4, SNT 4.0, EYE 4.3, IOD 3.5, TMP 3.5, TEY 2.5, TIB 22.4. Paratype: SVL 45.8, HDL 13.1, HDW 14.2, SNT 3.7, EYE 4.5, IOD 3.5, TMP 3.6, TEY 2.0, TIB 21.2.

Etymology. – The specific epithet from synoria (Greek) for borderland, referring to the proximity of the type locality to the Vietnam border.

Comparisons. – Ophyrophryne synoria differs from O. gerti Ohler, 2003, by having males with SVL 45.8-47.3 (male O. gerti 32.0-34.8), having the tympanum diameter 80% of the eye diameter (tympanum diameter about half the eye diameter in O. gerti), having the tympanum to eye distance 55-71% of Fig. 6. Female Bufo galeatus from Phnom Nam Lyr Wildlife the tympanum diameter (about equal in O. gerti), and having Sanctuary, Pichrada District, Mondolkiri Province, Cambodia.

136 THE RAFFLES BULLETIN OF ZOOLOGY 2006 evergreen forest, O Ngeugn Stream, near 12°18'58"N 107°05'59"E, marking; FMNH 262689 has an inverted V-shaped dark 550 m elev., 5 Nov.2003; FMNH 262763, hilly evergreen forest, O marking, and scattered round, black spots; and FMNH 262690 Doeung Por Stream, near 12°18'08"N 107°03'08"E, 500 m elev., has no distinct markings. All four specimens have a black 6 Nov.2003; FMNH 262764, hilly evergreen with bamboo forest, ocellar spot edged in white at the groin. 12°15'41.6"N 107°03'48.2"E, 480 m elev., 7 Nov.2003.

Siem Pang: FMNH 262758, bamboo mixed with evergreen forest, FMNH 262688 was collected at night (2030 hrs.) on the bare 14°17'39.5"N 106°37'42.3"E, 400 m elev., 2 Dec.2003. surface of a rock outcrop, and FMNH 262690 was taken on a forest trail in the morning (0930 hrs.), both away from water. Remarks. – These agree with Günther’s (1864) original description of a single specimen from Cambodia, and the Ohler et al. (2002) reported the species from the mountains expanded descriptions by Smith (1921) and Inger et al. (1999) of southwestern Cambodia. Parker (1934) and Bourret (1942) based on collections from Vietnam. The specimens have a reported the species from Cambodia, without specific bony ridge connecting the eye with the parotoid gland; large, localities. conical, pointed tubercles in an oblique band from the corner of the mouth to above the axilla; and similar but slightly smaller tubercles on the sides and limbs, the largest of these Microhyla berdmorei (Blyth) in a row from the parotoid to the groin. As reported by Inger et al. (1999), females have conspicuously longer and more Material examined. – Keo Seima: FMNH 262838, hilly evergreen pointed tubercles on the sides of the head and body than do mixed with deciduous and bamboo forest, near 12°10'34.9"N 106 57'47.6"E, 190 m elev., 9 Nov.2003; FMNH 262839-42, males, and females (SVL 75.3 – 88.5, mean ± SD 83.4 ± 7.1, ° bamboo mixed with evergreen forest, near 12°16'45.7"N N = 3) are considerably larger than males (SVL 49.9 – 60.2, 106°56'36.4"E, 180 m elev., 8 Nov.2003. mean ± SD 55.1 ± 3.4, N = 6). Males have a black nuptial pad on the dorsal and medial surface of the first and second O’Rang: FMNH 262837, hilly evergreen forest, near 12°16'24.6"N fingers. 107°03'53.1"E, 350 m elev., 10 Dec.2003.

Specimens were collected day and night on forest trails and Pichrada: FMNH 262653, Phnom Nam Lyr Mountain, evergreen at night along streams. FMNH 262759 was taken at night mixed with deciduous and bamboo forest, near 12°32'16"N 107°32'00"E, 600-700 m elev., 20 Jun.2000. (2015 hrs.) in shallow water of a 4 m wide seep running over a large rock face and FMNH 262760 was found at night (1840 Siem Pang: FMNH 262833-36, bamboo mixed with evergreen forest, hrs.) on a sand and boulder mid-stream bank in a 3 m wide tributary of O Chay Stream, 14°18'28.6"N 106°33'02.6"E, 310 m flowing stream. FMNH 261766-67 were in amplexus in June elev., 5 Dec.2003. on a log emerging from a small (<1 m wide) flowing stream with sandy substrate. Ta Veng: FMNH 262832, hilly evergreen forest mixed with bamboo forest, O Greng Mak Stream, 14°12'00.5"N 107°18'29.8"E, 200 m elev., 20 Nov.2003. This is the first report of the species in Cambodia since its original description. Remarks. – An adult female (SVL 41.7) and nine adult males (SVL 30.5 – 40.0, mean ± SD 35.1 ± 3.5, N = 9) have an obtusely pointed snout; dark throat; toes fully webbed, Microhylidae reaching base of expanded discs on toes; third and fifth toes equal in length; and outer and inner metatarsal tubercles. Kalophrynus interlineatus (Blyth) Males have a strong fold across the throat.

Material examined. – Keo Seima: FMNH 262687, along small river Specimens were collected day and night on sandy, gravel or in disturbed evergreen mixed with deciduous forest, 150 m elev., rocky banks of slow-moving streams, and on the forest floor coll. J. Walston, May.2000. away from water. Pichrada: FMNH 262688, rock hill at base of Phnom Nam Lyr Mountain, near 12°32'16"N 107°32'10"E, 700 m elev., 18 Jun.2000; Bourret (1942) reported the species from north-central FMNH 262689-90, Phnom Nam Lyr Mountain, evergreen mixed Cambodia, and both Bourret (1942) and Ohler et al. (2002) with deciduous and bamboo forest, near 12°32'16"N 107°32'00"E, reported the species from the mountains of southwestern 600-700 m elev., 21-22 Jun.2000. Cambodia.

Remarks. – These agree with the expanded descriptions by Parker (1934; as a subspecies of K. pleurostigma Tschudi) Ranidae and Matsui et al. (1996) by having toes less than one-third webbed, with the third toe webbing not extending beyond Limnonectes dabanus (Smith) the distal subarticular tubercle; and the free portion of the (Fig. 7) fifth toe longer than the distance from snout to nostril. The dorsal pattern is variable, as reported by Parker (1934): Material examined. – O’Rang: FMNH 262747, hilly evergreen FMNH 262687 has a narrow dark streak commencing on the forest, O Ronas Stream, 12°17'30.7"N 107°03'06.0"E, 450 m elev., tip of the snout; FMNH 262688 has a reticulated dark 3 Nov.2003.

137 Stuart et al.: Amphibians and reptiles from Cambodia

Pichrada: FMNH 261924-27, Phnom Nam Lyr Mountain, evergreen Specimens were collected in and along small streams. This mixed with deciduous and bamboo forest along O Nam Lyr Stream, was the most frequently encountered, forest-dwelling frog near 12°32'16"N 107°32'00"E, 600 m elev., 16 Jun.2000; FMNH species during our fieldwork. A developed male (FMNH 261929-32, 261935, 261937, 261942, 261944-45, Phnom Nam Lyr 261937) was collected at night (2015 hrs.) in June sitting in Mountain, evergreen mixed with deciduous and bamboo forest, near a depression about twice its body size in a sandy vertical bank 12°32'16"N 107°32'00"E, 600-700 m elev., 17-21 Jun.2000. with exposed tree roots, 15 cm above a stream pool. The Siem Pang: FMNH 262744-45, bamboo mixed with evergreen forest, depression appeared to have been constructed by the frog, near 14°16'03.7"N 106°37'44.6"E, 600 m elev., 1 Dec.2003; FMNH possibly for breeding activity. 262920, bamboo mixed with evergreen forest, O Chay Stream, 14°17'38.7"N 106°36'44.8"E, 370 m elev., 02 Dec.2003. This is the first report of the species in Cambodia since Bourret’s (1941b) description of its junior synonym R. Remarks. – Seventeen adult males fully agree with Smith’s toumanoffi from Mimot. (1922) brief original description of Rana macrognathus dabana from the Langbian Plateau of southern Vietnam by having distinctly enlarged heads, encompassing nearly half Limnonectes kuhlii (Tschudi) the SVL; a swelling consisting of dense fibrous tissue on top of the head, commencing at the interorbital region and Material examined. – Siem Pang: FMNH 262726-30, bamboo extending to or near to the level of the posterior border of the mixed with evergreen forest, near 14°16'03.7"N 106°37'44.6"E, 550- tympanum; the heels not or only slightly touching when the 600 m elev., 01 Dec.2003. hindlimbs are folded at right angles to the body; and in measurements (Table 4). The specimens also fully agree with Ta Veng: FMNH 262722-25, hilly evergreen mixed with bamboo forest, O Lopeung Stream, 14 11'16.3"N 107 17'36.1"E, 160 m the holotype (MNHN 1948.0126) of Rana toumanoffi ° ° elev., 19-21 Nov.2003. Bourret, 1941b, a single male from Mimot, Cambodia, which we have examined. There are no apparent differences in body Remarks. – Three males (SVL 60.3 – 66.5, mean ± SD 63.7 measurements among Smith’s syntypes, our Cambodian ± 3.1, N = 3) and six females (SVL 50.1 – 61.6, mean ± SD specimens, and Bourret’s holotype of R. toumanoffi (Table 55.5 ± 3.9, N = 6) have the tympanum hidden; a 1), although the degree of enlargement of the head and head supratympanic fold from the eye to behind axilla; low, swelling varies considerably among individual males, even radiating intersecting ridges on the dorsum; heavy wrinkling from the same locality. We treat Rana toumanoffi as a junior on flanks; rounded, white asperities on the dorsal surface of synonym of Limnonectes dabanus. the tibia and tarsus; and all toes fully webbed to disc. The head and odontoids at the front of the mandible are larger in Smith (1922) stated that juveniles and females of the species males than in females. belonging to the Rana doriae group, in which he included L. dabanus, resemble each other so closely that “it is practically Specimens were collected submerged in shallow water or on impossible to distinguish between them.” A large series the bank within 30 cm of rocky seeps, small rocky streams, (FMNH 261928, 261933-34, 261936, 261938-41, 261943, and side pools of rocky streams. 262665, 262669, 262743, 262746, 262902-19, 262921-31, 262933-38, 262940-51, N = 60) of females, juveniles, and This is the first report of the species from Cambodia. undeveloped males are tentatively assigned here to L. dabanus because they generally agree in skin texture and size (except head measurements) and were collected with the 17 developed Limnonectes poilani (Bourret) males. Nine of the 77 (11.7%) specimens have a light vertebral (Fig. 8) stripe from the tip of the snout to near the vent. Material examined. – O’Rang: FMNH 262741, hilly evergreen forest, O Rokhlong Stream, near 12°18'35.3"N 107°09'28.0"E, 500 m elev., 12 Dec.2003; FMNH 262742, hilly evergreen forest, O Kamen Stream, near 12°19'35"N 107°05'33"E, 500 m elev., 01 Nov.2003.

Pichrada: FMNH 262659-64, Phnom Nam Lyr Mountain, evergreen mixed with deciduous and bamboo forest along O Nam Lyr Stream, near 12°32'16"N 107°32'00"E, 600 m elev., 16 Jun.2000; FMNH 262666-68, Phnom Nam Lyr Mountain, evergreen mixed with deciduous and bamboo forest, near 12°32'16"N 107°32'00"E, 600- 700 m elev., 17 Jun.2000; FMNH 262670-71, gallery evergreen forest along stream, near 12°29'49"N 107°29'33"E, 700 m elev., 22 Jun.2000.

Remarks. – Bourret (1942) described poilani as a subspecies of Rana kohchangae (Smith, 1922) based on a single male Fig. 7. Male Limnonectes dabanus from Phnom Nam Lyr Wildlife from Dong Tam Ve, Vietnam. Ohler et al. (2002) moved Sanctuary, Pichrada District, Mondolkiri Province, Cambodia.

138 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Table 4. Measurements of male Limnonectes dabanus (Smith, 1922): syntypes of Rana macrognathus dabana, new Cambodian specimens, and the holotype of Rana toumanoffi. Measurements of Rana macrognathus dabana were taken from Smith (1922); all other measurements were taken by the present authors. Abbreviations are given in the text.

Measurement Rana macrognathus dabana FMNH Cambodian specimens Rana toumanoffi Smith, 1922 Bourret, 1941b 4845, 2637, 2547, 4844 MNHN 1948.0126 holotype syntypes Range; Mean ± S.D. Range; Mean ± S.D. (N=4) (N=17) (N=1) SVL 51.0 – 55.0; 53.0 ± 1.8 49.7 – 64.6; 57.6 ± 4.6 54.9 HDL 21.0 – 22.0; 21.8 ± 0.5 25.8 – 39.4; 32.4 ± 3.6 30.1 HDW 24.0 – 26.0; 25.0 ± 0.8 24.0 – 35.1; 29.6 ± 3.3 27.7 SNT 9.5 – 10.0; 9.9 ± 0.3 9.3 – 14.7; 12.3 ± 1.4 12.3 EYE 5.5 – 6.0; 5.8 ± 0.3 5.0 – 6.1; 5.7 ± 0.3 6.0 IOD 5.0 – 6.0; 5.6 ± 0.5 5.6 – 8.5; 7.3 ± 1.0 7.0 TMP 5.5 – 6.0; 5.8 ± 0.3 6.2 – 9.7; 8.0 ± 0.9 7.0 TIB 24.0 – 26.0; 25.3 ± 1.0 22.8 – 29.5; 26.1 ± 1.8 28.3 FOT 24.0 – 26.0; 25.3 ± 1.0 22.6 – 29.5; 26.5 ± 1.9 26.9 Range; Median (N = 4) Range; Median (N = 17) (N = 1) TIB:SVL 0.46-0.49; 0.48 0.43-0.49; 0.45 0.51

poilani to species rank and into the genus Limnonectes. We males have enlarged odontoids at the front of the mandible examined Bourret’s holotype of Rana kohchange poilani and enlarged heads with two temporal swellings. Bourret (MNHN 1948.0127) and it is clearly conspecific with adult (1942) described a median longitudinal ridge on the top of males in our collection from Cambodia. The Cambodian the head from the level of the posterior edge of the eyes to series also fully agrees with specimens of both sexes reported the level of the insertion of the forelimbs to the body; this as Rana cf. blythii from central Vietnam by Inger et al. (1999), character is apparent in the holotype, but not in the Cambodian and these should also be referred to L. poilani. Specifically, males. Two of four adult males and four of six adult females males (SVL 84.1 – 99.5, mean ± SD 90.0 ± 6.7, N = 4) and have a light vertebral stripe from the tip of the snout to the females (SVL 69.5 – 95.3, mean ± SD 80.2 ± 8.7, N = 6) vent. from Cambodia have a ridge of skin on the medial border of the outer three fingers, widest on the second finger; the first The colouration in life of the adult female FMNH 262741 and fifth toes have movable flaps of skin externally; the first follows. Brown above with dark brown and black spotting three toes and the fifth are fully webbed to the base of the on the dorsum and limbs; creamy-tan band followed swollen tips; the webbing on the fourth toe reaches the distal posteriorly by a black band between the upper eyelids; dark subarticular tubercle and continues to the tip as a movable brown lip bars; throat white with grey flecking, anterior half flap of skin; tubercles on the upper eyelid; two black bars of venter and underside of forelimbs white, posterior half of extend from the eye to lip; and a black streak covers the upper venter and underside of hindlimbs yellow; posterior surface half of the tympanum and the supratympanic fold. The adult of thigh with yellow and black reticulations; pinkish-orange wash on upper surface of fingers and toes.

The specimens were collected in shallow water or on dirt and leaf litter banks within 3 m of small streams. Orlov (1997) described the breeding behavior of this species (as Rana cf. blythii) in Vietnam.

This is the first report of the species in Cambodia.

Rana attigua Inger, Orlov & Darevsky

Material examined. – Pichrada: FMNH 261920-22, Phnom Nam Lyr Mountain, evergreen mixed with deciduous and bamboo, near 12°32'16"N 107°32'00"E, 600-700 m elev., 21 Jun.2000; FMNH Fig. 8. Limnonectes poilani from Phnom Nam Lyr Wildlife 261923, gallery evergreen forest along stream, near 12°29'49"N Sanctuary, Pichrada District, Mondolkiri Province, Cambodia. 107°29'33"E, 700 m elev., 22 Jun.2000.

139 Stuart et al.: Amphibians and reptiles from Cambodia

Remarks. – Four males with nuptial pads (SVL 34.0 – 36.6, O’Rang: FMNH 262739, 262899, hilly evergreen forest, O Chung mean ± SD 35.5 ± 1.3, N = 4) agree with the original Chry Stream, near 12°17'30"N 107°03'06"E, 500 m elev., 04 description (Inger et al., 1999) and the FMNH type series Nov.2003; FMNH 262892-98, hilly evergreen forest, O Ronas from Gia Lai Province, Vietnam, except the Cambodian males Stream, 12°17'30.7"N 107°03'06.0"E, 450 m elev., 03 Nov.2003; FMNH 262796, hilly evergreen forest, near 12 16'24.6"N are slightly smaller than males with nuptial pads in the type ° 107°03'53.1"E, 350 m elev., 10 Dec.2003; FMNH 262797-99, hilly series (SVL 36.2 – 45.5, mean ± SE 41.3 ± 0.3, N = 43; Inger evergreen forest, waterfall on O Ngeugn Stream, 12°18'50.5"N et al., 1999), and the nuptial pads have a distinct notch in the 107°06'03.4"E, 500 m elev., 11 Dec.2003; FMNH 262740, 262800- ventral margin. These discrepancies are noteworthy because 01, 262887-88, hilly evergreen forest, O Ngeugn Stream, near two of the characteristics used by Inger et al. (1999) to 12°18'58"N 107°05'59"E, 550 m elev., 30 Oct.-05 Nov.2003. distinguish R. attigua from the similar, co-occurring R. milleti Smith, 1921, at the type locality in Vietnam were body size Remarks. – These agree with Bain et al.’s (2003) original and condition of the nuptial pad. Specifically, Inger et al. description by having males with SVL 46.7 – 60.0 (mean ± (1999) reported that R. milleti males with nuptial pads have SD 55.4 ± 3.9, N = 10); males with gular pouches; females SVL 34.2 – 40.4 (mean ± SE 37.0 ± 0.2, N = 52), but only with uniformly pale yellow eggs; and weakly visible glandular four of 43 male R. attigua were less than 39.0, and the nuptial dorsolateral folds. Females are much larger than males, with pad in R. milleti has a distinct notch or constriction in the SVL 76.5– 108.7 (mean ± SD 96.0 ± 10.8, N = 10). ventral margin, but that of R. attigua is straight. We re- examined specimens assigned to R. milleti (FMNH 253425- Specimens were collected at night on banks, rock faces, 86) by Inger et al. (1999), and only about half of the adult boulders, tree branches, and tree roots along and above swift males have a notch or constriction in the ventral margin of rocky streams, often near cascades. The stomach of female the nuptial pad, rendering this character to be non-diagnostic. FMNH 262798 (SVL 104.5) contained a large centipede and The most conspicuous differences in preservative between large wasp. The ovaries of FMNH 262800 (SVL 108.7) R. attigua and R. milleti are those of colouration, as stated by contained 2,966 eggs of approximately 2-3 mm diameter. Inger et al. (1999): the underside of the thigh is immaculate whitish and the underside of the calf is immaculate or with This is the first report of the species since its description from only faint dark speckling in R. attigua, but the underside of Gia Lai Province, Vietnam. the thigh has dark speckling and the underside of the calf has dark spotting in R. milleti; and the rear of the thigh has only a light dusting of dark pigment in R. attigua, but the rear of Rana johnsi Smith the thigh has a distinct pattern of dark brown forming a network around small light spots or blotches in R. milleti. Material examined. – Pichrada: FMNH 261882, deciduous forest with grassy understory and bamboo, near 12°29'49"N 107°29'33"E, Three were taken in a flooded grassy clearing in the forest, 700 m elev., 23 Jun.2000. and one was on leaf litter 2 m from a small stream. Remarks. – A single mature female (SVL 56.5) agrees with This is the first report of the species from Cambodia. The Smith’s (1921) original description and Inger et al.’s (1999) similar R. milleti was recently reported from southwestern amplified description based on material from central and Cambodia (Chuaynkern et al., 2004). southern Vietnam in having long legs (TIB:SVL 0.69 in the Cambodian specimen; 0.62 – 0.68, median 0.64, N = 6 females, Smith 1921); a distinct, continuous dorsolateral fold; Rana banaorum Bain, Lathrop, Murphy, Orlov, & Ho full webbing to disc on toes III and V, webbing to distal subarticular tubercle on toe IV and continuing to disc as a Material examined. – Pichrada: FMNH 263255-58, gallery narrow fringe; an inverted, V-shaped glandular fold between evergreen forest along large stream, near 12°31'18"N 107°33'13"E, the shoulders; a dark streak below canthus from tip of snout 600 m elev., 19 Jun.2000; FMNH 263253-54, 263259, Phnom Nam to eye; and a dark patch from the eye to shoulder, enclosing Lyr Mountain, evergreen mixed with deciduous and bamboo, near the tympanum. 12°32'16"N 107°32'00"E, 600-700 m elev., 17-21 Jun.2000; FMNH 263247-52, Phnom Nam Lyr Mountain, evergreen mixed with The specimen was taken in the morning (0700 hrs.) in an deciduous and bamboo along O Nam Lyr Stream, near 12°32'16"N open grassy area. 107°32'00"E, 600 m elev., 16 Jun.2000.

Siem Pang: FMNH 262880, bamboo mixed with evergreen forest, This is the first report of the species from Cambodia. tributary of O Chay Stream, near 14°19'09.8"N 106°35'37.5"E, 325 m elev., 3 Dec.2003. Rana macrodactyla (Günther) Ta Veng: FMNH 262873, hilly evergreen mixed with bamboo forest, O Greng Mak Stream, 14°12'00.5"N 107°18'29.8"E, 200 m elev., Material examined. - Pichrada: FMNH 262061, Bou Sraa Village, 20 Nov.2003; FMNH 262870, 262874-75, 262878, hilly evergreen 12°32'05"N 107°25'55"E, 700 m elev., 14 Jun.2000. mixed with bamboo forest, O Lopeung Stream, 14°11'16.3"N 107°17'36.1"E, 160 m elev., 19-22 Nov.2003. Remarks. – A single female (SVL 38.5) has a very slender habitus; narrow, elongated head; fourth toe very long,

140 THE RAFFLES BULLETIN OF ZOOLOGY 2006 webbing on fourth toe reaching the middle subarticular Pichrada: FMNH 261971-74, evergreen mixed with deciduous and tubercle, continuing on postaxial side of toe to the base of bamboo forest, Phnom Nam Lyr Mountain, along O Nam Lyr disc as a movable flap of skin; distinct, broad, light-coloured Stream, near 12°32'16"N 107°32'00"E, 600 m elev., 16 Jun.2000; dorsolateral fold; dorsum olive with dark brown spots; light- FMNH 261975, Phnom Nam Lyr Mountain, evergreen mixed with deciduous and bamboo forest, near 12 32'16"N 107 32'00"E, 600- coloured medial longitudinal stripe from tip of snout to vent; ° ° 700 m elev., 17 Jun.2000; FMNH 261976-77, gallery evergreen and dark longitudinal stripes on posterior surface of thigh. forest along large stream, near 12°31'18"N 107°33'13"E, 600 m elev., 19 Jun.2000; FMNH 261978, gallery evergreen forest along stream, The specimen was taken on a log in a grassy area near a rice near 12°29'49"N 107°29'33"E, 700 m elev., 22 Jun.2000. paddy. O’Rang: FMNH 262858-61, hilly evergreen forest, O Ngeugn The species has been reported from localities throughout Stream, near 12°18'58"N 107°05'59"E, 550 m elev., 30-31 Oct.2003; Cambodia (Bourret, 1942; Ohler et al., 2002). FMNH 262862, hilly evergreen forest, O Kamen Stream, near 12°19'35"N 107°05'33"E, 500 m elev., 1 Nov.2003; FMNH 262863, hilly evergreen forest, O Ronas Stream, 12°17'30.7"N 107°03'06.0"E, 450 m elev., 3 Nov.2003; FMNH 262864, hilly Rana morafkai Bain, Lathrop, Murphy, Orlov, & Ho evergreen forest, O Chung Chry Stream, near 12°17'30"N 107°03'06"E, 500 m elev., 4 Nov.2003. Material examined. – Siem Pang: FMNH 262732-33, 262881-82, evergreen mixed with bamboo forest, near 14°18'28.9"N Siem Pang: FMNH 262851, bamboo mixed with evergreen forest, 106°33'32.1"E, 430 m elev.; 4 Dec.2003; FMNH 262883-85, O Chay Stream, 14°17'38.7"N 106°36'44.8"E, 370 m elev., 2 evergreen mixed with bamboo forest, waterfall of O Chhay Stream, Dec.2003; FMNH 262852-53, bamboo mixed with evergreen forest, tributary of O Chay Stream, 14°15'49.6"N 106°31'46.0"E, 200 m tributary of O Chay Stream, near 14°19'09.8"N 106°35'37.5"E, 325 elev.; 5 Dec.2003. m elev., 3 Dec.2003; FMNH 262854, evergreen mixed with bamboo forest, waterfall of O Chhay Stream, tributary of O Chay Stream, Ta Veng: FMNH 262731, hilly evergreen forest mixed with bamboo, 14°15'49.6"N 106°31'46.0"E, 200 m elev., 5 Dec.2003; FMNH O Greng Mak Stream, 14°12'00.5"N 107°18'29.8"E, 200 m elev., 262855, bamboo mixed with evergreen forest, tributary of O Chay 20 Nov.2003; FMNH 262871-72, 262876-77, 262879, hilly Stream, 14°18'28.6"N 106°33'02.6"E, 310 m elev., 5 Dec.2003; evergreen mixed with bamboo forest, O Lopeung Stream, FMNH 262856-57, evergreen mixed with bamboo forest, near 14°11'16.3"N 107°17'36.1"E, 160 m elev., 19-22 Nov.2003. 14°18'28.9"N 106°33'32.1"E, 430 m elev., 4 Dec.2003.

O’Rang: FMNH 262738, O Doeung Por Stream, hilly evergreen Ta Veng: FMNH 262843-44, 262850, hilly evergreen forest mixed forest, near 12°18'08.4"N 107°03'08.1"E, 500 m elev., 2 Nov.2003; with bamboo, O Lopeung Stream, 14°11'16.3"N 107°17'36.1"E, 160 FMNH 262886, hilly evergreen forest, near 12°16'24.6"N m elev., 19-22 Nov.2003; FMNH 262845-46, hilly evergreen forest 107°03'53.1"E, 350 m elev., 10 Dec.2003; FMNH 262734-37, mixed with bamboo, O Greng Mak Stream, 14°12'00.5"N 262803, hilly evergreen forest, O Ngeugn Stream, near 12°18’58"N 107°18'29.8"E, 200 m elev., 20 Nov.2003; FMNH 262847-49, hilly 107°05'59"E, 550 m elev., 31 Oct.-5 Nov.2003; FMNH 262802, evergreen forest mixed with bamboo, small tributary of O Lopeung 262804, 262889-91, 262900-01, hilly evergreen forest, O Doeung Stream, 14°11'16.3"N 107°17'36.1"E, 160 m elev., 21 Nov.2003. Por Stream, near 12°18'08"N 107°03'08"E, 500 m elev., 2-6 Nov.2003. Remarks. – These match the specimens from central Vietnam assigned by Inger et al. (1999) to this species by having males Remarks. – These agree with the original description (Bain and females not differing in SVL (males 48.1 – 60.0, mean et al., 2003) by having males with SVL 39.9 – 46.7 (mean ± ± SD 53.8 ± 3.2, N = 11; females 51.4 – 60.3, mean ± SD SD 42.3 ± 2.0, N = 8); males with gular pouches; females 55.9 ± 2.8, N = 13); the dark lateral band broken up by lighter with uniformly pale yellow eggs; and no dorsolateral folds. colouration; often dark pigmentation, usually in the form of Females are much larger than males, with SVL 74.0 – 85.5 mottling, on the venter; males without gular pouches but with (mean ± SD 81.3 ± 3.9, N = 8). wrinkled skin at the corners of the throat; males with gray nuptial pads on the dorsal and medial surfaces of the first Specimens were collected at night on banks, rock faces, finger; and a conspicuous, pigmented gland on the proximal boulders, and tree roots along and above swift rocky streams, one-third of the upper arm, larger in males than females. often near cascades. The ovaries of FMNH 262735 (SVL 79.3) contained 770 eggs of approximately 1.5 mm diameter. Specimens were collected in shallow water and on sand, The disparity in number and size of eggs between this species gravel and rocky banks of streams, usually in sections of and the very similar, co-occurring R. banaorum suggests they streams with slow-moving water. may have different reproductive strategies. This is the first report of the species from Cambodia. This is the first report of the species in Cambodia.

Rana taipehensis van Denburgh Rana nigrovittata (Blyth) Material examined. – Keo Seima: FMNH 262784, road through Material examined. – Keo Seima: FMNH 262865-69, bamboo grassland and disturbed evergreen forest, 12°08'22.9"N mixed with evergreen forest, near 12°16'45.7"N 106°56'36.4"E, 180 106°55'27.4"E, 165 m elev., 9 Dec.2003. m elev., 8 Nov.2003.

141 Stuart et al.: Amphibians and reptiles from Cambodia

Pichrada: FMNH 262064-65, deciduous dipterocarp forest with Rokhlong Stream, near 12°18'35.3"N 107°04'28.0"E, 500 m elev., grassy understorey, near 12°31'38"N 107°33'19"E, 600 m elev., 19 12 Dec.2003. Jun.2000. Pichrada: FMNH 262637-41, 262643-50, Phnom Nam Lyr Remarks. – A male (SVL 31.4, TMP:EYE 1.13) and two Mountain, evergreen mixed with deciduous and bamboo forest, near females (SVL 40.5-44.6; TMP:EYE 0.89-1.00) agree with 12°32'16"N 107°32'00"E, 600-700 m elev., 16-21 Jun.2000; FMNH van Denburgh’s (1909) original description. The specimens 262642, gallery evergreen forest along large stream, near 12°31'18"N 107 33'13"E, 600 m elev., 19 Jun.2000. have the interorbital distance greater than the width of upper ° eyelid; a distinct, broad dorsolateral fold; male with small Remarks. – Fifteen males (SVL 54.5 – 63.6, mean ± SD 59.5 white asperities on dorsum; finger I equal in length to finger ± 3.2, N = 15) and three females (SVL 70.5 – 79.4, mean ± II; male with whitish nuptial pad on medial and dorsal surface SD 74.6 ± 4.5, N = 3) agree with Smith’s (1924) original of finger I from its base to level of the distal edge of the description and Inger et al.’s (1999) expanded description of subarticular tubercle; toe IV webbed to distal subarticular the species. Males have a snout gently sloping in profile to tubercle; and an oval inner and round outer metatarsal a sharp point projecting beyond the lower jaw, and females tubercle. have a rounded snout in profile that is not sloping or projecting (Inger et al. 1999). The finger discs are rounded and those of In life, the dorsum was green and the dorsolateral folds the outer fingers larger than the tympanum, webbing extends yellow. to the distal edge of the subarticular tubercle of the first finger and to the discs of the outer three fingers, the toes are webbed The male and one female were collected at night (2040 hrs.) to the base of discs, and a low, oval inner but no outer in vegetation at the edge of a pond formed by water filling a metatarsal tubercle is present. Most of the specimens have bomb crater, and one female was collected at night (1813 two or three small papillae on each side just above the anus, hrs.) crossing a road. and four to six long white tubercles, rarely fused at the base, below the anus. Ohler et al. (2002) reported the species from the mountains of southwestern Cambodia. A juvenile (FMNH 262783, SVL 14.8) is grayish-white with large black spots on the dorsum with dark bars on the limbs, and fully agrees with the description of R. notater, a species Rhacophoridae described by Smith from a single metamorph in the same publication as R. annamensis. Orlov et al. (2002) also noted Chirixalus nongkhorensis (Cochran) the similarity of juvenile R. annamensis with R. notater. We treat Rhacophorus notater as a junior synonym of R. Material examined. – Pichrada: FMNH 263094-97, Bou Sraa Village, 12°32'05"N 107°25'55"E, 700 m elev., 14 Jun.2000; FMNH annamensis. 263098-101, deciduous dipterocarp forest with grassy understorey, near 12°31'38"N 107°33'19"E, 600 m elev., 19 Jun.2000. In O’Rang, adults were taken at night on tree branches 2.5- 4 m above the ground, and the juvenile was taken during the Remarks. – An adult female (SVL 31.7) and seven adult day (1550 hrs.) on a forest trail, all near large waterfalls. In males (SVL 21.2 – 25.5, mean ± SD 23.9 ± 1.5, N = 7) agree Pichrada, specimens were taken at night on vegetation, tree with Cochran’s (1927) original description. These have the branches, roots and rocks over small streams and seeps. There two outer fingers appearing to be opposable to the two inner in June, adults were calling, a foam nest was hanging over a ones; webbing at the base of the two outer fingers; interorbital small seep from tree leaves on the same branch where FMNH distance much greater than width of upper eyelid; dorsum brownish with irregular darker markings; and upper surface of hindlimb with dark spots but no complete crossbars.

Specimens were collected on vegetation at the edge of two ponds, one natural and the other formed by water filling a bomb crater.

This is the first report of the species from Cambodia.

Rhacophorus annamensis Smith (Fig. 9)

Material examined. - O’Rang: FMNH 262767-68, 262783, hilly evergreen forest, O Ngeugn Stream, near 12°18'58"N 107°05'59"E, 550 m elev., 31 Oct.2003; FMNH 262765, hilly evergreen forest, Fig. 9. Male Rhacophorus annamensis from Phnom Nam Lyr waterfall on O Ngeugn Stream, 12°18'50.5"N 107°06'’03.4"E, 500 Wildlife Sanctuary, Pichrada District, Mondolkiri Province, m elev., 11 Dec.2003; FMNH 262766, hilly evergreen forest, O Cambodia.

142 THE RAFFLES BULLETIN OF ZOOLOGY 2006

262645 was perched, and one pair (FMNH 262643-44) was dorsally, curving backward ventrally and forming a distinct taken in amplexus on a small rock mid-stream. angle mesially except posteriorly; four transverse folds interrupted by vent; tentacle near lip, closer to eye than nostril; This is the first report of the species from Cambodia. and small white asperities on the tail.

In life, the specimen had purplish-blue colouration; a broad, Theloderma asperum (Boulenger) golden yellow lateral stripe, one branch beginning at the middle of the lower jaw and one branch beginning under the Material examined. – Keo Seima: FMNH 262787, evergreen mixed eye, terminating at the tail tip; and a cream spot on tentacle with deciduous forest, UTM 0725385 1351814, coll. J. Walston, 18 and vent. Aug.2002; FMNH 261896, disturbed evergreen mixed with deciduous forest, 250 m elev., coll. J. Walston, May.2000. The specimen was collected during heavy rain at night (1835 hrs.) crawling on leaf litter and bare soil 2 m from a 1 m Remarks. – These two specimens (FMNH 262787 SVL 23.0, wide swift, rocky stream. FMNH 261896 SVL 22.3.) agree with Boulenger’s (1886) description by having scattered white asperities on the dorsal This is the first report of the Order Gymnophiona from surface; fingers without webbing; interorbital distance greater Cambodia. than width of upper eyelid; a grayish marking on top of the head, rear of lower back, and flanks; and a dark ventral surface with light reticulations. Bataguridae

Both were collected in artificial holes holding rainwater that Cyclemys atripons Iverson and McCord had been cut by resin-tappers in large dipterocarp trees. (Fig. 11)

Ohler et al. (2002) reported the species from the mountains Material examined. – O’Rang: FMNH 262710-11, hilly evergreen of southwestern Cambodia. forest, O Doeng Por Stream, near 12°18'08"N 107°03'08"E, 500 m elev., 6 Nov.2003.

Theloderma stellatum Taylor Pichrada: FMNH 259050, gallery evergreen forest, near 12°32'16"N 107°32'00"E, 600-700 m elev., 21 Jun.2000. Material examined. – Keo Seima: FMNH 262786, hilly evergreen forest mixed with deciduous & bamboo, near 12°10'34.9"N Remarks. – A male (CPL 181, CPW 142) and two females 106°57'47.6"E, 190 m elev., 9 Nov.2003. (CPL 200-214, CPW 161-162) have a mostly yellow plastron; densely pigmented bridge; nearly immaculate chin; and dark Remarks. – This single female (SVL 33.8) fully agrees with spotting on top of the head. FMNH 259050 has 8 dark and 7 Taylor’s (1962) description of the species from Chanthaburi, light, FMNH 262710 has 11 dark and 12 light, and FMNH Thailand. The specimen has the dorsal surface covered in 262711 has 10 dark and 11 light neck stripes. These match whitish asperities; fingers about 1/3 webbed, the third finger the descriptions of both C. atripons and C. pulchristriata, disc about equal to the diameter of tympanum; the interorbital two species that were described almost concurrently and that distance about equal to width of upper eyelid; and a dark have been considered the same taxon (Iverson in Guicking ventral surface with light reticulations. et al., 2002). These specimens from Cambodia are assigned to C. atripons following Stuart and Platt (2004); further The specimen was taken at night (1820 hrs.) on the trunk of a 1 m diameter at breast height tree, 20 cm above a water- filled cavity near the base of the tree.

This is the first report of the species in Cambodia.

Ichthyophiidae

Ichthyophis kohtaoensis Taylor (Fig. 10)

Material examined. – Ta Veng: FMNH 262775, hilly evergreen forest mixed with bamboo, small tributary of O Lopeung Stream, 14°11'16.3"N 107°17'36.1"E, 160 m elev., 21 Nov.2003.

Remarks. – The single specimen has body length 27 times Fig. 10. Adult Ichthyophis kohtaoensis (FMNH 262775) from the body width (TTL 320; BDW 11.8); pharyngeal region Virachey National Park, Ta Veng District, Ratanakiri Province, wider than head; 335 transverse body folds curving forward Cambodia.

143 Stuart et al.: Amphibians and reptiles from Cambodia taxonomic work is needed to determine the correct application brown with irregular black blotches on the dorsum (FMNH of these two names. The distribution record of FMNH 259050 263004, 263008). Only the second, blotched morph was was included in Stuart and Platt (2004). described in the type series of two adult males and two juveniles. Smith (1935) noted the same two colour morphs FMNH 259050 was found at night (2020 hrs.) at the base of in the similar D. siamensis (Boulenger) and stated that large boulder on a sandy bank 1.5 m from a swift, shallow, intergradation between them is rare. sandy-substrate stream. FMNH 262710-11, male and female, respectively, were found together at night (1830 hrs.) under Four adult males are present in the Cambodian series, and of 25 cm of water on the bottom of a 5 x 15 m stream pool with these one (FMNH 263007, SVL 42.6, uniform morph) has slow current and solid rock substrate, approximately 50 cm six preanal pores, two (FMNH 263004, SVL 46.0, and FMNH apart. 261844, SVL 46.5, blotched morphs) have eight preanal pores, and one (FMNH 261845, SVL 46.1, blotched morph) has nine preanal pores. Gekkonidae The specimens were active at night on a rocky outcrop and Dixonius vietnamensis Das riverbank near the edge of forest, and within 75 cm of the ground on large tree trunks (40-150 cm diameter at breast Material examined. – Pichrada: FMNH 261844-45, rocky outcrop height) in the forest. One (FMNH 263008) was found during at base of Phnom Nam Lyr Mountain, near 12°32'16"N 107°32'10"E, the day retreating under a board next to a wooden building. 700 m elev., 18 Jun.2000. This is the first report of the species since its original Keo Seima: FMNH 263008, Keo Seima camp, 12 08'16.0"N ° description from Tinh Khanh Hoa Province, Vietnam. 106°54'54.8"E, 160 m elev., 10 Nov.2003; FMNH 263003-07, hilly evergreen forest mixed with deciduous and bamboo, near 12°10'34.9"N 106°57'47.6"E, 190 m elev., 9 Nov.2003. Agamidae Siem Pang: FMNH 263002, evergreen forest mixed with bamboo, waterfall of O Chhay Stream, tributary of O Chay Stream, capra Günther 14°15'49.6"N 106°31'46.0"E, 200 m elev., 5 Dec.2003. (Figs. 12-13)

Remarks. – These specimens agree with Das’ (2004) original Material examined. – O’Rang: FMNH 262956, hilly evergreen description by having the adult head width (measured at angle forest near O Ngeugn Stream, near 12°18'58"N 107°05'59"E, 550 of jaws) greater than head length (measured from posterior m elev., 31 Oct.2003; FMNH 262957, hilly evergreen forest near O edge of posterior supralabial to snout tip); two supranasals Doeung Por Stream, near 12°18'08"N 107°03'08"E, 500 m elev., 6 in narrow contact; dark canthal stripe terminating at back of Nov.2003. head; ear opening less than one half eye diameter; and adult SVL 42.1-46.3 (mean ± SD 45.2 ± 1.7, N = 5). The Remarks. – These agree with Günther’s (1861) original Cambodian specimens differ from the original description description by having a long moveable horn at the posterior by having dark blotches on the lips in adults, and having two terminus of the supraciliary edge; no spine above the distinct colour morphs represented by both sexes: uniform tympanum or on the side of the neck; a diastema between the grayish-brown (FMNH 263002-03, 263005-07) and grayish- nuchal crest and the dorsal crest; and upper parts covered in small scales that are mostly uniform in size. In life, the male (FMNH 262957; SVL 110.5) had a green body with black spotting and a few larger yellow spots encircled with black;

Fig. 11. Male and female Cyclemys atripons (FMNH 262710-11) Fig. 12. Male (FMNH 262957) from the Seima from the Seima Biodiversity Conservation Area, O’Rang District, Biodiversity Conservation Area, O’Rang District, Mondolkiri Mondolkiri Province, Cambodia. Province, Cambodia.

144 THE RAFFLES BULLETIN OF ZOOLOGY 2006 yellow postorbital horn and nuchal crest; greenish-yellow mean ± SD 68.5 ± 1.6, N = 3). Acanthosaura coronata is head with a wide black band extending from around the eye morphologically distinct from A. lepidogaster (Cuvier) and to the nuchal crest, and continuing as a narrow band from the should be removed from its synonymy, where it was placed posterior end of the nuchal crest to the shoulder; black throat by Smith (1935). with a yellow and orange gular pouch; and reddish-brown iris. The gravid female (FMNH 262956; SVL 137.9) had a The colouration of FMNH 262964 follows. In life, upper parts dark olive-brown head and body; and a yellow supraciliary of body light green with gray-brown mottling; crown, nape, edge, postorbital horn, nuchal crest, upper lip and gular pouch. tympanum, and upper lip light green; distinct, light green (“yellowish-olive” according to Günther, 1861) bar edged Both were collected sleeping at night 1.5-2 m above the with black on crown between supraciliary edges; distinct ground on 2-3 cm diameter vertical saplings, 40-100 m from white and light green (“yellowish” according to Günther, swift streams. 1861) bar edged with black from eye to upper lip; five narrow, black, chevron-shaped markings on vertebral line, first at the This is the first report of the species in Cambodia since its level of insertion of forelimbs with body and fifth slightly original description. posterior to level of insertion of hindlimbs with body; limbs brown with dark brown bands; white spot on elbow and white stripe on outer surface of tibia; white band edged with black Acanthosaura coronata Günther on dorsal surface of hand and foot; tail with brown and (Fig. 14) pinkish-orange bands; orange-red bar on chest extending between forelimbs; throat and venter grayish-white with black Material examined. – O’Rang: FMNH 262964, hilly evergreen flecking; ventral surface of foot and fourth toe with brown forest near O Doeung Por Stream, 12°18'02.6”N 107°03'18.1"E, 500 bands; and iris reddish-brown. In preservative, the light green m elev., 2 Nov.2003; FMNH 262965, hilly evergreen forest with on the upper parts and the orange-red bar on the chest is lost. bamboo, 12°15'41.6”N 107°03'48.2"E, 480 m elev., 7 Nov.2003. Günther (1864) stated that “the ground-colour of the male is grey, of the female brownish red; irregular dark-brown bands Pichrada: FMNH 263034, Phnom Nam Lyr Mountain, hilly evergreen forest mixed with deciduous and bamboo, near across the back,” but the three males reported here vary 12°32'16"N 107°32'00"E, 600-700 m elev., 20 Jun.2000. considerably in preservative in ground-colour and the extent of dorsal dark markings. FMNH 263034 has reddish-brown Remarks. – Three males agree with the original description ground-colour but the others are grey-brown, and FMNH (Günther, 1861) and expanded descriptions (Günther, 1864; 262965 has heavy black chevron-shaped markings on the Boulenger, 1885) of A. coronata by having a prominently vertebral line but these are only slightly visible or absent in projecting, serrated supraciliary edge that obscures the eyelid the others. and orbit in dorsal view; a single short spine (sometimes broken) posterior to the supraciliary edge and separated from The species appears to live on or near to the ground. FMNH it by a notch; a single (sometimes two adjacent) short spine 262694 was observed during the day (1625 hrs.) on the ground on the back of head about midway between the tympanum and nuchal crest; a continuous nuchal and dorsal crest; a light- coloured band edged with black across the crown and from the eye to upper lip; and small body size (SVL 67.5 – 70.3,

Fig. 13. Female Acanthosaura capra (FMNH 262956) from the Fig. 14. Male Acanthosaura coronata (FMNH 262965) from the Seima Biodiversity Conservation Area, O’Rang District, Mondolkiri Seima Biodiversity Conservation Area, O’Rang District, Mondolkiri Province, Cambodia. Province, Cambodia.

145 Stuart et al.: Amphibians and reptiles from Cambodia in the forest and ran up a 15 cm diameter at breast height tree This is the first report of the species from Cambodia. when pursued. FMNH 263034 was taken during the day (1300 hrs.) on a rock near a small stream. FMNH 262965 was sleeping at night (1915 hrs.) on the twig of a small sapling 1 Calotes mystaceus Duméril & Bibron m above the ground. Material examined. – Pichrada: FMNH 262685, deciduous This is the first report of the species since its original dipterocarp forest with grassy understory, near 12°31'38"N description from Cambodia. 107°33'19"E, 600 m elev., 19 Jun.2000; FMNH 262686, Phnom Nam Lyr Mountain, hilly evergreen forest mixed with deciduous and bamboo, near 12°32'16"N 107°32'00"E, 600-700 m elev., 20 Jun.2000. Bronchocela smaragdina Günther Keo Seima: FMNH 262968, Keo Seima camp, 12°08'16.0"N Material examined. - Pichrada: FMNH 262309, deciduous 106°54'54.8"E, 160 m elev., 8 Nov.2003; FMNH 262969-70, hilly dipterocarp forest with grassy understorey, near 12°32'12"N evergreen forest, 12°11'29.3"N 106°59'52.3"E, 315 m elev., 10 107°32'53"E, 600 m elev., 19 Jun.2000. Nov.2003.

Remarks. – This adult female (SVL 110.5, TAL 365.0) fully O’Rang: FMNH 262967, hilly evergreen forest, near 12°16'24.6"N agrees with Günther’s (1864) original description, except the 107°03'53.1"E, 350 m elev., 11 Dec.2003. ventrolateral stripe was white in life rather than yellow. The specimen has the ventral scales in 12 rows, twice the size of Ta Veng: FMNH 262966, disturbed evergreen forest mixed with the lateral scales; small, uniform scales between the orbit and bamboo, near soldier outpost, 14°09'07.7"N 107°15'33.3"E, 130 m tympanum; the fourth toe about 1/4 longer than the third toe; elev., 23 Nov.2003. a weak nuchal crest; and no dorsal crest, gular pouch, or fold before the shoulder. Remarks. – These have one or two adjacent spines above the tympanum; no spine at the posterior end of the supraciliary In life, uniformly bright green, with head and venter edge; a spine on the occiput about midway between the yellowish-green; white ventrolateral stripe between insertion tympanum and nuchal crest; and a deep oblique skin fold in points of forelimb and hindlimb and on base of tail; brown front of the shoulder containing small, granular, darkly stripe on posterior surface of thigh; dorsal surface of proximal pigmented scales. 1/5 of tail green, lateral surface and distal 4/5 of tail brown with darker flecking; and tympanum brown. The specimens were collected during the day on a dirt road and around wooden buildings, or sleeping on trees at night The specimen was collected at dusk (1800 hrs.) on a tree 0.75-2 m above the ground in open forest and forest clearings. branch 3.5 m above the ground in the forest, away from any body of water. The species is probably arboreal and This is the first report of the species from Cambodia. infrequently observed; the three local hunters present at the time of capture stated that they had never seen this before. Draco indochinensis Smith Smith (1935) reported that only three specimens were known: Material examined. – O’Rang: FMNH 262713, hilly evergreen the two female types obtained by Mouhot in Cambodia, and forest, 12°20'05.8"N 107°02'26.2"E, 500 m elev., 12 Dec.2003; a third specimen obtained by Smith at Dalat on the Langbian FMNH 262714, hilly evergreen forest with bamboo, 12°15'41.6"N Plateau in southern Vietnam. 107°03'48.2"E, 480 m elev., 7 Nov.2003.

This is the first report of the species from Cambodia since Remarks. – A juvenile (FMNH 262713; SVL 70.8) and adult the original description. (FMNH 262714; SVL 107.8) fully agree with the original and expanded descriptions (Smith, 1928, 1935) of D. indochinensis, except the juvenile has 10 and the adult 11 Calotes emma Gray supralabials on each side (9 given by Smith for two females). The nostril is directed upwards; the tympanum is naked; the Material examined. - Ta Veng: FMNH 262954-55, hilly evergreen patagium is supported by five ribs; some of the four black forest mixed with bamboo, 14 11'52.8"N 107 18'18.0"E, 200 m ° ° transverse bands on the dorsal surface of the patagium elev., 20 Nov.2003. bifurcate as they approach the body; and a thick black Remarks. – These have a spine at the posterior end of the transverse band extends across the posterior gular region from supraciliary edge; a spine above the tympanum; a spine on one throat lappet to the other. Smith (1935) and McGuire the occiput about midway between the tympanum and nuchal and Heang (2001) distinguished both sexes of D. crest; and an oblique skin fold in front of the shoulder indochinensis from the similar D. blanfordii Boulenger by containing small, granular, darkly pigmented scales. the latter character.

Both specimens were collected at night sleeping on vegetation The juvenile was collected in the morning (0844 hrs.) on the in the forest 1 m or less above the ground. trunk of a tree 1.5 m above the ground. The adult was sleeping

146 THE RAFFLES BULLETIN OF ZOOLOGY 2006 at night (1835 hrs.) on the trunk of an 18 cm diameter at Ta Veng: FMNH 262989, soldier outpost, 14°09'07.7"N breast height tree 2.5 m above the ground. 107°15'33.3"E, 130 m elev., 19 Nov.2003.

This is the first report of the species in Cambodia since the Remarks. – Five specimens have tail length 2.50-4.30 holotype was obtained at Bokor, in the Elephant Mountains (median 4.08, N = 3) times the SVL; a single femoral pore; of southwestern Cambodia. four strongly keeled dorsal plates across the middle of the back, the keels forming continuous lines; keeled head shields; and no ocellate spots on flanks. Physignathus cocincinus Cuvier FMNH 262989-90 were collected in grassy areas close to Material examined. – Keo Seima: FMNH 262958-59, UTM wooden buildings, and FMNH 261856 was collected at night 0721041 1350686; Chen Chea, 12 Sept.2003. (1945 hrs.) sleeping on the leaf of a bush 15 cm above the ground. O’Rang: FMNH 262962, hilly evergreen forest along O Kamen Stream, near 12°19'35"N 107°05'33"E, 500 m elev., 1 Nov.2003; FMNH 262963, hilly evergreen forest along O Doeung Por Stream, This is the first report of the species from Cambodia. near 12°18'08"N 107°03'08"E, 500 m elev., 6 Nov.2003.

Pichrada: FMNH 262674-75, Phnom Nam Lyr Mountain, evergreen Scincidae mixed with deciduous and bamboo forest, near 12°32'16"N 107°32'00"E, 600-700 m elev., 18 Jun.2000. Lipinia vittigera (Boulenger) Siem Pang: FMNH 262961, hilly evergreen forest mixed with Material examined. – Ta Veng: FMNH 262975, 262984, disturbed bamboo, near 14°18'28.9"N 106°33'32.1"E, 430 m elev., 4 Dec.2003. evergreen forest mixed with bamboo near soldier outpost, 14 09'07.7"N 107 15'33.3"E, 130 m elev., 23 Nov.2003; FMNH Ta Veng: FMNH 262960, hilly evergreen forest mixed with bamboo ° ° 262986, hilly evergreen forest mixed with bamboo, O Lopeung along O Greng Mak Stream, 14°12'00.5"N 107°18'29.8"E, 200 m Stream, 14 11'16.3"N 107 17'36.1"E, 160 m elev., 19 Nov.2003. elev., 20 Nov.2003. ° °

Remarks. – Two adult males (SVL 176-201), four subadults, Remarks. – These have an acutely pointed snout nearly twice and two juveniles have strongly compressed tails, heavily the diameter of the eye; three distinct light-coloured keeled below; mostly green colouration, with two subadults (yellowish in life) longitudinal stripes across the back and one adult having four oblique, light-coloured body bands consisting of a vertebral stripe from the snout tip to tail and between the axilla and groin visible in preservative; well- a dorsolateral stripe from above the eye to tail, each flanked developed nuchal, dorsal, and caudal crests in the adults, by a black stripe; and a weak, light-coloured lateral stripe. In weakly visible in the young; and femoral pores, 6/5 (left/right) life the tail was bright orange. One is a juvenile (FMNH and 7/6 in adult males, 8/8 and 8/9 in two subadults. 262975, SVL 25.7) and two are larger and probably adults (FMNH 262984, SVL 37.5 and FMNH 262986, SVL 33.2). Four specimens (FMNH 262675, 262960, 262962-63) were collected at night sleeping on vegetation 1-4 m above the FMNH 262986 was taken during the day (1600 hrs.) on the ground, close to or overhanging forested streams. FMNH trunk of a 15 cm diameter at breast height tree 70 cm above 262674 was collected at night sleeping on a sapling in the ground, 8 m from a large stream. forest, 50 m from the nearest stream. FMNH 262961 drowned in a stream gill net set for fish. FMNH 262958-59 perished This is the first report of the species from Cambodia. from injuries after being confiscated by authorities from local hunters encountered in the forest at night. Lygosoma quadrupes (Linnaeus) Tirant (1885) reported the species from Cambodia, without a specific locality. Material examined. - Pichrada: FMNH 261866, on dirt road through deciduous forest with grassy understorey and bamboo, near 12°29'49"N 107°29'33"E, 700 m elev., 22 Jun.2000. Lacertidae Remarks. – A single specimen has an elongate, slender, Takydromus sexlineatus Daudin diminutive habitus; four very short limbs, the forelimb (3.8) 7.2% the SVL (52.4); five fingers, the three median fingers Material examined. – Keo Seima: FMNH 262990, 12°08'16.0"N about equal in length; five toes, the three median toes about 106°54'54.8"E, 160 m elev., Chen Chea, 27 Nov.2003. equal in length; the snout to forelimb distance (11.5) 31.9% the axilla to groin distance (36.0); a minute ear opening, partly Pichrada: FMNH 261856, Bou Sraa Village, 12°32'05"N 107°25'55"E, 700 m elev., 14 Jun.2000; FMNH 261857-58, Phnom covered with scales; a single frontoparietal; and brown Nam Lyr Mountain, evergreen mixed with deciduous and bamboo colouration with darker brown longitudinal lines on dorsum forest, near 12°32'16"N 107°32'00"E, 600-700 m elev., 20 Jun.2000. and flanks, continuing onto tail.

147 Stuart et al.: Amphibians and reptiles from Cambodia

The specimen was taken during the day (1200 hrs.) on a dirt Tropidophorus microlepis Günther road that formed part of the former Ho Chi Minh Trail. Material examined. – O’Rang: FMNH 262972, hilly evergreen This is the first report of the species from Cambodia. forest, O Doeung Por Stream, near 12°18'08.4"N 107°03'08.1"E, 500 m elev., 2 Nov.2003; FMNH 262973, hilly evergreen forest, O Ngeugn Stream, near 12°18'02.9"N 107°05'22.7"E, 440 m elev., 5 Nov.2003. maculatus (Blyth) Keo Seima: FMNH 262974, bamboo forest mixed with evergreen, Material examined. – O’Rang: FMNH 262992, hilly evergreen near 12°16'45.7"N 106°56'36.4"E, 180 m elev., 8 Nov.2003. forest, O Ngeugn Stream, near 12°18'58"N 107°05'59"E, 550 m elev., 31 Oct.2003; FMNH 262993, hilly evergreen forest, O Remarks. – These three specimens (SVL 57.0 – 69.0, mean Rokhlong Stream, 12°17'30"N 107°03'06"E, 500 m elev., 4 Nov.2003. ± SD 64.5 ± 6.5, N = 3) agree with Günther’s (1861) original description by having a single pair of frontonasals; keels on Siem Pang: FMNH 262982, bamboo mixed with evergreen forest dorsal tail scales terminating in elevated spines; keels on along tributary of O Chay Stream, 14°18'28.6"N 106°33'02.6"E, 310 dorsal body scales not terminating in elevated spines; smooth m elev., 5 Dec.2003; FMNH 262991, bamboo & grasses along O throat scales; median subcaudals wider than ventrals; and Kanome Stream, 14°15'08.0"N 106°37'58.8"E, 175 m elev., 11 three large preanals, flanked by two smaller scales. The keels Nov.2003. on the lateral body scales are arranged obliquely. FMNH 262972-73 have weak black spots on the chin scales and Remarks. – These have a concave rostral scale; all scales unmarked throats, but FMNH 262974 has distinct black spots smooth; scales on dorsum larger than those on side; bronze on the chin scales and black spots arranged into five above, with two median series of black spots on back; black longitudinal stripes on the throat. lateral band from tip of snout to tail, becoming lighter with jagged margin on tail; flank white with black speckling; and All were found in and along streams. Specifically, FMNH lower parts creamy-white. 262972 was taken at night (1840 hrs.) under a rock on a sandy bank 20 cm from a stream, FMNH 262973 was collected The specimens were collected on the banks of rocky streams. during the day (1515 hrs.) in a crevice of a wet rock face next to a 3 m high cascade of a 20 m wide stream, and FMNH This is the first report of the species from Cambodia. 262974 was found at night (1927 hrs.) clinging to a submerged branch under 15 cm of water next to the bank of a 3.5 m wide rocky stream. Sphenomorphus rufocaudatus Darevsky & Nguyen This is the first report of the species from Cambodia. Günther Material examined. – Keo Seima: FMNH 262999-3000, hill (1861) described this species in the same paper that he evergreen mixed with deciduous forest, near 12 18'02"N ° described other species collected by Mouhot in Cambodia. 107°03'18"E, 500 m elev., 2 Nov.2003. However, Smith (1935) placed the type locality as Khao O’Rang: FMNH 262997, hilly evergreen forest, 12°15'35.7"N Sebab in Chanthaburi Province, Thailand, which is close to 107°03'56.8"E, 542 m elev., 30 Oct.2003; FMNH 262998, evergreen the southwestern border with Cambodia. mixed with bamboo and deciduous forest, 12°16'50.2"N 107°03'31.0"E, 472 m elev., 1 Nov.2003; FMNH 263001, disturbed evergreen forest, 12°17'44.8"N 107°03'16.5"E, 450 m elev., Xenopeltidae 3 Nov.2003. Xenopeltis unicolor Reinwart in Boie Remarks. – These fully agree with the original description (Darevsky & Nguyen, 1983) and topotypes from Gia Lai Material examined. – Siem Pang: FMNH 263016, evergreen mixed Province, Vietnam (FMNH 252341-62). The specimens have with bamboo forest, near 14°18'28.9"N 106°33'32.1"E, 430 m elev., smooth scales; the dorsal and lateral scales about equal in 4 Dec.2003. size; 32-34 scale rows around body; four supraorbitals; two enlarged preanals; short limbs with five fingers; bronze Remarks. – A single female has smooth, highly iridescent colouration above, with weak, longitudinal, broken stripes scales; brown dorsal scales, the first row creamy-white edged formed by small black spots, those of the median stripe the in brown, the second to fourth rows brown edged in creamy- largest; and an irregular, broad, black dorsolateral stripe from white; creamy-white ventral scales, brown at outer edges nostril to tail. posteriorly; 15 longitudinal scale rows at mid-body; and a wedge-shaped head, indistinct from neck. Specimens were collected during the day on or under leaf litter, or under logs. The specimen was taken at night (1810 hrs.) on the leaf litter bank 1.5 m from a 3 m wide stream with slow current. This is the first report of the species from Cambodia. Saint Girons (1972a) reported the species from several localities in the lowlands of central Cambodia.

148 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Colubridae Remarks. – A single female closely agrees with Günther’s (1875) original description. The specimen has 17 scale rows Ahaetulla prasina (Reinwardt in Boie) at mid-body; the median four dorsal scale rows feebly keeled; 202 ventral scale rows; 80 subcaudal scale rows; an undivided Material examined. – Pichrada: FMNH 259198, evergreen mixed anal scale; loreal longer than deep; one preocular and two with deciduous and bamboo forest, Phnom Nam Lyr Mountain, near postoculars; eight supralabials, the third, fourth and fifth 12°32'16"N 107°32'00"E, 600-700 m elev., 20 Jun.2000. entering the orbit; black trunk with 26 white bands on the body, approximately 1-1.5 dorsal scale rows wide along the O’Rang: FMNH 263012, near 12°19'18"N 107°05'53"E, 7 Nov.2003. vertebral row at midbody, the first at about ventral scale row 23; 15 white bands on the tail; dark bands on posterior half Remarks. – These have an elongated snout, length about two of venter, subcaudals marbled with black. times eye diameter; rostral scale upturned, slightly projecting, but not as a nasal appendage; black markings on dorsal scale The specimen was taken at night (2005 hrs.) crawling on a 1 edges forming oblique lines anteriorly; and a uniform dark m diameter fallen tree, 2.2 m above a 3 m wide stream. gray venter. This is the first report of the species from Cambodia. The Pichrada specimen was in gray colour phase and found asleep at night (1855 hrs.) 1.5 m above the ground on the top of a large-leaved bush. The O’Rang specimen was in gold Homalopsis nigroventralis Deuve, new combination colour phase and killed by a local during the day (1500 hrs.) (Fig. 15) in herbaceous vegetation on the side of a road. Material examined. – Ta Veng: FMNH 263029, bamboo mixed Saint Girons (1972a) reported the species from several with deciduous forest, O Lopeung Stream, 14°10'39.2"N localities throughout Cambodia, including Ratanakiri. 107°17'25.1"E, 150 m elev., 19 Nov.2003.

Siem Pang: FMNH 263030, bamboo mixed with evergreen forest, cyanea (Duméril, Bibron & Duméril) O Kanome Stream, 14°13'33.7"N 106°36'16.0"E, 170 m elev., 30 Nov.2003; FMNH 263031-32, bamboo mixed with evergreen Material examined. – Keo Seima: FMNH 259177, on road through forest, O Kanome Stream, 14°15'08.0"N 106°37'58.8"E, 175 m elev., disturbed evergreen mixed with deciduous forest, 150 m elev., coll. 1 Dec.2003; FMNH 263033, bamboo mixed with evergreen forest, J. Walston and P. Davidson, May.2000. 14°12'53.8"N 106°35'51.3"E, 100 m elev., 29 Nov.2003.

O’Rang: FMNH 263014, hilly evergreen forest along O Rokhlong Remarks. – A juvenile male, subadult male, two adult males, Stream, near 12°18'35.3"N 107°04'28.0"E, 500 m elev., 12 and an adult female agree with Deuve’s (1970) description Dec.2003. of nigroventralis, as a subspecies of H. buccata (Linnaeus) from Laos, by having 11-13 supralabials; 15-16 infralabials; Remarks. – A male (FMNH 263014) has 21 scale rows at 35-38 longitudinal scale rows at midbody; 157-165 ventrals mid-body; enlarged vertebral scales; 257 ventral scale rows; (mean ± SD 160.8 ± 3.2, N = 5); and a dark venter with light and 124 subcaudal scale rows. FMNH 259177 is an spots. incomplete specimen consisting of only the head and the anterior two head-lengths of the body. Both specimens have eight supralabials; 10 infralabials; one preocular; and two In life, the juvenile male (FMNH 263033) had light orange postoculars. dorsal body bands; a broad tan band on the dorsal surface of head; a broken creamy-white ventrolateral stripe connecting In life, FMNH 263014 had a green dorsum and venter; white the dorsal body bands; a distinctive white X-like marking on chin with baby-blue infralabials; and bluish-white marbling the chin; and a black venter with scattered white spots. The on some anterior ventrals.

FMNH 259177 was killed on a road by a motor vehicle. FMNH 263014 was climbing at night (1925 hrs.) on the vertical trunk of a 10 cm DBH (diameter at breast height) sapling 4 m above the ground, 5 m from a 6 x 8 m stream pool at the base of a waterfall.

Saint Girons (1972a) reported the species from central and southwestern Cambodia.

Dinodon septentrionalis (Günther)

Material examined. – O’Rang: FMNH 263009, hilly evergreen Fig. 15. Male and female Homalopsis nigroventralis (FMNH forest, O Kamen Stream, near 12°19'35.3"N 107°05'33.4"E, 500 m 263031-32) from Virachey National Park, Siem Pang District, Stung elev., 1 Nov.2003. Treng Province, Cambodia.

149 Stuart et al.: Amphibians and reptiles from Cambodia three adults (FMNH 263030-32) had a brown dorsum with known. This is the first report of the species from Cambodia faint, lighter banding that disappeared posteriorly; an olive and represents a large range extension. brown head with black markings on the snout and over eyes; venter yellowish-olive (FMNH 263030, male), olive (FMNH 263031, female), or olive-brown (FMNH 263032, male), Oligodon cinereus (Günther) darkening posteriorly to dark gray; creamy-white spots on the ventrals and subcaudals, beginning on the throat as a single Material examined. – Pichrada: FMNH 259201, deciduous forest spot on each ventral forming a longitudinal line, becoming with grassy understory and bamboo, near 12°29'49"N 107°29'33"E, more scattered posteriorly but sometimes arranged in pairs; 700 m elev., 22 Jun.2000. creamy-white X-like marking on chin. Remarks. – A single juvenile agrees with Günther’s (1864) The juvenile was taken during the day (1400 hrs.) from the original description of a single specimen from Cambodia, and bottom of a 3 m wide swift stream flowing over solid rock the expanded description by Wagner (1975), by having 17 substrate, 50 cm from the base of a 30 cm high cascade. The longitudinal scale rows at mid-body, reducing to 15 specimen was under 25 cm of water, with its head and anterior posteriorly; eight supralabials, the fourth and fifth touching part of body emerging vertically from a small rock pile. The the eye; one preocular; two postoculars; 168 ventrals, with subadult was collected during the day (1255 hrs.) swimming an obtuse keel along each side; 26 divided subcaudals; and under 25 cm of water in a shallow, 8 m wide flat stream with a single anal scale. moderate current and a substrate of rocks. The adults drowned at night in gill nets set for catching fish in a 6 m wide stream In life the specimen was red above, with narrow light gray with moderate current and substrate composed of sand and crossbars bordered with black. small rocks. The stream in which the juvenile was found in was mostly covered by forest canopy, but the other streams The specimen was taken during the day (1100 hrs.) on a dirt had no canopy cover, and were lined with tall grass, bamboo, road through bamboo. and forest. Saint Girons (1972a) reported the species from Trapeang Homalopsis nigroventralis is easily distinguished from H. Chan, in central Cambodia. buccata by ventral colouration (buccata has a white venter with black spots) and habitat preference (buccata occurs in lentic and slow-moving bodies of water such as lakes, marshes Oligodon ocellatus (Morice) and large rivers), and we treat them as separate species. Deuve (Fig. 16) (1970) reported that buccata and nigroventralis do not co- occur in Laos. Material examined. – Ta Veng: FMNH 263011, hilly evergreen forest mixed with bamboo, 14°11'38.4"N 107°17'21.9"E, 200 m elev., 22 Nov.2003. This is the first report of the species from Cambodia. Remarks. – A single female agrees with Saint Girons’ (1972b) expanded description of Morice’s type specimens from Tay- Liopeltis stoliczkae (Sclater) Ninh, southern Vietnam by having 19 longitudinal scale rows at mid-body, reducing to 15 posteriorly; eight supralabials, Material examined. – Pichrada: FMNH 259193, deciduous forest the fourth and fifth touching the eye; two preoculars; and with grassy understory and bamboo, near 12°29'49"N 107°29'33"E, 700 m elev., 22 Jun.2000. two postoculars. Saint Girons (1972b) reported two Tay-Ninh females to have 161-165 ventrals and 27-29 subcaudals, but Remarks. – A single male (SVL 330; TAL 232) fully agrees the Cambodian female has 173 ventrals and 32 subcaudals. with Sclater’s (1891) original description and Smith’s (1943) expanded description of the species. The specimen has the head slightly distinct from neck; snout length twice the eye diameter; a small nostril in a long, undivided nasal; a square loreal; eight supralabials, the fourth and fifth touching the eye; 15 longitudinal scale rows at mid-body, reducing to 13 posteriorly; 155 ventrals; a brownish dorsum with a broad black stripe on the side of head that extends onto the anterior part of body before gradually disappearing; a gray stripe on the outer margins of the ventrals; and a thin gray stripe on the median of the ventrals.

The specimen was taken during the day (1200 hrs.) on bamboo 2 m above the ground at the edge of a dirt road.

Smith (1943) reported the species from northeastern India Fig. 16. Female Oligodon ocellatus (FMNH 263011) from Virachey and Myanmar, and to be rare, with only five specimens National Park, Ta Veng District, Ratanakiri Province, Cambodia.

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In life, the specimen was orange above, with dark orange This is the first report of the species from Cambodia. head markings edged in black; 11 dark orange dorsal body blotches, edged in black, the last above the vent, each separated by three narrow black crossbars; chin and anterior Rhabdophis chrysargos (Schlegel) 2/3 of ventrals yellowish-ivory; posterior 1/3 of ventrals salmon-pink; and subcaudals white. Material examined. – Siem Pang: FMNH 263020, evergreen forest, 14°16'34.8"N 106°32'10.5"E, 200 m elev., 5 Dec.2003. The specimen was collected during the day (1400 hrs.) in a cavity in the trunk of a 35 cm diameter at breast height tree Remarks. – A single juvenile has an olive-brown dorsum with 30 cm above the ground. whitish, lateral transverse bars connected across the vertebrals by black bars; a white upper lip with black triangular-shaped Saint Girons (1972a) reported the species from Kirirom, in markings pointing ventrally, white lip stripe extending southwestern Cambodia. posteriorly to form a white chevron-shaped marking outlined in black on the base of the neck; venter whitish with a black spot at the outer margin of the ventrals and subcaudals; 9 Pareas margaritophorus (Jan) supralabials, fourth, fifth and six touching the eye; 3 postoculars; and 2 anterior temporals. Material examined. – Siem Pang: FMNH 263022, bamboo forest, 14°18'52.8"N 106°35'49.9"E, 340 m elev., 3 Dec.2003. The specimen was collected during the day (1520 hrs.) on a rocky bank 1.5 m from a 3 m wide intermittent stream with Remarks. – A single juvenile has smooth scales; the head no current. distinct from neck; body not strongly compressed; vertical eye diameter slightly greater than the distance from the ventral Saint Girons (1972a) reported the species from Kirirom, in edge of eye to ventral edge of upper lip; vertebral scales not the mountains of southwestern Cambodia. enlarged; gray dorsum with lateral, black and white transverse bars comprised of black scales with a white spot on the anterior part of scale; underparts whitish with black speckling. Sibynophis triangularis Taylor & Elbel

The specimen was collected during the day (1440 h) on Material examined. – O’Rang: FMNH 263023, hilly evergreen bamboo leaf litter 10 m from an 8 m wide stream with no forest, 12°20'05.8"N 107°02'26.2"E, 500 m elev., 12 Dec.2003. current. Remarks. – The single male agrees with Taylor and Elbel’s This is the first report of the species in Cambodia. (1958) original description by having 10 supralabials, the tenth much the largest; the parietal in contact only with the upper of two postoculars, which is nearly square; a black Ptyas carinatus (Günther) triangular nuchal spot bordered by a white stripe continuous with the white supralabials; and a black spot on the outer Material examined. – O’Rang: FMNH 262712, hilly evergreen edge of the ventrals. forest, O Ngeugn Stream, Mok Romum Waterfall, near 12°18'58"N 107°05'59"E, 550 m elev., 5 Nov.2003. The specimen was killed in the morning (0900 hrs.) by a local resident on a human trail in the forest. Remarks. – A single female has large body size (SVL 189; TAL 62); 16 longitudinal scale rows at midbody, with weak This is the first report of the genus from Cambodia, and the keels on medial two rows; 207 ventrals; 111 subcaudals; and first report of the species outside of the type locality in Chon 9 supralabials. Buri Province, southeastern Thailand, although Taylor (1965) provisionally referred literature records from Annam (central In life, the specimen was uniform olive-brown above Vietnam) by Smith (1943) to this species. anteriorly, with indistinct yellowish-white bands formed by exposed interstitial skin; upper parts black posteriorly with five irregular, broken, yellowish-brown longitudinal stripes; Elapidae upper surface of tail black with a yellowish-brown spot in the two medial scale rows; venter creamy anteriorly, Bungarus candidus (Linnaeus) becoming gray posteriorly; venter with row of large, light- coloured spots on the outer edge of ventral and subcaudal Material examined. – O’Rang: FMNH 263015, hilly evergreen scales from mid-body to tip of tail, spots widely spaced forest, near 12°16'24.6"N 107°03'53.1"E, 350 m elev., 10 Dec.2003. anteriorly but on every scale row posteriorly. Remarks. – A single female has a white dorsum and venter The specimen was removed alive from the jaws of a 3.5 m with 27 black saddle-like markings on the dorsum, the first Ophiophagus hannah (Cantor) at night (1830 hrs.). Both in contact with the head, becoming narrower posteriorly; nine snakes were approximately 1 m from the water on the bank black saddle-like markings on the tail, and a black tail tip; of a swift, 5 m wide stream between two large waterfalls. head black above; supralabials and infralabials white; black

151 Stuart et al.: Amphibians and reptiles from Cambodia spotting on some white interspaces between dorsal black Trimeresurus macrops Kramer saddles; enlarged vertebral scales; 15 longitudinal scale rows at mid-body; 218 ventral scales; 45 subcaudal scales with Material examined. – O’Rang: FMNH 262717, hilly evergreen scattered, dark brown spotting; and the tail ending in a point. forest, waterfall on O Ngeugn Stream, 12°18'50.5"N 107°06'03.4"E, 500 m elev., 11 Dec.2003; FMNH 262718, hilly evergreen forest, The specimen was collected at night (1845 hrs.) crawling over O Kamen Stream, near 12°19'35.3"N 107°05'33.4"E, 500 m elev., large rocks 3 m from a small (<1 m wide) rocky stream. 1 Nov.2003; FMNH 262719, 262721, hilly evergreen forest, O Doeung Por Stream, near 12°18'08.4"N 107°03'08.1"E, 500 m elev., Saint Girons (1972a) reported the species from Bokor, in the 2-6 Nov.2003; FMNH 262720, hilly evergreen forest, O Ronas Elephant Mountains of southwestern Cambodia. Stream, 12°17'30.7"N 107°03'06.0"E, 450 m elev., 3 Nov.2003. Siem Pang: FMNH 262715-16, evergreen mixed with bamboo forest, Viperidae waterfall of O Chhay Stream, tributary of O Chay Stream, 14°15'49.6"N 106°31'46.0"E, 200 m elev., 5 Dec.2003. Calloselasma rhodostoma (Boie) Remarks. – These have short, broad heads; 9-11 supralabials, Material examined. – Pichrada: FMNH 259196, at base of Phnom the first partly fused with the nasal; internasals in contact; Nam Lyr Mountain on rock hill, near 12°32'16"N 107°32'10"E, the supraocular broad, equal in width to about three scales 700 m elev., 18 Jun.2000. on top of the head; the vertical eye diameter greater than the distance from the ventral edge of eye to ventral edge of upper Remarks. – A single subadult has smooth scales; two lip; 21 longitudinal scale rows at midbody; males with 161- prefrontals; two internasals; 21 longitudinal scale rows at 163 ventrals (mean ± SD 162.2 ± 0.9, N = 4), females with midbody; pointed, upturned snout tip; brownish-gray dorsum 164-173 ventrals (mean ± SD 170.0 ± 5.2, N = 3); and flecked with dark brown, with alternating, triangular-shaped hemipenis long, papillose at base. dark brown markings; light brown stripe edged in black from tip of snout, above eye, to rear of jaw; and underparts dirty- In life, male FMNH 262717 had a reddish-orange eye; baby- whitish heavily powdered with brown. blue wash on scales on side of head, especially infralabials; chin scales baby-blue, flanked with white; dark green dorsals; The specimen was found at night (2030 hrs.) curled at the scales of first body scale row on anterior half of body with base of a grass clump on a rock hill. white marking, those on posterior half of body also having a reddish-brown marking on the ventral edge, forming a Saint Girons (1972a) reported the species from several ventrolateral stripe; ventrals on anterior 1/3 of body baby- localities throughout Cambodia. blue, remaining ventrals bright green; and dorsal and lateral surface of tail rusty-red.

Trimeresurus albolabris Gray Specimens were collected at night (1810-2040 hrs.) on the soil bank, herbaceous plants, bushes, tree roots, and tree Material examined. – O’Rang: FMNH 263013, grass and disturbed branches within 10 m of swift, rocky streams, often near evergreen forest along road, 12°19'18.5"N 107°05'53.5"E, 550 m waterfalls. The species was collected together with T. vogeli elev., 9 Dec.2003. (see below); for example, FMNH 262717 was captured within 3 m of a T. vogeli (FMNH 263025). Remarks. – A single male has 11 supralabials; internasals in contact; supraocular narrow, equal in width to about two This is the first report of the species from Cambodia. scales on top of the head, 9 scales separating supraoculars; the vertical eye diameter less than the distance from the ventral edge of eye to ventral edge of upper lip; the side of the head Trimeresurus vogeli David, Vidal & Pauwels below eyes lighter in colouration than the rest of the head; 21 longitudinal scale rows at midbody; a uniformly yellow Material examined. – O’Rang: FMNH 263025, hilly evergreen (in life) lateral stripe; 163 ventrals; 68 subcaudals; dorsal forest, below waterfall on O Ngeugn Stream, 12°18'50.5"N 107 06'03.4"E, 500 m elev., 11 Dec.2003; FMNH 263026-28, hilly surface of tail reddish; and hemipenis long, papillose at base. ° evergreen forest, O Ngeugn Stream, Mok Romum Waterfall, near Smith (1943) stated that the first supralabial is “more or less 12°18'58"N 107°05'59"E, 550 m elev., 31 Oct.-5 Nov.003. completely united with the nasal,” but in this specimen the first supralabial on both sides is separated from the nasal by Remarks. – Four males agree with David et al.’s (2001) a weak suture. original description by having the first supralabial separated from the nasal; a short spinose hemipenis; fewer than 174 The specimen was taken at night (2000 hrs.) crawling on a ventrals (160-166, mean ± SD 163.7 ± 2.6, N = 4); white dirt road. vertebral spots (in three of four specimens); and a mostly green tail with a dark gray or brown tail tip. Saint Girons (1972a) reported the species from central and southwestern Cambodia. In life, FMNH 263025 had a tan-yellow eye; dark green dorsals, some with baby-blue outer edges; bright green

152 THE RAFFLES BULLETIN OF ZOOLOGY 2006 ventrals; baby-blue wash on supralabials and scales on side and all nine were reported from the Cardamom Mountains of head; chin scales green with baby-blue outer edges; small (Ohler et al., 2002; H. rugulosus as H. chinensis and O. white spots widely spaced on vertebrals; upper 2/3 of scale martensii as Phrynoglossus martensii in Ohler et al., 2002). in first body scale row white, lower 1/3 of scales reddish- Of the 21 species of frogs reported here as characteristic of brown, forming a ventrolateral stripe; baby-blue wash on undisturbed environments, 14 (66.7%) were reported from subcaudals and to a lesser extent on dorsal surface of tail; the Kontum Plateau (Inger et al. 1999; Ophryophryne hansi and tail tip dark gray-brown. as O. poilani, Limnonectes poilani as Rana cf. blythii, Rana banaorum as R. livida, and Rana morafkai as R. livida in All specimens were taken at night (1815-1900 hrs.) next to Inger et al., 1999), but only five (23.8%) were reported from a swift rocky stream with large waterfalls. Two were crawling the Cardamom Mountains by Ohler et al. (2002): Kalophrynus on vines 2-3 m above the ground and two were crawling on interlineatus, Microhyla berdmorei, Rana macrodactyla, rocks along the stream bank. The species was collected Rana taipehensis, and Theloderma asperum. There are together with T. macrops (see above). limitations in comparing these three collections owing to differences in the methods, lengths, and times of year of Malhotra et al. (2004) reported the species from Pursat and sampling. Nonetheless, two reasonably sound conclusions can Kampot Provinces in southwestern Cambodia. be made from these comparisons. First, the frog species characteristic of anthropogenically-modified environments are rather ubiquitous in southern Indochina. Second, intact DISCUSSION areas in eastern Cambodia contain a frog fauna that is similar to that of the mountains of adjacent Vietnam (of which the This collection of 30 species of amphibians and 42 species uplands in eastern Cambodia are the lower slopes), but very of reptiles contains major additions to the herpetofauna of different from that in the mountains of southwestern Cambodia. A new species each of Leptobrachium and Cambodia. Ophyrophryne are described. Of the remaining amphibians, 11 species (39.3%) are reported from Cambodia for the first This finding of two very different frog faunas in the two major time, including the first example of the Order Gymnophiona. upland areas of Cambodia has implications for national-level Of the snakes, seven species (33.3%) are reported from biodiversity conservation strategies. Recent publications on Cambodia for the first time. Most of the 20 species of lizards the herpetofauna of the Cardamom Mountains and their low- appear to be first definitive records for the country as well, lying river valleys (e.g. Daltry & Wüster, 2002; Ohler, et al., but the lack of any published treatises on the lizard fauna of 2002; Platt et al., 2003a, b) have attracted conservation Cambodia makes this statement difficult to quantify. attention. Owing to the stark faunal differences in at least the frogs, such attention to the uplands of eastern Cambodia is Four species in the collection are reported here for the first clearly warranted. time in Cambodia since Günther (1861, 1864) originally described them from Henri Mouhot’s Cambodian collection: Bufo galeatus, Bronchocela smaragdina, Acanthosaura capra ACKNOWLEDGMENTS and A. coronata. The distribution of the last species is unclear, as it has been treated in the literature as a junior synonym of The opportunity for BLS to work in Cambodia was made A. lepidogaster since Smith (1935). However, the first three possible by the Wildlife Conservation Society / Ministry of species are now known to occur in the Annamite Mountains Agriculture, Forestry and Fisheries / Ministry of Environment in adjacent central and southern Vietnam (Smith, 1921; Inger Collaborative Program. Financial support was provided by et al., 1999; Liu et al., 2000; Hallermann, 2004; Kalyabina- The John D. and Catherine T. MacArthur Foundation, the Hauf et al., 2004). It is very likely that these four species Wildlife Conservation Society, and the National Geographic were obtained by Mouhot during his single trip through Society (grant no. 6247-98 with Harold Heatwole), and eastern Cambodia in 1859 (to “Brelum,” located in extreme logistical support was provided by Joe Walston, Colin Poole, eastern Mondolkiri Province, Cambodia or just over the and the Wildlife Conservation Society. An Dara and Suon border in present-day Vietnam; see Introduction), and have Phalla assisted with fieldwork. Harold Voris, Alan Resetar, not been reported from Cambodia since their original and Jamie Ladonski facilitated examining specimens at the descriptions owing to a lack of herpetological study in the Field Museum of Natural History. Sean Bober constructed hilly eastern parts of the country. the map. Sophie Molia provided French translations. Annemarie Ohler (Muséum national d’Histoire naturelle, Two collections of frogs have been recently reported from Paris) loaned the holotypes of Limnonectes poilani and L. mountainous areas flanking hilly eastern Cambodia: a toumanoffi in her care. Jennifer Sheridan counted the eggs of collection from the Kontum Plateau of central Vietnam by female R. banaorum and R. morafkai. Robert Inger, Patrick Inger et al. (1999) and a collection from the Cardamom David, and John Murphy shared their taxonomic opinions. Mountains of southwestern Cambodia by Ohler et al. (2002). Natalia Ananjeva provided a copy and English translation of Of the nine species of frogs reported here as characteristic of Darevsky and Nguyen (1983), and Patrick David provided a anthropogenically-modified environments (Table 2), seven copy of Wagner (1975). Robert Inger, Raoul Bain, and an were reported from the Kontum Plateau (Inger et al., 1999) anonymous reviewer improved the manuscript.

153 Stuart et al.: Amphibians and reptiles from Cambodia

LITERATURE CITED Guicking, D., U. Fritz, M. Wink, & E. Lehr, 2002. New data on the diversity of the Southeast Asian leaf turtle genus Cyclemys Bell, Bain, R. H., A. Lathrop, R. W. Murphy, N. L. Orlov, & T. C. Ho, 1834. Molecular results (Reptilia: Testudines: Geoemydidae). 2003. Cryptic species of a cascade frog from Southeast Asia: Faunistische Abhandlungen Staatliches Museum für Tierkunde taxonomic revisions and descriptions of six new species. Dresden, 23(4): 75-86. American Museum Novitates, 3417: 1-60. Gray, J. E., 1861a. Description of a soft tortoise from Camboja. Boulenger, G. A., 1885. Catalogue of the Lizards in the British Proceedings of the Zoological Society of London, 1861(2): 41- Museum (Natural History). Second Edition. Vol. I. Geckonidae, 42. Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. British Gray, J. E., 1861b. List of Mammalia, tortoises and crocodiles Museum (Natural History), London. 436 pp. Vol. III. Lacertidae, collected by M. Mouhot in Camboja. Proceedings of the Gerrhosauridae, Scincidae, Anelytropidae, Dibamidae, Zoological Society of London, 1861(8): 135-140. Chamaeleontidae. 575 pp. Günther, A., 1861. Second list of Siamese reptiles. Proceedings of Boulenger, G. A., 1886. First report on additions to the batrachian the Zoological Society of London, 1861(10): 187-189. collection in the Natural-History Museum. Proceedings of the Günther, A. C. L. G., 1864. The Reptiles of British India. Taylor Zoological Society of London, 1886: 411-416. and Francis, London. 452 pp. Boulenger, G. A., 1903. Descriptions of three new batrachians from Günther, A., 1875. Second report on collections of Indian reptiles Tonkin. Annals and Magazine of Natural History, 7(12): 186- obtained by the British Museum. Proceedings of the Zoological 188. Society of London, 1875: 224-234. Bourret, R., 1936. Les Serpents de L’Indochine. Imprimerie Henri Hallermann, J., 2004. A new species of the genus Bronchocela from Basuyau & Cie., Toulouse. Vol. I, 141 pp.; Vol. II, 505 pp. the tropical rain forest of southern Vietnam. Russian Journal of Bourret, R., 1941a. Les tortues de l’Indochine. Institut Herpetology, 11(1): 30-34. Océanographique de l’Indochine, 38:1-235. Holloway, R. H. P., 2003. Domestic trade of tortoises and freshwater Bourret, R., 1941b. Notes herpétologiques sur l’Indochine française. turtles in Cambodia. Linnaeus Fund Research Report. Chelonian XXII. Reptiles et batraciens reçus au Laboratoire des Sciences Conservation and Biology, 4(3): 733-734. Naturelles de l’Université au cours de l’année 1941. Description Holloway, R. H. P. & S. Heng. 2004. Geographic distribution: d’une espèce et d’une variété nouvelles. Direction de Batagur baska. Herpetological Review, 35(3): 284. l’Instruction Publique: 1-29. Inger, R. F., N. Orlov, & I. Darevsky, 1999. Frogs of Vietnam: a Bourret, R., 1942. Les batraciens de l’Indochine. Mémoires de report on new collections. Fieldiana Zoology (New Series), 92: l’Institut Océanographique de l’Indochine, 6: 1-547. 1-46. Chuaynkern, Y., A. Ohler, C. Inthara, P. Kumtong, & A. Dubois, Iskandar, D. T., 1998. The Amphibians of Java and Bali. Research 2004. The recent distribution of Rana milleti Smith, 1921 in and Development Centre for Biology-LIPI, Indonesia. 117 pp. mainland Southeast Asia with the first record of Cambodia. The Natural History Journal of Chulalongkorn University, 4(1): 1- Kalyabina-Hauf, S., N. B. Ananjeva, U. Joger, P. Lenk, R. W. 13. Murphy, B. L. Stuart, N. L. Orlov, C. T. Ho, & M. Wink, 2004. Molecular phylogeny of the genus Acanthosaura (Agamidae). Cochran, D. M., 1927. New reptiles and batrachians collected by Current Herpetology, 23(1): 7-16. Dr. Hugh M. Smith in Siam. Proceedings of the Biological Society of Washington, 40: 179-192. Kou, Z., 1985. A new species of Ophryophryne from Yunnan. Acta Herpetologica Sinica, 4(1): 41-43. Daltry, J. & D. Chheang, 2000. Siamese crocodiles discovered in the Cardamom Mountains. Crocodile Specialist Group Lathrop, A., R. W. Murphy, N. L. Orlov, & C. T. Ho, 1998. Two Newsletter, 19(2): 7-8. new species of Leptobrachium (Anura: Megophryidae) from the Central Highlands of Vietnam with a redescription of Daltry, J. C. & W. Wüster, 2002. A new species of wolf snake Leptobrachium chapaense. Russian Journal of Herpetology, (Serpentes: Colubridae: Lycodon) from the Cardamom 5(1): 51-60. Mountains, southwestern Cambodia. Herpetologica, 58(4): 498- 504. Lehr, E. & R. Holloway. 2000. Geographic distribution: Manouria impressa. Herpetological Review, 31(2): 111. Darevsky, I. S. & S. V. Nguyen, 1983. New and little known lizard species from Vietnam. Zoologicheskii Zhurnal, 62(12): 1827- Lehr, E. & R. Holloway. 2002. Untersuchungen zum 1837 Schildkrötenhandel in Kambodscha. Reptilia, 7(5): 62-70. Das, I., 2004. A new species of Dixonius (Sauria: Gekkonidae) from Liu, W., A. Lathrop, J. Fu, D. Yang, & R. W. Murphy, 2000. southern Vietnam. Raffles Bulletin of Zoology, 52(2): 629-634. Phylogeny of East Asian bufonids inferred from mitochondrial DNA sequences (Anura: Amphibia). Molecular Phylogenetics David, P., N. Vidal, & O. S. G. Pauwels, 2001. A morphological and Evolution, 14(3): 423-435. study of Stejneger’s pitviper Trimeresurus stejnegeri (Serpentes, Viperidae, Crotalinae), with the description of a new species Malhotra, A., R. S. Thorpe, & B. L. Stuart, 2004. A morphometric from Thailand. Russian Journal of Herpetology, 8(3): 205-222. analysis of Trimeresurus vogeli (David, Vidal and Pauwels, 2001), with new data on diagnostic characteristics, distribution Deuve, J., 1970. Serpents du Laos. Mémoires O.R.S.T.O.M. (Office and natural history. The Herpetological Journal, 14: 65-77. de la Recherche Scientifique et Technique Outre-Mer), 39: 1- 251. Matsui, M., J. Nabhitabhata, & S. Panha, 1999. On Leptobrachium from Thailand with a description of a new species (Anura: Dubois, A. & A. Ohler, 1998. A new species of Leptobrachium Pelobatidae). Japanese Journal of Herpetology, 18(1): 19-29. (Vibrissaphora) from northern Vietnam, with a review of the taxonomy of the genus Leptobrachium (Pelobatidae, Matsui, M., T. Chan-ard, & J. Nabhitabhata, 1996. Distinct specific Megophryinae). Dumerilia, 4(1): 1-32. status of Kalophrynus pleurostigma interlineatus (Anura, Microhylidae). Copeia, 1996(2): 440-445.

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McGuire, J. A. & K. B. Heang, 2001. Phylogenetic systematics of Smith, M., 1922. The frogs allied to Rana doriae. Journal of the Southeast Asian flying lizards (Iguania: Agamidae: Draco) as Natural History Society of Siam, 4(4): 215-229. inferred from mitochondrial DNA sequence data. Biological Smith, M. A., 1924. New tree-frogs from Indo-China and the Malay Journal of the Linnean Society, 72: 203-229. Peninsula. Proceedings of the Zoological Society of London, Mouhot, M. H., 1864. Travels in the Central Parts of Indo-China 1924: 225-234. (Siam), Cambodia, and Laos, during the Years 1858, 1859, and Smith, M. A., 1928. Description of a new species of Draco from 1860. John Murray, London, Volume I, 303 pp., Volume II, the Indo-Chinese region. Annals and Magazine of Natural 301 pp. Reprinted in 1986 by White Lotus Co., Ltd., Bangkok. History, 10(2): 248. Murphy, J. C., H. K. Voris, B. L. Stuart, & S. G. Platt, 2002. Female Smith, M. A., 1935. The Fauna of British India, including Ceylon reproduction in the Rainbow Water Snake, Enhydris enhydris and Burma. Reptilia and Amphibia. Vol. II.-Sauria. Taylor and (Serpentes, Colubridae, Homalopsinae). The Natural History Francis, London. 440 pp. Journal of Chulalongkorn University, 1(2): 31-37. Smith, M. A., 1943. The Fauna of British India, Ceylon and Burma, Ohler, A., 2003. Revision of the genus Ophryophryne Boulenger, including the whole of the Indo-Chinese sub-region. Reptilia 1903 (Megophryidae) with description of two new species. and Amphibia. Vol. III.-Serpentes. Taylor and Francis, London. Alytes, 21(1-2): 23-44. 583 pp. Ohler, A., S. R. Swan, & J. C. Daltry, 2002. A recent survey of the Stuart, B. L., J. Smith, K. Davey, Prom Din, & S. G. Platt, 2000. amphibian fauna of the Cardamom Mountains, southwest Homalopsine watersnakes: The harvest and trade from Tonle Cambodia with descriptions of three new species. Raffles Sap, Cambodia. TRAFFIC Bulletin, 18(3): 115-124. Bulletin of Zoology, 50(2): 465-481. Stuart, B. L., D. An, & P. P. van Dijk, 2002. A record of the Ohler, A., A. Teynié, & P. David, 2004. A green-eyed leatherback sea turtle Dermochelys coriacea from Cambodia. Leptobrachium (Anura: Megophryidae) from southern Laos. Marine Turtle Newsletter, 96: 22. Raffles Bulletin of Zoology, 52(2): 695-700. Stuart, B. L. & S. G. Platt, 2004. Recent records of turtles and Orlov, N., 1997. Breeding behavior and nest construction in a tortoises from Laos, Cambodia, and Vietnam. Asiatic Vietnam frog related to Rana blythii. Copeia, 1997(2): 464-465. Herpetological Research, 10: 129-150. Orlov, N. L., R. W. Murphy, N. B. Ananjeva, S. A. Ryabov, & T. Taylor, E. H., 1962. The amphibian fauna of Thailand. The C. Ho, 2002. Herpetofauna of Vietnam, a checklist. Part I. University of Kansas Science Bulletin, 43(8): 265-599. Amphibia. Russian Journal of Herpetology, 9(2): 81-104. Taylor, E. H., 1965. The serpents of Thailand and adjacent waters. Parker, H. W., 1934. A Monograph of the Frogs of the Family The University of Kansas Science Bulletin, 45(9): 609-1096. Microhylidae. Jarrold and Sons Ltd., Norwich. 208 pp. Taylor, E. H., & R. E. Elbel, 1958. Contribution to the herpetology Platt, S. G., Heng Sovannara, Long Kheng, B. L. Stuart, & J. of Thailand. The University of Kansas Science Bulletin, 38(13): Walston, 2003a. Crocodylus siamensis along the Sre Ambel 1033-1189. River, southern Cambodia: habitat, nesting, and conservation. Herpetological Natural History, 9(2): 165-169. Tirant, G., 1885. Notes sur les Reptiles et les Batraciens de la Cochinchine et du Cambodge. Imprimerie du Gouvernement, Platt, S. G., B. L. Stuart, Heng Sovannara, Long Kheng, Kalyar, & Saigon, 104 pp. Heng Kimchay, 2003b. Rediscovery of the critically endangered river terrapin, Batagur baska, in Cambodia, with notes on Touch, T. S., H. L. Prak, T. Chul, S. Lieng, S. Chun, P. Hout, & K. occurrence, reproduction, and conservation status. Chelonian Heng, 2000. Overview of turtle trade in Cambodia. In: van Dijk, Conservation and Biology, 4(3): 691-695. P. P., B. L. Stuart, & A. G. J. Rhodin (eds.), Asian Turtle Trade: Proceedings of a Workshop on Conservation and Trade of Saint Girons, H., 1972a. Les serpents du Cambodge. Mémoires du Freshwater Turtles and Tortoises in Asia. Chelonian Research Muséum National d’Histoire Naturelle. Nouvelle Série A, Monographs 2. Chelonian Research Foundation, Lunenburg. Pp. Zoologie, 74: 1-170. 55-57. Saint Girons, H., 1972b. Etude de la collection de serpents de van Denburgh, J., 1909. New and previously unrecorded species of Cochinchine du Docteur Morice déposée en 1873 et 1874 au reptiles and amphibians from the island of Formosa. Proceedings Muséum d’Histoire Naturelle de Lyon. Nouveau Archives du of the California Academy of Sciences, 3: 49-56. Muséum d’Histoire Naturelle de Lyon, 9: 3-12. Wagner, F. W., II., 1975. A revision of the Asian colubrid snakes Sclater, W. L., 1891. Notes on the collection of snakes in the Indian Oligodon cinereus (Günther), Oligodon joynsoni (Smith), and Museum with descriptions of several new species. Journal of Oligodon cyclurus (Cantor). Unpublished Master’s Thesis, the Asiatic Society of Bengal, 60: 230-250. Louisiana State University, Baton Rouge: 1-97. Smith, M. A., 1921. New or little-known reptiles and batrachians from southern Annam (Indo-China). Proceedings of the Zoological Society of London, 1921: 423-440.

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THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 157-180 Date of Publication: 28 Feb.2006 © National University of Singapore

CHECKLIST OF THE HERPETOFAUNA OF THE SERIBUAT ARCHIPELAGO, WEST MALAYSIA WITH COMMENTS ON BIOGEOGRAPHY, NATURAL HISTORY, AND ADAPTIVE TYPES

L. Lee Grismer, Timothy M. Youmans, Perry L. Wood, Jr., Jesse L. Grismer Department of Biology, La Sierra University, Riverside, California 92515, USA

ABSTRACT. – The Seribuat Archipelago is an environmentally complex group of 62 islands which lie off the southeast coast of Peninsular Malaysia in the South China Sea. Its herpetofauna, which is documented herein for the first time, contains 23 species of frogs, one species of non-marine turtle, 43 species of lizards, 44 species of non-marine snakes, and represents a diverse array of adaptive types. Of these, one frog, 11 lizards and four snakes are endemic. The majority of the herpetofauna, including 14 of the 16 endemics, occurs in the outermost arc of islands which are the oldest, largest, and most environmentally diverse. Some of the species in this Outer Arc are more closely related to species from Sumatra, Java, and/or Borneo than they are to species from peninsular Malaysia, suggesting they had a vicariant origin.

KEY WORDS. – Seribuat Archipelago, Herpetofauna, Malaysia, Endemic, Biogeography.

INTRODUCTION 1966). Pulau Tioman continues to be an area of active biological research (Day, 1990; Sodhi et al., 1999) and The Seribuat Archipelago comprises an environmentally recently, the majority of published studies concerning this diverse array of islands located in the southern section of the island have focused on its herpetofauna (Das & Grismer, South China Sea. These islands flank the southeast coast of 2003; Das & Lim, 2000; Diaz et al., 2004; Escobar et al., peninsular Malaysia along the southern states of Pahang and 2003c; Hendrickson, 1966a, b; J. Grismer et al., 2003a, b, c, Johor (Fig. 1). The archipelago contains 62 islands ranging in size from 0.01–110 km2 (Table 1) and extends from Pulau Cebeh in the north 100 km southward to Pulau Tokong Yu. It has been demonstrated that these islands, along with the Anambas and Natuna Archipelagos of Indonesia, are part of a key phylogenetic link between the flora and fauna of peninsular Malaysia and other islands of the Sunda Shelf (Adura, 1998, L. Grismer, et al., 2003; 2004b; Kottelat, 1990; Leong et al., 2003; Tan & Mohamad, 1999; Ng, Tan and Lim, 1999; Sanders et al., 2005; Yeo & Ng, 1999; Yeo et al., 1999). Leong et al. (2003) provided preliminary checklists for the herpetofauna of the Anambas and Natuna archipelagos and we supplement these with the checklist provided herein.

Despite the extreme environmental diversity within and between the islands of this archipelago, biological studies have been relatively infrequent. In fact, there have been no archipelago-wide studies comparing broad taxonomic cross sections of any portion of the flora or fauna to one another or to taxa of the adjacent peninsula. Only the largest island, Pulau Tioman, has been the focus of several botanical studies (see Jaman & Latiff, 1999; Latiff et al., 1999 and references therein) as well as natural history expeditions which began Fig. 1. Islands of the Seribuat Archipelago. Island names in large in earnst during the mid 1960s (see Bullock & Medway, fonts represent island groups as delimited in the text.

157 Grismer et al.: Herpetofauna of Seribut Archipelago

Table. 1. Location, size, elevation, and period of the day of the survey of the islands of the Tioman Archipelago.

Island km2 elevation (m) location am/pm surveys INNER ARC Aceh group Aceh 1.32 150 02º 39.6' 103 46.3' am,pm Anak Aceh 0.01 ~25 02º 40.9' 103 45.9' am Tunas Utara 0.02 44 02º 41.4' 103 44.8' am Tunas Selantan 0.02 31 02º 41.2' 103 45.1' am Layak 0.24 69 02º 41.8' 103 44.4' am, pm Lalang 0.07 38 02º 39.2' 103 42.8' am Pelandok 0.04 ~30 02º 39.2' 104 45.0' am Tengah 0.16 58 02º 39.8' 103 45.3' am Mawar 0.33 72 02º 37.0' 103 47.0' am Lalang Selatan 0.02 ~10 02º 33.9' 103 49.7' am

Setindan group Setindan 3.8 105 02º 28.7' 103 51.0' am Batu Gajah 0.04 ~60 02º 29.5' 103 51.4' am Belanak 0.01 50 02º 21.0' 103 55.2' am MIDDLE ARC Seribuat group Seribuat 5.52 124 02º 41.5' 103 55.0' am, pm Sembilang 2.51 257 02º 41.5' 103 53.3' am, pm Mertang Barat 0.04 44 02º 39.3' 103 52.8' am Mertang Tengah 0.01 27 02º 39.1' 103 53.1' am Mertang Timur 0.03 38 02º 38.7' 103 53.4' am Tasu 0.04 70 02º 41.7' 103 55.7' am Rawa group Rawa 0.3 97 02º 31.6' 103 58.6' am Harimau 0.21 91 02º 33.5' 103 56.6' am Mensirip 0.08 32 02º 33.0' 103 57.5' am Gual 0.09 56 02º 32.1' 103 58.2' am Besar group Babi Besar 6.38 251 02º 25.9' 103 58.7' am, pm Babi Tengah 0.55 83 02º 28.6' 103 07.8' am Babi Hujong 0.66 99 02º 29.3' 103 57.0' am Tinggi group Tinggi 13.5 610 02º 18.0' 104 07.0' am, pm Apil 0.04 13 02º 16.7' 104 07.6' am Mentigi 0.01 3 02º 16.5' 104 07.1' am Nangka Kecil 0.04 ~15 02º 16.5' 104 07.5' am Nangka Besar 0.1 33 02º 16.2' 104 07.5' am Simbang 0.07 23 02º 15.0' 104 09.2' am Lanting 0.01 16 02º 16.5' 104 09.4' am Ibol 0.15 82 02º 18.4' 104 09.2' am Penyembong 0.04 17 02º 18.4' 104 08.0' am group Sibu 4.81 155 02º 13.0' 104 04.2' am, pm Sibu Tengah 0.45 74 02º 11.1' 104 06.1' am Papan 0.02 22 02º 11.3' 104 05.7' am Sibu Kukus 0.03 45 02º 10.3' 104 06.7' am Sibu Hujong 0.03 59 02º 10.9' 104 06.6' am Lima group Lima Besar 0.1 48 02º 13.2' 104 09.0' am, pm Lima Kecil 0.04 41 02º 13.4' 104 09.0' am Tokong Sanggol 0.01 22 02º 13.4' 104 08.1' am Tokong Raket 0.01 29 02º 13.0' 104 09.6' am Tokong Chondong 0.02 38 02º 11.6' 104 10.6' am Tokong Belalai 0.01 30 02º 11.4' 104 10.6' am Tokong Chupak 0.01 30 02º 10.6' 104 11.1' am Tokong Yu 0.01 28 02º 07.4' 104 14.8' am

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Table. 1. Location, size, elevation, and period of the day of the survey of the islands of the Tioman Archipelago. (Continued)

Island km2 elevation (m) location am/pm surveys OUTER ARC Tioman 110 1038 02 47.0' 104 10.0' am, pm Pemanggil 9.2 405 02 35.0' 104 19.0' am, pm Tulai group Tulai 0.87 98 02 54.5' 104 06.5' am, pm Labas 0.01 20 02 53.3' 104 04.0' am Sepoi 0.03 60 02 53.9 104 03.1' am, pm Cebeh 0.05 71 02 56.0' 104 05.9' am, pm Aur group Aur 12.75 521 02 27.0' 104 32.0' am, pm Dayang 2.09 158 02 28.3' 104 30.0' am Lang 0.05 38 02 27.5' 104 29.5' am Pinang 0.11 97 02 26.2' 104 33.3' am Tokong Burung group Tokong Burung Besar 0.36 54 02 47.0' 103 57.6' am Tokong Burung Condong 0.01 41 02 46.7' 103 57.6' am Tokong Burung Tengah >0.01 32 02 46.8 103 57.6' am Tokong Bahara 0.02 54 02 39.7' 104 03.8' am Jahat 0.05 30 02 39.8' 104 10.0' am

2004; L. Grismer, 2005a, Grismer, 2006b; L. Grismer et al., University Herpetological Collection (LSUHC). Material 2002, 2003, 2004a,b; Hien et al., 2001; Leong 2000; Leong collected from the latter two expeditions is temporarily & Grismer, 2004; Leong & Crane, 2001; Leong & Tan, 2000; deposited in the LSUHC pending further study. All tissues Lim & Lim, 1999; van Rooijen, 2001; Wood et al., 2003a, d; are deposited in the LSUHC and voucher photographs are Youmans et al., 2003). These reports provided the catalyst deposited in the La Sierra University Digital Photograph for a surge of herpetological exploration which has swept Collection (LSUDPC). Table 2 lists the species of the Seribuat over much of the Seribuat Archipelago (Escobar et al., 2003a, Archipelago and the islands on which they occur. Appendix b; L. Grismer, 2005a, b, 2006, a, b; Grismer & Das, 2006; L. I cross-references this table and provides species checklists Grismer et al., 2001a, b, 2003; Wood et al., 2003b, c; 2004a, for each island. Appendix II provides locality and catalogue b; Valdivia et al. 2004; Youmans & Grismer, 2006; Youmans numbers for all new island records not previously published. et al., 2002, 2003), resulting in the addition of several new island records and the discovery of at least 13 new species. The surface area (km2), elevation (meters above sea level [m]), However, even with the addition of these later efforts, only and locality for each island is provided (Table 1). To form 10 of the 62 islands had ever been explored. Therefore, we the basis for future ecological studies, a general description report here 171 new island records resulting from our surveys of each island is given which includes topography, vegetative of the remaining 52 islands and additional surveys made on structure, and habitat types. Island names and spelling follow previously explored islands. those proposed by the Director of National Mapping, Malaysia, 1995–1997.

MATERIALS AND METHODS Abbreviations used are as follows: G. = Gunung (mountain); Kg. = Kampung (village); P. = Pulau (island) S. = Sungai Two primary surveys augmenting the previous works cited (river); Tk. = Telok (bay). above, took place from 19–25 August 2003 and from 7–15 September 2004 in an effort to generate baseline data from the unexplored islands. All islands were visited at least once VEGETATION ZONES OF THE SERIBUAT during the day by seven or eight people. Some of the more ARCHIPELAGO heavily forested islands were surveyed at night as well (Table 1). Specimens were collected by hand and/or blow pipe, Vegetation zones generally serve to highlight broad photographed, tissued for liver samples (for future molecular categorical differences between habitats across sizable studies), fixed in 10% formalin, and subsequently transferred geographic areas. These categorical differences lack well- to 70% ethanol. Material collected from previous expeditions defined geographic boundaries (Ashton, 1995) and, with the is deposited at the Forest Research Institute of Malaysia exception of mangrove communities, each zone transitions (FRIM), The Raffles Museum of Biodiversity Research, The smoothly, continuously, and usually imperceptibly into one National University of Singapore (ZRC), the Malaysian another along an altitudinal transect. The vegetation zones Wildlife Department (PERHILITAN), and the La Sierra used here follow those of Latiff et al. (1999).

159 Grismer et al.: Herpetofauna of Seribut Archipelago

Table 2. Island checklist of species in the Tioman Archipelago. Asterisk indicates the species is endemic.

Amphibians Megophryidae Leptolalax kajangensis*Tioman Megophrys nasuta Tioman Bufonidae Ansonia tiomonica*Tioman Bufo asper Tioman Bufo melanostictus Tioman Bufo parvus Tioman Pelophryne brevipes Aur, Tioman Microhylidae Chaperina fusca Tioman Kalophrynus pleurostigma Tioman Kaloula baleata Aceh, Tioman Kaloula pulchra Tioman Microhyla borneensis Sibu Rhacophoridae Nyctixalus pictus Tioman Philautus petersi Tioman Polypedates leucomystax Aceh, Seribuat, Sembilang, Babi Besar, Babi Tengah, Sibu, Tioman Theloderma horridum Tioman Ranidae Fejervarya cancrivora Seribuat, Sembilang, Babi Besar, Sibu, Tioman, Limnonectes blythii Tinggi, Tioman, Aur, Pemanggil Rana erythraea Tioman Rana hosii Tinggi, Tioman Rana picturata Tioman Rana raniceps Tioman Taylorana hascheana Tioman Ichthyophiidae Ichthyophis sp. Tioman Reptiles Trionychidae Dogania subplana Tinggi, Tioman Agamidae Acanthosaura armata Aur, Tioman Aphaniotis fusca Aceh, Seribuat, Sembilang, Babi Besar, Babi Hujong, Tinggi, Sibu, Tioman Bronchocela cristatella Babi Besar, Tinggi, Sibu, Sibu Tengah, Tioman, Tulai, Pemanggil, Aur Draco fimbriatus Tioman Draco formosus Tinggi Draco haematopogon Tioman Draco melanopogon Tinggi, Sibu, Tioman Draco sumatranus Babi Besar, Tinggi, Tioman Gonocephalus chamaeleontinus Tioman Gonocephalus grandis Tioman Gekkonidae Cnemaspis baueri* Aur, Dayang Cnemaspis kendallii Aceh, Seribuat, Sembilang, Babi Besar, Babi Hujong, Tinggi, Ibol, Sibu, Sibu Tengah, Tioman, Tulai Cnemaspis limi*Tioman, Tulai Cnemaspis pemanggilensis* Pemanggil Cosymbotus craspedotus Tinggi, Tioman Cosymbotus platyurus Tioman, Aur Cyrtodactylus aurensis* Aur Cyrtodactylus quadrivirgatus Tioman Cyrtodactylus seribuatensis* Seribuat, Sembilang, Nangka Kecil, Mentigi, Sibu, Sibu Tengah, Lima Besar Cyrtodactylus tiomanensis*Tioman Gehyra mutilata Rawa, Babi Besar, Babi Tengah, Nangka Besar, Sibu, Lima Besar, Lima Kecil, Tioman, Aur Gekko monarchus Aceh, Layak, Tunas Selatan, Pelandok, Batu Gajah, Seribuat, Sembilang, Mertang Tengah, Mertang Timur, Babi Besar, Babi Hujong, Babi Tengah, Tinggi, Penyambong, Ibol, Simbang, Nangka Kecil, Mentigi, Sibu, Lima Besar, Tioman, Tulai, Cebeh, Jahat, Pemanggil, Aur, Dayang Gekko smithii Tioman, Tulai

160 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Table 2. Island checklist of species in the Tioman Archipelago. Asterisk indicates the species is endemic. (Continued)

Hemidactylus frenatus Aceh, Aceh Anak, Layak, Tunas Selatan, Tunas Utara, Lalang Selatan, Lalang Utara, Pelandok, Mawar, Setindan, Belanak, Seribuat, Semilang, Tasu, Mertang Timur, Rawa, Harimau, Gual, Babi Besar, Babi Hujong, Babi Tengah, Tinggi, Penyambong, Nangka Besar, Nangka Kecil, Mentigi, Sibu, Sibu Tengah, Sibu Kukus, Lima Besar, Lima Kecil, Chondong, Belalai, Tioman, Tulai, Sepoi, Pemanggil, Aur, Dayang, Lang, Tokong Burung Besar, Tokong Burung Chondong, Tokong Burung Tengah, Tokong Bahara Hemiphyllodactylus typus Sibu Lepidodactylus lugubris Tunuas Selatan, Mawar, Babi Tengah, Nangka Kecil, Mentigi, Sibu, Papan, Sanggol, Raket, Tioman, Ceben, Sepoi, Jahat Ptychozoon kuhli Aceh, Sembilang, Babi Besar, Tinggi, Tioman, Tulai Ptychozoon lionatum Sibu, Tioman Scincidae Dasia olivacea Babi Besar, Babi Tengah, Tinggi, Sibu, Tioman, Aur Emoia atrocostata Seribuat, Simbang, Nangka Besar, Sibu, Papan, Sibu Tengah, Sibu Kukus, Tulai, Aur Eutrophis longicaudata Tioman Eutrophis multifasciata Aceh, Seribuat, Sembilang, Rawa, Babi Besar, Babi Tengah, Tinggi, Sibu, Sibu Kukus, Tioman, Tulai, Sepoi, Pemanggil, Aur, Dayang, Lang, Tokong Burung Besar, Tokong Burung Chondong, Tokong Burung Tengah, Tokong Bahara Larutia seribuatensis*Tioman, Tulai Lipinia surda Tioman, Aur Lipinia vittigera Babi Besar, Tioman, Aur Lygosoma bowringii Aceh, Tunus Selatan, Mawar, Setindan, Seribuat, Sembilang, Mertang Tengah, Mertang Timur, Rawa, Harimau, Mensirip, Babi Besar, Babi Tengah, Ibol, Nangka Kecil, Mentigi, Sibu, Papan, Sibu Tengah, Lima Besar, Tioman, Pemanggil, Aur Sphenomorphus ishaki*Tioman Sphenomorphus sibuensis* Sibu Sphenomorphus scotophilus Tioman, Tulai, Pemanggil, Aur Varanidae Varanus nebulosus Tioman, Aur Varanus salvator Aceh, Anak Aceh, Layak, Tunas Selatan, Tunas Utara, Tengah, Pelandok, Mawar, Setindan, Batu Gajah, Seribuat, Sembilang, Tasu, Mertang Barat, Mertang Tengah, Rawa, Harimau, Mensirip, Babi Besar, Babi Hujong, Simbang, Sibu, Sibu Tengah, Sibu Hujong, Sibu Kukus, Tokong Belalai, Tioman, Tulai, Pemanggil, Aur, Dayang Dibamidae Dibamus tiomanensis*Tioman, Tulai Typhlopidae Ramphotyphlops albiceps Tioman Ramphotyphlops braminus Aceh, Tunas Selatan, Babi Besar, Sibu, Sibu Tengah, Tioman, Aur, Dayang Pythonidae Python reticulatus Tioman, Tulai Colubridae Ahaetulla prasina Babi Besar, Tinggi, Tioman, Tulai, Aur Asthenodipsas vertebralis Tioman Boiga cynodon Tioman Boiga dendrophila Babi Besar, Sibu Boiga drapiezii Tinggi, Tioman Boiga nigriceps Tioman, Aur Calamaria ingeri*Tioman Clamaria lovii Aur Calamaria lumbricoidea Tioman Calamaria pavimentata Tioman Cerberus rynchops Seribuat, Tioman Chrysopelea paradisii Sibu Tengah Chrysopelea pelias Tioman Dendrelaphis caudolineatus Aceh, Babi Besar, Tinggi, Sibu, Sibu Tengah, Tioman Dendrelaphis cyanochloris Tinggi, Tengah, Tioman Dendrelaphis pictus Aceh, Mawar, Babi Besar, Tioman Dendrelaphis striatus Tioman Dryocalamus subannulatus Tinggi, Sibu, Tioman Dryophiops rubescens Tioman Elaphe flavolineata Tioman Elaphe taeniura Tioman Enhydris enhydris Tioman

161 Grismer et al.: Herpetofauna of Seribut Archipelago

Table 2. Island checklist of species in the Tioman Archipelago. Asterisk indicates the species is endemic. (Continued)

Enhydris plumbea Tioman Fordonia leucobalia Tioman Gongylosoma mukutense*Tioman Gonyosoma oxycephalum Tioman Lepturophis albofuscus Tioman Liopeltis tricolor Tioman Lycodon capucinus Rawa, Babi Besar, Babi Hujong Oligodon booliati*Tioman Oligodon purpurescens Tioman Psammodynastes pulverulentus Tioman Pseudorabdion longiceps Sibu Rhabdophis chrysargos Tioman Sibynophis melanocephalus Tioman Zaocys carinatus Tioman Elapidae Bungarus flaviceps Tioman intestinalis Tioman Ophiophagus hannah Tioman Viperidae Popeia sp. Tioman Tropidolaemus wagleri Aceh

Mangroves (0 m). – Mangrove swamps are usually disjunctly Ridge forest (hill top summits between 950–1035 m). – distributed in thin belts along many of the islands’ coastlines. Ridge forest occurs on summits where mosses, ferns, lichens, Characteristic plant species include Bakan Minyak and bryophytes predominate. Due to increased exposure to (Rhizophora apiculata), Tumu (Bruguiera gymnorhiza), sun and wind, trees are relatively short. At this altitude, Buta-buta (Excoecaria agallocha), and Api-api Putih species such as Garcinia penangiana, Licuala tiomanensis, (Avicennia alba) which, in some localities on P. Tioman and and Scleria sumatrensis dominate and presumably have P. Mawar, are unusually tall with large girth, attesting to the adapted to live in the damp, wind-blown environment typical antiquity of the grove. of ridge forests.

Coastal vegetation (0–80 m). – Coastal vegetation forms a relatively narrow zone between the mangrove swamps (when GEOGRAPHY OF THE SERIBUAT ARCHIPELAGO present) and the lower reaches of the lowland dipterocarp forest. It is characterized by palms, Screw Pine (Pandanus The environmental diversity of the Seribuat Archipelago is dubius), and moderately-sized trees such as Healing Plant unparalleled by any other island groups off the coast of (Scaevola taccada), Beauty Leaf (Calophyllum inophyllum), peninsular Malaysia. This is paramount to the generation and Arabian Lilac (Vitex trifolia). Dipterocarp trees are and maintanence of the herpetofaunal diversity and its broad noticeably absent. array of adaptive types. The large islands (> 4.0 km2) such as Aur, Babi Besar, Pemanggil, Tinggi, Tioman, and Sibu Lowland dipterocarp forest (usually 80–300 m). – Lowland generally maintain extensive tracts of primary dipterocarp dipterocarp forest occurs on the alluvial slopes between forest (Fig. 2). The largest of these islands, Tioman and coastal vegetation and hill dipterocarp forest and is usually Tinggi, also maintain significant sources of permanent fresh dominated by large, non-dipterocarp trees such as Kabong water in the form of streams (Fig. 3), which harbor aquatic Tukas (Arenga pinnata), Fishtail Palm (Caryota mitis), and and riparian species. Pulau Tioman even has a river system, Feather Palm (Nenga macrocarpa). A few large dipterocarp S. Mentawak (Fig. 3), which serves as the only locality in species such as Mersawa Kuning (Anisoptera curtisii) and the archipelago for the frog Rana picturata. Conversely, Chengal (Neobalanocarpus heimii) exist as emergents. many of the very small islands (> 0.03 km2) such as the “Tokongs” are little more than wind-blown, arid, rocky Hill dipterocarp forest (300–950 m). – This zone is situated outcrops rising precipitously out of the ocean which support immediately above and adjacent to the lowland dipterocarp only grasses and low-growing shrubs (Fig. 4). But here too, forest with which it is continuous. The transition from potentially endemic species have been found which occur lowland dipterocarp forest to hill dipterocarp forest at nowhere else in the archipelago (e.g., Eutropis cf. approximately 300 m is almost imperceptible and many plant multifasciata on P. Tokong Bahara, see below). The species common to the lowland dipterocarp forest occur at intermediate-sized islands may be barren (P. Layak), forested lower elevations in the hill dipterocarp forest. Hill dipterocarp (P. Babi Hujong), or combinations of both such as in the forest is dominated throughout by the larger dipterocarp Seribuat group (Fig. 5) and may or may not support ephemeral species of the genera Shorea and Dipterocarpus. bodies of fresh water. Many of the islands in the archipelago,

162 THE RAFFLES BULLETIN OF ZOOLOGY 2006 both large and small, are fringed with broad stands of The Inner Arc mangroves (Fig. 6). The westernmost set of islands is referred to here as the Inner Arc (Fig. 7). This arc contains 13 islands lying immediately The islands of the Seribuat Archipelago are landbridge islands offshore in water less than 10 m deep. It extends 45 km from which formerly comprised a series of at least three low P. Layak in the north to P. Belanak in the south. The three mountain ranges paralleling the more extensive ranges largest islands of this arc, P. Aceh, P. Mawar, and P. Setindan, running the length of the adjacent Malay Peninsula. The represent eastward extensions of prominent points of land summits of these ranges became transformed into a series of which have become separated from the adjacent Malay three, parallel island arcs (Fig. 7) following the most recent Peninsula. Pulau Aceh in the north (1.32 km2; 150 m) is rise in sea level which inundated the Greater Sunda Shelf 8– flanked by the smaller satellite islands of Anak Aceh, Layak, 18,000 years before present (Inger & Voris, 2001; Voris, Tunas Selatan, Tunas Utara, Tengah, Pelandok, and Lalang 2000). Thus, it is likely that much of the flora and fauna of (Fig. 1). Together with P. Aceh, these islands are collectively these islands had a vicariant origin, being effectively isolated referred to as the Aceh group. Anak Aceh, Lalang, Tunas on these summits prior to their isolation by rising sea levels. Selatan, and Tunas Utara are small, rocky islands (> 0.24 This isolation, coupled with genetic drift and the new selection km2) reaching no more than 69 m in elevation (Table 1). They pressures attendant with living on islands, likely promoted have small patches of beach habitat (P. Anak Aceh being the evolutionary change in a number of these newly isolated exception) backed by depauperate, coastal vegetation and populations. As a result, a significant portion of the flora barren, grassy, wind-blown interiors. Pulau Pelandok and P. and fauna in this archipelago are endemic (see Sodhi et al., Tengah are steep-sided and maintain healthy tracts of coastal 1999). vegetation and lowland dipterocarp forest. Pulau Mawar (0.33 km2; 72 m) lies 4.2 km south of P. Aceh. It is a low, elongate The Seribuat Archipelago is divisible into three island arc island supporting extensive stands of mangroves along its systems corresponding to the different mountain ranges to coastline and a thick primary coastal forest in its interior. which each of these islands belonged prior to the mid- Pulau Lalang (0.02 km2; ~10 m), 6.7 km to the south, is a Pleistocene rise in sea level (Voris, 2000). These island arcs small rocky island lacking beach habitat and supporting only are separated from one another by a sequence of marine low-growing shrubs in its interior. The Setindan group is valleys whose depths increase eastwardly, indicating that the composed of the larger P. Setindan (3.8 km2; 105 m) and the outer or easternmost arc of islands is the oldest (i.e., the first much smaller P. Batu Gajah (0.04 km2; ~60 m). Pulau to be formed) and the inner or westernmost arc of islands is Setindan has a system of beaches running nearly the entire the youngest (i.e., latest to be formed). length of its eastern side and its interior maintains coastal

Fig. 2. Upper.–Lowland dipterocarp forest on P. Tioman. Lower.– Fig. 3. Upper–Sungai Mentawak on P. Tioman. Lower.–Small Hill dipterocarp forest on P. Tioman. stream in closed canopy forest on P. Tioman.

163 Grismer et al.: Herpetofauna of Seribut Archipelago

and lowland dipterocarp forest fringed by groves of coconut palms. Much of the low-lying portion of the forest floor has been disturbed by pigs. Pulau Batu Gajah is little more than a large rock with a small west-facing beach backed by fragmented coastal vegetation. The southernmost island of the Inner Arch is Belanak, a small (0.01 km2; 50 m) rocky island 16 km south of P. Setindan. It has a steep interior with coastal vegetation cover and a rocky shoreline lacking beaches.

The Middle Arc Lying approximately 12 km east of the Inner Arc is the central set of islands referred to here as the Middle Arc (Fig. 7). This arc contains 35 islands lying in a water depth of 10–19.5 m. It extends 79 km from P. Sembilang and P. Seribuat in the north to P. Tokong Yu in the south (Fig. 1). The islands of this arc show great diversity in size, geological make-up, and vegetative structure and are divisible into six separate island groups. Each group is composed of a large, usually centrally located island flanked by a number of much smaller satellite islands.

The Seribuat group is the northernmost (Fig. 1). The two largest islands of this group, Sembilang and Seribuat are separated by a shallow, mangrove-lined channel less than 800 m wide (Fig. 5). The largest island, P. Seribuat (5.52km2; 124 m), is steep-sided, rocky, and volcanic in composition. Much of its interior is arid and covered with scrub vegetation on exposed slopes although a number of deeper canyons maintain dense lowland and coastal forest. The hilly eastern portion of the island is separated from the western low-lying section by an estuary lined with an extensive mangrove system. Pulau Sembilang (2.51 km2; 207 m) is circular and approximately 2 km in diameter. Like P. Seribuat, some of its interior is covered with low scrub and recently cut secondary forest. Both islands maintain ephemeral freshwater streams which support small permanent pools during the dry season. Wood et al. (2003b) reported on the herpetofauna of these islands.

Sembilang and Seribuat are flanked to the southwest by the small satellite islands Mertang Barat (Achi), Mertang Tengah, and Mertang Timur and to the east by P. Tasu. The Mertang

Fig. 4. Upper.–Pulau Tokong Bahara. Lower.–Pulau Tokong Chupak looking northward towards P. Tinggi in the far background. Right.– Pulau Tokong Burung Condong looking northward towards Pulau Tokong Burung Besar.

164 THE RAFFLES BULLETIN OF ZOOLOGY 2006 islands are small (> 0.04 km2; > 44 m), rocky islands with small beaches edged by coastal vegetation (Fig. 5). The interiors are exposed and grassy except for Pulau Mertang Barat which maintains a low coastal forest. Pulau Tasu (0.04 km2; 70 m) is a steep-sided island lacking a sandy beach and coastal vegetation. Its interior is composed of tall grass, low shrubs, and scattered rocky outcrops.

Twenty kilometers southeast of the Seribuat group is a string of four, small volcanic islands referred to here as the Rawa

Fig. 5. Upper.–Pulau Seribuat looking westward towards Pulau Sembilang. Lower.–Pulau Mertang Barat looking southward towards Pulau Mertang Tengah (middle) and Pulau Mertang Timur Fig. 6. Upper.–Mangrove root system on Pulau Sibu. Lower.– (background). Mangrove forest at Telok Nipah, Pulau Tioman.

165 Grismer et al.: Herpetofauna of Seribut Archipelago group. This group extends 6 km from P. Harimau in the north which comprise the Tinggi group (Fig. 1). Pulau Tinggi, the to P. Rawa in the south (Fig. 1). All are steep-sided islands dominant island, is one of the largest islands of the Seribuat with rocky, volcanic bluffs punctuated by narrow, sandy Archipelago. It is approximately 6.4 km long and 3.4 km beaches. Two of these islands, P. Harimau (0.21 km2; 91 m) wide (13.5 km2). The centre of the island is dominated by a and P. Gual (0.09 km2; 56 m), are covered with grasses and 610 m high conical volcanic peak from which the island gets low-growing shrubs. Pulau Mensirip (0.08 km2; 32 m) has its name. Pulau Tinggi is mountainous and crossed by several a long, low profile with the coastal forest on the western slope small streams. Its coastline has wide stretches of sandy opening up onto a narrow beach. Pulau Rawa (0.3 km2; 97 beaches, mangroves, and rocky caves, but unlike many other m) is the southernmost and largest island of the group. It is islands, outcroppings of granite boulders are absent. With 1.8 km in length and dominated by a central ridge whose the exception of minor cutting on the lower periphery of the western slope is well forested, giving way to a wide beach. west coast, the island harbors large tracts of closed-canopy, Much of its coastal vegetation has been disturbed for the lowland dipterocarp and bamboo forests (Turner et al., 1993a). construction of tourist lodges. Its peak is dominated by arid-adapted grasses (Turner et al., 1993a). Escobar et al. (2003b) reported on the herpetofauna Four kilometers southwest of the Rawa group lies a chain of of this island. three islands referred to here as the Besar group (Fig. 1). The geological composition of these islands is granitic and all Flanking the eastern and southern shores of P. Tinggi are maintain large tracts of lowland dipterocarp forest and eight satellite islands of varying size. The islands of Apil, numerous small beaches. The largest of these islands, P. Babi Mentigi, Nangka Kecil, and Nangka Besar edge the southern Besar (6.38 km2; 251 m), is 5 km in length and dominated by end of P. Tinggi. Pulau Nangka Besar (0.1 km2; 33 m) and a steep-sided ridge extending the length of the island that P. Nangka Kecil (0.04 km2; ~15 m) are the most prominent opens up onto alluvial beaches. The island’s interior is of this cluster. Pulau Nangka Besar is a low, forested hill covered with primary and reinvaded secondary forest with a ringed by granitic boulders whereas P. Nangka Kecil is more few granitic boulder outcrops. Two large peaks occur along flat with sandy beaches and scrub vegetation. Apil (0.04 km2; its central ridge: Bukit Berot in the south, reaching 251 km 13 m) is little more than a small, forested hill and P. Mentigi and Bukit Atap Zink in the north, reaching 206 km. Wood (0.01 km2; 3 m) is low with coral beaches and scrub et al. (2004a) reported on the herpetofauna of this island. vegetation. Off the southeast end of P. Tinggi lies P. Simbang (0.07 km2; 23 m). This island has a rocky coastline punctuated Pulau Babi Tengah (0.55 km2; 83 m) and P. Babi Hujong by small beaches and an interior composed of low forest. Off (0.66 km2; 99 m) to the north are much smaller and do not its northern coast lies P. Lanting (0.01 km2; 16 m) which is reach the elevation of Pulau Babi Besar. Both maintain little more than a pile of rocks with one tree growing out of healthy tracts of primary coastal forest, lowland dipterocarp the top. Penyembong (0.04 km2; 17 m) and P. Ibol (0.15 forests in their interiors, and numerous beaches. km2; 82 m) lie off the eastern coast of P. Tinggi. Penyembong is a small hump emerging above the waterline as a pile of Sixteen kilometers southeast of the Rawa group are 10 islands boulders with a low forest. Ibol is much larger, steep-sided, and supports dense tracts of coastal forest in its interior.

The Sibu group is a chain of five islands lying 6 km southwest of the Tinggi group, extending 8 km from P. Sibu in the north southward to P. Sibu Kukus (Fig. 1). The largest island of the group, P. Sibu (4.81 km2; 155 m) is an hourglass-shaped island composed of coastal forest, coconut plantations, and scattered mangrove swamps (Turner et al., 1993b). The southern end reaches an elevation of 155 m and gives way to an extensive alluvial beach to the west dominated by coconut plantations and smaller beach scrub. Wood et al. (2004b) reported on the herpetofauna of this island.

Flanking the southern end of P. Sibu is a chain of four smaller satellite islands. Of these, P. Sibu Tengah is the largest (0.45 km2; 74 m). It has a precipitous, rocky, mangrove-lined eastern side and a steep, forested, mountainous interior that gives way to beaches on its western coast. The remaining three islands, Papan (km2; 91 m), Sibu Kukus (0.03 km2; 45 m), and Sibu Hujong (0.03 km2; 59 m) are little more than heavily forested rocky humps. Of the three, only P. Sibu Kukus has beach habitat.

Fig. 7. The location of the Island Arcs showing the 10 m and 19.5 The southernmost island group of the Middle Arc (and of the m depth contours. Island names represent the island groups as delimited in the text. Seribuat Archipelago) is the Lima group. This group lies 6

166 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Table 3. Checklist of species occurring on the island arcs. Asterisk indicates the species is endemic.

Inner Arc Middle Arc Outer Arc Amphibians Megophryidae Leptolalax kajangensis* X Megophrys nasuta X Bufonidae Ansonia tiomonica* X Bufo asper X Bufo melanostictus X Bufo parvus X Pelophryne brevipes X Microhylidae Chaperina fusca X Kalophrynus pleurostigma X Kaloula baleata XX Kaloula pulchra X Microhyla borneensis X Rhacophoridae Nyctixalus pictus X Philautus petersi X Polypedates leucomystax XX X Theloderma horridum X Ranidae Fejervarya cancrivora XX Limnonectes blythii X Rana erythraea X Rana hosii X Rana picturata X Rana raniceps X Taylorana hascheana X Ichthyophiidae lchthyophis sp. X Reptiles Trionychidae Dogania subplana XX Agamidae Acanthosaura armata X Aphaniotis fusca XX X Bronchocela cristatella XX Draco fimbriatus X Draco formosus X Draco haematopogon X Draco melanopogon XX Draco sumatranus XX Gonocephalus chamaeleontinus X Gonocephalus grandis X Gekkonidae Cnemaspis baueri* X Cnemaspis kendallii XX X Cnemaspis limi* X Cnemaspis pemanggilensis* X Cosymbotus craspedotus XX Cosymbotus platyurus XX Cyrtodactylus aurensis* X Cyrtodactylus quadrivirgatus X Cyrtodactylus seribuatensis*X Cyrtodactylus tiomanensis* X Gehyra mutilata XX Gekko monarchus XX X Gekko smithii X Hemidactylus frenatus XX X Hemiphyllodactylus typus X Lepidodactylus lugubris XX X Ptychozoon kuhli XX X Ptychozoon lionatum XX

167 Grismer et al.: Herpetofauna of Seribut Archipelago

Table 3. Checklist of species occurring on the island arcs. Asterisk indicates the species is endemic. (Continued)

Inner Arc Middle Arc Outer Arc Scincidae Dasia olivacea XX Emoia atrocostata XX Eutrophis longicaudata X Eutrophis multifasciata XX X Larutia seribuatensis* X Lipinia surda X Lipinia vittigera XX Lygosoma bowringii XX X Sphenomorphus ishaki* X Sphenomorphus sibuensis*X Sphenomorphus scotophilus X Varanidae Varanus nebulosus X Varanus salvator XX X Dibamidae Dibamus tiomanensis* X Typhlopidae Ramphotyphlops albiceps X Ramphotyphlops braminus XX X Pythonidae Python reticulatus X Colubridae Ahaetulla prasina XX Asthenodipsas vertebralis X Boiga cynodon X Boiga dendrophila X Boiga drapiezii XX Boiga nigriceps X Calamaria ingeri* X Clamaria lovii X Calamaria lumbricoidea X Calamaria pavimentata X Cerberus rynchops XX Chrysopelea paradisii X Chrysopelea pelias X Dendrelaphis caudolineatus XX X Dendrelaphis cyanochloris XX Dendrelaphis pictus XX X Dendrelaphis striatus X Dryocalamus subannulatus XX Dryophiops rubescens X Elaphe flavolineata X Elaphe taeniura X Enhydris enhydris X Enhydris plumbea X Fordonia leucobalia X Gongylosoma mukutense* X Gonyosoma oxycephalum X Lepturophis albofuscus X Liopeltis tricolor X Lycodon capucinus X Oligodon booliati* X Oligodon purpurescens X Psammodynastes pulverulentus X Pseudorabdion longiceps X Rhabdophis chrysargos X Sibynophis melanocephalus X Zaocys carinatus X Elapidae Bungarus flaviceps X Calliophis intestinalis X Ophiophagus hannah X Viperidae Popeia sp. X Tropidolaemus wagleri X

168 THE RAFFLES BULLETIN OF ZOOLOGY 2006 km south of P. Tinggi and constitutes a chain of eight islands rugged topography and its hillsides are cut by a number of beginning with P. Tokong Sanggol in the north, extending boulder-strewn, fast-flowing streams in both open and closed 16.4 km southward to P. Tokong Yu (Fig. 1). With the canopy forest (Fig. 3). The herpetofauna of this island has exception of P. Lima Besar (0.10 km2; 48 m) and P. Lima been extensively studied (see J. Grismer et al., 2004 and Kecil (0.04 km2; 41 m), these are very small (ca. 0.04 km2), references therein). volcanic, rocky, precipitous islands that rise abruptly 22–38 m out of the water (Fig. 4). They are covered with arid- The Tulai group is the northernmost cluster of islands in the adapted, low-growing, wind-swept vegetation with little soil. Outer Arc. It lies 5 km northwest of P. Tioman and is Three of these islands, Tokong Sanggol and Tokong Raket dominated by the central, large island of Tulai flanked by (Raket is two islands side by side) support large nesting three smaller satellite island, Labas and Sepoi to the colonies of terns whereas P. Tokong Chondong, P. Tokong southwest, and Cebeh to the north (Fig. 1). Pulau Tulai is a Belalai, and P. Tokong Chupak have only a few nests. The small (0.87 km2; 98 m), T-shaped island extending 1.6 km in southernmost island of this group is P. Tokong Yu. This length along an east-west axis and 1 km in width along a island was heavily bombed by the United States during World north-south axis. Its interior is hilly and rocky and lacks War II and its interior is covered with shrapnel and exploded permanent fresh water. Most of the island was cleared for bomb casings. The possible existence of unexploded coconut plantations prior to 1960 (Medway & Bullock, 1966) ordinance makes field work on this island dangerous. The but some large dipterocarp trees were left and secondary bombing changed the topography of the island significantly, growth is now extensive. Its herpetofauna was discussed by leaving it covered with large, deep craters. Grasses and small L. Grismer et al. (2001b). shrubs are the only vegetation present. Pulau Lima Besar is 2 the largest island of the group. This elongate, steep-sided Pulau Labas (0.01 km ; 20 m) is little more than a large pile island has an eroded rocky backbone with five conspicuous of granitic boulders cresting the surface of the sea. It humps. The western side of the island has a large beach essentially lacks soil and small plants and supports only a bordered by a healthy stand of coastal vegetation. Its eastern few coconut palms. Its low profile subjects it to a constant side is steep and rocky. The much smaller P. Lima Kecil to buffeting by seawater during storms. The nearby P. Sepoi (0.03 km2; 60 m) is a conical island composed of large granitic the north is ringed by a rocky coastline but maintains a boulders. It lacks beaches and its steep, rocky bluffs rise forested, steep-sided interior. precipitously out of the water. The interior of the island is composed of an open coastal forest maintaining moderately- Two maps, one published in 1996 under the superintendence sized, drought resistant trees and a deep soil base. Pulau of the Royal Malaysian Navy and another published in 1997 Cebeh (0.05 km2; 71 m) is a well-vegetated, small granitic by the Director of National Mapping, report the presence of pile of huge boulders lying just to the north of P. Tulai. Its a small island named P. Tokong Gantang between P. Tokong interior maintains a dense coastal forest with large trees and Chondong and P. Tokong Raket. We were unable to locate a deep leaf-litter and soil base. this island on either visit to the Lima Group and believe it does not exist. Pulau Pemanggil (9.2 km2; 405 m) is a smaller rocky island lying between P. Tioman and P. Aur (Fig. 1). Its rugged The Outer Arc coastline and interior are dominated by large granitic boulder The easternmost island arc in the Seribuat Archipelago is outcroppings which have created a number of extensive cave- referred to here as the Outer Arc. This arc is situated like systems. Only one small patch of primary forest remains approximately 20 km east of the Middle Arc (Fig. 7) and near the island’s summit. The remainder of the island is contains the oldest and largest group of islands in the Seribuat covered by heavily degraded secondary forest. One small Archipelago. These islands lie in water 20–64 m deep. This freshwater stream exists on the northeastern side. Youmans system is composed of 10 islands extending 75 km from P. et al. (2002) is the only published report on this island’s Cebeh in the north to P. Aur in the south (Fig. 1). The herpetofauna. topography of all the islands in this arc is dramatic, including steep interiors dominated by extensive outcroppings of huge The Aur group consists of the island Aur and its smaller granitic boulders. The coastlines are ringed by massive satellite islands, Dayang, Lang, and Pinang (Fig. 1). These granitic rocks with intermittent beaches backed by heavily are the southernmost islands in the Outer Arc. Like P. forested interiors. Pemanggil, P. Aur is smaller than P. Tioman (12.75 km2; 41 m), steep-sided, and rugged. Its interior has limited permanent The largest islands in the arc are Tioman, Pemanggil, and running water and is characterized by large granitic outcrops. Aur. Pulau Tioman, the largest of the three (110 km2; 1038 Two peaks, one at each end of the island, form the dominant m), extends approximately 22 km in length and 11 km in topographical features with the southernmost peak, Bukit maximum width at its southern end. Characterized by steep- Makum, reaching an elevation of 521 m. Much of the forest sided mountainous terrain reaching 1038 m in elevation at along the low-lying coastal periphery of P. Aur has been cut G. Kajang, the island supports lowland dipterocarp forest on and replanted with coconut palms. However, dense lowland its alluvial foothills and hill dipterocarp forest above 300 m. dipterocarp forest with closed canopy remains in the island’s Its coastline and low-lying periphery are dominated by interior on the steeper slopes at higher elevations. L. Grismer mangrove and coastal forests. Exposed granitic outcroppings et al. (2001a) and Escobar et al. (2003a) report on the consisting of large boulders define much of the island’s herpetofauna of this island.

169 Grismer et al.: Herpetofauna of Seribut Archipelago

Of the three satellite islands, P. Dayang (2.09 km2; 158 m), Biogeography. – Surveys from other archipelagos and islands located 0.08 km north of P. Aur, is the most significant. It surrounding the Malay Peninsula are still not complete. Only consists of a low range of granitic rock outcroppings with an preliminary work has been done on the Langkawi Archipelago elevated middle section consisting of a large granitic peak. (L. Grismer et al., 2006; Ibrahim et al., in press), Penang (Das The majority of P. Dayang is covered with degraded coastal and Lim, in prep) and work on Redang and Perhentian is still vegetation. Small streams occur at its northern end. Wood preliminary (Tamblyn et al., 2005.). Thus any meaningful et al. (2003c) reports on the herpetofauna of this island. comparisons of species richness between these areas and the Pinang (0.11 km2; 38 m) and Lang (0.05 km2; 38 m) are small, Seribuat Archipelago would be premature. vegetated granitic outcrops rising out of the water. The herpetofauna of the Seribuat Archipelgo is composed of There are four sets of very small (>0.05 km2), isolated, 23 species of frogs, one species of caecilian, one species of outlying islands that occur in the 20 km wide deep-water non-marine turtle, 43 species of lizards and 44 species of non- channel separating the Middle and Outer Arcs. The marine snakes. The relationships of these species are diverse. northernmost of these, the Tokong Burung group consists of Some, such as Gonocephalus chamaeleontinus, Cnemaspis three small islands (0.005–0.36 km2; 32–54 m) lying midway limi, Calamaria ingeri, Gongylosoma mukutense, and Popeia between P. Seribuat and P. Tioman (Fig. 1). These islands sp. occurring on the Outer Arc, are most closely related to are separated from each other by less than 100 m. They are Bornean, Sumatran, or Javan species (see L. Grismer et al., precipitous rocky peaks which emerge out of the ocean from 2003; 2004b) rather than species from Peninsular Malaysia, underwater sea mounts (Fig. 4). They are covered with short indicating their geographic regions of origin are from areas grasses and arid-adapted, windblown vegetation. Various much further to the south or east as opposed to just 42 km to species of terns use these islands as rookeries. The grasses the west in Peninsular Malaysia. This biogeographical pattern are periodically burned off by egg collectors in order to kill is not restricted to amphibians and reptiles but has been the hatchlings and near-term eggs. This stimulates the reported in mosses (Tan and Mohamad, 1999), a balitorid females to lay fresh eggs which the collectors gather one week loach (Kottelat, 1990), freshwater gobies (Ng, Tan and Lim, later, ensuring the eggs contain only yolk and no embryos. 1999), decapods (Yeo and Ng, 1999; Yeo et al., 1999), and Pulau Tokong Bahara (0.02 km2; 54 m) is also isolated, lying mouse deer (Adura, 1998). This indicates these species had midway between P. Seribuat and P. Tioman south of the a vicariant origin in the Outer Arc as a result of rising sea Tokong Burung group (Fig. 1). Its shape and interior are levels separating an ancient mountain system which extended like those islands of the Tokong Burung group and the grasses from the Riau Archipelago northeastward through the on this island are also periodically burned to facilitate egg Seribuat, Anambas, and Natuna archipelagos to Borneo collecting. The remaining island, P. Jahat (0.05 km2; 30 m), (Voris, 2000; Inger, pers. comm., 2003). Species in the lies 7 km south of P. Tioman (Fig. 1). It is a low pile of huge Seribuat Archipelago that are most closely related to granitic boulders supporting a low-growing forest with a peninsular Malaysian forms may also have had a vicariant modest soil base. origin but overwater dispersal cannot be ruled out.

Sixteen species in the Seribuat Archipelago are endemic: two RESULTS AND DISCUSSION species of frogs (Leptolalax kajangensis and Ansonia tiomanica); 10 species of lizards (Cnemaspis baueri, C. limi, It is certain that the results reported herein are an C. pemanggilensis, Cyrtodacylus tiomanensis, Crytodactylus underestimate of the herpetofaunal diversity of the Seribuat seribuatensis, Cyrtodactylus aurensis, Larutia seribuatensis, Archipelago and more species will be found on many of the Sphenomorphus ishaki, Sphenomorphus sibuensis and islands. We expect snakes to be the most under represented Dibamus tiomanensis); and four species of snakes (Calamaria group of our collecting efforts. Most of the islands were ingeri, Gongylosoma mukutense, Oligodon booliati, and visited only during the day (Table 1) which biased collecting Popeia sp). Two of these, Cyrtodactylus seribuatensis and towards lizards. Had we spent equal time on these islands at Sphenomorphus sibuensis, are endemic to the Middle Arc night we could have found many more snakes. Even diurnal whereas the remaining 14 species are endemic to the Outer snakes are most easily found at night while sleeping in Arc. This is in accord with the Outer Arc islands’ older age vegetation where their camouflage is ineffective in torch light. and more complex environment. With the exception of L. Another problem was that during our second collecting season seribuatensis, Calamaria ingeri, O. booliati, and Popeia sp., from 7–15 September 2004 the weather was extremely dry. the relationships, and hence geographic origins, of these No appreciable precipitation had fallen in the Seribuat species are yet unknown. Archipelago since 4 July 2004. As a result, we found fewer species than we expected on many of the larger, well- The majority of the species in the Seribuat Archipelago occur vegetated islands such as Mawar, Tengah, Tinggi, and on islands of the Outer Arc. Here, 96% of the frogs (22 Setindan. These islands still require much more intesive field species), 100% of the caecilians (one species), 88% of the research. We found no species on P. Labas, P. Lanting, and lizards (38 species), and 95% of the snakes (42 species) are P. Tokong Yu. Nonetheless, the completeness of these found (Table 3). This high diversity is maintained by the surveys provides a foundation for the following environmental heterogeneity of these islands which is in part, biogeographical and ecological observations. related to their larger size and extreme topography. This is

170 THE RAFFLES BULLETIN OF ZOOLOGY 2006 most notable on the largest island of Tioman which has 75% to those of the smaller species of Draco such as D. (96 species) of the species found in the archipelago. The melanopogon. On the islands of Babi Besar, Babi Tengah, Outer and Middle Arc share two species of frogs, one turtle, Babi Hujong, and Aceh, A. fusca occurs in unusually high 17 species of lizards, and five species of snakes (Table 3). densities. Although the forest is dense and closed canopy, Only one species of frog, nine species of lizards, and three D. melanopogan is absent from the interiors of these islands species of snakes are shared by all three arcs. Much more (D. sumatranus occurs in the coastal, coconut groves on P. will be revealed about the historical biogeography and Babi Besar). On islands where Draco and Aphaniotis both historical patterns of dispersal and colonization upon the exist (i.e. Tioman, Tinggi, Sibu), A. fusca occurs in much completion of our molecular studies. lower densities. Given the similar ecologies of A. fusca with some of the smaller species of Draco, it is likely that A. fusca MacArthur & Wilson (1967) provided a model of island competitively excludes Draco from the Besar group and P. biogeography which attempts to explain insular diversity as Aceh. a function of rates colonization and extinction based on an island’s size and distance from a continental source. Gekkonidae However, their model considers only a single continental Gekko monarchus. – Gekko monarchus expresses a modest source of origin for colonizing taxa whereas the Seribuat degree of morphological and colour pattern variation on some Archipelago has multiple continental sources as demonstrated of the small islands. On Mertang Barat, Mertang Tengah, and by the phylogenetic studies that have been done on a handfull Mertang Timur, G. monarchus is nearly melanistic with large, of species. Considering only a single source of origin will white tubercles. On P. Cebeh, G. monarchus is the only inflate species index values for islands with multiple sources species of gekko occurring in the interior of the island. Its of origins (L. Grismer, 1994b). Until the relationships of density is extremely high and it occupies every conceivable this herpetofauna are better understood, entertaining the niche (both terrestrial and arboreal). Specimens from this MacArthur & Wilson (1967) model of island biogeography population generally lack the paired, paravertebral blotches may obscure more knowledge than it would reveal. which diagnose this species and their tubercles are unusually numerous and large. Investigations of this population’s Natural History Observations. – Reported here are anecdotal morphology, colour pattern, and genetics are currently natural history observations made on a number of species at underway to determine its species status and relationship to the time of their collection. Often, insular populations of other G. monarchus. Like Hemidatlylus frenatus on P. Sepoi essentially continental species will expand the repertoire of (see below), G. monarchus appears to restrict the distribution what is considered to be their general behavior (see L. of Lepidodactylus lugubris to large boulders in the splash Grismer, 1990, 1994a, 2002; and references cited therein). zone of the intertidal. This is usually the result of adapting to a new environment with different combinations of predators, competitors, and Hemidactylus frenatus. – Hemidactylus frenatus is an resources or absences thereof. The observations here are extremely adept over-water disperser, human facilitated offered as qualitative rather than quantitative bookmarks for (Bauer, 1994) or otherwise. Many authors have documented the formulation and testing of future hypotheses. this species’ relatively recent arrival all over the world. Its colonizing abilities are illustrated well within the Seribuat Amphibia Archipelago, being that it occurs on at least 41 of the 62 Microhylidae islands. Some of these islands are little more than barren Kaloula baleata. – This species is generally active following rocks (i.e., P. Tokong Bahara) whereas others are large and periods of precipitation when individuals are found environmentally diverse (i.e., P. Tioman). Two other adept congregating in the vicinity of temporary water. During colonizing geckos, Gehyra mutilata and Lepidodactylus intermittent dry periods, this species usually resides in lugubris, also occur within the Seribuat Archipelago. The burrows, treeholes, or beneath surface objects. No rain had latter shows some interesting distribution patterns with respect fallen on P. Aceh within weeks of our first collection period to H. frenatus which suggests that on some small islands, and in 2004, no rain had fallen for two months prior to our competition between these two species may be intense. In visit. On 29 August 2003, two specimens (LSUHC 5712– the Lima group, eight small islands are inhabited by one or 13) were collected from the trunks of separate trees sitting the other of these two species (Table 2) and they basically 1.5 m above the forest floor facing head-up. During the alternate in occurrence in a north to south directional trend evening, both were pale grey and both had turned dark brown (P. Sanggol – L. lugubris; P. Lima Kecil and P. Lima Besar by morning. On 9 September 2004, a single specimen – H. frenatus; P. Tokong Raket – L. lugubris; P. Tokong (LSUHC 6954) was found in a small tree cavity filled with Condong and P. Tokong Belalai – H. frenatus; and P. Tokong putrid water 2.5 m above the ground. Chupak – L. lugubris) suggesting that whichever species colonizes the island first will eventually exclude the other Reptilia species. On P. Sepoi, this apparent phenomenon is even more Agamidae striking. Here, H. frenatus occupies every conceivable niche Aphaniotis fusca. – This species is an arboreal, diurnal lizard (terrestrial and arboreal) within the island’s interior and occurs commonly found in undisturbed closed canopy forests where in an extremely high density. Lepidodactylus lugubris, it perches on the sides of trees and vines. Its inability to however, occurs only on the periphery of the island amongst glide notwithstanding, it occupies microhabitats very similar the larger boulders in the splash zone within the intertidal

171 Grismer et al.: Herpetofauna of Seribut Archipelago areas. On P. Layak, H. frenatus is also found in the intertidal throat are yellow. The body is relatively narrow (especially areas and L. lugubris has not been found on that island. when compared to those of P. Cebeh) and original tails are Similar observations have been made on islands in the South nearly twice the length of the body. Morphological Pacific (Petren & Case, 1996). Yet on P. Mawar, a relatively investigations indicate that each one of these populations large island (Table 1) with a high degree of environmental differ significantly from one another and from mainland diversity, H. frenatus and L. lugubris can be found populations (A. Ponce, unpublished). Molecular analyses are syntopically beneath the bark of dead trees in the mangrove currently underway to determine their species status and forest. relationships.

Scincidae All the living on the three islands of the Tokong Emoia atrocostata. – Emoia atrocostata is a specialized Burung group and P. Tokong Bahara reside within the tern species found in intertidal areas, usually in the vicinity of rookeries, taking refuge in the rocks, cracks, and grass mangroves. It occurs on at least nine islands in the Seribuat hummocks. Other insular lizard populations living within Archipelago and although its distribution appears random, it seabird rookeries derive a significant portion of their resources is predictable. For this species to be present, a rocky shore from the in the form of regurgitated food and the backed by vegetation is required. The rocks must be of attracted to the nests (e.g., Barrett et al., 2005; L. moderate size (~ 1 m diamater) usually not giant boulders Grismer, 2002; Hews, 1990; Rounsevell et al., 1985). We (with the exception of P. Tulai) and the vegetation does not suspect the same is happening with Seribuat Archipelago necessarily have to be mangrove. Therefore, this species’ populations. One of us (LLG) observed a eating a absence from the large island of Tioman and others, which portion of regurgitated fish from a nest on Tokong Burung at first seems perplexing, is explainable because these islands Condong. lack the two fundamental characteristics (with the exception of a few small isolated places on the east coast at Tk. Dalam Lygosoma bowringii. – Lygosoma bowringii is another very on P. Tioman). Yet its presence on very small islands such successful overwater colonizer of the Seribuat Archipelago. as P. Sibu Kukus is easily explainable. Unlike Hemidactylus frenatus which is a renowned human commensal, the colonization of Lygosoma bowringii is most Eutropis multifasciata. – Eutropis multifasciata is a common, likely the result of natural events. This species occurs on 23 diurnal, forest-floor skink that spends much of its time basking of the 62 islands. Some of the islands are extremely small and foraging in relatively open areas with direct sunlight. In (i.e., P. Tunas Selatan) whereas others are large and diverse the Seribuat Archipelago, many populations manifest (i.e., P. Tinggi and P. Tioman). What all these islands have significant departures from this behavior and/or morphology. in common is sandy beaches. Islands with rocky shorelines On P. Cebeh and P. Sepoi, this species occurs in high densities lacking sandy beach habitats are not inhabited by L. bowringii. and is active at night. Specimens on both islands were Sandy beaches likely serve as an appropriate habitat for initial observed crawling and running through the leaf litter during colonization. Although this species occurs in the leaf-litter the evening hours. It was originally believed these lizards of some of the islands’ interiors, it is extremely common were frightened out of hiding by our lights. But with lights beneath the surface debris and flotsam found along beaches. off, lizards could be heard moving through the leaf litter. The Its short limbs, elongate body, and smooth shiny scales skinks from P. Cebeh are inordinately thick and robust and facilitate this species ability to “sand-swim” to escape capture. from a distance, do not even appear to be E. multifasciata. On several occasions we have observed this species escaping On P. Sepoi and the Tokong Burung group, E. multifasciata by burrowing into and swimming through the sand. On P. occur in the intertidal, living and foraging in and among the Ibol, L. bowringii were also found to be common above rocks and crevices. At first glance we mistook these skinks ground moving about in the leaf litter following periods of for Emoia atrocostata. Eutropis multifasciata has a relatively precipitation. modest degree of colour pattern variation in its continental forms (see Chan-Ard et al., 1999). On small islands in the Varanidae Seribuat Archipelago, this variablilty is often fixed and taken Varanus salvator. – Varanus salvator (or the tracks thereof) to extremes. On the large islands of Tioman, Aur, Tinggi, were observed on 31 islands ranging in size from 0.01–110 Sibu, and Babi Besar, E. multifasciata has the typical pattern km2. The presence of this large, top predator on small, isolated of a bronze-coloured dorsum, dark flanks with black-edged islands relatively far out at sea suggests monitors are white spots, and a cream-coloured venter. On P. Sepoi, adult periodically swimming to them because there are not enough males are uniformly grey in colour whereas females and resources on these small islands to support permanent juveniles are greyish with light green flanks bearing yellow populations of the large carnivores. We observed a 2 m spots. On P. Seribuat and P. Sembilang all skinks have a specimen on P. Tokong Belalai (0.01 km2) where the only grey dorsum and bright orange flanks. On the Tokong Burung conceivable food source were terns and their chicks and eggs. group, males and females are a uniform greenish gold. The On another small island, P. Layak (0.24 km2), we observed juveniles, however, have metallic green flanks. Skinks from the tracks of V. salvator near a large circular depression in P. Tokong Bahara are the most distinctive of all. The head, the sand where it had been digging up sea turtle eggs. We forepart of the body, and forelimbs are coppery orange; the suggest that V. salvator have knowledge of the food resources posterior of the body, hind limbs, and legs are olive green. on these islands and make periodic visits to exploit them. The posterior of the jaws are bright orange, and the chin and

172 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Adaptive types. – Another way to underscore environmental Cyrtodactylus aurensis, C. quadrivirgatus, C. tiomanensis, diversity of an area is to look at it in terms of the number of Gekko monarchus, Lepidodactylus lugubris, and adaptive types present (as opposed to a simple tally of the Sphenomorphus scotophilus). One nearly limbless species, number of species). Logically, regions with high Dibamus tiomanensis, barely surpasses 4 cm in length and environmental diversity will support a greater range of species spends most of its time underground eating the eggs of ants manifesting a broader array of adaptive types. In contrast, and termites. Another nearly limbless species, Larutia more homogeneous environments will have less diversity and seribuatensis, forages through the leaf litter. There is a fewer adaptive types. The relatively high species diversity montane, leaf litter specialist restricted to G. Kajang on P. in the Seribuat Archipelago coupled with its environmental Tioman (Sphenomorphus ishaki) and one all-female species heterogeneity lends itself well to this type of qualitative of gecko (Lepidodactylus lugubris) found on 13 islands that interpretation. The observations below are restricted to spontaneously lays eggs, producing genetic clones. Lastly, Seribuat Archipelago populations. there are general terrestrial species that exist in a range of habitats (Eutropis multifasciata, Lygosoma bowringii, and The herpetofauna of the Seribuat Archipelago contains a Varanus nebulosus). number of species with diverse ecologies and adaptations as well as many species that are microhabitat specialists. Most Snakes show an equally impressive diversity of adaptive of the frogs, (Megophrys nasuta, Bufo parvus, Chaperina types. Many are arboreal shrub and trunk-specialists (diurnal fusca, Kalophrynus pleurostigma, Kaloula baleata, Microhyla and nocturnal) that prey on lizards and frogs (Ahaetulla borneensis, and Taylorana hascheana) are forest floor prasina, Boiga, Chrysopelea, Dendrelaphis, Dryocalamus species. Of these, some are highly specialized. Megophrys subannulatus, Dryophiops rubescens, Lepturophis nasuta is a leaf-mimic; C. fusca breeds in tree and root holes albofuscus, Psammodynastes pulverulentus, Popeia sp., and and palm fronds; and K. baleata often resides in tree holes. Tropidolaemus wagleri). Ahaetulla prasina even has evolved Leptolalax kajangensis and Ansonia tiomanica are principally a concave, elongate, pointed snout and horizontal pupils that cave dwelling, cloud forest inhabitants; whereas Bufo asper, allow it to see forward. Two species are confined to the Rana hosii, and Rana picturata are riparian species. The four interface between land and sea (Cerberus rynchops and species of rhacophorids on the island are arboreal, of which Fordonia leucobalia). Other species (Calamaria and Nyctixalus pictus and Theloderma horridum are tree-hole Pseudorabdion longiceps) are small, leaf-litter semi-fossorial breeders. Additionally, T. horridum is a tree bark mimic and specialists that emerge only at night to feed on earthworms Philautus petersi only occurs in the cloud and ridge forests. (Stuebing and Inger, 1999). Conversely, Python reticulatus One species, Fejervarya cancrivora, can live and breed in reaches 6 m in length and eats mouse deer and monkeys. brackish water along the coast and eats crabs. Chrysopelea launch themselves from branches, expand their rib cages, and glide for considerable distances to the next All frogs have toxic skin secretions but the secretions of some tree. Dietary specialists include an estuarine species, (Ansonia tiomanica, Rana picturata, and Rana hosii) can kill Fordonia leucobalia, that preys on mud lobster and crabs; a other frogs when kept in close proximity. Rana hosii has montane species, Asthenodipsas vertebralis, that eats slugs secretions that can even be dangerous to humans. and pulls snails out of their shells; and two fossorial species, Kalophrynus pluerostigma uses its extremely adhesive skin Ramphotyphlops albiceps and R. braminus, which eat only secretions as a defense mechanism. the larvae of ants and termites. The genus Oligodon specializes in eating primarily reptile eggs by slicing them Many of the archipelago’s arboreal lizards (nocturnal and open with an enlarged, razor-sharp tooth on each side of the diurnal) live almost exclusively on trees, coming down to upper jaw. the ground only to nest or get from one tree to the next, the exception of course being the gliding species of Draco and Ptychozoon. These include Acanthosaura armata, Aphaniotis CONSERVATION fusca, Broncochela cristatella, all five species of Draco, Gonocephalus chamaeleontinus, Gehyra mutilata, Gekko Although the majority of the Seribuat Archipelago is smithii, Hemiphyllodactylus typus, Dasia olivacea, and uninhabited, all the larger islands support a rapidly growing Lipinia vittigera. One species of lizard, Gonocephalus tourism industry which is largely inspired by their natural grandis, is a riparian specialist which jumps from rocks and beauty. The need to responsibly manage these islands’ trees into the water to escape. Other species live only on biodiversity and maintain their ecotourism value is apparent; vegetation and rocks in the intertidal zone (Cyrtodactylus but this can only be achieved with an informed understanding seribuatensis; Emoia atrocostata). One small species of of their flora and fauna. The basis for this understanding gecko (Gehyra mutilata) has tear-away skin that remains in will ultimately come from field studies that augment the predator’s grasp while the lizard runs off and yet another university and museum collections with adequate series of species (Varanus salvator) inhabits coastal waterways, specimens and the careful examination and study of those reaches nearly 3 m in length, and is one of the archipelago’s collections. These specimens serve as permanent records of top predators. All but one species of Cnemapsis, C. kendallii, what was on the islands at a particular time, and where these are specialized for living on the flat surfaces of large boulders. species occurred. Molecular studies, which is the next phase Many other species are generally scansorial and are found of our analysis, will identify unique, insular, gene pools, on rocks and vegetation (i.e., Cnemaspis kendallii, advance our knowledge of the history of this faunas

173 Grismer et al.: Herpetofauna of Seribut Archipelago colonization and subsequent inter-island dispersal patterns, Das, I. & L. J. Lim, 2000. A new species of Cyrtodactylus (Sauria: and document their phylogenetic relationships and centres Gekkonidae) from Pulau Tioman, Malaysia. The Raffles Bulletin of geographic origin. of Zoology, 48: 223-231. Day, M., 1990. Zoological research. Pp. 25-43 in: Day, M. and T. Mowbray (eds.). University of Bristol Tioman Archipelago ACKNOWLEDGEMENTS Expedition, Peninsular Malaysia, 1998, Final Report. Unpublished. Many people have made contributions to this study in a Diaz, R. E., T.-M. Leong, L. L. Grismer, & N. S. Yaakob, 2004. A myriad of ways. We wish to thank officials of the new species of Dibamus (: Dibamidae) from West Environmental Planning Unit, Prime Minister’s Department Malaysia. Asiatic Herpetological Research, 10: 1-4. for permission to do research in Malaysia and for the issuance Escobar, R. A. III, J. Castro, D. Morgan, S. M. Hover, T. R. Szutz, of a research permit (40/200/19 SJ.1105). We would like to K. McCloskey, & R. Gregory, 2003a. Additions to the thank all the participants of the 2001, 2002, 2003, 2004, and herpetofauna of Pulau Aur, Johor, West Malaysia. Hamadryad, 2005 Tropical Field Biology classes from La Sierra University 27: 287-288. who worked hard, made significant discoveries, and Escobar, R. A. III, J. L. Grismer, T. M. Youmans, P. L. Wood, S. contributed to the utility and completeness of this study. For D. Kendall, J. Castro, T. Magi, C. Rasmussen, T. R. Szutz, S. comments on early drafts of the manuscript we thank I. Das, M. Hover, D. Morgan, C. Raynor, K. McCloskey, N. Izvernari, Norsham, Y., R. F. Inger, T. M. Leong, K. K. P. Lim, B. A. Hunter, J. M. Bernard, N. Hinojosa,, T. Dyer, J. Anlauf, J. Martinez, S. Andreiko, R. Gregory, L. S. Yeen, W. Wuertz, H. Jones, S. Wight, R. Johnson, and J. Sukumaran. We are Kaiser, & L. L. Grismer, 2003b. First report on the herpetofauna indebted to B. L. Lim, I. Das, T. M. Leong, M. Ryan, N. of Pulau Tinggi, Johor, West Malaysia. Hamadryad, 27: 259- Elias, J. Senawi, S. Yeen, J. Sukumaran, J. McGuire, W. 267. Grossmann, F. Tilack, J., B. Jones, A. Ponce, R. Gregory, B. Escobar, R. A. III, J. L. Grismer, & T. M. Youmans, 2003c. Jones, N. Triska, and M. and J. van Rooijen for their help in Kalophrynus pleurostigma. Geographic distribution. the field. For accommodations on Pulau Tioman, logistic Herpetological Review, 33: 317-318. support, and kindness we are indebted to the family of Wak Grismer, J. L., R. A. Escobar III, & T. M. Youmans, 2003a. and Zainah Rahim. This research was supported in part by a Theloderma horridum. Geographic distribution. Herpetological Franklin grant from the American Philosophical Society, a Review, 33: 22. College of Arts and Sciences grant from La Sierra University Grismer, J. L., T.–M. Leong, & Y. Norsham, 2003b. Two new awarded to LLG and Rychman grants from La Sierra Sundaland skinks of the genus Larutia and their phylogenetic University awarded to JLG, PLW and TMY. relationships. Herpetologica, 59: 554-566. Grismer, J. L., C. Rasmussen, & J. M. McGuire, 2003c. Sibynophis melanocephalus. Herpetological Review, 34: 266. LITERATURE CITED Grismer, J. L., L. L. Grismer, I. Das, Y. Norsham, B. L. Lim, T.– M. Leong, T. M. Youmans, & H. Kaiser, 2004. Species diversity Adura, M. A., 1996. Mitochondrial DNA variation of Malaysian and checklist of the herpetofauna of Pulau Tioman, Peninsular Chevrotans (Tragulus sp.). Unpublished Ph.D. dissertation. Malaysia with a preliminary overview of habitat utilization, Universiti Kabangsaan Malaysia. 134 pp. 2004. Asiatic Herpetological Research, 10: 244-276. Ashton, P. S., 1995. Biogeography and ecology. In: Soepadmo, E. Grismer, L. L., 1990. A new long nosed snake (Rhinocheilus & Wong, K. M. (eds.) Tree Flora of Sabah and Sarawak. Vol. lecontei) from Isla Cerralvo, Baja California Sur, México. 1. Forestry Research Institute Malaysia, Sabah Forest Proceedings of the San Diego Society of Natural History, 4: 1- Department and Sarawak Forest Department, Kuala Lumpur. 7. Pp. XLIII–LI. Grismer, L. L., 1994a. Three new side-blotched lizards (genus Uta) Barrett, K., W. B. Anderson, D. A. Wait, L. L. Grismer, G. A. Polis, from the Gulf of California. Herpetologica, 50: 451-474. and M. D. Rose, 2005. Marine subsidies alter the diet and abundance of insular and coastal lizard populations. Oikos 109: Grismer, L. L., 1994b. Geographic origins for the reptiles on islands 145–153. in the Gulf of California, México. Herpetological Natural History, 2: 17-40. Bauer, A. M., 1994. Familia Gekkonidae (Reptilia, Sauria). Part I Australia and Oceania. Pp. 1–306 in: Wermuth, H and M. Grismer, L. L., 2002. The Amphibians and Reptiles of Baja Fischer (eds.). Das Tierreich. The Animal Kingdom. New York. California, its Pacific Islands, and the Islands in the Sea of Cortés, México. The University of California Press, Berkeley, Bullock, J. A. & L. Medway, 1966. Observations of the fauna of California. 785 pp. Pulau Tioman and Pulau Tulai. 1. General introduction. Bulletin of the National Museum, Singapore 34: 1–8. Grismer, L. L., 2005a. A New Species of Bent-Toed Gecko (Cyrtodactylus Gray 1827) from Pulau Aur, Johor, West Chan-Ard, T., W. Grossmann, A. Gumprecht, & K.-D. Schulz, 1999. Malaysia. Journal of Herpetology, 39: 424-432. Amphibians and Reptiles of Peninsular Malaysia and Thailand. An Illustrated Checklist. Bushmaster Publications, Wuerselen, Grismer, L. L. 2005b. Tioman Archipelago: evolution’s natural Germany. 240 pp. laboratory. Malaysian Naturalist, 59: 12-23. Das, I. & L. L. Grismer, 2003. Two new species of Cnemaspis Grismer, L. L. 2006a. The Amphibians and Reptiles of the Tioman Strauch, 1887 (Squamata: Gekkonidae) from the Seribuat Archipelago. Malaysia. Forestry Department of Pahang. Kuala Archipelago, Pahang and Johor states, West Malaysia. Lumpur, Malaysia (in press). Herpetologica. 59: 546–554.

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Grismer, L. L. 2006b. Two new species of skinks (Genus Kottelat, M., 1990. New species and populations of cave Sphenomorphus Fitzinger 1843) from the Seribuat Archipelago. nemacheilines in South and South-east Asia (Osteichthyes: Herpetological Natural History, 12: 124-133. Balitoridae). Memiors Biospéologia, 17: 49-55. Grismer, L. L., J. A. McGuire, R. A. Sosa, & H. Kaiser, 2002. Latiff, A., I. F. Hanum, A. Z. Ibrahim, M. W. K. Goh, A. H. B. Loo, Revised checklist and comments on the terrestrial herpetofauna & H. T. W. Tan., 1999. On the vegetation and flora of Pulau of Pulau Tioman, Peninsular Malaysia. Herpetological Review, Tioman, Peninsular Malaysia. Raffels Bulletin of Zoology 33: 26-29. Supplement Number 6, 11-72. Grismer, L. L., N. S. Yaakob, B. L. Lim, T.–M. Leong, I. Das, R. Leong, T.-M., 2000. Limnonectes hascheanus. Geographic A. Sosa, J. L. Grismer, K. M. Crane, R. E. Diaz, S. V. Figueroa, distribution. Herpetological Review, 31: 182. C. A. Ledbetter, S. C. Newbold, S. R. Newbold, C. P. Patel, J. Leong, T.-M., L. L. Grismer, & Mumpuni, 2003. Preliminary Castro, R. A. Escobar III, S. Guerrero, J. W. Pinedo, J. Trujillo checklists of the Anambas and Natuna Islands (South China Sea). & H. Kaiser, 2001a. First report on the herpetofauna of Pulau Hamadryad, 27: 165-174. Aur, Johor, West Malaysia. Hamadryad, 26: 322–325. Leong, T.-M. & L. L. Grismer, 2004. A new species of Kukri Snake, Grismer, L. L., N S. Yaakob, B. L. Lim, T.–M. Leong, I. Das, R. A. Oligodon (Colubridae) from Pulau Tioman, West Malaysia. Sosa, J. L. Grismer, K. M. Crane, R. E. Diaz, S. V. Figueroa, C. Asiatic Herpetological Research, 10: 12-16. A. Ledbetter, S. C. Newbold, S. R. Newbold, C. P. Patel, J. Castro, R. A. Escobar III, S. Guerrero, J. W. Pinedo, P. Jones, Leong, T. M. & K. M. Crane, 2001. Nyctixalus pictus. Geographic & H. Kaiser, 2001b. Report of the herpetofauna of Pulau Tulai, distribution. Herpetological Review, 33: 62. West Malaysia. Hamadryad, 26: 341-342. Leong, T. M. & H. H. Tan, 2000. Enhydris enhydris. Geographic Grismer, L. L., T. M. Youmans, P. L. Wood, Jr., A. Ponce, S. B. distribution. Herpetological Review, 31: 114. Wrught, B. S. Jones, R. Johnson, K. L. Sanders, D. J. Gower, Lim, K. K. P. & L. J. Lim., 1999. The terrestrial herpetofauna of Norsham S. Y., and K. K. P. Lim, 2006. Checklist of the Pulau Tioman, Peninsular Malaysia. Raffles Bulletin of Zoology herpetofauna of Pulua Langkawi, Malaysia, with comments on Supplement Number 6, 131-156. Taxonomy. Hamadryad, 30: 61-74. MacAurthur, R. H. & E. O. Wilson, 1967. The Theory of Island Grismer, L. L., I. Das, & T.–M. Leong, 2003. A new species of Biogeography. Princeton University Press. Princeton. 203 pp. Gongylosoma (Squamata: Colubridae) from Pulau Tioman. Medway, L. & J. A. Bullock, 1966. Observations of the fauna of Herpetologica, 59: 567-574. Pulau Tioman and Pulau Tulai. 1. Bulletin of the National Grismer, L. L., J. L. Grismer, & T. M. Youmans, 2004a. A new Museum, Singapore 34: 1-158. species of Leptolalax (Anura: Megophryidae) from Pulau Ng, H. H., H. H. Tan & K. K. P. Lim, 1999. The inland fishes of Tioman, West Malaysia. Asiatic Herpetological Research, 10: Pulau Tioman, Peninsular Malaysia. Raffles Bulletin of Zoology 8-11. Supplement Number 6, 169-188. Grismer, L. L., H. Kaiser, & N. S. Yaakob., 2004b. A new species Petren, K. & T. J. Case, 1996. An experimental demonstration of of reed snake of the genus Calamaria H. Boie, 1827 from Pulau exploitative competition in an ongoing invasion. Ecology, 149: Tioman, Pahang, West Malaysia. Hamadryad, 28: 1-6. 118-132. Grismer, L. L. & I. Das, 2006. A new species of Cnemaspis Rounsevell, D. N. Brothers, & M. Holdsworth, 1985. The status (Squamata: Gekkonidae) from Pulau Pemanggil, Johor, West and ecology of the Pedra Branca Skink Pseudoemoia Malaysia. Herpetological Natural History, 12: 1-7. palfreymani. Pp. 477-480 in Grigg, C. G., R. Shine, & H. F. W. Hendrickson, J. R., 1966a. Observations of the fauna of Pulau Ehmann (eds.) The Biology of Australian Frogs and Reptiles. Tioman and Pulau Tulai. 5. The reptiles. Bulletin of the Surrey Beatty and Sons with Zoological Society. New South National Museum, Singapore 34: 53-71. Wales, Sydney. Hendrickson, J. R., 1966b. Observations of the fauna of Pulau Sanders, K. L., A. Malhotra, & R. S. Thorpe, 2005. Combining Tioman and Pulau Tulai. 6. The amphibians. Bulletin of the molecular, morphological and ecological data to infer species National Museum, 34: 74-84. boundaries in a cryptic tropical viper. Biological Journal of the Hews, D. K., 1990. Resource, defense, sexual selection, and sexual Linnean Society (in press). dimorphism in the lizard Uta palmeri. Ph.D. dissertation. Sodhi, N. S., H. S. Yong, & P. K. L. Ng, 1999 (eds.). The University of Texas, Austin. Biodiversity of Pulau Tioman, Peninsular Malaysia. Raffles Hien, P., W. Grossmann, & C. Schäfer, 2001. Beitrag zur Kenntnis Bulletin of Zoology Supplement Number 6, 288 pp. der landbewohnenden Reptilienfauna von Pulau Tioman, West- Stuebing, R. B. & R. F. Inger, 1999. A Field Guide to the Snakes Malaysia. Sauria, 23: 11-28. of Borneo. Natural History Publications (Borneo), Sabah, Ibrahim, J., Shahrul Anuar, M. S., Norhayati, A., Shukor, M. N., Malaysia. 254 pp. Shahriza, S., Nurul’Ain, E., Norzalipah, M. and Mark Ryan, Tamblyn, A., C. Turner, R. O’Malley, N. Weaver, T. Hughes, S. D., in press. An annotated checklist of herpetofauna of Langkawi Hardingham, & H. Roberts, 2005. Malaysia Tropical Forest Island, Kedah, Malaysia. Malayan Nature Journal. Conservation Project Report of the Perhentian Phase. Coral Cay Inger, R. F. & H. K. Voris, 2001. The biogeographical relations of Conservation, London. the frogs and snakes of Sundaland. Journal of Biogeography, Tan, B. C. & H. Mohamed, 1999. Apreliminary checklist of mosses 28: 863-891. of Pulau Tioman, Peninsular Malaysia. Raffles Bulletin of Jaman R. & A. Latiff., 1999. The pteryophyte flora of Pulau Tioman, Zoology Supplement Number 6: 73-76. Peninsular Malaysia. Raffles Bulletin of Zoology Supplement Turner, I. M., H. T. W. Tan, K. S. Chua, P. T. Chew & H. S. B. H. Number 6, 77-100. Ahmad, 1993a. A botanical reconnaissance of Pulau Tinggi, Johore. Malayan Nature Journal, 46: 157-168.

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Turner, I. M., H. T. W. Tan, P. P. Kumar, K. S. Chua & H. S. B. H. Islands are listed in alphabetical order. Significant portions of the Ahmad., 1993b. The vegetation of Pulau Sibu, Johore. Malayan large island of Tioman are currently proposed for development. Nature Journal, 46: 169-188. Therefore, the checklist of this island’s species is given for each Valdivia, C. M., L. Escobar, N. Padilla, J. Wheatly, S. Wright, S. locality from which specimens are known. The documentation for Y. Teen, and R. Rosser, 2004. Gekko monarchus. Geographic these localities can be found in J. Grismer et al. (2004) and references distribution. Herpetological Review, 35: 286. cited therein. van Rooijen, J., 2001. Herpetologische waarneiningen in Maleisie Pulau Aceh (1): Pulau Tioman. Lacerta, 59: 97-103. Kaloula baleata, Aphaniotis fusca, Cnemaspis kendallii, Gekko Voris, H. K., 2000. Maps of Pleistocene sea levels in Southeast monarchus, Hemidactylus frenatus, Ptychozoon kuhli, Eutropis Asia: shorelines, river systems, time durations. Journal of multifasciata, Lygosoma bowringii, Varanus salvator, Biogeography, 27: 1153-1167. Ramphotyphlops braminus, Dendrelaphis caudolineatus, Dendrelaphis pictus, Tropidolaemus wagleri Wood, P. L., N. A. Elias, & D. Morgan, 2003a. Dendrelaphis striatus. Geographic distribution. Herpetological Review, 34: Pulau Aceh Anak 264. Hemidactylus frenatus, Varanus salvator Wood, P. W., T. R. Szutz, J. L. Grismer, T. M. Youmans, & L. L. Grismer, 2003b. First report on the herpetofauna of Pulau Pulau Aur Sembilang and Pulau Seribuat, Pahang, West Malaysia. Pelophryne brevipes, Limnonectes blythii, Acanthosaura armata, Hamadryad, 27: 281-284. Bronchocela cristatella, Cnemaspis baueri, Cosymbotus platyurus, Wood, P. L., T. M. Youmans, C. Raynor, J. M. Bernard, N. Hinojosa, Cyrtodactylus aurensis, Gehyra mutilata, Gekko monarchus, T. Dyer, S. Andreiko, P. P. van Dijk, W. Wuertz, & L. S. Yeen, Hemidactylus frenatus, Dasia olivacea, Emoia atrocostata, Eutropis 2003c. First report on the herpetofauna of Pulau Dayang, Johor, multifasciata, Lipinia surda, Lipinia vittigera, Lygosoma bowringii, West Malaysia. Hamadryad, 27: 284-285. Sphenomorphus scotophilus, Varanus nebulosus, Varanus salvator, Ramphotyphlops braminus, Ahaetulla prasina, Boiga nigriceps, Wood, P. L., T. M. Youmans, & T. R. Szutz, 2003d. Elaphe Calamaria lovii flavolineata. Geographic distribution. Herpetological Review, 34: 388. Pulau Babi Besar Wood, P. L., H. Kaiser, S. Looper, T. M. Youmans, J. L. Grismer, Polypedates leucomystax, Fejervarya cancrivora, Aphaniotis fusca, & L. L. Grismer, 2004a. A first report on the herpetofauna of Bronchocela cristatella, Draco sumatranus, Cnemaspis kendallii, Pulau Besar, Johor, West Malaysia. Hamadryad, 28: 106-109. Gehyra mutilata, Gekko monarchus, Hemidactylus frenatus, Wood, P. L., T. M. Youmans, J. L. Grismer, J. Wheatly, S. Wright, Ptychozoon kuhli, Dasia olivacea, Eutropis multifasciata, Lipinia C. Valdivia, A. Ponce, L. Escobar, S. Amin, P. Baker, J. Bernard, vittigera, Lygosoma bowringii, Varanus salvator, Ramphotyphlops S. Looper, N. Marsh, L. Martin, N. Padilla, R. Rosser, A. braminus, Ahaetulla prasina, Boiga dendrophila, Dendrelaphis Srivastava, V. Srivastava, X. Wright, L. S. Yeen, H. Kaiser, & caudolineatus, Dendrelaphis pictus, Lycodon capucinus L. L. Grismer, 2004b. First report on the herpetofauna of Pulau Sibu, Johor, West Malaysia. Hamadryad, 28: 116-119. Pulau Babi Hujong Aphaniotis fusca, Cnemaspis kendallii, Gekko monarchus, Yeo, D. C. J., Y. Cai & P. K. L. Ng, 1999. The freshwater and Hemidactylus frenatus, Varanus salvator, Lycodon capucinus terrestrial decopod crustacea of Pulau Tioman, Peninsular Malaysia. Raffles Bulletin of Zoology Supplement Number 6, Pulau Babi Tengah 197-244. Polypedates leucomystax, Gehyra mutilata, Gekko monarchus, Yeo, D. C. J. & P. K. L. Ng., 1999. A new species of Geosesarma Hemidactylus frenatus, Lepidodactylus lugubris, Dasia olivacea, (Crustacea: Decopoda: Brachyura: Graspidae) from Pulau Eutropis multifasciata, Lygosoma bowringii Tioman, Peninsular Malaysia. Raffles Bulletin of Zoology Supplement Number 6, 189-196. Pulau Batu Gajah Gekko monarchus, Varanus salvator Youmans, T. M., & L. L. Grismer, 2006. Description of a new species of Cyrtodactylus (Reptilia: Squamata: Gekkomidae) Pulau Belanak from the Seribuat Archipelago, West Malaysia. Herpetological Hemidactylus frenatus Natural History, 12: 289-297. Youmans, T. M., J. L. Grismer, R. A. Escobar III, R. Johnson, & L. Pulau Cebeh L. Grismer, 2002. First report on the herpetofauna of Pulau Gekko monarchus, Eutropis multifasciata, Lepidodactylus lugubris Pemanggil, Johor, West Malaysia. Hamadryad, 27: 148-149. Youmans, T. M., P. L. Wood, & T. R. Szutz, 2003. Pareas Pulau Dayang vertebralis. Geographic distribution. Herpetological Review, Cnemaspis baueri, Gekko monarchus, Hemidactylus frenatus, 34: 38. Eutropis multifasciata, Varanus salvator, Ramphotyphlops braminus

Pulau Gual Hemidactylus frenatus APPENDIX I Pulau Harimau The following are species-checklists of all islands in the Seribuat Hemidactylus frenatus, Lygosoma bowringii, Varanus salvator Archipelago from which specimens have been recorded. The species are listed in the order of frogs, caecilians, turtles, lizards, and snakes. Pulau Ibol Cnemaspis kendallii, Gekko monarchus, Lygosoma bowringii

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Pulau Jahat Pulau Simbang Gekko monarchus, Lepidodactylus lugubris Gekko monarchus, Emoia atrocostata, Varanus salvator

Pulau Lalang Pulau Sembilang Hemidactylus frenatus Polypedates leucomystax, Fejervarya cancrivora, Aphaniotis fusca, Cnemaspis kendallii, Cyrtodactylus seribuatensis, Gekko Pulau Lalang Selatan monarchus, Hemidactylus frenatus, Ptychozoon kuhli, Eutropis Hemidactylus frenatus multifasciata, Lygosoma bowringii, Varanus salvator

Pulau Lang Pulau Sepoi Hemidactylus frenatus, Eutropis multifasciata Hemidactylus frenatus, Lepidodactylus lugubris, Eutropis multifasciata Pulau Layak Gekko monarchus, Hemidactylus frenatus, Varanus salvator Pulau Seribuat Polypedates leucomystax, Fejervarya cancrivora, Aphaniotis fusca, Pulau Lima Besar Cnemaspis kendallii, Cyrtodactlus sp. 1, Gekko monarchus, Cyrtodactylus seibuatensis, Hemidactylus frenatus, Gehyra mutilata, Hemidactylus frenatus, Emoia atrocostata, Eutrophis multifasciata, Gekko monarchus, Lygosoma bowringii Lygosoma bowringii, Varanus salvator, Cerberus rynchops

Pulau Lima Kecil Pulau Setindan Gehyra mutilata, Hemidactylus frenatus Hemidactylus frenatus, Lygosoma bowringii, Varanus salvator

Pulau Mawar Pulau Sibu Hemidactylus frenatus, Lepidodactylus lugubris, Lygosoma Microhyla borneensis, Polypedates leucomystax, Fejervarya bowringii, Varanus salvator, Dendrelaphis pictus cancrivora, Aphaniotis fusca, Bronchocela cristatella, Draco melanopogon, Cnemaspis kendallii, Cyrtodactylus seribuatensis, Pulau Mensirip Gehyra mutilata, Gekko monarchus, Hemidactylus frenatus, Lygosoma bowringii, Varanus salvator Hemiphyllodactylus typus, Lepidodactylus lugubris, Ptychozoon lionatum, Dasia olivacea, Emoia atrocostata, Eutropis multifasciata, Pulau Mentigi Lygosoma bowringii, Sphenomorphus ishaki, Varanus salvator, Cyrtodactylus seribuatensis, Gekko monarchus, Hemidactylus Ramphotyphlops braminus, Boiga dendrophila, Dendrelaphis frenatus, Lepidodactylus lugubris, Lygosoma bowringii caudolineatus, Dryocalamus subannulatus, Pseudorabdion longiceps Pulau Mertang Barat Gekko monarchus, Varanus salvator Pulau Sibu Hujong Lepidodactylus lugubris, Varanus salvator Pulau Mertang Tengah Gekko monarchus, Lygosoma bowringii Pulau Sibu Kukus Hemidactylus frenatus, Emoia atrocostata, Eutropis multifasciata, Pulau Mertang Timur Varanus salvator Gekko monarchus, Hemidactylus frenatus, Lygosoma bowringii Pulau Sibu Tengah Pulau Nangka Besar Bronchocela cristatella, Cnemaspis kendallii, Cyrtodactylus Gehyra mutilata, Hemidactylus frenatus, Emoia atrocostata seribuatensis, Hemidactylus frenatus, Emoia atrocostata, Lygosoma bowringii, Varanus salvator, Chrysopelea paradisi Pulau Nangka Kecil Cyrtodactylus seribuatensis, Hemidactylus frenatus, Emoia Pulau Tasu atrocostata Hemidactylus frenatus, Varanus salvator

Pulau Papan Pulau Tengah Lepidodactylus lugubris, Emoia atrocostata, Lygosoma bowringii Varanus salvator

Pulau Pelandok Pulau Tinggi Gekko monarchus, Hemidactylus frenatus, Varanus salvator Polypedates leucomystax, Limnonectes blythii, Rana hosii, Dogania subplana, Aphaniotis fusca, Bronchocela cristatella, Draco Pulau Pemanggil formosus, Draco melanopogon, Draco sumatranus, Cnemaspis Limnonectes blythii, Bronchocela cristatella, Cnemaspis kendallii, Cosymbotus craspedotus, Gekko monarchus, pemanggilensis, Gehyra mutilata, Gekko monarchus, Hemidactylus Hemidactylus frenatus, Ptychozoon kuhli, Dasia olivacea, Eutropis frenatus, Dasia olivacea, Eutropis multifasciata, Lygosoma multifasciata, Ahaetulla prasina, Boiga drapiezii, Dendrelaphis bowringii, Sphenomorphus scotophilus, Varanus salvator caudolineatus, Dendrelaphis cyanochloris, Dryocalamus subannulatus Pulau Penyambong Gekko monarchus, Hemidactylus frenatus Pulau Tioman Cross-island trail (near 300 m). Leptolalax kajangensis, Pulau Rawa Megophrys nasuta, Kalophrynus pleurostigma, Ansonia tiomanica, Gehyra mutilata, Hemidatylus frenatus, Eutropis multifasciata, Bufo melanostictus, Bufo parvus, Pelophryne brevipes, Chaperina Lygosoma bowringii, Varanus salvator, Lycodon capucinus fusca, Kaloula pulchra, Nyctixalus pictus, Polypedates leucomystax,

177 Grismer et al.: Herpetofauna of Seribut Archipelago

Theloderma horridum, Limnonectes blythii, Taylorana hascheana, longicaudata, Lygosoma bowringii, Varanus nebulosus, Rana raniceps, Rana hosii, Dogania subplana, Acanthosaura Ramphotyphlops braminus, Python reticulatus, Boiga cynodon, armata, Aphaniotis fusca, Bronchocela cristatella, Draco fimbriatus, Chrysopelea pelias, Dendrelaphis caudolineatus, Dendrelaphis Draco melanopogon, Draco sumatranus, Gonocephalus cyanochloris, Dendrelaphis pictus, Dendrelaphis striatus, chamaeleontinus, Gonocephalus grandis, Cnemaspis kendallii, Dryophiops rubescens, Elaphe taeniura, Gonyosoma oxycephalum, Cnemaspis limi, Cosymbotus craspedotus, Cyrtodactylus Rhabdophis chrysargos, Sibynophis melanocephalus quadrivirgatus, Cyrtodactylus tiomanensis, Gehyra mutilata, Gekko Kg. Lalang. Fejervarya cancrivora, Cerberus rynchops monarchus, Gekko smithii, Ptychozoon kuhli, Ptychozoon lionatum, Kg. Mukut. Megophrys nasuta, Bufo melanostictus, Chaperina Dasia olivacea, Eutropis multifasciata, Larutia seribuatensis, fusca, Limnonectes blythii, Aphaniotis fusca, Bronchocela Lipinia vittigera, Sphenomorphus scotophilus, Varanus nebulosus, cristatella, Draco melanopogon, Draco sumatranus, Cnemaspis Dibamus tiomanensis, Ramphotyphlops albiceps, Ramphotyphlops kendallii, Gehyra mutilata, Lygosoma bowringii, Eutropis braminus, Ahaetulla prasina, Boiga cynodon, Boiga drapiezii, Boiga multifasciata, Varanus nebulosus, Ahaetulla prasina nigriceps, Calamaria ingeri, Calamaria lumbricoidea, Calamaria Kg. Paya. Bufo melanostictus, Limnonectes blythii, Rana erythraea, pavimentata, Chrysopelea pelias, Dendrelaphis caudolineatus, Rana hosii, Draco melanopogon, Gonocephalus chamaeleontinus, Dendrelaphis cyanochloris, Dendrelaphis pictus, Dryocalamus Cnemaspis kendallii, Cyrtodactylus tiomanensis, Gekko monarchus, subannulatus, Dryophiops rubescens, Gongylosoma mukutense, Dasia olivacea, Eutropis multifasciata, Lygosoma bowringii, Lepturophis albofuscus, Liopeltis tricolour, Oligodon booliati, Varanus nebulosus, Varanus salvator, Dibamus tiomanensis, Oligodon purpurascens, Psammodynastes pulverulentus, Ramphotyphlops braminus, Python reticulatus, Dendrelaphis Rhabdophis chrysargos, Zaocys carinatus, Bungarus flaviceps, formosus, Enhydris enhydris, Enhydris plumbea, Lepturophis Calliophis intestinalis, Popeia sp. albofuscus G. Kajang (near the summit or at Gua Tengkuk Air at 980- Kg. Salang. Acanthosaura armata, Draco melanopogon, Draco 1035m). sumatranus, Gonocephalus chamaeleontinus, Gonocephalus Leptolalax kajangensis, Megophrys nasuta, Ansonia tiomanica, grandis, Cnemaspis kendallii, Cnemaspis limi, Cosymbotus Pelophryne breviceps, Kalophrynus pleurostigma, Philautus petersi, platyurus, Dasia olivacea, Eutropis multifasciata, Lipinia vittigera, Acanthosaura armata, Draco haematopogon, Draco melanopogon, Varanus nebulosus, Varanus salvator Cnemaspis kendallii, Cnemaspis limi, Cyrtodactylus quadrivirgatus, Kg. Tekek. Bufo melanostictus, Bufo parvus, Kaloula baleata, Cyrtodactylus tiomanensis, Sphenomorphus ishaki, Sphenomorphus Kaloula pulchra, Polypedates leucomystax, Fejervarya cancrivora, scotophilus, Asthenodipsas vertebralis, Ahaetulla prasina, Limnonectes blythii, Rana raniceps, Rana erythraea, Bronchocela Calamaria lumbricoidea, Pareas vertebralis, Psammodynastes cristatella, Draco melanopogon, Draco sumatranus, Cosymbotus pulverulentus, Calliophis intestinalis, Popeia sp. platyurus, Cyrtodactylus tiomanensis, Gehyra mutilata, Gekko G. Kajang (below 980m to S. Mentawak). Leptolalax kajangensis, monarchus, Hemidactylus frenatus, Lepidodactylus lugubris, Dasia Megophrys nasuta, Ansonia tiomanica, Bufo parvus, Pelophryne olivacea, Eutropis longicaudata, Lygosoma bowringii, Varanus brevipes, Chaparina fusca, Kalophrynus pleurostigma, Limnonectes nebulosus, Varanus salvator, Ramphotyphlops braminus, Python blythii, Rana picturata, Nyctixalus pictus, Acanthosaura armata, reticulatus, Boiga cynodon, Cerberus rynchops, Dendrelaphis Aphaniotis fusca, Draco melanopogon, Gonocephalus caudolineatus, Dryocalamus subannulatus, Elaphe flavolineata, chamaeleontinus, Gonocephalus grandis, Cnemaspis kendallii, Enhydris plumbea, Fordonia leucobalia Cnemaspis limi, Gekko smithii, Eutropis multifasciata, Larutia Monkey Bay. Limnonectes blythii, Acanthosaura armata, seribuatensis, Lipinia vittigera, Sphenomorphus ishaki, Aphaniotus fusca, Bronchocela cristatella, Draco fimbriatus, Draco Sphenomorphus scotophilus, Calamaria lumbricoidea, Chrysopelea melanopogon, Draco sumatranus, Cnemaspis kendallii, Cnemaspis pelias, Dryophiops rubescens, Liopeltis tricolour, Oligodon limi, Cyrtodactylus tiomanensis, Gehyra mutilata, Ptychozoon kuhli, purpurascens, Rhabdophis chrysargos, Calliophis intestinalis Dasia olivacea, Eutropis multifasciata, Lipinia vittigera, G. Rokam. Cnemaspis limi Sphenomorphus scotophilus, Varanus nebulosus, Ahaetulla prasina, Kg. Air Batang. Rana erythraea, Bronchocela cristatella, Draco Chrysopelea pelias, Dendrelaphis striatus, Elaphe taeniura, sumatranus, Cyrtodactylus tiomanensis, Gekko monarchus, Oligodon booliati Lygosoma bowringi, Varanus nebulosus, Varanus salvator, Dibamus Sedagong. Limnonectes blythii, Rana raniceps, Aphaniotis fusca, tiomanensis, Python reticulatus, Cerberus rynchops, Dendrelaphis Draco melanopogon, Draco sumatranus, Gonocephalus grandis, caudolineatus, Dryophiops rubescens, Enhydris plumbea, Eutropis multifasciata, Varanus nebulosus, Varanus salvator, Psammodynastes pulverulentus, Calliophis intestinalis, Rhabdophis chrysargos Ophiophagus hannah S. Asah (behind K. Asah). Gonocephalus grandis, Dasia olivacea, Kg. Asah. Aphaniotis fusca, Draco melanopogon, Gonocephalus Lipinia vittigera grandis, Dasia olivacea, Sphenomorphus scotophilus, Varanus S. Air Besar. Megophrys nasuta, Dogania subplana, salvator Sphenomorphus scotophilus Kg. Berjaya. Bufo melanostictus, Kaloula pulchra, Polypedates S. Air Raja (behing K. Genting). Limnonectes blythii leucomystax, Rana raniceps, Rana erythraea, Rana hosii, Aphaniotis S. Air Surin. Megophrys nasuta fusca, Bronchocela cristatella, Draco melanopogon, Draco S. Baharu. Limnonectes blythii, Dogania subplana sumatranus, Gonocephalus chamaeleontinus, Cnemaspis kendallii, S. Benuang. Chaperina fusca, Aphaniotis fusca, Draco Cnemaspis limi, Gekko monarchus, Hemidactylus frenatus, Eutropis melanopogon, Gonocephalus chamaeleontinus, Gonocephalus multifasciata, Lygosoma bowringii, Sphenomorphus scotophilus, grandis, Cnemaspis kendallii, Sphenomorphus scotophilus Varanus salvator, Ramphotyphlops braminus, Python reticulatus, S. Besar. Taylorana hascheana Ahaetulla prasina, Cerberus rynchops, Dendrelaphis caudolineatus, S. Durian Kallang (behind K. Paya). Limnonectes blythii, Rana Dendrelaphis cyanochloris, Oligodon purpurascens, hosii Psammodynastes pulverulentus, Rhabdophis chrysargos, Zaocys S. Keliling. Megophrys nasuta, Bufo parvus, Limnonectes blythii, carinatus Rana raniceps, Ichthyophis sp., Dogania subplana, Gekko Kg. Genting. Limnonectes blythii monarchus, Dendrelaphis formosus Kg. Juara. Bufo melanostictus, Bufo parvus, Polypedates S. Mentawak. Megophrys nasuta, Bufo asper, Pelophryne brevipes, leucomystax, Rana raniceps, Draco sumatranus, Cyrtodactylus Chaperina fusca, Limnonectes blythii, Taylorana hascheana, Rana tiomanensis, Hemidactylus frenatus, Dasia olivacea, Eutropis raniceps, Rana hosii, Rana picturata, Nyctixalus pictus,

178 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Acanthosaura armata, Aphaniotis fusca, Bronchocela cristatella, Pulau Tulai Draco melanopogon, Draco sumatranus, Gonocephalus Bronchocoela cristatella, Cnemaspis kendallii, Cnemaspis limi, chamaeleontinus, Gonocephalus grandis, Cnemaspis kendallii, Gekko monarchus, Gekko smithii, Hemidactylus frenatus, Cnemaspis limi, Gekko smithii, Ptychozoon kuhli, Dasia olivacea, Ptychozoon kuhli, Emoia atrocostata, Eutropis multifasciata, Larutia Eutropis multifasciata, Lipinia vittigera, Sphenomorphus seribuatensis, Sphenomorphus scotophilus, Varanus salvator, scotophilus, Varanus nebulosus, Varanus salvator, Ahaetulla Dibamus tiomanensis, Ahaetulla prasina, Python reticulatus prasina, Chrysopelea pelias, Dryocalamus subannulatus, Lepturophis albofuscus, Oligodon purpurescens, Rhabdophis Pulau Tunas Selatan chrysargos, Bungarus flaviceps, Calliophis intestinalis Gekko monarchus, Hemidactylus frenatus, Lepidodactylus lugubris, S. Pasal. Limnonectes blythii, Eutropis multifasciata Lygosoma bowringii, Varanus salvator, Ramphotyphlops braminus S. Paya. Bufo parvus, Limnonectes blythii, Taylorana hascheana, Ichthyophis sp., Hemidactylus frenatus, Cerberus rynchops, Pulau Tunas Utara Enhydris enhydris Hemidactylus frenatus, Varanus salvator S. Raya (2 km east of Kg. Mukut). Rana raniceps, Rana hosii, Cnemaspis limi, Eutropis multifasciata, Lipinia vittigera, Sphenomorphus scotophilus, Enhydris plumbea, Gongylosoma APPENDIX II mukutense, Rhabdophis chrysargos, Zaocys carinatus Tk. Dalam. Bufo parvus, Fejervarya cancrivora, Taylorana Catologue numbers for voucher specimens and non-vouchered site hascheana, Aphaniotis fusca, Bronchocela cristatella, Draco records for all new island records from the Tioman Archipelago, melanopogon, Draco sumatranus, Cnemaspis kendallii, Cnemaspis West Malaysia are reported below. All records for Varanus salvator limi, Ptychozoon kuhli, Eutropis multifasciata, Varanus nebulosus, are sight records (or track records) because the legal status of this Lepturophis albofuscus, Python reticulatus, species does not allow its collection. Tk. Dungun. Bufo parvus, Polypedates leucomystax, Taylorana hascheana, Rana raniceps, Rana hosii, Draco melanopogon, Draco Kaloula baleata — Pahang, P. Aceh; LSUHC 5712–13, 6954. sumatranus, Bronchocela cristatella, Gonocephalus Polypedates leucomystax — Pahang, P. Aceh calls heard. Dogania chamaeleontinus, Gonocephalus grandis, Cnemaspis kendallii, subplana — Johor, P. Tinggi LSUHC 6332. Aphaniotis fusca — Cnemaspis limi, Hemidactylus frenatus, Dasia olivacea, Eutropis Pahang, P. Aceh LSUHC 5699–702; Johor, P. Babi Hunjong multifasciata, Lipinia vittigera, Varanus nebulosus, Python LSUHC 5743–47. Cnemaspis kendallii –—Pahang, P. Aceh reticulatus, Ahaetulla prasina, Oligodon purpurascens, Rhabdophis LSUHC 5703; Johor, P. Babi Hujong LSUHC 5749–52; P. Ibol chrysargos LSUHC 6380–83; P. Sibu Tengah seen on side of tree. Tk. Nipah. Bufo asper, Chaperina fusca, Rana raniceps, Aphaniotis Crytodactlyus seribuatensis — Johor, P. Nangka Kecil LSUHC fusca, Draco melanopogon, Draco sumatranus, Gonocephalus 6348–54; P. Mentigi LSUHC 6348; P. Sibu LSUHC 6398–99; P. grandis, Cnemaspis limi, Cyrtodactylus tiomanensis, Hemidactylus Sibu Tengah LSUHC 6410; P. Lima Besar LSUHC 6413-21. frenatus, Dasia olivacea, Eutropis multifasciata, Lipinia surda, Cyrtodactylus aurensis — Johor, P. Aur LSUHC 7015, 7022–23. Lipinia vittigera, Sphenomorphus scotophilus, Varanus nebulosus, Gehyra mutilata — Johor, P. Rawa seen on side of building. P. Varanus salvator, Python reticulatus, Lepturophis albofuscus, Babi Tengah LSUHC 5754–59; P. Nanka Besar LSUHC 6334; P. Calliophis intestinalis Lima Besar LSUHC 5834, 5837–38; P. Lima Kecil LSUHC 5825; Tk. Penut. Bufo parvus Pulua Pemangggil LSUHC 7518, 7521. Gekko monarchus — U. Lalang. Ansonia tiomanica, Pelophryne brevipes, Philautus Pahang, P. Aceh LSUHC 6955-57; P. Layak LSUHC 6976–79; P. petersi, Oligodon booliati, Oligodon purpurascens Tunas Selatan LSUHC 6967; P. Pelandok LSUHC 6982–83; Johor, P. Batu Gajah LSUHC 6997; P. Seribuat LSUHC 5214–16; P. Pulau Tokong Bahara Sembilang LSUHC 5588; P. Mertang Tengah LSUHC 6950; P. Hemidactylus frenatus, Eutropis multifasciata Mertang Timur LSUHC 6944–49; P. Babi Hujong LSUHC 5740– 42; P. Penyambong LSUHC 7006–09; P. Ibol LSUHC 6384–87; P. Pulau Tokong Belalai Simbang LSUHC 6367–76; P. Nangka Kecil LSUHC6360; P. Hemidactylus frenatus, Varanus salvator Mentigi LSUHC 6338–41; P. Lima Besar LSUHC 5839–41; P. Cebeh LSUHC 6908–15; P Jahat LSUHC 6285–6288. Hemidactylus Pulau Tokong Burung Besar frenatus — Pahang, P. Aceh LSUHC5703; Aceh Anak LSUHC Hemidactylus frenatus, Eutropis multifasciata 6960–62; P. Layak LSUHC 6972–75; P. Tunas Selatan LSUHC 6963–66; P. Tunas Utara LSUHC 6971; Johor, P. Lalang Selatan Pulau Tokong Burung Condong LSUHC 6994; Pahang, Lalang Utara LSUHC 6980–81; P. Pelandok Hemidactylus frenatus, Eutropis multifasciata LSUHC 6984–85; Johor, P. Mawar LSUHC 6989–90; P. Setindan LSUHC 6999; P. Belanank LSUHC 7000–02; P. Tasu LSUHC Pulau Tokong Burung Tengah 6935–36; P. Mertang Timur LSUHC 6937–39; P. Rawa seen on Hemidactylus frenatus, Eutropis multifasciata houses; P. Harimau seen on radar tower; P. Gual seen in rock crack; P. Babi Hujong LSUHC 5738; P. Babi Tengah LSUHC 5753; P. Pulau Tokong Chondong Penyambong LSUHC 7010–11; P. Nangka Besar LSUHC 6335– Hemidactylus frenatus, Varanus salvator 36; P. Nangka Kecil LSUHC 6355–57; P. Mentigi LSUHC 6342– 44; P. Sibu Tengah LSUHC 6409; P. Sibu Kukus LSUHC 5802; P. Pulau Tokong Chupak Lima Besar seen on trees; P. Lima Kecil LSUHC 5824; P. Chondong Lepidodactylus lugubris LSUHC 5821–23; P. Belalai LSUHC 5827–29; Tuali seen on trees; P. Sepoi LSUHC 5685–86; P. Lang seen in bungalow; Pahang, P. Pulau Tokong Raket (two islands) Tokong Burung Besar LSUHC 5691; P. Tokong Burung Condong Lepidodactylus lugubris LSUHC 5695–96; P. Tokong Burung Tengah LSUHC 6933; P. Tokong Bahara LSUHC 5843–47. Lepidodactylus lugubris — Pulau Tokong Sanggol Pahang, Pulau Tunas Selatan LSUHC 6968; Johor, P. Mawar Lepidodactylus lugubris LSUHC 6991–92; P. Babi Tengah LSUHC 5761; P. Nangka Kecil

179 Grismer et al.: Herpetofauna of Seribut Archipelago

LSUHC 6358–59; P. Mentigi LSUHC 6347; P. Papan LSUHC 5819; Setindan LSUHC 6998; Mertang Tengah LSUHC 6950–51; P. P. Sibu Hujong LSUHC 5803–04; P. Sanggol LSUHC 5830–31; P. Mertang Timur LSUHC 6940–43; Rawa LSUHC 5723–25; P. Raket LSUHC 5826; Pahang, P. Cebeh LSUHC 6924–28; P. Sepoi Harimau LSUHC 5717; P. Mensirip LSUHC 5718–19; P. Babi LSUHC 6891–99; P. Jahat LSUHC 6289. Ptychozoon kuhli — Tengah seen beneath surface debris on beach; P. Ibol LSUHC 6377– Pahang, P. Aceh LSUHC 5708; P. Tinggi LSUHC 6321. Ptychozoon 79; P. Nangka Kecil LSUHC 6361; P. Mentigi LSUHC 6345–46; lionatum — Johor, P. Sibu LSUHC 5539, 5583, 6697; Pahang, P. P. Papan LSUHC 5817–18; P. Sibu Tengah LSUHC 5810–11; P. Tioman LSUHC 6437. Dasia olivacea — Johor, Pulau Babi Besar Lima Bear LSUHC 5835–36. Ramphotyphlops braminus — Pahang, LSUHCPC 0246; Pulau Babi Tengah LSUHC 5748; P. Pemanggil P. Aceh LSUHC 5705; P. Tunas Selatan LSUHC 6969; Johor, P. LSUHC 7513-14. Emoia atrocostata — Johor, P. Nangka Besar Sibu Tengah LSUHC 6411–12. Python reticulatus — P. Tulai LSUHC 6333; P. Papan LSUHC 5814–16; P. Sibu Tengah LSUHC LSUDPC 266-69. Boiga drapiezii — Pahang, P. Tioman LSUHC 5806; P. Sibu Kukus LSUHC 5800–01. Eutropis multifasciata 6400. Boiga nigriceps — Johor. P. Aur LSUHC 2020–21. Pahang, P. Aceh LSUHC 5716; Johor, P. Rawa seen on forest floor; Chrysopelea paradisii — Johor, P. Sibu Tengah LSUHC 5848. P. Babi Tengah LSUHC seen at edge of forest; P. Sibu Kukus seen Dendrelaphis caudolineatus — Pahang, P. Aceh LSUHC 6958; on forest floor, Pahang, P. Sepoi LSUHC 5867–68; Johor, P. Lang Johor, P. Sibu Tengah LSUHC 6408. Dendrelaphis cyanochloris seen on forest floor; Pahang, P. Tokong Burung Besar LSUHC — Johor, P. Tinggi LSUHC 6401. Dendrelaphis pictus — Pahang, 5692–94; P. Tokong Burung Condong LSUHC 6929; P. Tokong P. Aceh LSUHC 6959; Johor, P. Mawar LSUHC 6993. Lycodon Burung Tengah LSUHC 6933; P. Tokong Bahara LSUHC 6134– capucinus — Johor, P. Rawa LSUHC 5725; P. Babi Besar LSUHC 35. Larutia seribuatensis — Pahang, P. Tioman LSUHC 7074. 5768; P. Babi Hujong LSUHC 5767. Pseudorabdion longiceps — Lygosoma bowringii — Pahang, P. Aceh LSUHC 5697–98; P. Tunas Johor. P. Sibu LSUHC 6388. Tropidolaemus wagleri — Pahang, Selatan LSUHC 6970; Johor, P. Mawar LSUHC 6986–88; P. P. Aceh LSUHC 5714–15.

180 THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 181-197 Date of Publication: 28 Feb.2006 © National University of Singapore

THE CURRENT AND HISTORIC DISTRIBUTION OF TOMISTOMA SCHLEGELII (THE FALSE GHARIAL) (MÜLLER, 1838) (CROCODYLIA, REPTILIA)

Robert B. Stuebing Division of Amphibians and Reptiles, Field Museum of Natural History, 1400 S. Lakeshore Dr., Chicago, IL 60605-2496 USA

Mark R. Bezuijen P.O. Box 183, Ferny Creek, Victoria, Australia 3786

Mark Auliya Zoologisches Forschungsinstitut & Museum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

Harold K. Voris Division of Amphibians and Reptiles, Field Museum of Natural History, 1400 S. Lakeshore Dr., Chicago, IL 60605-2496 USA

ABSTRACT. – Tomistoma schlegelii the “False Gharial” (Müller, 1838) is one of the largest yet least-known of the world’s 23 crocodilian species, restricted to Indonesia and Malaysia. We compiled and mapped 210 records and overlaid these against geological river systems and current vegetation, in order to assess historic and current distribution, relationship to key habitats and potential centres of current abundance. The current distribution of T. schlegelii extends over lowland regions of eastern Sumatra, Kalimantan and western Java (Indonesia), Sarawak and Peninsular Malaysia, Vietnam and Thailand, within 5 degrees north and south of the equator. Additional records from Sabah (Borneo) and Thailand are unconfirmed, and early records from Thailand may have originated from regions currently within Peninsular Malaysia. The Pliocene to early Pleistocene distribution of T. schlegelii was apparently much broader than its historical range, and extended over lowland river systems of Indonesia, Malaysia and into southern China. Since the discovery of the species in 1838, a significant decline in the density of populations throughout range states has occurred, more markedly since the 1940s. Ironically, the broad outlines of the original distribution seem to have remained. A large portion of the records appear to be associated with peat swamp forest, a highly threatened category of tropical forest, suggesting that a decline in abundance is related to habitat loss. Current areas in which the species appears to still maintain reasonable levels of abundance are in southern Sumatra, and south- , although some smaller protected populations occur in central Sarawak. Current T. schlegelii breeding habitats are under threat throughout range states as a result of the progressive draining, logging, burning and clearing of swamp forests.

KEY WORDS. – Tomistoma, false gharial, peat swamp, crocodilian.

INTRODUCTION assessments conducted between 1985 and 2004 (Bezuijen, et al., 1995, 1997, 2001b, 2002a, b; Bezuijen, 2004; Cox and Tomistoma schlegelii, or “False Gharial” (Müller, 1838) is a Gombek, 1985; Frazier, 1994; Frazier and Maturbongs, 1990; large (4-5 m total length), freshwater crocodilian whose extant Muin and Ramono, 1994; Ramono, 1994; Ross et al., 1998; distribution is restricted to Indonesia (east Sumatra, west Java Sebastian, 1993a,b; 1994; Simpson et al., 1997; Simpson, and Kalimantan) and Malaysia (Sarawak and Peninsular 2004). Key findings from these limited data indicate that the Malaysia). The species was first described from “southern preferred nesting habitat of T. schlegelii is peat or freshwater Borneo” (currently Kalimantan, Indonesia). It is one of the swamp forest, and that the species appears to occur at low least-known of the world’s 23 extant crocodilian species and densities throughout most of its range (Bezuijen et al., 1998, is listed as globally “Endangered” by the IUCN-World 2001a; Stuebing et al., 2004). Conservation Union (Thorbjarnarson, 1992; Ross, 1998b; www.redlist.org). To date, there have been no long-term Peat swamp forest is one of the most threatened tropical forest autecological studies of the species, and most available ecosystems in South-east Asia and under intensive loss or information is from a small number of rapid status degradation from indiscriminate harvest of wood, forest fires

181 Stuebing et al.: Distribution of Tomistoma schlegelii and swamp reclamation (Rieley and Page, 1997; Stibig et Tomistoma schlegelii records al., 2002). In Borneo, <50% (95,000km2) of original swamp Records were assigned to one of three categories reflecting forest is estimated to remain (WCMC, 2000). This has their validity: “1” - museum specimens with information on resulted in extensive loss of documented and potential T. date, collector and full or partial locality data, e.g. river or schlegelii nesting habitat (Bezuijen et al., 2002a, b; Bezuijen, district name; “2” - first-hand field observation by the authors; 2004). In this paper, we map the current and historic global or, an observer who related the record directly to one of us distribution of T. schlegelii and evaluate these data to identify and correctly described the species, locality and date; or, areas where viable populations of the species may still occur, captive specimen observed by the authors in the source region and to make an initial assessment of whether there have been and for which information on capture location and date was significant changes in the overall geographic distribution of given by the original collector; “3” - first- or second-hand the species in recent (e.g., post 1940) times. field observation by a non-scientist, usually a local villager, in which a general location and/or date is given but few other details, and which was related to the authors or another METHODS scientist. Other records (second- and third-hand information) were excluded from compilation. Categories 1 and 2 were Study area regarded as “confirmed” records and category 3 records as Sumatra, Borneo and Peninsular Malaysia are located within “unconfirmed”. Wherever possible, reliability, location, the biogeographic region “Sundaland”, referring to South- collector, date and geographic coordinates (latitude/longitude) east Asian countries located on the shallow (<100 m depth) were collected for each record. Multiple records for a single Sunda shelf of the South China Sea (Voris, 2000). Mean site were treated as independent records if the authors of annual ambient temperatures experience a low range and survey reports had accounted for and removed potential humidity is high (e.g. 23-31°C and 85% respectively in repeat-sightings (i.e. four separate sightings in a single river Sumatra; Whitten et al., 1987). A distinct “wet” (November- were compiled as four separate records). If this was not clear, February) and “dry” (March-September) season occurs, then multiple records for a single site from a single survey, although the wet season is extended in Sumatra, lasting from were compiled as a single record. Authors of recent reports October-April. In Sumatra, Java, Borneo, and Peninsular describing T. schlegelii sightings were contacted for GPS or Malaysia mean annual rainfall ranges from 2,500-3,500 mm, map coordinates of records. For records with adequate with peak rainfall from December to March (Whitmore, 1984; descriptions but no map coordinates (especially museum Whitten et al., 1996; Meteorological Department, Sarawak; specimens), a map coordinate was obtained from regional Malaysian Meteorological Department, Kuala Lumpur). topographic maps. For these records, coordinates were Large, meandering rivers dominate the lowlands, and tidal assigned to a location above the limit of saltwater intrusion, influence extends up to 100 km inland and well into totally as we assumed that the T. schlegelii only occurs in freshwater freshwater areas. Tidal range is highly variable between habitats. regions, and there is generally one tidal cycle per day, but can be two per day during neap tides, with a tidal range of 5 The following records were excluded from compilation: all m in some areas (Hadi et al., 1977; ISDP, 1997). records (including museum specimens) with no locality information or vague locality descriptions (e.g. a record only Most lowland regions of Sumatra, Borneo and Peninsular stating “Perak River, Malaysia” – this river is >200 km long); Malaysia historically have contained extensive peat and captive specimens of uncertain origin (usually those observed freshwater swamp forest habitats, e.g., forests that are in trade or on crocodile farms some distance from a potential inundated for a significant portion of the year, and because source region); “third-hand” reports; and, “eyeshine” records of their acidity and anaerobic character, often contain layers (where a crocodilian was sighted in a survey but the species of organic detritus in the form of peat. Borneo and Sumatra, was not confirmed). where the largest area of these habitats are located, historically supported approximately 220,000 km2 of peat swamp forest Mapping and data analysis (including 154,000 km2 peat swamp forest), which has been Records were plotted in a ArcView 3.2 geographic reduced to approximately 95,000 km2 due to development information system and overlaid on a 1: 5 500 000 scale activities (a loss of <50%); substantial remnants (>20,000 vegetation map of south-east Asia (source “cartographic km2) remain in the middle Sg [= River] Mahakam, in central digital products” at http://www-gvm.jrc.it/Forest/ and western Kalimantan (WCMC, 2002). In Sumatra, most defaultForest.htm). Records were analysed for: (1) extent of swamp forests are located in a large belt of low-lying, tidal global coverage and distribution within south-east Asia; (2) floodplains, termed the ‘Eastern Lowlands’, which extend extent of overlap in records, at sites with repeat records; (3) from southern to northern Sumatra. These lowlands comprise implications of any habitat preference by T. schlegelii. some 18% (88,000 km2) of the island, mostly in the provinces of Riau, Jambi and South Sumatra (Claridge, 1994), and consist mainly of alluvial and marine deposits, frequently RESULTS overlain by a layer of peat that may reach depths of 20 m (Danielsen and Verheugt, 1990). Historic distribution A total of 210 T. schlegelii records were compiled, dating from 1838-2004 (Table 1; Appendix 1): 41 category “1”, 117

182 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Table 1: Summary of False Gharial presence records 1838 - 2004

Region Records (categories 1-3)* Total specific Admin. regions Total Records localities 123 INDONESIA Sumatra Aceh Province 0 0 0 0 North Sumatra Province 5 0 0 3 Riau Province 2 8 4 8 Jambi Province 1 20 2 9 South Sumatra Province 4 57 0 12 Lampung Province 0 3 1 2 West Sumatra Province 00(2) (2)* Total 12 88 9 36 7 109 Kalimantan Province 1 2 18 20 West Kalimantan Province 8 3 5 10 Central Kalimantan Province 1 12 0 3 Province 2 0 0 1 Total 11 17 23 34 4 51 Java 1 Banten Province 1 1 0 2 Total 110 222 MALAYSIA Peninsular Malaysia Pahang State 3 0 9 10 Perak State 2 2 0 4 Selangor State 1 0 1 2 Trengganu State (1) 1 Total 7210 17 4 18 Sarawak Division 1 7 2 4 Division 1 1 0 2 Semarahan Division 8 1 0 3 Division 0 0 6 5 Total 10 9 8 14 4 26 Sabah (2)* West Coast Residency 0 0 (1) (1) (1) (1) Sandakan Residency 0 0 (1) (1) (1) (1) *see text category “2” and 52 category “3” records (four, including the basis of museum records and geological river systems, two from Sabah and two from West Sumatra), were not the historic distribution of T. schlegelii may have considered further for evaluation of Tomistoma distribution encompassed most low lying river systems in Borneo (within because of problems either of species confirmation or Sarawak and Kalimantan), east Sumatra, Java and Peninsular verification of locality. Museum records (category “1”) were Malaysia. usually the earliest published records. All records were from Indonesia (Sumatra, Kalimantan and Java) and Malaysia In Sumatra, records and interviews with local people (Sarawak and Peninsular Malaysia). “Historic” distribution (Bezuijen et al., 1995, 1997) indicate that T. schlegelii was was defined in this study as pre-1940, because the human historically widely distributed in eastern Sumatra, from south- activities which may have influenced T. schlegelii distribution east Aceh to Lampung Provinces, and was absent from and abundance became much more significant in the 1950s western Sumatra (even though there are some recent second (extensive regional commercial crocodile hunting from the hand reports from that region, these have not been verified) 1950s-70s) (Bezuijen et al., 1997; Simpson et al., 1997) and (Fig. 1). Possible reasons for the absence of T. schlegelii from 1960s (start of large-scale land clearing and development of western Sumatra include the existence of extensive mountain swamp forest; Barber & Schweithelm, 2000). Geologically, ranges dividing west and east Sumatra, which may have the global distribution of T. schlegelii corresponds with that stopped westward expansion; a lack of swamp forest habitats of historic river systems in south-east Asia (Voris, 2000). On in most of western Sumatra; and the lack of physical

183 Stuebing et al.: Distribution of Tomistoma schlegelii connections between the rivers of western Sumatra with the Rivers, with many records from western and a few from Great Sunda River (Voris, 2000). In Borneo, rivers in western central areas of the State (Fig. 1). Kalimantan originally formed tributaries of the North Sunda River, while those of southern and eastern Kalimantan were Current distribution part of a south-eastern drainage flowing towards the Arafura A total of 180 T. schlegelii records were compiled for the Sea (Hanebuth et al., 2000; Voris, 2000). Rivers in the period 1980 – 2003. The current range of T. schlegelii northeastern regions of East Kalimantan Province and eastern comprises regions of Borneo (Sarawak, central, west and east Sabah were not connected to either of the latter river systems, Kalimantan), Sumatra and Pensinsular Malaysia. Brief which drained vast lowland (and perhaps swampy) terrain. regional summaries on current T. schlegelii is given below. Tomistoma schlegelii was probably widespread in Kalimantan during the Pleistocene (<1 million years ago), and has been Sumatra (Indonesia) reported by local people in the far uppermost parts of the Sg Confirmed (e.g. categories 1 & 2) records are from lowland, Barito north of Muara Teweh in Central Kalimantan coastal regions of eastern Sumatra, in Riau (10 records), Jambi (Stuebing, et al., 2003). In Sarawak, most records are from (21) and South Sumatra (61) Provinces (Table 1; Appendix swampy upstream areas of the Sadong and (Batang) Lupar 1). The western limits of T. schlegelii distribution are probably

Fig 1. Historical peat swamp distribution in Southeast Asia (after Andriesse, 1974).

Legend (for Fig. 1 only) BORNEO SUMATRA JAVA 1. 2. Sg Batanghari complex 3. Sg Lalan complex 4. Sg Melawai 5. Sg Kapuas 6. Sg Sekunyir 7. Sg Barito 8. Sg Mahakam (& Mahakam lakes complex) 9. Sg Sebuku 10. Sg Pahang 11. Sg Bera (& Tasek Bera wetland) 12. Sg Perak – Sg Bernam area 13. Ujung Kulon

184 THE RAFFLES BULLETIN OF ZOOLOGY 2006 the foothills of the Barisan mountains, where unconfirmed Central and West Kalimantan (Table 1; Appendix 1). In South reports were recorded (Bezuijen et al., 1997). Photographs Kalimantan Province, extensive loss of swamps due to rice of T. schlegelii eggs and hatchlings said to originate from and plantation agriculture has probably eliminated any historic western Riau Province were shown to one of the authors (MB) T. schlegelii populations. There are no confirmed records from in 2002. The most recent confirmed records (2002) are from Sabah, where northward extension from Kalimantan may the Merang River (South Sumatra Province) and Berbak have been constrained by the replacement of slow, swampy National Park (Jambi Province) (Bezuijen et al., 2002a, b). waterways with shorter, swift-flowing rivers. Unconfirmed There are three T. schlegelii records since 1980 in Lampung reports of T. schlegelii occur from the Sebuku River (East Province, and no records from North Sumatra and Aceh Kalimantan Province) and Klias and Kinabatangan Rivers Provinces since 1980, although local reports indicate the (Sabah) (Appendix 1). species may historically have been widespread in some parts of these provinces (Bezuijen et al., 1997). In Lampung Kalimantan. In West Kalimantan Province, T. schlegelii Province, of 13 reported sightings since the 1980s, only one records are from the southern, central and northern regions report was of a nest (Bezuijen et al., 1997). Surveys in the of the province (Galdikas & Yeager, 1984; Yeager, 1991; largest remaining protected lowland forest in eastern Simpson, 2004). Confirmed records are from Gunung Palung, Lampung Province provided no evidence of current Danau Sentarum and Betung-Kerihun National Parks, with occurrence of T. schlegelii (Bezuijen et al., 2002a, b). In scattered reports from other locations in the province Sumatra, Riau, Jambi and South Sumatra Provinces are the (Bezuijen et al., 2004). In South Kalimantan Province, there strongholds for T. schlegelii, and the species is now probably is one confirmed record (a museum specimen from the rare or locally extinct in many regions of Lampung, North “Banjarmasin” area, De Rooij, 1917). This specimen was Sumatra and Aceh Provinces. probably collected north of Banjarmasin town, which is located in tidal habitat rarely occupied by T. schlegelii. In Borneo (Kalimantan, Sarawak, Sabah) Central Kalimantan Province, the most recent confirmed Confirmed (categories 1 & 2) records are from the lowlands record of T. schlegelii is in the Simpang Kanan and Kuma of western and west-central Sarawak (19 records) and West, Rivers in/near Tanjung Puting National Park (Simpson, 2004). South, Central and East Kalimantan (28 records) (Table 1). In addition, the type specimen for the species was “Lake Records in Kalimantan are from a large geographic area, from Lamoeda, southern Kalimantan” (Müller, 1838), which was western Kalimantan, south to Gunung Palung and Tanjung probably in the Dusun River (a tributary of the ) Puting National Parks, and north to Muara Teweh in the upper near Muara Teweh town (M. Hoogmoed, pers. comm.). In Barito/Mahakam Rivers, the Mahakam Lakes and eastern part East Kalimantan Province, confirmed records indicate T. of East Kalimantan (Fig. 2). Most records in Borneo are in

Fig 2. Current forested habitats remaining in the known range of Tomistoma, with some important locality records indicated.

185 Stuebing et al.: Distribution of Tomistoma schlegelii schlegelii occur in large sections of the middle Barito River, were made during surveys. The species was listed in the and the Mahakam Lakes (Appendix 1; Ross Park’s fauna by Blower and Zon (1977) and Clarbrough et al., 1996). Unconfirmed reports are from the Pari River (1990), although it was not mentioned by Hoogerwerf (1970) (Stuebing, et al., 2003; B. Simpson, 2003 pers. comm.) and in a detailed description of the Park’s crocodilian fauna. Based Sebuku River (KSDA Forest Ranger, 2003 pers. comm.). on land connections between Sumatra and Java during the Pleistocene (<1 million years ago; Voris, 2000), it is possible Sarawak. Historic accounts give contrasting reports of T. that the species was found on the western portion of the island. schlegelii abundance, variously described as “quite common” Most of Java’s natural habitats are now altered or destroyed, in Borneo (Müller and Schlegel, 1845), but much less and any remnant population in Ujung Kulon National Park common than C. porosus (Hornaday, 1885), or very locally is probably small and isolated. distributed. Shelford (1916) viewed its distribution as “local”, and that the species seemed abundant only in the Sadong River Peninsular Malaysia (a river just east of in western Sarawak) with few All (category 1-3) T. schlegelii records originate from the specimens having been taken elsewhere in Sarawak. Most central interior and western coast of Peninsular Malaysia, museum specimens, field records and approximately 25 including an unconfirmed record from the Sg Jelud in northern individuals currently in the Jong Crocodile Farm, Kuching, Perak State (that was not included in Appendix 1; Sukumaran, were obtained or are the offspring of animals taken from the 2002). There are four museum specimens for the species from upstream tributaries of the Sadong and Batang Lupar Rivers 1896-1940 in the Perak and Selangor States, and all (Appendix 1). Other confirmed records are from the Mayeng, subsequent records are from the Tasek Bera region in the Sarang and Penyilam Rivers (Lading & Stuebing, 1997; central and eastern parts of Pahang State (Appendix 1). Stuebing, et al, 1998, 2004) and Loagun Bunut lake (K. Simpson et al. (1997, 1998) reported that based on local Jensen, 2004, pers. comm.). There are few confirmed records reports, T. schlegelii was present in Tasek Bera in the central from northern Sarawak, except for a record from the Baram and western parts of Pahang State, Peninsular Malaysia in River (B. Simpson, pers. comm.). A crocodile farm owner in 1998, but the team was unable to confirm this despite two Miri, near the Baram River, declared in 2003 that several T. weeks of intensive surveys over much of the area. schlegelii at his farm were obtained from Kalimantan, but In the Bahau district Negri Sembilan, during the early years this could not be verified. (1950-1970’s) T. schegeli was quite common in Sg Triang and Sg Pertang that adjoin the main Pahang river. At least Brunei Darussalam. Until April, 2005, there were no three specimens of T. schlegelii were collected by hunters in confirmed records from Brunei Darussalam, although there these rivers for food (B.L. Lim, pers. comm.). The latest was were some anecdotal reports of its existence from this a large specimen caught in the in Sg Triang - Sg Pertang area geographical area1 . On 12 April 2005, one of us (RBS) visited by hunters in Oct 1999, and picked up by animal dealers (LBL Bandar Seri Begawan, and was shown a clear photograph of witnessed the specimen). A relative of LBL in Bahau a subadult Tomistoma schlegelii obtained from the Tutong confirmed that at least one more Tomistoma schlegelii was River in late 2002. caught in September 2002 and was sold to a restaurant in Bahau. Sabah. There are no confirmed records from Sabah. No T. schlegelii were recorded during crocodile surveys by In 2003, two T. schlegelii were captured in Perak State: a Whitaker (1984) or R. Stuebing (unpubl. data, June-July 2002, large male from the Sg Sungkai (near the town of the same from surveys in the Klias River, Sabah). Local residents in name) a tributary of the Sg Perak, and a subadult from the Sg the upper Klias River reported seeing a crocodile resembling Erung (Appendix 1). There are also local reports of T. a T. schlegelii in April 2002, and a second-hand sighting was schlegelii from the Sg Kinta, another tributary of the Sg Perak, reported by a tourist in the Kinabatangan River (RBS unpubl. just south (downstream) of the 1894 Pulau Tiga locality. data). These records were not considered suitable for inclusion Residents of this area maintain that Tomistoma can still be in the discussion due to vagueness of the reports. found there (Ahmad Darubi, pers. comm.).

Java (Indonesia) Thailand The current status of T. schlegelii in Java is unclear. The The status, and even the presence of T. schlegelii in Thailand species was reported to occur in Java by Klein (1863), on the is unclear. One museum specimen is known (UKNHM basis of a crocodile skull described as “Rhamphostoma specimen no. 157993), whose label states “collected in Schlegelii (Crocodilus Schlegeli Müller and Schlegel)”, Thailand”, but with no other supporting information provided although apparently the locality data have been lost (Auliya, (this record was not included in the database). There are 2003). This report was later cited by Strauch (1866) and De unconfirmed reports of T. schlegelii from Hala-Bala Wildlife Rooij (1917), although Koningsberger (1913) and Mohr Sanctuary and Kaeng Krachen National Park (P. P. van Dijk, (1921) noted there were no confirmed records of occurrence 2003 pers. comm.). Some local crocodile farms have many on the island and that further information was required. T. schlegelii but the origin of these animals could not be Iskandar (2001) does not record this species as confirmed in determined. Smith (1916) lists T. schlegelii as occurring in Java. Local reports correctly describing this species were the “Inland Sea (Peninsular Siam)” (Thailand), but gives no collected from Ujung Kulon National Park (western Java) in further data, yet Chan-ard et. al. (1999) did not list T. schlegelii 2002 (Auliya, 2002a, b, 2003), although no direct sightings as occurring in Thailand. It is possible that T. schlegelii did

186 THE RAFFLES BULLETIN OF ZOOLOGY 2006 not historically occur in Thailand, unless northern areas dearth of good records, both fossil and recent (especially in previously under Thai sovereignty (now the Malaysian States view of the difficulty in conducting effective surveys). of Kedah and Perlis) are included. Approximately 95% of the current geographic distribution of T. schlegelii appears to be confined to a region about five Vietnam degrees north and south of the equator. Recent records do In the January-March 2005 issue of its Newsletter, the IUCN/ seem to cluster within one degree of the equator, but this may SSC Crocodile Specialist Group carried a letter from Mr. J. be an artefact of increased survey effort in specific regions, Mucelli, who stated that he had witnessed the killing of a such as in the Danau Sentarum and the Mahakam Lakes. The Tomistoma schlegelii by South Vietnamese soldiers in 1967, overall presence for the species seems rather widely scattered “in the vicinity of Dai Quay bridge, Lam Dong Province” over the ten degree latitudinal band. Kalimantan and Sarawak (along highway 20, the Ho Chi Minh City (formerly Saigon) are considered to potentially support the largest global to Dalat road). This is the first and only report of the species populations of T. schlegelii, on the basis of land mass and (that we are aware of) from Vietnam, and requires further large remnants of potential swamp forest nesting and foraging investigation. The author of the article was convinced that habitat. the species did once inhabit the area, a “lowland forest with large tracts of bamboo” (Mucelli, 2005). In Sumatra, the species may now be rare or absent from regions of the north-east and south-east of the island, i.e. Lampung, Aceh and North Sumatra Provinces. These areas DISCUSSION may have historically supported smaller regions of swamp forest than the central-eastern provinces of Riau, Jambi and Geologic and current distribution South Sumatra. It is likely that habitat loss has contributed to Tomistoma-like crocodilians were widely distributed in the this apparent range decline. In Lampung Province, the most Americas, Europe, Africa and Asia from the beginning of intensively developed and populated of Sumatran provinces, the Tertiary to the Miocene (approximately 60- 30 mya), and large-scale habitat loss and disturbance has occurred in sites are well-represented in the fossil record. In north and east where local people formerly reported sightings and nesting Asia, at least one genus of Tomistoma-like crocodilian, (Bezuijen et al., 1997, 2002a). Similarly, much of eastern Tomistomine, occurred until the Pliocene or early Pleistocene Sumatra has been subjected to land clearing or burning (C. Brochu, 2003). More recently, the fossil distribution of (Barber and Schweithelm, 2000; Bezuijen et al., 2001a, b). Tomistoma may have included southern China, where sub- Local people interviewed in eastern Sumatra reported that T. fossil remains of a Tomistoma crocodilian were reported, schlegelii was previously “common” or “often seen” 20 years apparently dating from 1,300-1,400 CE (Ross et al., 1996). ago, but had declined over the last 15-20 years, particularly Tomistomines, whose fossil record is far more obscure, in areas which had been burnt (Bezuijen et al., 1995, 1997). occurred in Japan and Taiwan as late as the Pliocene or early It is possible that T. schlegelii populations are reduced or Pleistocene, but their range appears to have progressively absent from a substantial number of the river systems of contracted as the climate appropriate for them deteriorated, eastern Sumatra. resulting in their present distribution (Steel, 1989). Pleistocene maps of the Sunda region showing the outline of In Peninsular Malaysia, scattered populations of T. schlegelii major river systems in Peninsular Malaysia, Sumatra, Java still exist, and small numbers are seen in any given year in and Borneo (Voris, 2000) do indicate that T. schlegelii Perak State and possibly also in Pahang State. Historic centres probably occupied a greater range than it does today. The of distribution included Pahang, Perak and Selangor States. vast interlinked river systems of the Sunda region (Great These sites, however, have been seriously disturbed by tin Sunda River basin) were undoubtedly associated with some mining, the opening of rubber and oil palm plantations, and peat swamp habitats (long since submerged by rising seas). urban development. Furthermore, Malaysia’s human This ecological scenario would imply that T. schlegelii could population has nearly doubled since 1970, and currently have been distributed broadly throughout ancient Sundaland, exceeds 25 million inhabitants (Government of Malaysia, from northern and eastern Sumatra and adjacent parts of 2004). Given the limited land area and increasing Peninsular Malaysia, southwards to western Java and development in the peninsular, it seems likely the populations eastwards into large areas of Borneo. Nearly all records of T. schlegelii have declined significantly, becoming rare, examined in this paper can be seen to lie within the geologic or absent in many waterways where they formerly occurred. watersheds of several ancient and low-lying river systems depicted in Voris (2000). Western Borneo and south-central Habitat Sumatra seem to form one biogeographical region (Yap, Peat swamps apparently made up 12% of the area of Sarawak 2002), where the majority of False Gharial records are located. (about 14,500km2) and about 22.6% of Brunei (Anderson, Outlier records, most of which are unconfirmed, are from 1964). Sabah’s peat swamp and kerangas habitats are limited Vietnam, Thailand, extreme East Kalimantan, northern to a small corner in its extreme southwest in a triangle between Sarawak and south-west Sabah. the towns of Membakut, Beaufort and Kuala Penyu encompassing an insignificant portion of the land area of that Compared to its apparent prehistoric range, the current state. Peat swamps cover significant areas of the western and distribution of the False Gharial seems to have become more southern Kalimantan, southeastern Sumatra and a band in fragmented, though this can only be surmised, given the Peninsular Malaysia running from central Perak, to Malacca,

187 Stuebing et al.: Distribution of Tomistoma schlegelii and although interrupted by the Main Range, eastwards from (Auliya, 2002a, b, 2003), a highly visited and surveyed region. Negeri Sembilan and the upper Sg Pahang drainage to Confirmed records (in 2003) of individuals in Perak State extensive coastal areas south of the town of Kuantan (Fig. and Kenyir Lake (Terengganu State) in Peninsular Malaysia 2). Over the last 35 years, oceanographers have found peat indicate that at least some individuals continue to survive and swamp remnants in cores taken from sites as disparate as the breed, even in relatively developed areas. In general, these Malacca Straits, off the coast of Singapore, off the east coast records indicate the species may remain widespread, but in of Peninsular Malaysia and along an extensive NE/SW reduced densities, in many areas where it was formerly transect in the South China Sea (Inger et al., 2005). In both abundant. In reality, however, there are few areas, perhaps Sumatra and Borneo, peat swamps extend well inland from with the exception of Tanjung Puting National Park in coastal areas, and are found as far inland as the Sg Penyilam southern Kalimantan (Central Kalimantan Province) where and Loagan Bunut in Sarawak, and the Sg Maruwai in the it can be stated with any confidence that T. schlegelii is extreme northern portion of Central Kalimantan Province actually “abundant”. Until more effective survey approaches (Stuebing, et al., 2003). Based on carbon-14 dates from the and techniques can be developed, any assertions on abundance above peat cores, the age of the Sunda region peat swamps or rarity will remain of questionable value, for all but the ranges from approximately 44,200 - 3530 yBP, corresponding most obviously altered habitats. to a period of large fluctuations in the level the South China Sea during the last temperate-zone glaciation (Anderson, Threats and conservation priorities 1964; Geyh et al., 1979; Hesp et al., 1998; Hanebuth et al., Key anthropogenic threats to the long-term survival of T. 2000). schlegelii include regional loss of nesting habitat due to forest fires, incidental or commercial harvest for food, illegal The geographical records for T. schlegelii are associated with logging or the development of industrial tree plantations, peat low lying areas such as peat swamps, freshwater swamps and swamp clearance and drainage for irrigated agriculture, oil flood plains. It is clear from the location of most records of and gas exploitation in swamp forests, uncontrolled regional T. schlegelii, that the species seems to favor that habitat (Fig. settlement of swamp forests, and also incidental mortality of 2). Most museum or field records of T. schlegelii nests are breeding individuals in fishing nets (Bezuijen et al., 2002a, from peat swamp or freshwater swamp forest, and the species 2004; Frazier, 2000). Small, fragmented and isolated has generally been regarded as a peat swamp specialist populations (particularly those in Peninsular Malaysia, where (Bezuijen et al., 1998, 2001a; Ross et al., 1998). Müller and most regions are subject to intensive development) are Schlegel (1845) described the species habitat as “quiet/ probably vulnerable to demographic or environmental isolated and calm water bodies, from which it occasionally stochasticity, and may eventually become locally extinct due visits slow-moving tributaries and swamps with black smelly to a lack of suitable nesting habitat or disturbance. In Sarawak, water. … it rarely enters rivers with strong currents.” Surveys peat swamp tributaries of the Sg Sadong, including the Sg by the authors and interviews with local people in range states Ensengai, where T. schlegelii was formerly known to be have consistently reported similar habitats for this species. reasonably abundant in the early 1980s (Cox and Gombek, Unfortunately, the wide variation in the manner in which these 1985), have now been drained or the main waterway data have been collected precludes a more systematic analysis “straightened” with a permanent drainage system (J. Jong, than has been possible for this paper. pers. comm.). Based on the records and mapping conducted for this paper, T. schlegelii still appears to occupy large parts Although peat swamp forest appears to be the preferred of its global distribution, although many populations are habitat of this species, Tomistorna has also been recorded probably at substantial risk from the threats given above. From from other lowland forest habitats, e.g. impounded water this data, it appears that although insufficient population data catchments in Peninsular Malaysia (Chan-Ard et al., 1999), is available to confidently assess whether the IUCN freshwater swamp forests in central Sarawak (R. Stuebing, “Endangered’ status of the species as is warranted, “Critically photographic evidence from Bukit Sarang in Ulu Kakus/ Endangered” status is at least, unlikely. ; see Appendix) and the Sg Sibau in West Kalimantan (Bezuijen et al., 2004). Conservation activities of high-priority for this species were identified by Bezuijen et al. (2003), and include baseline Abundance surveys in specific regions of eastern Sumatra, Borneo and Effective assessment of T. schlegelii abundance is hindered Peninsular Malaysia, as well as the implementation of by its cryptic coloration and its secretive nature. Spotlight population monitoring programs in three sites for which there surveys in areas where T. schlegelii are known to occur, are repeat surveys: the Sg Merang and Berbak National Park frequently do not result in sightings (authors pers. obs. in (eastern Sumatra) and Tanjung Puting National Park (central Sumatra, Kalimantan, Sarawak). Ross et al. (1996) noted that Kalimantan). In this review, we add the following sites to in many areas of Kalimantan, the species was regarded as that list for baseline surveys: Sg Sibau and Gunung Palung the “common” crocodilian compared with Crocodylus National Park (West Kalimantan), and also the Lingga- species. The species appears to be abundant in some rivers Simunjan area and Wildlife Sanctuary in Sarawak. close to Tanjung Puting National Park in Central Kalimantan To assist in the objective prioritisation of these areas, a simple Province (Simpson, 2004). The difficulty in assessing the scoring system to prioritise each area is recommended, based local status of the species is illustrated by its recent on the following criteria: A) whether T. schlegelii appears to “rediscovery” in Ujung Kulon National Park in western Java be abundant in a site; B) the site supports suitable nesting

188 THE RAFFLES BULLETIN OF ZOOLOGY 2006 and foraging habitat (based on field records, etc); C) the site’s LITERATURE CITED potential for effective protection and management. Individual sites would be classified as “high priority” for conservation Anderson, J. A. R., 1964. The structure and development of the (scoring in all three categories), “medium priority” (a site peat swamps of Sarawak and Brunei. Journal of Tropical with only two of these variables), or “low priority” (a site Geography, 18:7-16. supporting only variable). Identification and classification Andriesse, J. R., 1974. Tropical lowland peats in Southeast Asia, of sites could begin by examining geological maps in the main Kon. Inst. Voor de Tropen, Amsterdam, 63 pp. areas of abundance (Sumatra and Borneo) to identify old, Auliya, M., 2000. Record of Tomistoma schlegelii in West low lying alluvial basins that contain peat swamp remnants Kalimantan. Crocodile Specialist Group Newsletter 19: 8-9. or flooded “blackwater” forests where the species seems to Auliya, M., 2002a. Tomistoma on Java. Crocodile Specialist Group reside within some fairly distinct vertebrate communities Newsletter 21 (3): 5-6. (Inger et al., 2005). Auliya, M., 2002b. Two crocodile species in West Java? Crocodile Specialist Group Newsletter 21 (1): 11-13. Current centres of distribution Auliya, M., 2003. Entdeckung des Sunda-Gavials (Crocodylia: Current strongholds for the species are few, but some remain Tomistoma schlegelii) im Ujung-Kulon National Park (Java, in several (restricted) regions of Borneo, including Central, Indonesien). ZGAP Mitteilungen 19: 2-10. East and West Kalimantan Provinces, in western Sarawak Barber, C. V. & J. Schweithelm, 2000. Trial by Fire. Forest Fires and in central and eastern Sumatra. The species appears to and Forestry Policy in Indonesia’s Era of Crisis and Reform. be relatively abundant (although difficult to detect) in some World Resources Institute (Forest Frontiers Initiative), sites in southern Borneo (such as Tanjung Puting National Washington. Park in Central Kalimantan), while in other regions in Borneo Bezuijen, M. R., 2004. Preliminary assessment of the status of the and Sumatra its abundance may be underestimated. In areas False Gharial (Tomistoma schlegelii) in the Mayeng Sarang where the species appears to have declined or become rare River, Grand Perfect SDN. BHD. Planted Forest Area, Bintulu (north-eastern and south-eastern Sumatra, Peninsular Division, Sarawak. Internal report for Grand Perfect SDN. BHD, Malaysia), this decline appears to have occurred relatively Kuching. recently, i.e. within the last 30-40 years. Local people along Bezuijen, M. R., P. Cannucciari, S. C. Manolis, Samedi, R. lowland forested waterways in eastern Sumatra report the Kadarisman & B. K. Simpson, 1995. Project Tomistoma. Field species to be ‘common’, but believe it has declined in the Expedition to the Lalan River and its tributaries, South Sumatra, last 20 years (Bezuijen, et al. 1997, 1998), because of Indonesia, August-October 1995: Assessment of the Distribution, extensive habitat loss, including regional forest fires such as Abundance, Status and Nesting Biology of the False Gharial in the 1990s (Barber and Schweithelm, 2000). (Tomistoma schlegelii). Unpublished Report, Wildlife Management International Pty Limited, Darwin. Bezuijen, M. R., P. Hartoyo, M. Elliott & B. A. Baker, 1997. Project ACKNOWLEDGMENTS Tomistoma. Second Report on the Ecology of the False Gharial (Tomistoma schlegelii) in Sumatera. Unpublished Report, We thank the following personnel and institutions for Wildlife Management International Pty Limited, Darwin. providing information on T. schlegelii specimens held in their Bezuijen, M. R., F. Hasudungan, R. Kadarisman, S. Wardoyo, G. J. collections (full names of institutions are given in Appendix W. Webb, Samedi & S. C. Manolis, 2002a. False Gharial 1): W. Böhme (ZFMK); J. Boundy (LDWF); N. Gilmore (Tomistoma schlegelii) surveys in southeast Sumatra, Indonesia (ANSP); Hamburg Museum; J. Hanken (MCZ); M. (1995-2002). Unpublished Report, Wildlife Management International Pty Limited, Darwin. Hoogmoed (NNM); B. Lardner (NHM); K. Lim (RMBR); U. Manthey (for Senckenberg and Berlin Museums); J. Bezuijen, M. R., R. Kadarisman, Yanto, Jamhari, F. Hasudungan, Rosado (MCZ); C. McCarthy (BMNH); A. Schmitz (MDH); K. Rauf, Samedi & G. J. W. Webb, 2001a. Rapid appraisal of the conservation status of the False Gharial (Tomistoma G. Schneider (UMMZ); J.E. Simmons (UKNHM); A. Snider schlegelii) in two locations in southeast Sumatra, Indonesia. (DZ); K. Tighe (USNM); J. Vindum (CAS); K. Zyskowski Unpublished report, Wildlife Management International Pty (PM); H.-J. Stibig (GVM, Global Vegetation Monitoring Limited, Darwin. Unit, Italy); R. Melisch (WWF Germany); GIS Unit (ZFMK). Bezuijen, M. R., R. B. Stuebing, M. Auliya & S. C. Manolis, 2003. We also thank the following persons for providing field False Gharial (Tomistoma schlegelii) conservation: priorities for records or other contributions in the field towards compilation action. Unpublished report, vers. 2 (June 2004). IUCN of this records: Aman; Jasmi Abdul; C. Brochu; A. Darubi; Tomistoma Task Force www.tomistoma.org. I. Das; V. Dinets; Eddie; A. Etin; R. F. Inger; Z. A. Jaafar; Bezuijen, M. R., G. J. W. Webb, P. Hartoyo & Samedi, 2001b. Peat K. Jensen; J. Jong; D. M. N Khalid; A. Kong; E. Lading; K. swamp forest and the false gharial (Tomistoma schlegelii) in Lazarus; W. Landong; B. L. Lim; A. Long; G. MacNair; S. the Merang River, eastern Sumatra, Indonesia. Oryx, 35: 301- C. Manolis; A. Resetar; S. A. M. Sah; B. Simpson. Professor 307. G. J. W. Webb and two anonymous reviewers gave critical Bezuijen, M. R., G. J. W. Webb, P. Hartoyo, Samedi, W. S. Ramono comments on a draft manuscript. This study was partially & S. C. Manolis, 1998. The False Gharial (Tomistoma schlegelii) funded by an award from the Field Museum of Natural in Sumatra. In: Crocodiles. Proceedings of the 14th Working History Bass Fund for Visiting Scholars, for which the authors Meeting of the Crocodile Specialist Group, IUCN-The World are grateful. Conservation Union. Pp. 10-31. IUCN/SSC Crocodile Specialist Group, Gland.

189 Stuebing et al.: Distribution of Tomistoma schlegelii

Bezuijen, M. R., P. Wibowo & H. Wirawijaya, 2002b (eds). Frazier, S. & R.A. Maturbongs, 1990. Report on an initial series of Prosiding Workshop Buaya Senyulong (2002b). Pengkajian crocodile surveys in East and Central Kalimantan, Indonesia Rencana Pengelolaan dan Konservasi Wilayah Sungai Merang (1 August-22 October 1990). Unpublished Report, FAO/PHPA sebagai Habitat Buaya Senyulong (Tomistoma schlegelii). Project No. GCP/INS/060/JPN, Jayapura. (Proceedings of the 2002 False Gharial Workshop: Assessment Galdikas, B. M. F. & C. P. Yeager, 1984. Crocodile Predation on of the Management and Conservation of the Merang River as a Crab-eating Macaque in Borneo. American Journal of habitat for the False Gharial (Tomistoma schlegelii). Wildlife Primatology 6: 49-51. Management International Pty Limited, Darwin & Wetlands International-Indonesia Program, Palembang. Geyh, M. A., H. R. Kudrass & H. Streif, 1979. Sea-level changes during the late Pleistocene and Holocene in the Strait of Malacca. Blower, J. H. & A. P. M. van der Zon, 1977. Proposed Ujung Kulon Nature, 278: 441-443. National Park including Gunung Honje, Pulau Peucang & Pulau Panaitan. Field Report of UNDP/FAO. Nature Government of Malaysia, 2004. Department of Statistics, Malaysia Conservation & Wildlife Management Project INS/73/013. FAO, web portal, http://www.statistics.gov.my/ Rome, 90 pp. Hadi, S., A. J. Hanson, Koesoebiono, M. Mahlan, M. Purba & S. Boulenger, G.A., 1889. Tomistoma. In: Catalogue of the Chelonians, Rahardjo, 1977. Tidal patterns and resource use in the Musi- Rhynchocephalians and Crocodiles in the British Museum Banyuasin coastal zone of Sumatra. Marine Research in (Natural History). Pp. 276-277. British Museum, London. Indonesia, 19: 109-135. Boulenger, G.A., 1896. On the occurrence of Schlegel’s Gavial Hanebuth, T., K. Stattegger & P. M. Grootes, 2000. Rapid flooding (Tomistoma schlegeli) in the Malay Peninsula, with Remarks of the Sunda Shelf: a late-glacial sea-level record. Science, 288: on the Atlas and Axis of the crocodilians. Proceedings of the 1033-1035. Zoological Society, XLI: 628-633. Harshman, J., C. J. Huddleston, J. P. Bollback, T. J. Parsons & M. Brochu, C. A., 2003. Phylogenetic approaches toward Crocodylian J. Braun, 2003. True and false gharials: A nuclear gene phylogeny of Crocodylia. Systematic Biology, 52: 386-402. history. Annual Review of Earth and Planetary Sciences, 31: 357-397. Hesp, P. A., C. H. Chang, M. Hilton, L. M. Chou & I. M. Turner, 1998. A first tentative Holocene sea-level curve for Singapore. Chan-Ard, T., W. Grossmann, A. Gumprecht, and K.-D. Schulz, Journal of Coastal Research, 14: 308-314. 1999. Amphibians and Reptiles of Peninsular Malaysia and Thailand: an illustrated checklist/Amphibien und Reptilien der Hoogerwerf, A., 1970. Udjung Kulon. The land of the last javan Halbinsel Malaysia and Thailands: eine illustriete Checkliste. rhinoceros. E. J. Brill, Leiden. Würselen: Bushmaster Publications. 240 pp. Hornaday, W. T., 1885. Two Years in the Jungle (Parts II & IV, Clarbrough, M. L., 1990. Ujung Kulon – Indonesia’s National Park, Malaya and Borneo). Oxford University Press, Singapore (1993). Handbook. Ministry of Forestry and Directorate General of 199 pp. Forest Protection and Nature Conservation, Indonesia & Hose, C., 1929. The Field Book of a Jungle-Wallah. Oxford Ministry of Foreign Affairs and Trade and Department of University Press, Singapore (1985 reprint). 216 pp. Conservation, New Zealand, 72 pp. Inger, R. F., R. B. Stuebing & Zainudin, R., 2005. Peat Swamp Claridge, G., 1994. Management of coastal ecosystems in eastern Frogs of Borneo. Proceedings of the International Conference: Sumatra: the case of Berbak Wildlife Reserve, Jambi Province. Wallace in Sarawak – 150 Years Later, 13-15 July, Kuching, Hydrobiologia 285: 287-302. Sarawak. Cox, J. H. & F. Gombeck, 1985. A Preliminary Survey of the Iskandar, D., 2001. Turtles and Crocodiles of Insular Southeast Asia Crocodile Resource in Sarawak, East Malaysia. IUCN/WWF and New Guinea. Institut Teknologi Bandung, Bandung. 191 Project No. MAL 74/85. Unpublished Report, World Wildlife pp. Fund Malaysia & the National Parks and Wildlife Office, Forest von Klein, 1863. Beiträge zur Osteologie der Crocodilschädel. II. Department, Sarawak. Aufsätze und Abhandlungen, pp. 70-100. In: Mohl v. H., H. v. Danielsen, F. & W. J. M. Verheugt, 1990. Integrating conservation Fehling, O. Fraas, F. Krauss & W. Menze (eds.) Jahreshefte and land-use planning in the coastal region of South Sumatra. des Vereins für vaterländische Naturkunde in Württemberg. XIX With contributions from H. Skov, R. Kadarisman, U. Suwarman Jahrgang. Verlag Ebner & Seubert, Stuttgart. & A. Purwoko. PHPA/Asian Wetland Bureau – Indonesia, Koningsberger, J., 1913. Java Zoologisch en Biologisch. Hoofdstuk Bogor. 17: De Zoetwaterfauna en de Detritusfauna van het De Rooij, N., 1917. The Reptiles of the Indo-Australian Archipelago. Cultuurgebied. Aflevering 8-9: 373-413. Drukkerij, Buitenzorg. Vol. I. Lacertilia, Chelonia, Embydosauria. E.J. Brill Ltd, (The saltwater and detrital fauna of rural areas. Chapter 17 in Leiden. Edition 8-9 of: The Zoology and Biology of Java. pp 373-413.). Endert, F.H., 1927. Een en ander over Krokodillen en Lading, E. & R. B. Stuebing, 1997. Nest of a False Gharial from Krokodillennesten in Koetai. De Tropische Natuur 16: 172-177. Sarawak. Crocodile Specialist Group Newsletter 16: 12-13. Frazier, S., 1994. A preliminary Dry Season Crocodile Survey of Mohr, Van der Meer, J.C., 1921. Een excursie in Krokodillenland. Suaka Margasatwa Danau Sentarum (Lake Sentarum Wildlife De Tropische Natuur, 10: 129-134. Reserve) in Kalimantan Barat, Indonesia. Unpublished Report Mucelli, J., 2005. Tomistoma in Vietnam. Crocodile Specialist for the Directorate-General of Forest Protection and Nature Group Newsletter, 24(1): 12. Conservation and the Asian Wetland Bureau, UK-Indonesia Tropical Forest Management Project, Bogor. Muin, A. & W. Ramono, 1994. Preliminary survey of Buaya Sumpit (Tomistoma schlegelii) and Buaya Kodok (Crocodylus Frazier, S., 2000. The crocodiles of Danau Sentarum, West siamensis) in East Kalimantan. Unpublished Report, Asian Kalimantan. Borneo Research Bulletin 31: 307-322. Conservation and Sustainable Use Group and Crocodile Specialist Group, Jakarta.

190 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Müller, S., 1838. Waarnemingen over de Indische Krokodillen en Simpson, B. K., A. Lopez, S. Latif & A. Yusoh, 1998. Tomistoma Beschrijving van Eene Nieuwe Soort. (Observations of the (Tomistoma schlegelii) at Tasek Bera, Peninsular Malaysia. In: Indonesian crocodiles and description of a new species). Crocodiles. Proceedings of the 14th Working Meeting of the Tijdschrift voor Natuurlijke Geschiedenis en Physiologie, 5: 61- Crocodile Specialist Group, IUCN-The World Conservation 87. Union. Pp. 32-45. IUCN/SSC Crocodile Specialist Group, Müller, S. & H. Schlegel 1845. De Krokodillen van het Indischen Gland. Archipel. 1-26, Plaat I-III. Smith, M. A., 1916. A list of the crocodiles, tortoises, turtles and Ramono, W. S., 1994. Tomistoma schlegelii in the provinces of lizards at present known to inhabit Siam. Journal of the Natural Sumatra Selatan and Jambi. Unpublished Report, Asian History Society of Siam, 2: 48-57. Conservation and Sustainable Use Group, Jakarta. Stibig, H.-J. & J.-P. Malingreau, 2003. Forest Cover of Insular Rieley, J. O. & S. E. Page (Eds), 1997. Biodiversity and Southeast Asia. Ambio, 32: 469-475. Sustainability of Tropical Peatlands. Proceedings of the Stibig, H-J., R. Beuchle & P. Janvier, 2002. Forest cover map of International Symposium on Biodiversity, Environmental insular southeast Asia at 1:5 500 000 derived from SPOT- Importance and Sustainability of Tropical Peat and Peatlands, Satellite images. TREES Publications Series D: Thematic Palangka Raya, Central Kalimantan, Indonesia 4-8 September outputs no.3. Institute for Environment and Sustainability, 1995. Samara Publishing Limited, Samara House, UK. European Commission. Ross, C. A. (ed), 1989. Crocodiles and Alligators. Facts on File., Steel, R., 1989. Crocodiles. Christopher Helm, London. 198 pp. Inc. New York. 240 pp. Strauch, A., 1866. Synopnis der gegenwärtig lebenden Crocodiliden Ross, C. A., J. Cox & H. Kurniati, 1996. Preliminary Survey of nebst Bemerkungen über die im Zoologischen Museum der Palustrine Crocodiles in Kalimantan. Project Progress Report kaiserlichen Akademie der wissenschaftlichen vorhandenen Phase I – 1995. Pusat Penelitian dan Pengembangan Biologi Repräsentanten dieser Familie. Mémoires de l’académie des (LIPI), Bogor & Smithsonian Institution, Washington. sciences de St. Pétersbourg, VII Série, 10: 1-120. Ross, C. A., J. H. Cox, H. Kurniati, & S. Frazier, 1998. Preliminary Stuebing, R. B., et al., 2003. Baseline Fauna survey at PT Maruwai Surveys of Palustrine Crocodiles in Kalimantan. In: Crocodiles. Coal Deposits (Pari, NE Lampunut and Lampunut Blocks), East Proceedings of the 14th Working Meeting of the Crocodile Kalimantan. Unpublished technical report, Wildlife Specialist Group, IUCN-The World Conservation Union. Pp. Managememtn International Pty. Ltd., Darwin (NT). 68pp. 46-79. IUCN/SSC Crocodile Specialist Group, Gland. Stuebing, R.,B., E. Lading & J. Jong, 1998. The Status of the False Ross, J. P. (ed), 1998a. Crocodiles. Status Survey and Conservation Gharial (Tomistoma schlegelii Mueller) in Sarawak. In: Action Plan. Second Edition. IUCN/SSC Crocodile Specialist Proceedings of the 14th Working Meeting of the Crocodile Group, Gland. Specialist Group, Singapore. Pp. 1-9. IUCN/SSC Crocodile Ross, J. P., 1998b. Meetings (notes of the 14th Working Meeting of Specialist Group, Gland. the CSG). Crocodile Specialist Group Newsletter 17: 3-7. Stuebing, R.,B., S. A. Mohd. Sah, E. Lading & J. Jong, 2004. The Sebastian, A. C., 1993a. The Crocodilians of Malaysia. A Review. Status of the False Gharial (Tomistoma schlegelii Mueller) in th In: Crocodiles. Proceedings of the 2nd Regional (Eastern Asia, Malaysia. In: Proceedings of the 16 Working Meeting of the Oceania, Australasia) Meeting of the Crocodile Specialist Crocodile Specialist Group, Darwin, NT, Australia. (in press). Group, IUCN-The World Conservation Union (not numbered). IUCN/SSC Crocodile Specialist Group, Gland. IUCN/SSC Crocodile Specialist Group, Gland. Sukumaran, J., 2002. The amphibian fauna of a forested area in Sebastian, A. C., 1993b. The Tomistoma or False Gharial Tomistoma Temengor, Perak, Malaysia, with the first record of Philautus schlegelii. The Need for its Conservation in South East Asia. parvulus (Boulenger, 1893) in the Malay Peninsular. In: Crocodiles. Proceedings of the 2nd Regional (Eastern Asia, Hamadryad, 27: 1-10. Oceania, Australasia) Meeting of the Crocodile Specialist Thorbjarnarson, J. B., Compiler, 1992. Crocodiles. An Action Plan Group, IUCN-The World Conservation Union (not numbered). for their Conservation. IUCN, Gland. IUCN/SSC Crocodile Specialist Group, Gland. WCMC, 2000. World Conservation Monitoring Centre, http:// Sebastian, A. C., 1994. The Tomistoma Tomistoma schlegelii in www.unep-wcmc.org. June 2004. Southeast Asia, a Status Review and Priorities for its th Weber, M., 1890. Reptilia from the Malay Archipelago. 1. Sauria, Conservation. In: Crocodiles. Proceedings of the 12 Working Crocodilidae, Chelonia. In: Weber, M. (Ed). Zoologische Meeting of the Crocodile Specialist Group, IUCN-The World Ergebnisse einer Reise in Niederländisch Ost-Indien. Pp 159- Conservation Union. Vol.1. Pp 98-112. IUCN/SSC Crocodile 177. E. J. Brill., Leiden. Specialist Group, Gland. Whitaker, R., 1984. Preliminary survey of crocodiles in Sabah, East Shelford, R. W. C., 1916. A Naturalist in Borneo. T. Fisher Unwin, Malaysia. IUCN/WWF Project No. 3127. World Wildlife Fund, London. xxvii (+ 1) + 331 pp + 32 pls. Kuala Lumpur. 69 pp. Simpson, B. K., 2004. False Gharials (Tomistoma schlegelii) in Whitmore, T. C., 1984. Tropical Rain Forests of the Far East. Tanjung Puting National Park, Kalimantan, Indonesia. In: Oxford University Press, Walton street, Oxford. Proceedings of the 16th Working Meeting of the Crocodile Specialist Group, Darwin, NT, Australia. (in press). IUCN/SSC Whitten, A. J., S. J. Damanik, J. Anwar & N. Hisyam, 1987. The Crocodile Specialist Group, Gland. Ecology of Sumatra. Gadjah Mada University Press, i-xvii, 583 pp. Simpson, B. K., A. Lopez, S. Latif & A. Yusoh, 1997. Project Tomistoma. Report on the Malayan False Gharial (Tomistoma Whitten, A. J., R. E. Soeriaatmadja & S. A. Ariff, 1996. Ecology of schlegelii) at Tasek Bera, Peninsular Malaysia. Unpublished Java. Periplus, Singapore. 969 pp. Report prepared for Wetlands International-Asia Pacific. Witkamp, H., 1925. Een en ander over Krokodillen in Koetai. De Wildlife Management International Pty Limited, Darwin. Tropische Natuur, Jaargung XIV. Aflevering II, Buitenzorg.

191 Stuebing et al.: Distribution of Tomistoma schlegelii

Voris, H. K., 2000. Maps of Pleistocene sea levels in Southeast Asia: CAS California Academy of Sciences, Berkeley shorelines, river systems and time durations. Journal of DZ Detroit Zoo Biogeography, 27: 1153-1167. HM Hamburg Museum Yap, S. Y., 2002. On the distributional patterns of Southeast-East FMNH Florida Museum of Natural History, Unversity of Asian freshwater fish and their history. Journal of Biogeography, Florida, Gainesville 29: 187-1199. LSUMZ Museum of Vertebrate Zoology, Louisiana State University, Baton Rouge Yeager, C.P., 1991. Possible antipredator behavior associated with MCZ Museum of Comparative Zoology, Harvard river crossings by proboscis monkeys Nasalis larvatus. University American Journal of Primatology, 24: 61-66. MDH Museum d’histoire naturelle, Genèva Zulkifli & Albertus, 2003. Laporan perjalanan survei oranghutan NNM Nationaal Natuurhistorisch Museum Leiden Nang Potan, das Sibau, 15-17 Ocktober 2003. World Wildlife NRM Naturhistoriska Riksmuséet, Stockholm Fund-Indonesia, Pontianak. (Report on an Oranghutan survey PM Peabody Museum, Yale University, New Haven in the Sibau River region). RMBR Raffles Museum of Biodiversity Research, Natonal University of Singapore SMF Senckenberg Museum UKNHM University of Kansas Natural History Museum APPENDIX 1 (Kansas, USA) UMMZ University of Michigan Museum of Zoology USNM National Museum of Natural History, Smithsonian Appendix 1. A gazetteer of localities for Tomistoma schlegelii based Institution, Wash. D.C. on museum specimens, field observations and local reports. ZFMK Zoologische Forschungsinstitut & Museum Alexander Koenig, Bonn Key (for appendix 1only) ZMB Zoologische Museum, Berlin ANSP Academy of Natural Sciences, Philadelphia BMNH British Museum of Natural History, London

192 THE RAFFLES BULLETIN OF ZOOLOGY 2006

eweh

T

Collector: W.L. Abbot Collector: W.L.

1937 Collector: unknown.

Ramono, 1994 Along Mahakam River

& Ramono, 1994

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

et al., 1996

Rooij, 1917 Collector: N. De Rooij

Rooij, 1917 Collector: N. De Rooij

Rooij, 1917 Collector: N. De Rooij

itkamp, 1925itkamp, “Alan” Creek, Mahakam

eager, 1991 eager,

Auliya, 2002a, b2002a, Auliya, Seen at Nyiur Swamp

Müller 1838Müller specimen. Near Muara Type

Simpson, 2004Simpson, near Simpan Kanan Creek Simpson, 2004Simpson, near Simpan Kanan Creek Simpson, 2004Simpson, near Simpan Kanan Creek Simpson, 2004Simpson, near Simpan Kanan Creek Simpson, 2004Simpson, Camp Leakey as mapping point Simpson, 2004Simpson, Camp Leakey as mapping point Simpson, 2004Simpson, Camp Leakey as mapping point

Simpson, 2004Simpson, Camp Leakey as mapping point Simpson, 2004Simpson, Camp Leakey as mapping point

Simpson, 2004Simpson, Camp Leakey as mapping point

Simpson, 2004Simpson, Camp Leakey as mapping point

Simpson, 2004Simpson, Camp Leakey as mapping point

W

Muin & Ramono, 1994

Ross et al., 1996River Barito Along

Muin & Ramono, 1994River? Mahakam of Tributary

Muin & Ramono, 1994

Ross et al., 1996 Ross et al., 1996

Endert, 1927Endert, E. Kalimantan Telen, Nest with eggs. Sg

Frazier, 2000 Frazier, Giessen in litt. to Frazier (2000) W.

Ross et al., 1996

NMNH, 1915 Collector: H.C. Raven.

2002 1937 NMNH,

1917 De

1996 Ross

Category Date Source Notes

E1

13'00"E 3

°

20'30"E 2

°

2108"E 3 1996 Ross

17'04"E 3 1996 Ross 17'04"E 3 1994 Muin

19'58"E 3 1996 Ross

46'00"E 3 1996 Ross

42'49"E 3 1996 Ross

42'49"E 3 1994 Muin &

47'50"E 3 1996 Ross

48'32"E 3 1996 Ross

45'27"E 3 1996 Ross

17'08"E 3 1996 Ross

55'00"E 2 2002 55'00"E 2 2002 55'00"E 2 2002 55'00"E 2 2002 50'00"E 2 2002 50'00''E 2 2002 50'00"E 2 2002

50'00"E 2 2002 50'00"E 2 2002

50'00"E 2 2002

50'00"E 2 2002

50'00"E 2 2002

4147" 3 1996 Ross

°

° °

°

°

°

°

°

°

°

°

°

° ° ° ° ° ° °

° °

°

°

°

°

42'E 2 1927

40'E 3 1996

25'E 1 NMNH

°

04'E 1 1838

20'E 1 SMF Collector: Schlegel?

20'E 1 1917 De

40'E 1 1991 Y

31'E 2 1925

15'E 3 1994

57'E 3 1996

30'E 3 1994

15'E 3 1994

15'E 3 1996

30'E 1 1917 De

00'E 3 1994

00'E 3 1996

15'E 1 1915

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

E1

S 115

°

°

S 105

°

40'00"S 105

48'S 105

50'00"S 111 50'00"S 111 50'00"S 111 50'00"S 111 47'00"S 111 47'00"S 111 47'00"S 111

47'00"S 111 47'00"S 111

47'00"S 111

47'00"S 111

47'00"S 111

30'S 116

51'S 116 27'21"S 116 27'21"S 116

15'00"S 116

3106"S 116

55'48"S 112

14'29"S 116

30'S 116

58'S 114

09'28"S 116

09'28"S 116

59'35"S 114

02'16"S 114

10'S 116

20'S 116

52'00"N 112

05'52"S 116

30'S 117

30'S 116 26'N 116

25'S 114

58'S 112

58'S 112 00'S 109

00'S 109

30'S 117 10'N 110 39'53"S 112

°

°

° ° ° ° ° ° °

° °

°

°

°

°

° ° °

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

° °

°

°

° °

°

° ° °

Latitude / Longitude

0 0

anjung Puting NP 4

Sg Dusun 1

Locality

Sg Sekonyer 2 Sg Sekonyer 2 Sg Sekonyer 2 Sg Sekonyer 2 Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2 Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2 Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2

Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2 Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2

Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2

Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2

Sg Simpan Kanan (Tanjung Puting NP)Sg Simpan Kanan (Tanjung 2

T

Atan Creek 0

Danau Belibis Lake / Sg Bongon Danau Belibis Lake / Sg Bongon Danau Anggelam 0

Danau Lamuda approx 1

Danau Mesangat 0

Danau Semati (Meliau) 0

Danau Tebrau 0

Danau Tempatung 0

Lahe 0

Muara Kaman 0

Muara Kaman 0

Sg Batampang 1

Sg Batilap 2

Sg Belayan 0

Sg Kahata 0

Sg Leboyan 0

Sg Liang Buaya 0

Sg Mahakam 0

Sg Sedulang 0 Sg Telen 0

Banjarmasin 3

Danau Sentarum NP 0

Pontianak 0

Samarinda 0

Sg Jongkong 0

Province

Province

Province Ujung Kulon NP 6

al Kalimantan Province

ral Kalimantan Province

ral Kalimantan Province

ral Kalimantan Province

ntral Kalimantan Province

est Java Province Batavia (= Jakarta) 06

est Kalimantan Province Danau Sentarum NP 0 est Kalimantan Province

est Kalimantan Provincetown Pontianak 0 est Kalimantan

est Kalimantan

est Kalimantan Province

est Kalimantan Province(Kapuas) Sanggau 0

entral Kalimantan Province

W

Political unit

Central Kalimantan Province Central Kalimantan Province Central Kalimantan Province Central Kalimantan Province

Central Kalimantan Province

Central Kalimantan Province

Central Kalimantan Province

Central Kalimantan Province

East Kalimantan Province

East Kalimantan Province East Kalimantan Province East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province

East Kalimantan Province East Kalimantan Province

South Kalimantan Province

W

W

Java Banten

Kalimantan

Kalimantan

Kalimantan C

Kalimantan

Kalimantan Kalimantan Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan

Kalimantan Kalimantan

Kalimantan

Kalimantan W

ndonesia ndonesia Java ndonesia Kalimantan Cent

ndonesia Kalimantan ndonesia Kalimantan ndonesia Kalimantan

ndonesia Kalimantan Centr

Indonesia Kalimantan

Indonesia Kalimantan Cent

Indonesia

Record Country Region 1I 2I 3I 4

5I 6I 7I 8 9I 10 Indonesia Kalimantan Cent 11

12 Indonesia Kalimantan Ce 13 Indonesia

14 Indonesia Kalimantan

15 Indonesia

16 Indonesia Kalimantan

17 Indonesia

18 Indonesia 19 Indonesia 20 Indonesia

21 Indonesia

22 Indonesia

23 Indonesia

24 Indonesia

25 Indonesia

26 Indonesia

27 Indonesia

28 Indonesia

29 Indonesia

30 Indonesia

31 Indonesia

32 Indonesia

33 Indonesia

34 Indonesia

35 Indonesia

36 Indonesia 37 Indonesia

38 Indonesia

39 Indonesia 40 Indonesia Kalimantan

41 Indonesia Kalimantan W 42 Indonesia Kalimantan W

43 Indonesia Kalimantan W

45 Indonesia Kalimantan

44 Indonesia Kalimantan W

193 Stuebing et al.: Distribution of Tomistoma schlegelii

near Tanjung Lhasa near Tanjung

Nest

Collector: L.P. De Bussy Collector: L.P. Collector: L.P. De Bussy Collector: L.P.

, 1996, photographed & reported 4.1 m indiv.

1997 Nest

1997 Nest

al., 1997

et al., 1996

Rooij, 1917 Collector: N. De Rooij

Rooij, 1917 Collector: N. De Rooij. near Sg Sambas

Cox unpubl. data, 1990

Ross et al., 1996

Auliya, 2000; ZFMK 66591

Ross, et al., 1996

Auliya, 2000 Lhasa Above Tanjung Zulkifli & Albertus, 2003 Zulkifli &

RMBR, 1905RMBR, Collector: unknown? - ROB?

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 2001

Bezuijen et al., 2002

Bezuijen et al., 1997specimen Captive

Bezuijen et al., 1997

Bezuijen et al.,

Independent Newspaper

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2002

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al.,

NMNH NMNH

Bezuijen et al., 1997

NMNH, 1901 Collector: J. Hurter

Bezuijen et al., 1997

2001

Category Date Source Notes

10'E 1 NRM Collector: E. Mjˆberg 00'E 1 HM, 1884/85Siemsen Collector:

15'E 1 1901

° °

°

45' E 1 SMF Collector: unknown? - ROB?

45'E 1 1906 45'E 1 1906

45'E 1 SMF Collector: unknown? - ROB?

°

° °

°

23'04.7"E 2 1996

22'03.6"E 2 1996

22'03.7"E 2 2001

22'03.7"E 2 1990 J.

20'47.2"E 2 2002

20'43.3"E 2 2001

20'43.3"E 2 2001

20'43.3"E 2 2001

21'02.3"E 2 2001

21'02.3"E 2 2001

20'46.6"E 2 2002

°

°

°

°

°

°

°

°

°

°

°

00'30"E 3

00'30"E 3 2001

09'00"E 2 1996 Ross

°

°

°

36'E 2 1996

57'E 1 1995

05'E 3 1996

00'E 1 1917 De

57'E 2 1996 58'E 2 2003

40'E 1 1917 De

40'E 3 1996

45'E 1 1905

15'E 2 1996

15'E 2 1996

15'E 2 1994

15'E 2 Bezuijen et al., 1997

30'E 2 1996

30'E 2 Bezuijen et al., 1997

40'E 2 1995

40'E 2 Bezuijen et

22'E 2 1993

22'E 2 Bezuijen et al., 1997

25'E 2 1985

25'E 3 1980

45'E 3 1996

45'E 3 Bezuijen et al., 1997

05'E 2 1985

°

°

°

°

° °

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

30'S 98

30'S 98

30'S 98

30'S 98

30'S 102

00'N 100 50'N 102

°

°

°

°

° °

°

1'S 102

25'S 109

25'S 109

48'00"S 112

58'N 112

49'N 119

55'N 119

58'N 112 02'N 112

40'S 103

20'20"S 104

20'20"S 104

22'55.1"S 104

23'43.5"S 104

23'49.0"S 104

23'49.0"S 104

25'37.2"S 104

25'S 102

25'S 102

25'S 102

25'S 102

28'S 102

00'S 102

05'S 102

05'S 102

1 22'52.4"S 104

22'52.4"S 104

22'52.4"S 104

23'41.7"S 104

23'41.7"S 104

22'43.9"S 104

48'S 105

48'S 105

42'S 105

42'S 105

45'S 102

45'S 102

55'N 100

°

°

°

°

°

°

° °

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

° °

°

°

°

°

°

°

°

°

°

°

°

°

Latitude / Longitude

approx 3 approx 3

Locality Sg Kapuas 0

Sg Kapuas 0

Sg Sekolat 0

Sg Sibau 0

Sg Tempuran 0

Singkawang 0

Jambi 1

Medan Perbaungan

est Kalimantan Province

est Kalimantan Province

est Kalimantan Province

est Kalimantan Province

est Kalimantan Province

est Kalimantan Province

est Kalimantan Province Sg Sibau 0 est Kalimantan Province Sg Sibau 1

Political unit W

W

W

W

W

W

Jambi Province

Jambi ProvinceJambi Air Hitam Dalam Sg 1

Jambi ProvinceJambi Air Hitam Dalam Sg 1

Jambi ProvinceJambi Air Hitam Laut Sg 1

Jambi ProvinceJambi Air Hitam Laut Sg 1

Jambi ProvinceJambi Air Hitam Laut Sg 1

Jambi ProvinceJambi Air Hitam Laut Sg 1

Jambi ProvinceJambi Air Hitam Laut Sg 1

Jambi Province Sg Alai 1

Jambi Province Sg Alai 1

Jambi Province Sg Alai 1

Jambi Province Sg Alai 1

Jambi ProvinceJambi Sg Batang Hari 1

Jambi Province Sg Belango 2

Jambi Province Sg Mentawak 2

Jambi Province Sg Mentawak 2

Jambi Province Sg Merangin 2 Jambi ProvinceJambi Sg Simpang Melaka 1

Jambi ProvinceJambi Sg Simpang Melaka 1

Jambi ProvinceJambi Sg Simpang Melaka 1

Jambi ProvinceJambi Sg Simpang Melaka 1

Jambi ProvinceJambi Sg Simpang Melaka 1

Jambi ProvinceJambi Sg Simpang Melaka 1

Lampung Province Sg Mesuji 3

Lampung Province Sg Mesuji 3

Lampung Province Sg Pengbuan 4

Lampung Province Sg Pengbuan 4

North Sumatra Province Deli (= Medan) approx 3

North Sumatra Province North Sumatra Province

North Sumatra Province Deli (= Medan) approx 3

North Sumatra ProvinceBalei Tandjung approx 3 Riau ProvinceEstate Indrapura approx 0 Riau Province Sg Gangsal 0

Riau Province Sg Gangsal 0

Riau Province Sg Indragiri approx 0 Riau Province Sg Kubu 1

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Record Country Region 46 Indonesia Kalimantan

47 Indonesia Kalimantan

48 Indonesia Kalimantan

49 Indonesia Kalimantan

50 Indonesia Kalimantan

51 Indonesia Kalimantan

52 Indonesia Kalimantan W 53 Indonesia Kalimantan W 54 Indonesia

55 Indonesia

56 Indonesia

57 Indonesia

58 Indonesia

59 Indonesia

60 Indonesia Sumatra

61 Indonesia Sumatra

62 Indonesia Sumatra

63 Indonesia Sumatra

64 Indonesia Sumatra

65 Indonesia Sumatra

66 Indonesia Sumatra

67 Indonesia Sumatra

68 Indonesia Sumatra

69 Indonesia Sumatra

70 Indonesia Sumatra 71 Indonesia Sumatra

72 Indonesia Sumatra

73 Indonesia Sumatra

74 Indonesia Sumatra

75 Indonesia Sumatra

76 Indonesia Sumatra

77 Indonesia Sumatra

78 Indonesia Sumatra

79 Indonesia Sumatra

80 Indonesia Sumatra

81 Indonesia

83 Indonesia Sumatra 84 Indonesia Sumatra

82 Indonesia

85 Indonesia 86 Indonesia Sumatra 87 Indonesia

88 Indonesia Sumatra

89 Indonesia Sumatra 90 Indonesia

194 THE RAFFLES BULLETIN OF ZOOLOGY 2006

1997 Nest

1997 Nest

Kadarisman pers. comm.

Cox unpubl. data

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al.,

Bezuijen et al., 1997

Bezuijen et al.,

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1995 Bezuijen et al., 1995 Bezuijen et al., 1995

Bezuijen et al., 1995 Bezuijen et al., 1995 Bezuijen et al., 1995 Bezuijen et al., 1995 Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 1997

Bezuijen et al., 2001a, b

Category Date Source Notes

08'42.9"E 2 1995

03'52.7"E 2 1995 03'52.7"E 2 1995 02'06.7"E 2 1995 02'06.7"E 2 1995 02'06.7"E 2 1995

00'49.4"E 2 1995

00'49.4"E 2 1995

00'49.4"E 2 1995

00'49.4"E 2 1995

58'54.5"E 2 1995

00'12.8"E 2 1995

00'12.8"E 2 1995

00'12.8"E 2 1995

01'12.7"E 2 1995

01'12.7"E 2 1995

03'52.7"E 2 1996

59'55.3"E 2 1996

00'12.8"E 2 1996

01'12.7"E 2 1996

01'12.7"E 2 1996

01'12.7"E 2 1996

00'37.9"E 2 2001

°

° ° ° ° °

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

14'21"E 2 1995

52'05"E 2 1995 52'05"E 2 1995

°

° °

36'E 2 1996

36'E 3 1996

38'E 2 1995

19'E 3 1996

37'E 2 1986

22'E 2 1996

22'E 2 1995

22'E 2 1993

22'E 2 1985

21'E 2 1996

21'E 2 1996

21'E 2 1996

21'E 2 1993

05'E 2 1995

05'E 2 1995

37'E 2 1995

45'E 2 1990 J.

37'E 2 Bezuijen et al., 1995

30'E 2 1993 R.

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

55'N 100

55'N 100

37'S 101

28'S 102

00'S 101

00'S 101

00'S 101

00'S 101

26'N 100

40'S 104

40'S 104

40'S 104

40'S 104

02'39"S 104

10'S 104

10'S 104

05'S 103

04'S 103

05'S 103

15'S 105

01'20"S 103 01'20"S 103 08'38.9"S 104

06'01.3"S 104 06'01.3"S 104 00'07.8"S 104 00'07.8"S 104 00'07.8"S 104

58'49.2"S 104

58'49.2"S 104

58'49.2"S 104

58'49.2"S 104

55'37.3"S 103

53'17.1"S 104

53'17.1"S 104

53'17.1"S 104

51'07.1"S 104

51'07.1"S 104

06'01.3"S 104

53'37.9"S 103

53'17.1"S 104

51'07.1"S 104

51'07.1"S 104

51'07.1"S 104

51'56.2"S 104

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

° ° °

° ° ° ° °

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

Latitude / Longitude

Locality

Political unit Riau Province Sg Kubu 1

Riau Province Sg Kubu 1

Riau Province Sg Siak 0

Riau ProvinceSibayang-bayang Sg 0

Riau Province Sg Teso 0

Riau Province Sg Teso 0

Riau Province Sg Teso 0

Riau Province Sg Teso 0

Riau ProvinceRiau Putih Pujud Tanah 1

South Sumatra Province Sg Benu 1

South Sumatra Province Sg Benu 1

South Sumatra Province Sg Benu 1

South Sumatra Province Sg Benu 1

South Sumatra Province Sg Kepahyang 2

South Sumatra Province Sg Lalan 2

South Sumatra Province Sg Lalan 2

South Sumatra Province Sg Lalan 2

South Sumatra Province Sg Lalan 2

South Sumatra Province Sg Lalan 2

South Sumatra Province Sg Lumpur 3

South Sumatra Province Sg Medak 2 South Sumatra Province Sg Medak 2 South Sumatra Province Sg Merang 2

South Sumatra Province Sg Merang 2 South Sumatra Province Sg Merang 2 South Sumatra Province Sg Merang 2 South Sumatra Province Sg Merang 2 South Sumatra Province Sg Merang 2

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 2

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

Sumatra

Sumatra

Sumatra Sumatra Sumatra

Sumatra Sumatra Sumatra Sumatra

Sumatra

Sumatra

Sumatra

ndonesia ndonesia ndonesia

ndonesia ndonesia ndonesia ndonesia Sumatra

Indonesia Sumatra

Indonesia

Indonesia

0

2I 3I 4

5I 6I 7I 8I 9

1I

Record Country Region 91 Indonesia Sumatra

92 Indonesia Sumatra

93 Indonesia Sumatra

94 Indonesia Sumatra

95 Indonesia Sumatra

96 Indonesia Sumatra

97 Indonesia Sumatra

98 Indonesia Sumatra

99 Indonesia Sumatra

100 Indonesia Sumatra

101 Indonesia

102 Indonesia

103 Indonesia Sumatra

104 Indonesia Sumatra

105 Indonesia Sumatra

106 Indonesia Sumatra

107 Indonesia Sumatra

108 Indonesia Sumatra

109 Indonesia Sumatra

11

11

11 11 11

11 11 11 11 11

120 Indonesia Sumatra

121 Indonesia Sumatra

122 Indonesia Sumatra

123 Indonesia Sumatra

124 Indonesia Sumatra

125 Indonesia Sumatra

126 Indonesia Sumatra

127 Indonesia Sumatra

128 Indonesia

129 Indonesia Sumatra

130 Indonesia Sumatra

131 Indonesia

132 Indonesia Sumatra

133 Indonesia Sumatra

134 Indonesia Sumatra

135 Indonesia Sumatra

195 Stuebing et al.: Distribution of Tomistoma schlegelii

fice

fice

Collector: Chancellor & Stuart

Photos of eggs & hatchlings shown to M.B.

Photos at State wildlife of

, 2003), Photos at State wildlife of

1905 Eggs

Hasan pers. comm., 1994

Cox unpubl. data

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Bezuijen et al., 2001a, b

Ramono, 1994

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995

Bezuijen et al., 1995 Branch of Calik River FMNH FMNH Collector: Chancellor & Stuart FMNH Collector: Chancellor & Stuart Bezuijen et al., 1997

RMBR, 1940unknown Collector:

BMNH, 1896Wray Collector: L.

Boulenger, 1896Boulenger, Skull, skin

Butler, Butler,

Simpson et al., 1997

Category Date Source Notes

44'E 1 1929 44'E 1 1929 44'E 1 1929

° ° °

03'06.2"E 2 2001

00'44.1"E 2 2001

03'06.2"E 2 2001

03'06.2"E 2 2001

03'06.2"E 2 2001

00'12.8"E 2 2001

00'12.8"E 2 2001

00'12.8"E 2 2001

00'12.8"E 2 2001

00'12.8"E 2 2001

58'54.5"E 2 2001

58'54.5"E 2 2001

58'54.5"E 2 2001

58'54.5"E 2 2001

°

°

°

°

°

°

°

°

°

°

°

°

°

°

36'48"E 3 Simpson et al., 1997

36'50"E 3 Simpson et al., 1997

52'17"E 2 1995

52'17"E 2 1995

22'00"E 2 1995

22'00"E 2 1995

°

°

°

°

°

°

30'E 1 1899

°

38'E 3 Simpson et al., 1997

35'E 3 Simpson et al., 1997

16'E 3 Simpson et al., 1997

28'E 3 Simpson et al., 1997

38'E 1 Simpson et al., 1997photographed jaw Lower

38'E 3 Simpson et al., 1997

20'E 3 Simpson et al., 1997

32'E 1 1940

32'E 1 Simpson et al., 1997photographed Skull

50'E 1 1896

57'E 2 2003 (Stuebing

03'E 1 1896

07'E 2 2003 (Stuebing, 2003)+I33

10'E 3 1994

34'E 3 Simpson et al., 1997

03'E 2 1990 J.

14'E 1 SMF Collector unknown. Locality “near Nuni”(?)

50'E 2 1994

12'E 2 1995

20'E 3 1996 43'E 3 1994 Mr

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

° °

01'S 104 01'S 104 01'S 104

° ° °

25'N 101

1'N 102

°

51'37.7"S 104

50'43.8"S 104

50'43.8"S 104

50’43.8"S 104 50'43.8"S 104

53'17.1"S 104

53'17.1"S 104

53'17.1"S 104

53'17.1"S 104

53'17.1"S 104

55'37.3"S 103

55'37.3"S 103

55'37.3"S 103

55'37.3"S 103

06'S 104

05'S 103

15'S 105

44'27"S 103

44'27"S 103

29'00"S 104

29'00"S 104

35'S 104

55'S 101 15'S 101

09'50"N 102

31'N 102

57'N 102

45'N 103

32'N 103

57'N 102

57'N 102

53'N 103

1 09'16"N 102

13'N 102

13'N 102

13'N 100

48'N 100

20'N 101

59'N 101

23'N 101

°

°

°

° °

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

°

° °

°

°

°

°

°

°

°

°

° °

°

°

°

°

°

°

°

Latitude / Longitude

Locality

est Sumatra Province Sg Pangean 0 est Sumatra ProvinceTunjuan Sg 1

Political unit South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1 South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 1

South Sumatra Province Sg Merang 2

South Sumatra Province Sg Muara Klingi 3

South Sumatra Province Sg Pedada 3

South Sumatra Province Sg Pejudian 1

South Sumatra Province Sg Pejudian 1

South Sumatra Province Sg Serdang 2

South Sumatra Province Sg Serdang 2

South Sumatra ProvinceTungkal Sg 2

South Sumatra Province upper Sg Ogan approx 3 South Sumatra Province upper Sg Ogan approx 3 South Sumatra Province upper Sg Ogan approx 3 W W

Malaysia Pahang State Sg Jelai 3

Malaysia Pahang State Sg Jengka 3

Malaysia Pahang State Sg Lepar 4

Malaysia Pahang State Sg Luit 4

Malaysia Pahang State Sg Pahang 3

Malaysia Pahang State Sg Rasau 2

Malaysia Pahang State Sg Rasau 2

Malaysia Pahang State Sg Rompin 2

Malaysia Pahang State Sg Serting 3 Malaysia Pahang State Sg Tasek 3

MalaysiaState Pahang Bera NP Tasek 3

MalaysiaState Pahang Bera NP Tasek 3

Malaysia Perak State Pulau Sg Tiga, Perak 4

Malaysia Perak State Sg Erung 3

Malaysia Perak State Sg Kinta 4

Malaysia Perak State Sg Sungkai 3

Malaysia Selangor State Parit Champedak ~ 3 Malaysia Selangor State Selangor Sg Tengi, 3

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra

Sumatra Sumatra Sumatra

Record Country Region 136 Indonesia Sumatra

137 Indonesia Sumatra

138 Indonesia

139 Indonesia Sumatra 140 Indonesia Sumatra

141 Indonesia Sumatra

142 Indonesia Sumatra

143 Indonesia Sumatra

144 Indonesia

145 Indonesia Sumatra

146 Indonesia Sumatra

147 Indonesia Sumatra

148 Indonesia Sumatra

149 Indonesia Sumatra

150 Indonesia Sumatra

151 Indonesia

152 Indonesia Sumatra

153 Indonesia

154 Indonesia Sumatra

155 Indonesia Sumatra

156 Indonesia Sumatra

157 Indonesia

158 Indonesia 159 Indonesia 160 Indonesia 161 Indonesia Sumatra 162 Indonesia Sumatra 163 Malaysia P.

164 Malaysia P.

165 Malaysia P.

166 Malaysia P.

167 Malaysia P.

168 Malaysia P.

169 Malaysia P.

170 Malaysia P.

171 Malaysia P. 172 Malaysia P.

173 Malaysia P.

174 Malaysia P.

175 Malaysia P.

176 Malaysia P.

177 Malaysia P.

178 Malaysia P.

179 Malaysia P. 180 Malaysia P.

196 THE RAFFLES BULLETIN OF ZOOLOGY 2006

Juvenile

Unconfirmed

Collector: C. Hose

Collector: W.T. Hornaday Collector: W.T.

, 1889 Collector: E.R. Stilwell

1999

& Gombek, 1985

& Gombek, 1985

& Gombek, 1985 & Gombek, 1985 & Gombek, 1985

Rooij, 1917 Collector: N. De Rooij

Rooij, 1917 Collector: N. De Rooij

Stuebing, pers. data 2002

Stuebing, pers. obs. 2004

Lading, pers. comm.

Das & K. Jensen, pers. obs

Chan-ard et al.,

Zainal Abidin, pers.comm. 2003 Zainal

Bezuijen, 2004 Bezuijen, 2004

Bezuijen, 2004

Bezuijen, 2004

Bezuijen, 2004 Bezuijen, 2004

Bezuijen, 2004

SMF Collector: unknown

RMBR, 1905”unknown Collector:

BMNH, 1893A. Everett Collector:

BMNH, 1900BMNH, Collector: C. Hose BMNH,1900

ANSP, 1897 ANSP, Collector: C. Hose

BMNH, 1897; Boulenger

Sebastian, 1993a, b Sebastian, 1993a, b

Category Date Source Notes

1'E 3 2003

45'E 1 1999

39'E 3 2002 R.

15'E 1 1917 De

26'E 3 2004 R.

03'E 2 2000 03'E 2 2000

03'E 2 2001

03'E 2 2003

03'E 2 2003 03'E 2 2004

03'E 2 2004

35'E 2 2004 I.

00'E 1 pre-1900

40'E 2 1985 Cox 40'E 1 1905

40'E 1 1893

40'E 1 1900 40'E 1 1900

40'E 1 1897

40'E 1 1897

40'E 1 1917 De

27'E 1 NMNH

45'E 3 1985 Cox

1

30'E 3 1997 Cox

25'E 3 1985 Cox 20'E 3 1985 Cox

00'E 3 1993 05'E 3 1985 05'E 3 1998 E.

°

° °

°

°

°

° °

°

° °

°

°

°

°

° °

°

° °

°

°

°

°

°

° °

° ° °

55'N 102

29'N 118 26'N 115

45'N 112

55'N 113

39'N 113

39'N 113 39'N 113

39'N 113

39'N 113

39'N 113 39'N 113

20'N 111 55'N 114

40'N 114

25'N 110

30'N 110 30'N 110

30'N 110

30'N 110 30'N 110

30'N 110

30'N 110

38'N 110 15'N 111 20'N 111 15'N 110 30'N 111 40'N 111 40'N 111

°

° °

°

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Latitude / Longitude

asek Kenyir 5

Locality

Mukah 2

est Coast Residency Sg Klias 5

rengganu T

Political unit

Sandakan Residency Sg Kinabatangan 5

W

Bintulu Division

Bintulu Division Sg Penyilam 2

Bintulu Division Sg Sarang 2

Bintulu Division Sg Sarang 2 Bintulu Division Sg Sarang 2

Bintulu Division Sg Sarang 2

Bintulu Division Sg Sarang 2

Bintulu Division Sg Sarang 2 Bintulu Division Sg Sarang 2

Bintulu Division Sg Tisak 1 Loagan Bunut 3

Miri Division Sg Baram 4

Semarahan Division Sg Ensengai 1

Semarahan Division Sg Sadong 1 Semarahan Division Sg Sadong 1

Semarahan Division Sg Sadong 1

Semarahan Division Sg Sadong 1 Semarahan Division Sg Sadong 1

Semarahan Division Sg Sadong 1

Semarahan Division Sg Sadong 1

Semarahan Division Sg Sarawak 1 Sg Dor 1 Sri Aman Division Sg Kelauh 1 Sri Aman Division Sg Kerang 1 Sri Aman Division Sg Lupar 1 Sri Aman Division Sg Maludam 1 Sri Aman Division Sg Maludam 1

Malaysia T

Record Country Region 181 Malaysia P.

182 Malaysia Sabah

183 Malaysia Sabah

184 Malaysia Sarawak

185 Malaysia Sarawak

186 Malaysia Sarawak

187 Malaysia Sarawak 188 Malaysia Sarawak

189 Malaysia Sarawak

190 Malaysia Sarawak

191 Malaysia Sarawak 192 Malaysia Sarawak

193 Malaysia Sarawak 194 Malaysia Sarawak

195 Malaysia Sarawak

196 Malaysia Sarawak

197 Malaysia Sarawak 198 Malaysia Sarawak

199 Malaysia Sarawak

200 Malaysia Sarawak 201 Malaysia Sarawak

202 Malaysia Sarawak

203 Malaysia Sarawak

204 Malaysia Sarawak 205 Malaysia Sarawak 206 Malaysia Sarawak 207 Malaysia Sarawak 208 Malaysia Sarawak 209 Malaysia Sarawak 210 Malaysia Sarawak

197

THE RAFFLES BULLETIN OF ZOOLOGY 2006

THE RAFFLES BULLETIN OF ZOOLOGY 2006 54(1): 199-202 Date of Publication: 28 Feb.2006 © National University of Singapore

BOOK REVIEW

A Photographic Richly illustrated with almost all living material, Guide to the photographed from various angles showing different life Inlandwater Fishes of stages (e.g. Metzia mesembrina) and developmental stages Taiwan. Volume 1. of tubercle formation (e.g. Candidia barbata). The Cypriniformes. photographers are highly commended for their patience to By Chen, I-S. & Y.-C. get properly acclimatized fish and some species even in their Chang, 2005. Managing breeding colours. Most species accounts are also editor: Shao, K.-T. The accompanied by a habitat picture. Shueichan Press, Taiwan R.O.C., xx+284 pages. ISBN: 957-8596-72-3, obtainable This publication also includes 2 appendices, covering 10 from www.scppress.com, priced at USD50. introduced and 3 doubtful species. There are a total of 5 species with uncertain identifications, highlighting the current This is a medium sized (26 x 20 cm) bilingual (in Chinese taxonomical status. and English) volume lavishly filled with colour plates, covering the cypriniform fishes of Taiwan. This is the first One shortcoming of this vivid volume is the lack of a general of five volumes covering all the inland freshwater fish species map or individual maps showing distribution of the fish of Taiwan. Volume 1 covers only the cypriniform fishes, species. with a total of 45 species (35 cyprinids, 6 balitorids and 4 cobitids). Volume 2 covering the Gobiidae, volume 3 the This is a delightful volume to read and browse through for miscellaneous taxa, volume 4 the anguillid and synbranchid the layperson, hobbyist and scientist. eels, and volume 5 the phylogeny of Taiwanese freshwater fish.

There are between 4 to 6 full pages devoted to each species. Each species is listed by both scientific and vernacular names, including sub-family designation. Each species account is Tan Heok Hui further divided into three sections, covering morphology, Department of Biological Sciences distribution, and ecology and remarks. The authors had spent National University of Singapore much time here and list some very valuable ecological Kent Ridge 119260 observations of the fish species. Republic of Singapore

199 Book Review

A Guide to Gobies of Singapore. This book is a cumulative effort of 10 years Singapore. between the two authors, one based in Darwin, Australia, and By Larson, H. K. & K. K. the other in Singapore. P. Lim, 2005. Singapore Science Centre, 164 The habitats of the gobies in general and a fold-out map of pages. ISBN: 981-05- Singapore are provided, giving readers an idea of the diversity 3407-8. of habitats gobies are found in. Another fold-out of the general morphology of gobies is included to give readers an This is a small pocket- idea of the variable body forms of gobies. sized volume richly filled with colour plates, A total of 106 species is covered with one species per page, covering the freshwater, most with life coloration or preserved specimens depicted, brackish water and along with numerous line drawings. Each species is listed marine gobies of with both vernacular and scientific names; followed by a brief Singapore. This is the diagnosis, description and distribution. 40th book in the series of guides to Singapore This is a most compact and comprehensive volume to peruse sponsored by BP. for the layperson, hobbyist and gobiologist.

This book covers the 3 families from the sub-order Gobioidei found in Singapore – Eleotridae, Gobiidae and Tan Heok Hui Microdesmidae. The family Gobiidae is the largest in the Department of Biological Sciences sub-order and most gobies are typified by having two dorsal National University of Singapore fins and fused pelvic fins. The authors had provided a Kent Ridge 119260 comprehensive taxonomic key to the 60 genera located from Republic of Singapore

A first look at the fish Over the course of the surveys, a total of 47 fish species was species of the middle obtained. Fishes from the Cyprinidae family consist the bulk Malinau. Taxonomy, of catch (68%). There were also possible new species ecology, vulnerability obtained (e.g. Puntius and Gastromyzon). Species potentially and importance. vulnerable to logging activities were highlighted. Impacts By Rachmatika, I., R. of logging on the fish fauna were also discussed. Logged Nasi, D. Sheil & M. habitats yield lower diversity and abundance of fish species. Wan, 2005. Center for International Forestry Two appendices were included. Appendix 1 listing the Research (CIFOR), collection stations. Appendix 2 provided an annotated list of Bogor, Indonesia. vi+34 fish species obtained. The systematics used is not updated, pages. e.g. Belontiidae had been synonymised with Osphronemidae, www.cifor.cgiar.org Barbonymus is a senior synonym to Barbodes. The species ISBN: 979-3361-67-0. list is listed alphabetically by family name and not in the taxonomic order. This is an A4-sized volume covering the Despite the few shortcomings, this is a timely publication to freshwater fishes of the middle Malinau (tributary of the illustrate the real state of knowledge we have of the tropical Sesayap River), East Kalimantan, Indonesia. biodiversity in Southeast Asia. It is a commendable effort for the authors to publish this despite restricted access to This is basically a technical report based on field collections updated publications and taxonomical help. conducted in 1999 and 2000. The site was chosen because it was adjacent to the Kayan Mentarang National Park. Tan Heok Hui Surveys were conducted in both pristine and logged habitats Department of Biological Sciences of the Seturan catchment for comparisons. The local Punan, National University of Singapore Merap and Kenyah communities were also interviewed and Kent Ridge 119260 socio-economic data gathered. Republic of Singapore

200 THE RAFFLES BULLETIN OF ZOOLOGY 2006

An Atlas of parts – toxic, medicinal and dangerous fishes. The first part Ichthyotoxic, Medicinal deals with ciguatoxic fishes, covering 9 sections – flesh-toxic, and Dangerous Fishes. tetraodontoxic, ichthyootoxic, gall bladder-toxic, By Wu, H.-L., D.-H. ichthyohemotoxic, ichthyohepatotoxic, pelagic species-toxic, Chong, Y. Mou, Y.-H. gempylotoxic and carchatoxic species. The second part deals Chen & Y. Ni, 2005. with fishes of medicinal properties. The third part deals with Chief Editor: Wu, H.-L. dangerous fishes, covering 4 sections – biting, electrical, saw Shanghai Scientific & snouts and parasitic species. Technical Publishers, Shanghai, v+482 pages. All species come with a one page account, starting with the ISBN: 7-5323-7996-5 [in Chinese vernacular and scientific names, then describing Chinese]. morphology, habit, first aid technique or medicinal benefits. All species accounts are illustrated with line drawings. This is a small hard- cover volume (15 x 21 This is a very interesting and informative volume for the cm) dealing with all toxic layperson, hobbyist, scientist and gastronomist. fishes, but also covering fishes consumed for medicinal purposes. Tan Heok Hui Department of Biological Sciences The colour plates are situated at the start, with 8 pages National University of Singapore depicting 31 species. This mini-tome covers 422 species of Kent Ridge 119260 freshwater and marine fishes. The atlas is divided into 3 main Republic of Singapore

Development of Lanjak Several tables are provided listing the fish obtained from the Entimau Wildlife abovementioned areas. Table 2 provides a fish list from Sanctuary as a Totally Lanjak Entimau Wildlife Sanctuary, of which several species Protected Area. Fish are of questionable identities; as follows: Resources Assessment Gastromyzon contractus – as from illustrations from plate Study of Lanjak 25, it is G. fasciatus; Entimau Wildlife Gastromyzon ridens – from current understanding (and Sanctuary and Batang unpublished data), it is restricted to the Kapuas basin; Ai National Park. All three species of Glaniopsis – from current By Ivy S Abdullah, 2004. understanding (and unpublished data), they are all International Tropical restricted to their respective type localities and basins. Timber Organization. However, these cannot be confirmed till the actual Forestry Department, specimens are examined. Sarawak, Malaysia. ITTO project PD16/99 Table 4 provides a fish list from Batang Ai National Park, of rev. 2(F), phase III, which several species are of questionable identities; as vii+70 pages. follows: Akysis baramensis – this is a rather southern distribution This is a well illustrated medium sized volume covering the from the type locality, more detailed examination required freshwater fishes of the catchment area of the Lanjak Entimau to clarify the identity; Wildlife Sanctuary (Rejang basin) and Batang Ai National Gastromyzon borneensis - from current understanding Park (Lupar basin). (and unpublished data), it is restricted to western Sabah; Gastromyzon ridens – from current understanding (and This publication covers mainly the fish diversity, unpublished data), it is restricted to the Kapuas basin; recommendations for conservation and possible applications All three species of Glaniopsis – from current in aquaculture and aquarium trade. The fish list draws from understanding (and unpublished data), they are all several earlier unpublished reports and substantiates the restricted to their respective type localities and basins. findings with more records and lists possible new species. However, these cannot be confirmed till the actual specimens are examined. The maps provided are very detailed contour maps of poor resolution. Simpler line drawing maps would have been At least one reference is missing, e.g. Roberts, 1980, in which sufficient. the year could be wrongly quoted. This publication suffers

201 Book Review from outdated taxonomical names and identifications, as This publication also lists good recommendations for the follows: conservation of the areas mentioned. Also providing avenues Plate 3 – should be Puntius banksi juvenile for local people’s involvement through aquaculture of local Plate 5 – should be Lobocheilos sp. fish species, e.g. Tor spp. Plate 6 – should be Osteochilus sarawakensis Plate 10 – should be Auriglobus nefastus Despite these shortcomings, this is a timely publication to Plate 13 – should be Rasbora agyrotaenia illustrate the real state of knowledge we have of the tropical Plate 15 – should be Rasbora caudimaculata biodiversity in Southeast Asia. It is a commendable effort Plate 18 – should be Glyptothorax major for the author to publish this despite restricted access to Plate 20 – should be Clarias leiacanthus updated publications and taxonomical help. Plate 21 – should be Barbonymus collingwoodi Plate 24 – should be Schismatorhynchos holorhynchos Plate 25 – should be Gastromyzon fasciatus Plate 33 – should be Homaloptera nebulosa Plate 35 – should be Nemacheilus saravacensis Plate 36 – should be Neogastromyzon cf. pauciradiatus Tan Heok Hui Plate 37 – should be Leiocassis poecilopterus Department of Biological Sciences Plate 38 – should be Pseudomystus rugosus National University of Singapore Plate 45 – should be Macrognathus circumcinctus Kent Ridge 119260 Plate 47 – should be Schismatorhynchos holorhynchos Republic of Singapore

202