taxonomy and biogeography of the subgenus eopaederus of the

Ann. soc. entomol. Fr. (n.s.), 2008, 44 (3) : 327-344

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ARTICLE

Taxonomy and biogeography of the subgenus Eopaederus of the genus Paederus Fabricius (Coleoptera: Staphylinidae: Paederinae) from China

Abstract. The subgenus Eopaederus Scheerpeltz 1957 belongs to the genus Paederus sensu stricto (Coleoptera: Staphylinidae: Paederinae) and exhibits a wide geographical distribution from the far east of Asia through West Europe. In this paper, we describe two new species: Paederus (Eopaederus) noncurvatus n. sp. and P. (E.) curvatus n. sp. from Hubei in the middle of China. The species P. (E.) basiventris Bernhauer 1933 is resurrected to a valid species and not anymore a synonym of P. (E.) nigricornis Bernhauer 1911. Other two species, P. (E.) birmanus Fauvel 1895 and P. (E.) alternans Walker 1858 are designed to the subgenus Eopaederus based on the materials collected respectively from Yunnan and Hainan, South China, whereas the former P. (E.) birmanus Fauvel 1895 is the fi rst-time-record species to the Chinese fauna. A key to nine Chinese species is presented. Zoogeographic analyses show that the subgenus track can be divided into three parts (namely Asia, Middle East and Europe) and each one possesses its own endemic species. Likewise, endemic species ranges can also designed as Europe type, Middle East type, or South and East Asia type. Along the south boundary of Himalayas (Qinghai-Tibet Plateau) through South China, species richness and endemism are obviously higher than in other regions.

Résumé. Taxonomie et biogéographie du genre Paederus Fabricius, sous-genre Eopaederus (Coleoptera : Staphylinidae : Paederinae) de Chine. Le sous-genre Eopaederus Scheerpeltz 1957 appartient au genre Paederus sensu stricto (Coleoptera : Staphylinidae : Paederinae) et présente une vaste répartition géographique depuis l’Extrême-Orient jusqu’en Europe Occidentale. Dans cet article, nous décrivons deux nouvelles espèces : Paederus (Eopaederus) noncurvatus n. sp. et P. (E.) curvatus n. sp. du Hubei, au centre de la Chine. L’espèce P. (E.) basiventris Bernhauer 1933 est restaurée comme espèce valide et non plus comme un simple synonyme de P. (E.) nigricornis Bernhauer 1911. Deux autres spèces, P. (E.) birmanus Fauvel 1895 et P. (E.) alternans Walker 1858, sont assignées au sous-genre Eopaederus sur la base de matériel, respectivement, du Yunnan et du Hainan, au sud de la Chine. P. (E.) birmanus Fauvel 1895 est signalé pour la première fois de la faune de Chine. Une clé des 9 espèces de Chine est présentée. L’analyse zoogéographique montre que la route du sous-genre peut être subdivisée en 3 parties, qui correspondent à l’Asie, au Moyen-Orient et à l’Europe, avec chaque fois des espèces endémiques. On peut donc distinguer des espèces de type européen, de type moyen-oriental et de type sud- et est-asiatique. Tout le long de la limite sud de l’Himalaya (plateau du Qinghai-Tibet) jusqu’au sud de la Chine, la richesse en espèces et l’endémisme sont de toute évidence plus élevés que dans d’autres régions. Keywords: Paederus, Eopaederus, taxonomy, biogeography, China.

Xiao-Yan Li (1, 2) & Hong-Zhang Zhou (1)*

(1) Institute of Zoology, Chinese Academy of Sciences, Da-Tun-Lu, Chao-Yang, 100101 Beijing, P.R.China(2) Graduate School of the Chinese Academy of Science, 19 Yuquan Rd., Shijingshan, 100039 Beijing, P. R. China

* Correspondence author

E-mail: [email protected]é le 17 avril 2008

The genus Paederus s. str. Fabricius 1775 is one of very diverse and large groups in the rove beetle

subfamily Paederinae (Coleoptera: Staphylinidae). Th is genus is a successful group well adapted to habitats associated with moist environment. Currently, eleven subgenera were erected to accommodate the species of the Paederus s. l. (Scheerpeltz 1957, 1976; Coiff ait 1982; Boháč 1985; Frank 1988; Newton et al. 2000; Newton 2003; Willers 2001; Herman 2003; Löbl & Smetana 2004; Li & Zhou 2007).

Th e subgenus Eopaederus was erected by Scheerpeltz (1957) and Paederus (E.) caligatus Erichson 1840 from France was designated as the type species originally. Before our study, nine species were included in this subgenus: P. (E.) balachowskyi (Jarrige 1971) from Iran, P. (E.) debilior Eppelsheim 1892 from Turkestan, P. (E.) limnophilus Erichson 1840 from Saxonia, P. (E.) mesopotanicus Eppelsheim 1889 from Iraq, P. (E.) basalis Bernhauer 1914 from India, P. (E.) nigricornis Bernhauer 1911 from Nepal, P. (E.) pubescens Cameron 1914 from India, and P. (E.) tamulus Erichson 1840 from Singapore; the latter four species were recorded in China.

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X.-Y. Li & H. -Z. Zhou

In this paper, we describe two new species: P. (E.) noncurvatus n. sp. and P. (E.) curvatus n. sp., from Hubei province in the middle of China. Th e species P. (E.) basiventris Bernhauer 1933, recorded in Sichuan and Xizang (=Tibet), Southwestern China, is now resurrected to a valid species and not anymore a synonym of P. (E.) nigricornis Bernhauer 1911. Other two species, P. (E.) birmanus Fauvel 1895 and P. (E.) alternans Walker 1858 are now designed to the subgenus Eopaederus based on the materials collected respectively from Yunnan and Hainan, South China, whereas the former, P. (E.) birmanus Fauvel 1895 is the fi rst-time-record species to the Chinese fauna of the subgenus. Th us, the species number of this subgenus is increased to fourteen and nine of them recorded in China. A key to nine Chinese species is presented.

Zoogeography is of great signifi cance in studies of species origin and diversifi cation. Track analyses based on geographical records of species distributions, or frankly, comparing species ranges (scales, patterns and degree of endemism), always give us valuable cues when we try to distinguish close-related species, to recognize true monophyletic grades or taxa, and to establish really “natural” and wide acceptable classifi cations. As a matter of fact, zoogeography of Paederus was poorly studied and even totally a gap with respect to the subgenus Eopaederus, or to the Chinese fauna, even though the subgenus has a rather wide geographical distribution from the far east of Asia through West Europe. Here, we analyzed distribution patterns of the subgenus and hope that these general patterns and some other zoogeographic characteristics may benefi t our future studies. Th e result can also off er us in return a kind of support for the new species reported here.

Materials and MethodsSpecimens were relaxed in warm water (60 °C) for about 5–8 hours. It was cleared 5 minutes in 10% KOH, and then put in 75% alcohol (ready for dissection). Specimens were dissected to observe morphology of mouthparts, 8th-9th abdominal segments in both sexes, and aedeagus in males. After observations, diff erent parts of the dissected specimens were stored permanently in glycerin for future studies. Observations and drawings were done under a compound microscope (LEICA MZ-APO). Measurements (in millimeter, signed as mm) were done at 40× magnifi cation. Most of the specimens used in this study, including the types of the new species, are deposited in IOZ-CAS. Th e types of new taxa are deposited in the IOZ-CAS (Institute of Zoology, Chinese Academy of Sciences, Beijing, China).Th e distribution of Eopaederus Scheerpeltz was studied and analyzed using the GIS method according to the geographic distribution database of museum specimens and published literatures. We also used data of quantitative of specimens which were borrowed from MNHUB, NHMW, NHMG, DEI. In addition, because many geographic data mentioned are not

precise geographic coordinates but locality name, we basically substitute the geographical coordinate with locality names.We compiled the species information and distribution data with coordinate (geographical locations with longitudes and latitudes) into a database (Microsoft Access), then transmit and input it into a Geographic Information System program (ArcView 3.2) for the mapping process. Biogeographic analysis is conducted and distribution maps were compiled.

Abbreviations used in the text

HL= Head length; EL= Eye length; AL= Antennal length; PL= Pronotum length; ELL= Elytral lateral length; ESL=Elytral sutural length; HW= Head width (including eyes); PW= Pronotum width; ELW= Elytral width; ABW= Widest segment of abdomen.

Collections

MNHUB= Museum für Naturkunde, Humboldt- Universität, Berlin, Germany (Dr. J. Frisch); NHMG= Entomology Muséum d’Histoire Naturelle, Genève, Switzerland (Dr. G. Cuccodoro); NHMW= Naturhistorisches Museum, Wien, Austria (Dr. H. Schillhammer); ZALF (DEI) = Deutsches Entomologisches Institute, Müncheberg, Germany (Dr. L. Zerche).

Taxonomy

Subgenus Eopaederus Scheerpeltz 1957Scheerpeltz 1957: 458; Szujecki 1965: 29; Coiff ait 1982: 32; Boháč 1985:

370; Löbl & Smetana 2004: 612.Type species: Paederus (Eopaederus) caligatus Erichson 1840, by original designation.

Diagnosis. Th is subgenus can be easily distinguished from other subgenera of Paederus Fabricius by the black tibiae and primitive aedeagus. Antennae from third segment onwards, tibiae and tarsi black. Elytra about 1/5 longer than broad. Pronotum distinctly narrowed posteriorly, posterior margin attenuated. Median lobe of aedeagus is as primitive as in the genus Paederidus Mulsant & Rey 1878. Th is group is diff erent from the subgenus Paederus s. str. by the black tibiae and antenna and the primitive internal armatures of aedeagus.

Key to species of the subgenus Eopaederus Scheerpeltz from China

1. Base of 1st segment of abdomen black or blackish brown .......................................................................... 2

- Base of 1st segment of abdomen brownish red or brownish yellow .......................................................... 5

2. Dorsal plate of aedeagus and median lobe regular, its apex shorter than median lobe ..................................... 3

- Dorsal plate of aedeagus and median lobe irregular, its apex longer than median lobe ... P. (E.) alternans Walker 1858

3. Head and apical two abdominal segments blackish blue with metallic shining ...... P. (E.) pubescens Cameron 1914

- Head and apical two abdominal segments black without metallic shining ........................................................... 4

4. Median notch of labrum acute and rather protruding, median lobe of aedeagus with lateral margin nearly parallel ............................... P. (E.) tamulus Erichson 1840

Taxonomy and biogeography of Subgenus Eopaederus from China

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Figure 1Paederus (E.) noncurvatus n. sp. from Hubei, China. A. labrum, male. B. labrum, female. C. left mandible, male. D. 8th tergite, male. E. 8th sternite, male. F. 9th sternite, male. G. 8th tergite, female. H. 8th sternite, female. I. 9th sternite, female. J-K. aedeagus. Scale = 0.5 mm.

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- Median notch of labrum weakly developed and rather short, median lobe of aedeagus with lateral margin broadly rounded ................. P. (E.) birmanus Fauvel 1895

5. Aedeagus rather simple and basal part rather bulgy ....... 6- Aedeagus slightly complicated and basal part slightly

bulgy ........................................................................... 76. Head black without metallic shining, median notch

of labrum with one obtusely small tooth ................... ........................................... P. (E.) basalis Bernhauer 1914

- Head blackish blue with metallic shining, median notch of labrum without such tooth ..................... .................................................. P. (E.) noncurvatus n. sp.

7. Dorsal plate of median lobe irregular with apical part slender and S-shaped ................................................... 8

- Dorsal plate of median lobe regular with apical part rounded ..................... P. (E.) nigricornis Bernhauer 1911

8. Anterior margin of labrum sharply sinuate, the protruding of both sides small and acute ............... ........................................................ P. (E.) curvatus n. sp.

- Anterior margin of labrum slightly sinuate, the protruding of both sides large and obtuse .................. ..................................... P. (E.) basiventris Bernhauer 1933

1. Paederus (Eopaederus) noncurvatus n. sp. (Fig. 1A–L).

Description of male. Length: 8.00–8.85 mm; width: 1.60–1.63 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.29:2.91:1.31:1.86 mm; width: HW:PW:ELW:ABW = 1.24:1.05:1.55:1.61 mm. Glossy, head black with blue shining, labrum black with anterior margin brownish red. Mandibles black with inner margin brown. Labial palpi blackish brown. Antennae with basal two segments brownish yellow, and the rests black. Pronotum brownish red. Scutellum black, elytra blackish blue with metallic shining. Legs black with apex of coxae slightly brownish yellow. First four abdominal segments brownish red with base of 1st segment and apical two black. Head nearly hexagonal shape, HL/HW = 1.04, lateral margin relatively straight. Eyes large, sharply protruded laterally, HL/EL = 3.04, diameter of eye longer than gena and as long as temple (gena:eye:temple = 0.34:0.43:0.43 mm). Punctuation on head irregular, of diff erent size, dense and deep, but denser laterally. Each puncture with a black or brown oblique hair of diff erent length. Vertex and middle front with punctures sparse, deep and large. Gular sutures separated; gular plate broader in anterior 1/3, gently narrowed posteriorly, parallel in middle 1/3 but broadened in basal 1/3. Antennae fi liform, 11 segmented, extending back to basal 1/4 of elytra; each segment longer than wide, thickened apically, covered with brownish yellow pubescence and setae on apex; 1st segment largest and thickest than others, 2nd slightly longer than half of 1st, 3rd slightly longer than 1st. Mouthparts. Labrum (Fig. 1A) transverse, anterior margin slightly sinuate with a pair of sharp and acute angles besides the middle notch. Mandibles (Fig. 1C) long and robust, apex curved inward, outer surface near base covered with hairs and punctures, inner margin with two sharp teeth, the notch behind it arched. Length of mentum about 2.5 times as long as prementum. Th orax. Pronotum nearly oviform, PL/PW = 1.25, widest in basal 2/3, anteriolateral margin broadly rounded and posteriolateral margin relatively straight. Disc rather convex, longitudinal media area smooth and glabrous, lateral areas covered with dense and large setiferous punctures,

denser laterally. Mesosternum black and small-sized, sternum uneven with very large and dense setiferous punctures, surface of posterior part slightly convex, covered with fi ne sculpture and large and sparse punctures, outside of it with irregularly fi ne sculpture and rather sparse punctures. Metasternum black and broad, broadly convex, surface with dense and fi ne setiferous punctures, a longitudinal nail-like groove in the middle of posterior 1/2, gradually tapered anteriorly. Elytra broad and lateral margin nearly parallel, ELL\ELW = 1.20, ESL/ELL = 0.78, humeral angles well developed, posterior margin slightly sinuate. Punctures on elytra larger and denser than those on head or pronotum, arranged irregularly. Interval between punctures slightly longer than diameter of a puncture, each puncture with an obliquely blackish brown hair. Hind wings well developed. Legs long, covered with pale pubescence and blackish brown setae. Tibiae furnished with pectinate pubescence on inner apex and a pair of setae on both sides of apex. First three segments of tarsi with two setae on anteriolateral apex, densely ciliate below; protarsi broadened, 2nd segment broadest, 1st, 3rd and 4th gradually narrowed apically. Meso- and metatarsi with fi rst four segments gradually shorted apically. Abdomen elongate, widest at 7th (5th visible) segment, where wider than head or pronotum but slightly narrower than elytra. Pleurotergite tapered gradually in 7th segment backward. Setiferous punctures on tergites fi ne and dense, closer laterally. Base of each tergite with transverse depression where punctures very large, deep and dense. Sternite punctures fi ne and dense. 7th sternite with hind margin broadly but shallowly emarginated, front of the emargination with a semicircular impunctate area. 8th sternite (Fig. 1E) with hind margin deeply notched. Th e notch slightly shorter than 1/2 of the whole sternite length, its bottom rounded and inner sides nearly parallel. Aedeagus (Figs. 1J–L) thick and symmetrical, length/width = 2.51. Median lobe (without dorsal plate) shorter than parameres, basal part rather bulgy and gradually narrowed apically. Dorsal plate of median lobe elongate, slightly longer than median lobe but shorter than parameres, lateral margin broadly rounded, posterior portion thin and apex acute. Internal armatures (Figs. 1J) strongly chitinized with apex S-shaped and acute. Female. Th e labrum (Fig. 1B) with anterior margin sharply sinuate, both sides with acute angles. 8th sternite (Fig. 1H) with apex triangular, both sides slightly emarginated. 9th sternite (Fig. 1I) length/width = 2.26, about 0.41× length of male, middle of anterior margin with a small triangular notch, apex of the sternite slightly rounded.Type Material. Holotype ♂, China, Hubei Province, Shengnongjia County, Dongxi (31.58°N, 110.13°E), 600 m, 4.VIII.1998, collected by Zhou Hongzhang (deposited in IOZ-CAS). Paratypes. 19 ♀♀, 21 ♂♂, same data as holotype. Etymology. Th e species name is from Latin words non (means no), and curvatus (means curve).

Comment. Th is new species is similar to P. (E.) curvatus n. sp., but diff er from the latter by the bulbous base of aedeagus, less sinuate and protrudent labrum margin in male. It can be easily distinguished from P. (E.) basalis Bernhauer by the rather slender body and the median notch of labrum without very acute tooth.


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Figure 2 Paederus (E.) curvatus n. sp. from Hubei, China. A. labrum, male. B. left mandible, male. C. 8th tergite, male. D. 8th sternite, male. E 9th sternite, male. F. 8th tergite, female. G. 8th sternite, female. H. 9th sternite, female. I–K. aedeagus. Scale = 0.5 mm.

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2. Paederus (Eopaederus) curvatus n. sp. (Fig. 2A–K).

Description of male. Length: 7.75-8.50 mm; width: 1.45-1.48 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.13:3.46:1.21:1.51 mm; width: HW:PW:ELW:ABW = 1.15:1.08:1.36:1.46 mm. Glossy, head and labrum black. Mandibles black with inner margin brownish red. Labial palpi blackish brown with 4th segment brownish yellow. Antennae with basal two segments brownish yellow, the rests black. Pronotum brownish red. Scutellum black, elytra blackish blue with metallic shining. Legs black with apex of coxae and base of trochanter slightly brownish yellow. First four abdominal segments brownish red with base of 1st and apical two segments black. Head hexagonal, HL/HW = 0.98, straight laterally. Eyes large, sharply protruded laterally, HL/EL = 3.05, diameter of eye slightly shorter than gena but longer than temple (gena:eye:temple = 0.39:0.37:0.32 mm). Punctuation on head irregular, of diff erent size, dense and deep, but denser laterally. Each puncture with a black or brown oblique hair of diff erent length. Vertex and middle front with punctures sparse, deep and large. Gular sutures separated; gular plate broad in anterior 1/3, slightly narrowed posteriorly, parallel in middle 1/3 but broadened in basal 1/3. Antennae fi liform, 11 segmented, extending back to basal 1/3 of elytra; each segment longer than wide, thickened apically, covered with brownish yellow pubescence and brown setae on apex; 1st segment largest and thickest than others, 2nd slightly longer than half of 1st, 3rd slightly longer than 1st. Mouthparts. labrum (Fig. 2A) transverse, anterior margin slightly sinuate with two pairs of angles: the inner ones short and sharp besides the middle notch and the outer ones sharply acute. Mandibles (Fig. 2B) long and robust, apex curved inward, outer surface near base covered with hairs and punctures, inner margin with two sharp teeth, the notch behind it arched. Length of mentum about 4 times as long as prementum. Th orax. Pronotum oviform, PL/PW = 1.12, widest in basal 2/3, anteriolateral margin broadly rounded and posteriolateral margin relatively straight. Disc rather convex, longitudinal media area smooth and glabrous, lateral areas covered with large and moderately dense punctures, denser laterally, each puncture with a black hair long or short. Mesosternum black and small-sized, sternum uneven with very large and dense punctures, surface of posterior part slightly convex, covered with fi ne sculptures and large and sparse setiferous punctures, outside of it with irregularly fi ne sculpture and rather dense punctures. Metasternum black and broad, broadly convex, surface with dense and fi ne setiferous punctures, a longitudinal nail-like groove in the middle of posterior 1/2, gradually tapered anteriorly. Elytra broad and nearly parallel, ELL\ELW = 1.11, ESL/ELL = 0.87, humeral angles well developed, posterior margin slightly sinuate. Punctures on elytra denser than those on head or pronotum, of diff erent size, arranged irregularly. Interval between punctures slightly longer than diameter of a puncture, each puncture with an obliquely brown hair. Hind wings well developed. Legs long, covered with slightly brown pubescence and brown setae. Tibiae furnished with pectinate pubescence on inner apex and a pair of setae on both sides of apex. First three segments of tarsi with two setae on anteriolateral apex, densely ciliated below; protarsi broadened, with 2nd segment broadest, 3rd segment narrowest, 1st and 4th nearly equal in width. Meso- and metatarsi with fi rst three segments gradually shorted apically, 4th shorter than 2nd but slightly longer than 3rd. Abdomen elongate, widest at 7th

(5th visible) segment, where wider than head or pronotum but slightly narrower than elytra. Pleurotergite gradually tapered in 7th segment backward. Setiferous punctures on tergites fi ne and dense, denser laterally. Base of 1st tergite with transverse depression where punctures very large, deep and dense, the following segments without such punctures. Sternite punctures larger and denser than punctures on tergites. 7th sternite with hind margin shallowly emarginated, front of the emargination with a longitudinally half-oviform impunctate area. 8th sternite (Fig. 2D) with hind margin deeply notched. Th e notch slightly shorter than 2/5 of the whole sternite length, its bottom rounded and inner sides nearly parallel. 9th sternite (Fig. 2E) with a longitudinally shallow groove near base and broadened posteriorly. Aedeagus (Figs. 2I-K) slender and symmetrical, length/width = 3.42. Median lobe (without dorsal plate) with basal portion slightly bulbous and gradually narrowed apically, lateral margin sinuate, posterior margin broadly emarginated. Dorsal plate of median lobe irregular and elongate, longer than median lobe but shorter than parameres, lateral margin broadly rounded and sinuate. Posterior portion of dorsal plate narrow and acute, apex S-shaped. Internal armatures (Figs. 2K) strongly chitinized, apex thin and sinuate.Female. Th e labrum broader than male. Anterior margin sharply sinuate and without distinct teeth on both sides of middle notch. 9th sternite (Fig. 2H) length/width = 1.92, about 0.48× length of that in male. Basal margin with a triangular notch, apex of the sternite obtusely curved.Type Material. Holotype ♂, China, Hubei Province, Shengnongjia County, Guanmen Mountain (31.43°N, 110.36°E), 1400–1720 m, 23-30.VII.1998, collected by Zhou Hongzhang, Ye Chanjuan and He Junjian (deposited in IOZ-CAS). Paratypes. 10 ♀♀, 18 ♂♂, same data as holotype; 5 ♀♀, 6 ♂♂, China, Hubei Province, Shengnongjia County, Muyu town (31.47°N, 110.39°E), 1150 m, 5. VIII. 1998, collected by Zhou Haisheng; 1 ♀, 2 ♂♂, China, Hubei Province, Shengnongjia County, Banqiao (31.46°N, 110.10°E), 5. VIII. 1998, collected by Zhou Hongzhang.Etymology. Th e species name is from Latin words curvatus (means curve).

Comment. Th is new species can be easily distinguished from P. (E.) noncurvatus n. sp. according to the content mentioned before this. It can be easily separated from other Eopaederus species by the sharp-sinuated anterior margin of labrum, extraordinary-curved dorsal plate of median lobe of aedeagus, and the slightly S-shaped internal armatures.

3. Paederus (Eopaederus) basiventris Bernhauer 1933 (Fig. 3A–H), status resurrectus

Bernhauer 1933: 28 (type locality: China: Xizang [=Tibet]); Scheerpeltz 1957: 474; Frank 1988: 102; Willers 1999: 155 (syn. of P. (E.) nigricornis Bernhauer); Löbl & Smetana 2004: 611.

Description of male. Length: 8.10–9.00 mm; width: 1.64–1.68 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.41:3.51:1.46:1.86 mm; width: HW:PW:ELW:ABW = 1.38:1.32:1.66:1.41 mm. Glossy, head and labrum black. Mandibles black with inner margin brownish red. Labial palpi blackish brown. Antennae black with 1st segment brownish red. Pronotum brownish red. Scutellum black, elytra blackish


333

blue with metallic shining. Legs black with base of procoxae brownish yellow. Abdomen with 1st and apical two segments black and the rests brownish red. Head nearly rhombic, HL/HW = 1.02, lateral margin relatively straight. Eyes relatively small, moderately protruded laterally, HL/EL = 3.44, diameter of eye slightly shorter than gena or temple (gena:eye:temple = 0.45:0.41:0.51 mm). Labrum (Fig. 3A) transverse, anterior margin sinuate, both sides of middle notch with two pairs of sharply protruding angles. Mandibles (Fig. 3C) with two sharp teeth on inner margin. Length of mentum slightly longer than prementum. Th orax. Pronotum nearly oviform, PL/PW = 1.11, widest in basal 2/3, rounded laterally. Disc covered with moderately dense and large punctures, slightly denser laterally, each puncture with a black hair long or short. Elytra broader than head or pronotum and lateral margin nearly parallel, ELL\ELW = 1.12, ESL/ELL = 0.69, humeral angles well developed. Punctures on surface dense and large, interval between punctures shorter than diameter of a puncture, each puncture with a black hair or setae. Hind wings well developed.

Legs long, protarsi with 1st segment slightly narrower than 2nd, 2nd to 4th gradually narrowed apically. Meso- and metatarsi with fi rst four segments gradually shorted apically. Abdomen elongate, widest at 7th (5th visible) segment, where wider than head or pronotum but narrower than elytra. Pleurotergite gradually tapered in 7th segment backward. Setiferous punctures on tergites fi ne and dense, denser laterally. Base of 1st segment with transverse depression where punctures rather dense, large and deep. Sternite punctures slightly fi ner and denser than tergite punctures. Hind margin of 7th sternite with shallow and narrow emargination. 8th sternite with hind margin deeply notched. Th e notch about 1/3 length of the whole sternite, its bottom slightly rounded and inner sides parallel. 9th sternite with a longitudinally narrow line, behind the line with a broad groove to apex. Aedeagus (Figs. 3F-H) relatively short, length/width = 3.33. Median lobe (without dorsal plate) with basal portion bulbous and gradually narrowed apically, lateral margin broadly rounded and sinuate, apex slightly emarginated. Dorsal plate of median lobe with

Figure 3Paederus (E.) basiventris Bernhauer 1933. A. labrum, male. B. labrum, female. C. right mandible, male. D. 8th tergite, male. E. 8th tergite, female. F-H. aedeagus. Scale = 0.5 mm

334


S-shaped apex and longer than median lobe but shorter than parameres. Internal armatures (Figs. 3F, 3H) strong chitinized and slightly S-shaped.Female. 9th sternite length/width = 2.48, about 0.49× length of that in male, with a V-shaped notch in basal margin.Specimens Examined. 1 ♂, China, Sichuan Province, Wolong Wassuland, Sankiankou (31.29°N, 103.36°E), 2000 m (det. Willers 2006), 7.X.1934 (deposit in NHMW); 7 ♀♀, 12 ♂♂, China, Sichuan Province, Baoxing (30.36°N, 102.84°E), 1300-1400 m, 14.VIII.1995, collected by Yu Peiyu; 2 ♂♂, China, Sichuan Province, Baoxing Fengtongzhai (30.36°N, 102.84°E), 1500–1535 m, 17-18.VIII.2003, collected by Zhao Caiyun (deposited in IOZ-CAS).Comment. Th e species is resurrected as a valid species and not a synonym of P. (E.) nigricornis Bernhauer 1911 anymore.

4. Paederus (Eopaedus) birmanus Fauvel 1895 (Fig. 4A–H), newly designed to the subgenus,

fi rst record from China.Fauvel 1895: 233 (type locality: Burma: Carin Cheba); Bernhauer &

Schubert 1912: 204; Cameron 1931: 47; Frank 1988: 103; Willers 1999: 157; Löbl & Smetana 2004: 614.

Description of male. Length: 6.7–7.5 mm; width: 1.34–1.39 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.17:3.02:1.16:1.59 mm; width: HW:PW:ELW:ABW = 1.02:0.96:1.36:1.18 mm. Glossy, head and labrum black. Mandibles brown with inner margin brownish red. Labial palpi blackish brown with base of 2nd segment brownish yellow. Antennae with fi rst two segments brownish yellow, the rests black. Pronotum and fi rst four abdominal segments brownish red whereas apical two black. Scutellum black, elytra blackish blue with metallic shining. Legs black with procoxae brownish yellow. Head nearly rhombic, HL/HW = 1.15, lateral margin relatively straight. Eyes large, sharply protruded laterad, HL/EL = 3.15, diameter of eye longer than gena but slightly shorter than temple (gena:eye:temple = 0.31:0.37:0.39 mm). Labrum (Fig. 4A) transverse, anterior margin sinuate with two pairs of obtuse angles on sides of middle notch. Mandibles (Fig. 4C) with two sharp teeth of inner margin. Length of mentum about 3 times as long as prementum. Th orax. Pronotum nearly oviform, PL/PW = 1.21, widest in basal 2/3, broadly rounded laterally. Disc rather convex, longitudinal media area smooth and glabrous, lateral areas covered with moderately large and sparse punctures, slightly denser laterally, each puncture with a black hair long or short. Mesosternum black and medium-

Figure 4 Paederus (E.) birmanus Fauvel 1895. A. labrum, male. B. labrum, female. C. right mandible, male. D. 8th tergite, male. E. 8th tergite, female. F-H. aedeagus. Scale = 0.5 mm.


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sized. Elytra broad and nearly parallel, ELL\ELW = 1.17, ESL/ELL = 0.83, with humeral angles well developed, posterior margin slightly sinuate. Punctures on elytra fi ner than those on pronotum and moderately dense, interval between punctures slightly longer than diameter of a puncture, each puncture with a brown or black hair. Hind wings well developed. Legs long, protarsi with 1st slightly narrower than 2nd, 2nd to 4th gradually narrowed apically, meso- and metatarsi with fi rst four segments gradually shorted apically. Abdomen elongate, widest at 7th (5th visible) segment, where wider than head or pronotum but narrower than elytra. Pleurotergite gradually tapered in 7th segment backward. Setiferous punctures on tergites fi ne and dense, denser laterally. Base of each tergite with transverse depression where punctures rather sparse. Sternite punctures sparser than tergite punctures. Hind margin of 7th sternite with shallow and narrow emargination. 8th sternite with hind margin deeply notched. Th e notch about 1/2 length of the whole sternite, its bottom slightly rounded and inner sides nearly parallel. 9th sternite with a longitudinally shallow groove. Aedeagus (Figs. 4F–H) slender and symmetrical, length/width = 3.20. Median lobe (without dorsal plate) with basal

portion bulbous and gradually narrowed apically, lateral margin broadly rounded, apex slightly rounded. Dorsal plate of median lobe longitudinally elliptic and apex shorter than median lobe or parameres. Internal armatures (Figs. 4F) irregularly shaped.Female. Anterior margin of labrum (Fig. 4B) sharply sinuate with obtuse angles on both sides. 9th sternite length/width = 2.40, about 0.49× length of that in male, basal margin with a triangular notch.Specimens Examined. 1 ♂, China, Yunnan Province, Xishuangbannan, NW Mengla (21.48°N, 101.56°E), ca. 700 m (CWBS 364), 8.XI.1999, collected by Jäch, et al.; 1 ♀, China, Yunnan Province, Xishuangbannan, NW Mengla (21.48°N, 101.56°E), ca. 600 m (det. Willers, 8. 04; CWBS 377), 12.XI.1999, collected by Jäch, et al. (deposit in NHMW); 7 ♀♀, 17 ♂♂, China, Yunnan Province, Xishuangbannan (22.01°N, 100.48°E), 630 m, 1.II.2005, collected by Wu Jie; 5 ♀♀, 4 ♂♂, China, Yunnan Province, Xishuangbannan, Mengla (21.48°N, 101.56°E), 620-650 m, 18-19.XI.1958, collected by Pu Fuji and Zhang Yiran; 1 ♂, China, Yunnan Province, Menghai (21.95°N, 100.50°E), 630 m, 18.IV.1982, collected by Yu Peiyu (deposited in IOZ-CAS).

Figure 5 Paederus (E.) alternans Walker 1858 A. labrum, male. B. left mandible, male. C. 8th sternite, female. E-F. aedeagus. Scale = 0.5 mm.

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5. Paederus (Eopaederus) alternans Walker 1858 (Fig. 5A–F), newly designed to the subgenus.

Walker 1858: 205 (type locality: Ceylon: Colombo: Maskeliya); Cameron 1931: 41; Willers 2001: 21; Löbl & Smetana 2004: 614.

Paederus rufi coxis Kraatz 1859: 151; Fauvel 1904: 54 (syn. of P. alternans Walker); Bernhauer & Schubert 1912: 209; Cameron 1931: 41; Frank 1988: 129; Löbl & Smetana 2004: 614.

Description of male. Length: 5.7–6.5 mm; width: 1.19–1.23 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.05:2.41:0.96:1.44 mm; width: HW:PW:ELW:ABW = 1.04:0.85:1.21:0.91 mm. Glossy, head and labrum black. Mandibles brownish red with outer margin blackish brown. Labial palpi with fi rst two segments brownish yellow, apex of 3rd segment blackish brown. Antennae with fi rst two segments brownish yellow, the rests black. Pronotum and fi rst four abdominal segments brownish red whereas apical two black. Scutellum brownish yellow with margins black, elytra blackish blue with metallic shining. Legs black with procoxae brownish yellow. Head nearly hexagonal, HL/HW = 1.01, lateral margin nearly straight. Eyes large, sharply protruded laterally, HL/EL = 2.84, diameter of eye longer than gena but slightly shorter than temple (gena:eye:temple = 0.26:0.37:0.30 mm). Labrum (Fig. 5A) transverse, anterior margin slightly sinuate with one pair of short angles on both sides, middle notch with a small and obtuse denticle in its bottom. Mandibles (Fig. 5B) with an acute bicuspidate tooth on inner margin. Length of mentum about 1.5 times as long as prementum. Th orax. Pronotum nearly oviform, PL/PW = 1.13, widest in basal 2/3, lateral margin broadly

rounded. Disc rather convex, longitudinal media area smooth and glabrous, lateral areas covered with moderately large and sparse punctures, slightly denser laterally, and each puncture with a brown short hair. Mesosternum black and small sized. Elytra broad and nearly parallel, ELL\ELW = 1.19, ESL/ELL = 0.79, with humeral angles well developed, posterior margin slightly sinuate. Punctures on elytra fi ner and shallower than those on pronotum, but moderately dense. Interval between punctures longer than diameter of a puncture, each puncture with a black hair. Hind wings well developed. Legs long, protarsi with 2nd segment broadest, 1st and 3rd equal in width, 4th narrowest. Meso- and metatarsi with fi rst four segments gradually shorted apically. Abdomen elongate, widest at 6th (4th visible) segment, where wider than pronotum but narrower than head or elytra. Pleurotergite gradually tapered in 7th segment backward. Setiferous punctures on tergites fi ne and dense, denser laterally. Base of each tergite with transverse depression where punctures rather dense and large. Sternite punctures denser and fi ner than tergite punctures. 7th sternite with hind margin shallowly and narrowly emarginated. 8th sternite with hind margin deeply notched, about 1/2 length of the whole sternite, its bottom slightly rounded and inner sides nearly parallel. 9th sternite with a transverse fi ne line and a longitudinally shallow groove in anterior 1/2. Aedeagus (Figs. 5D-F) slender and symmetrical, length/width = 3.76. Median lobe (without dorsal plate) with basal portion slightly bulbous and broadly sinuate laterally, apex slightly rounded. Dorsal plate of median lobe longitudinally elliptic with apex shorter than median lobe or parameres. Internal armatures (Figs. 5E) irregularly shaped.

Figure 6 Paederus (E.) basalis Bernhauer 1914. A. labrum, male. B. labrum, female. C. 8th tergite, male. D 8th sternite, female. E-G. aedeagus. H. Internal armature. Scale = 0.5 mm.


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Female. 9th sternite length/width = 1.78, about 0.50× length of that in male, with a V-shaped notch in basal margin.Specimens Examined. 1 ♀, China, Hainan (202) E. Jianfeng (18.36°N, 108.71°E) 60 m (det. Willers, 7. 06), 21.I.1996, collected by Jäck; 1 ♂, Th ail, 150km, NW Bankok (18.59°N, 101.10°E) 90 m, Whan-Kha, VII.1990, collected by Th ielen (deposit in NHMW); 13 ♀♀, 16 ♂♂, China, Hainan Province, Tongshen (18.46°N, 109.31°E), 340 m, 22.IV.1960, collected by Zhang Xuezhong and Li Changqing; 6 ♀♀, 15 ♂♂, China, Hainan Province, Jianfeng (18.36°N, 108.71°E), 70 m, 16-26.III.1980, collected by Pu Fuji; 1 ♀, 5 ♂♂, China, Hainan Province, Jianfeng (18.36°N, 108.71°E), 750 m, 17.III.1980, collected by Wang Shuyong and Pu Fuji; 1 ♀, China, Hainan Province, Shuiman (18.38°N, 108.35°E), 640 m, 29.V.1960, collected by Li Xianfu (deposited in IOZ-CAS).

6. Paederus (Eopaederus) basalis Bernhauer 1914 (Fig. 6A–H).

Bernhauer 1914: 98 (type locality: East India: Siwaliks); Cameron 1931: 54; Scheerpeltz 1933: 1221; Scheerpeltz 1957: 474; Frank 1988: 102; Willers 1999: 154; Löbl & Smetana 2004: 611.

Paederus ledouxi Coiff ait 1978: 561; Coiff ait 1982: 33, 52; Frank 1988: 117; Willers 1999: 157 (syn. of P. basalis Bernhauer); Löbl & Smetana 2004: 611.

Paederus kuluensis Bernhauer 1914: 99; Cameron 1931: 55; Scheerpeltz 1957: 474; Coiff ait 1982: 51; Frank 1988: 117; Willers 1999: 154 (syn. of P. basalis Bernhauer); Löbl & Smetana 2004: 611.

Paederus punjabensis Cameron 1945: 68; Frank 1988: 128; Willers 1999: 154 (syn. of P. basalis Bernhauer); Löbl & Smetana 2004: 611.

Measurements of male. Length: 8.80–10.40 mm; width: 1.52–1.62 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.35:3.44:1.30:1.96 mm; width: HW:PW:ELW:ABW = 1.21:1.06:1.57:1.32 mm.Specimens Examined. 1 ♀, 5 ♂♂, Nepal (27.43°N, 85.19°E) (Kolekondo, Kornoli R. 700 m det. G. de. Rougemont 1998), XI.1987, collected by P. Morvan (deposit in NHMG); 1 ♀, 1 ♂, Lachiwala Siwaliks (31.00°N, 78.00°E), 10.IV.1921, collected by Cameron (deposit in MNHUB); 44 ♀♀, 33 ♂♂, China, Sichuan Province, Baoxing Yaoji (30.67°N, 102.73°E), 2100–2200 m, 20-21.VIII.2003, collected by Zhou Hongzhang and Zhao Caiyun; 1 ♂, China, Sichuan Province, Baoxing Yaoji (30.67°N, 102.73°E), 1300 m, 16.VIII.1995, collected by Yu Peiyu; 1 ♂, China, Sichuan Province, Baoxing Fengtongzhai (30.36°N, 102.84°E), 29.VII.2006, collected by Bai Xiaoshuan; 2 ♀♀, 1 ♂, Guangxi Province, Chacheng Xijiaodian (22.69°N, 109.20°E), 30.VII.1985, collected by Liao Subai; 1 ♀, China, Guangxi Province, Miao’er Mountain (25.90°N, 110.30°E) 2140 m, 1.VII.1985, no collector (deposited in IOZ-CAS).

7. Paederus (Eopaederus) nigricornis Bernhauer 1911 (Fig. 7A–D).

Bernhauer 1911: 61 (type locality: Nepal: Soondrijal); Bernhauer & Schubert 1912: 209; Cameron 1931: 47; Scheerpeltz 1957: 474;

Figure 7 Paederus (E.) nigricornis Bernhauer 1911. A. labrum, male. B. left mandible, male. C-D. aedeagus and internal armature. Scale = 0.5 mm.

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Coiff ait 1982: 33; Frank 1988: 123; Willers 1999: 157; Willers 2001: 21; Willers 2001a: 308; Löbl & Smetana 2004: 611.

Measurements of male. Length: 8.8–9.4mm; width: 1.60–1.68 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.44:3.76:1.37:2.17 mm; width: HW:PW:ELW:ABW = 1.26:1.10:1.66:1.43 mm.Specimens Examined. 3 ♀♀, 1 ♂, China, Xizang Province, Chayu County, Xiachayu town (28.62°N, 97.49°E), 1600 m, 25.VIII.2005, collected by Wang Xuejian; 1 ♀, China, Xizang Province, Linzhi County, Pailong town (29.59°N, 94.25°E),

2050 m, 1.IX.2005, collected by Wu Jie; 1 ♀, 1 ♂, Nepal (27.43°N, 85.19°E), 1300 m, VI.1989, collected by P. Morvan (deposited in IOZ-CAS).

8. Paederus (Eopaederus) pubescens Cameron 1914 (Fig. 8A–H).

Cameron 1914: 539 (type locality: India: Assam); Cameron 1925: 35 (syn. of P. extraneus Wiedemann); Cameron 1931: 44 (species); Coiff ait 1982: 35, 50; Frank 1988: 102; Willers 1999: 157; Löbl & Smetana 2004: 611.

Figure 8P. (E.) pubescens Cameron. from Taiwan, China. A. labrum, male. B. right mandible, male. C. labrum, female. D. 8th sternite, male. E 8th sternite, female. F-H. aedeagus. Scale = 0.5 mm.


339

Measurements of male. Length: 7.2–8.2 mm; width: 1.18–1.25 mm. Length of diff erent body parts: HL:AL:PL:ELL = 1.11:3.06:1.35:1.64 mm; width: HW:PW:ELW:ABW = 1.15:1.16:1.54:1.41 mm.Specimens Examined. 1♂, China, Taiwan (Anping Formosa (22.97°N, 120.22°E); det. Willers. 12.02; coll. DEI; Müncheberg), 7.VI.1912, collected by H. Sauter; 1♂, China, Taiwan (Kosempo Formosa (25.05°N, 121.52°E); det. Willers. 12. 02; coll. DEI, Muncheberg), IV.1912, collected by H. Sauter; 1♂, China, Taiwan (Tainan Formosa (22.97°N, 120.22°E); det. Willers. 12. 02; coll. DEI, Müncheberg), 7.VI.1912, collected by H. Sauter; 1♂, China, Taiwan (Tainan Formosa (25.05°N, 121.52°E); det. Willers. 12. 02; coll. DEI, Müncheberg), 13.VI.1912, collected by H. Sauter; 1♀, China, Taiwan (Nanshanxi Formosa (25.05°N, 121.52°E); det. Willers. 12. 02; coll. DEI, Eberanalde), 24. V. 1912, collected by K. Masumoto (deposited in DEI).

9. Paederus (Eopaederus) tamulus Erichson 1840 (Fig. 9A–F).

Erichson 1840: 661 (type locality: Singapore); Bernhauer & Schubert 1912: 211; Cameron 1931: 43; Scheerpeltz 1957: 459; Coiff ait 1982: 36, 50; Willers 1999: 157; Willers 2001: 21; Willers 2001a: 308; Löbl & Smetana 2004: 611.

Paederus dubius Kraatz 1859: 151; Fauvel 1869: 491 (syn. of P. tamulus Erichson); Scheerpeltz 1957: 459; Löbl & Smetana 2004: 611.

Paederus mixtus Sharp 1874: 75; Fauvel 1903: 153 (syn. of P. tamulus Erichson); Scheerpeltz 1957: 459; Löbl & Smetana 2004: 611.

Paederus rugipennis Motschulsky 1860: 74; Fauvel 1903: 153 (syn. of P. tamulus Erichson); Scheerpeltz 1957: 459; Löbl & Smetana 2004: 611.

Measurements of male. Length: 6.0–7.1mm; width: 1.08–1.13 mm. Length of diff erent body parts: HL:AL:PL:ELL = 0.95:2.28:1.01:1.47mm; width: HW:PW:ELW:ABW = 0.95:0.88:1.21:1.10 mm.

Specimens Examined. 9 ♀♀, 13 ♂♂, China, Beijing, Miyun County, Jiudaowan (40.37°N, 116.85°E), 14-15.V.2005, collected by Zhou Hongzhang and Li Xiaoyan; 5 ♀♀, 2 ♂♂, China, Fujian Province, Huangkeng (27.55°N, 117.7°E), 270-410 m, 6.VI.1960, collected by Ma Chenglin; 2 ♂♂, China, Hainan Province, Haikou City (20.02°N, 110.35°E), 100 m, 15.IV.1980, collected by Wang Shuyong; 4 ♀♀, 3 ♂♂, Tonkin, Hoa-Binh (20.49°N, 105.20°E), VII.1939, collected by A. de Cooman; 1 ♀, France, Chateauroux (46.50°N, 1.40°E), VI.1939, no collector (deposited in IOZ-CAS).

Figure 9P. (E.) tamulus Erichson 1840. A. labrum, male. B. labrum, male. C. left mandible, male. D-F. aedeagus and internal armature. Scale = 0.5 mm

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Table 1. Geographical distribution of the species of the subgenus Eopaederus.

Species Geographical distribution

alternans China (Guangxi, Hainan); Burma (Kawkareik, Pegu), Vietnam (Tonkin), Sri Lanka (Colombo, Maskeliya), India (Nilgiri Hills, Kanara, Himalayas).

balachowskyi Iran (Chah Bahar), Turky.

basalis China (Guangxi, Sichuan, Shaanxi); Nepal (Karnali), India (Siwaliks, Dehra-Dun-U, Mussoorie District, Himachal Pradesh, Uttaranchal, Uttar Pradesh, Kulu), Afghanistan, Pakistan.

basiventris China (Sichuan, Xizang); Nepal (Karnali, Koshi), India (Kulu), Pakistan, Afghanistan.

birmanus China (Yunnan); Birma (Carin Cheba; Carin Asciuii Cheba), North India, Nepal.

caligatus Austria, Belgium, Bosnia-Herzegovina, Bulgaria, Czech Republic, Denmark, France (Corsica, Monaco, Paris), Great Britain, Germany, Hungay, Ireland, Italy, Th e Netherlands, Poland, Portugal, Romania, Russia (Central and South European Territory), Slovakia, Spain, Switzerland.

curvatus China (Hubei).

debilior Turkmenistan, Turkestan, Uzbekistan.

limnophilus Austria, Balkans, Bosnia-Herzegovina, Bulgaria, Czech Republic, France (Corsica, Monaco), Germany, Hungay, Italy (Sardegna, Sicily, San Manno, Turin), Latvia, Poland, Romania, Slovakia, Russia (South and Central Territory), Switzerland, Yugoslavia (Montenegro).

mesopotanicus Iran, Iraq (Baghdad), Syria.

nigricornis China (Hubei, Sichuan, Xizang); Sikkim, Nepal, Indian (Uttaranchal, Garhwal hills, Simla-Hills, Darjeeling), Afghanistan, Pakistan.

noncurvatus China (Hubei).

pubescens China (Taiwan); Bhutan, Nepal (Ilam, Chitwan, Bheri), Bengalen, India (North-India, Punjab), Pakistan, Afghanistan.

tamulus China (Hongkong, Taiwan); Sri Lanka, Nepal, Vietnam (Tonkin; Hoa-Binh), Burma, India (Delhi, Okhla Dam), Japan, Sumatra, Indonesia, Singapore, Algfi anistan, Iran.

Figure 10Distribution patterns of the species of the subgenus Eopaederus Shcheerpeltz. P. (E.) alternans Walker; P. (E.) balachowskyi (Jarrige); P. (E.) basalis Bernhauer; P. (E.) basiventris Bernhauer; P. (E.) birmanus Fauvel; P. (E.) caligatus Erichson; P. (E.) curvatus n. sp.; P. (E.) debilior Eppelsheim; P. (E.) limnophilus Erichson; P. (E.) mesopotanicus Eppelsheim; P. (E.) nigricornis Bernhauer; P. (E.) noncurvatus n. sp.; P. (E.) pubescens Cameron;

P. (E.) tamulus Erichson. … River; — National boundaries.


341

Discussion

Zooogeography: Distribution patternsTh e map (Fig. 10) shows the general patterns of

geographical distributions of 14 species (Table 1) recorded worldwide in the subgenus Eopaederus. It is a result after summarizing existing geographical distribution data, so far we found in specimen labels and literatures, of all the species found in the world. Obviously, the subgenus Eopaederus distributes mainly in Eurasian continent, with the west part of the subgenus track mainly in temperate zones, whereas the east part shifts to some extent southward and is expanding to the whole tropical Asia. Th is subgenus is thus a faunistic element shared by the Oriental and the Palaearctic region.

Analyzing the general track (as showed in the map, Fig.10) of the subgenus from a point of perspective view, we were able to divide it into three parts: the Europe part, the Middle East part and the South and East Asia part. In these three regions, each has their own endemic species which do expand their species ranges to others. Here we designed the species distributions, according to their geographical ranges, to three diff erent types: Europe type, Middle-East type and South and East Asia type.

1. Europe type (Fig. 11): Two species can be categorized to this type: P. (E.) caligatus Erichson and P. (E.) limnophilus Erichson. Th eir geographical distributions cover mainly Europe, where they were more frequently recorded than in other areas, and expande eastward to Russia (Europe part through Siberia and Caucasia). It is interesting to see that, though both species exhibit a general pattern same in zoogeography and overlap most their species tracks, P. (E.) caligatus Erichson has a large range and distributes to the coastal zone of Europe continent and even to England and Ireland, whereas P. (E.) limnophilus Erichson westward not far beyond the line A (Fig. 11) and only to the central Europe. P. (E.) caligatus prefers therefore the Atlantic marine climate more than P. (E.) limnophilus. Th e latter is rather a species well addated to continental climate. Th e range diff erence can also indicates that the geographical distributions of the subgenus were strongly infl uenced by the monsoon climate of the Atlantic Ocean and the resulted seasonality of weather changes.

2. Middle-East type (Fig. 10): in this subgenus, three species, P. (E.) debilior, P. (E.) mesopotanicus Eppelsheim and P. (E.) balachowskyi (Jarrige), distribute only in the Middle East, a region in the middle of the subgenus track. In point of faunistic view, these three

Figure 11Distribution patterns of European type. P. (E.) caligatus Erichson; P. (E.) limnophilus Erichson.

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species can be considered as a special type, for they are obviously diff erent from Europe type as well South and East Asia type. We can also consider this narrow area (between line A and B in the map, Fig. 10) as boundary that isolates the west and the east parts of the subgenus track. Th at is to say, the species of Europe type do not expand eastward, and those of South and East Asia type do not expand westward, their species ranges to cross over this boundary.

3. South and East Asia type (Figs 10, 12, 13): nine species of the subgenus Eopaederus display a geographical pattern of species ranges which can be categorized to this type. Th e distributions of these species are mainly in the Oriental regions and only some of them expand northward, with the north limit to North China (Beijing around) and Japan. Th ey cover almost the whole of the region, South and East Asia. In viewpoint of species richness and endemism, this region is more important than others, for nine of 14 species of Eopaederus occurred there. Th e species ranges

and their variations are thus relatively complicate. A remarkable result was obtained after we analyzed the range variations in this region and distinguished them further into diff erent subcategories.

Of the species with a geographical ranges as South and East Asia type (Figs 10, 12, 13), they distribute mostly in a narrow band along the south boundary of Himalayas (Qinghai-Tibet Plateau), and eastward, their ranges are widely expanded along latitude with the north limit reaching North China and Japan and the south to Indonesia. Th is forms the most general patterns of species ranges in the subgenus (Figs 10, 13). Moreover, the narrow band along the south boundary of Himalayas exhibits very high richness and endemism. Th is fact supports the global biodiversity hotspot patterns proposed by Myers et al. (2000), namely Southwest China, the Himalayas and Indo-Burma, which are of great signifi cance in international conservation of global biodiversity and nature reserves.

Figure 12Distribution patterns of South and East Asia type. P. (E.) basalis Bernhauer; P. (E.) basiventris Bernhauer; P. (E.) curvatus n. sp.; P. (E.) nigricornis Bernhauer; P. (E.) noncurvatus n. sp.; P. (E.) pubescens Cameron.


343

As a matter of fact, only one species P. (E.) tamulus Erichson has a really wide range of distributions in the monsoon region of East Asia (Fig. 13). Other two species P. (E.) alternans Walker and P. (E.) birmanus Fauvel only expand their ranges southward but not northward and not cross over the line B in map (Fig. 13). Many other species, however, display even narrower species ranges which are limited within a narrow latitudinal range between 27.02°N- 31.10°N.(Fig. 12): they are P. (E.) basalis Bernhauer, P. (E.) basiventris Bernhauer, P. (E.) nigricornis Bernhauer, P. (E.) pubescens Cameron, P. (E.) noncurvatus n. sp., and P. (E.) curvatus n. sp.

Obviously, the subgenus Eopaederus is a grade which distributes in North hemisphere. South and East Asia possesses high species richness and endemism, and thus can be considered as a center of distribution, diversifi cation and speciation, as was supported by many other studies (Hewitt 1996; Johnson & Cicero 2004; Weir & Schluter 2004; Huang et al. 2006). Our results may be infl uenced by unbalanced collections and insuffi cient data in such

a large geographical region. Our study is nevertheless one that synthesizes worldwide data and the analysis was conducted more comprehensively than other likes. Th us, main conclusions here are convincing. Th e results will form a fundamental base for further studies on phylogeography and species diversifi cation of the genus Paederus in the future.Acknowledgments. We are indebted to Mr. Joachim Willers, Mr. Johannes Frisch and Dr. Manfred Uhlig (MNHUB), Dr. Harald Schillhammer (NHMW), Dr. Giulio Cuccodoro (NHMG) and Dr. Matthias Hartmann (Naturkunndemuseum Erfurt, Deutschland), Dr. Lothar Zerche (DEI) for sending literatures and specimens and off ering many kinds of help. We are also grateful to Dr. Alfred F. Newton (Field Museum of Natural History Chicago, USA) and Dr. Young Bok Cho (Natural History Museum Hannam University, Korea) for off ering important message. Two anonymous referees gave valuable suggestions to improve the manuscript. Th is study was supported by National Natural Science Foundation of China (NSFC-30570215 and 30670285), CAS Innovation Program (KSCX2-YW-Z-017) and National Science Fund for Fostering Talents in Basic Research (Special subjects in animal taxonomy, NSFC-J0630964/J0109).

Figure 13Distribution patterns of South and East Asia type. P. (E.) alternans Walker; P. (E.) birmanus Fauvel; P. (E.) tamulus Erichson.

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