in Southeast Asia

Zootaxa 3992 (1): 001–094 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press

Monograph

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3992.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:E0335F03-4449-409A-8C60-8FB52AA04A8B

ZOOTAXA 3992

Revision of the genus Oxytelus Gravenhorst (Staphylinidae: Oxytelinae) in Southeast Asia LIANG LÜ1 & HONG-ZHANG ZHOU1,* 1

Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1 Beichen West Rd., Chaoyang District, Beijing 100101, P. R. China * Corresponding author. E-mail: [email protected]; Tel: +86-10-64807252; Fax: +86-10-64807099.

Magnolia Press Auckland, New Zealand

Accepted by J. Klimaszewski: 2 Jun. 2015; published: 29 Jul. 2015

LIANG LÜ & HONG-ZHANG ZHOU Revision of the genus Oxytelus Gravenhorst (Staphylinidae: Oxytelinae) in Southeast Asia (Zootaxa 3992) 94 pp.; 30 cm. 29 Jul. 2015 ISBN 978-1-77557-753-9 (paperback) ISBN 978-1-77557-754-6 (Online edition)

FIRST PUBLISHED IN 2015 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2015 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326

(Print edition)

ISSN 1175-5334

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2 · Zootaxa 3992 (1) © 2015 Magnolia Press

LÜ & ZHOU

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 Key to species from Southeast Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 1. Oxytelus armiger Fauvel, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7 2. Oxytelus bengalensis Erichson, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 3. Oxytelus nigriceps Kraatz, 1859 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 4. Oxytelus javanus Cameron, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 5. Oxytelus ruptus Fauvel, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 6. Oxytelus piceus (Linné, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12 7. Oxytelus varipennis Kraatz, 1859 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 8. Oxytelus migrator Fauvel, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13 9. Oxytelus incisus Motschulsky, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 10. Oxytelus finitimus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15 11. Oxytelus bellicosus Fauvel, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 12. Oxytelus castaneus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 13. Oxytelus lividus Motschulsky, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 14. Oxytelus insulanus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 15. Oxytelus sublividus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 16. Oxytelus punctipennis Fauvel, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20 17. Oxytelus ferreirai Scheerpeltz, 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 18. Oxytelus lucens Bernhauer, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 19. Oxytelus lompobatangensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23 20. Oxytelus poecilopterus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 21. Oxytelus barbatus Fauvel, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 22. Oxytelus mandibularis Cameron, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 23. Oxytelus puncticeps Kraatz, 1859 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 24. Oxytelus lucidulus Cameron, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27 25. Oxytelus subferrugineus Cameron, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28 26. Oxytelus megaceros Fauvel, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 27. Oxytelus fallax Fauvel, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30 28. Oxytelus grandiculus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31 29. Oxytelus subsculptus Cameron, 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32 30. Oxytelus subincisus Cameron, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 Species excluded from Oxytelus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Anotylus transversipennis (Scheerpeltz, 1978) comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Some species from adjacent regions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36

Abstract Oxytelus species are widespread over all continents except Antarctica. Southeast Asia is one of the biodiversity hotspots in the world. In this paper, we review the Oxytelus species currently known from Southeast Asia. Seven species are described as new to science: O. castaneus sp. nov. (Vietnam), O. finitimus sp. nov. (Laos), O. grandiculus sp. nov. (Malaysia, Indonesia, and Thailand), O. insulanus sp. nov. (Malaysia and Indonesia), O. lompobatangensis sp. nov. and O. poecilopterus sp. nov. (Indonesia), and O. sublividus sp. nov. (Vietnam and Laos). Seven new synonymies are proposed: O. ruptus Fauvel = O. sublucidus Cameron, O. lucens Bernhauer = O. malaisei Scheerpeltz, O. puncticeps Kraatz = O. (Anotylus) micantoides Scheerpeltz, O. subferrugineus Cameron = O. kedirianus Cameron, O. megaceros Fauvel = O. kalisi Bernhauer, O. subincisus Cameron = O. fruhstorferi Cameron, O. antennalis Fauvel = O. cheesmani Bernhauer. Lectotypes are designated for the following names: O. armiger Fauvel, O. bellicosus Fauvel, O. discalis Cameron, O. gigantulus Fauvel, O. ginyuenensis Bernhauer, O. javanus Cameron, O. kalisi Bernhauer, O. kedirianus Cameron, O. lucens Bernhauer, O. lucidulus Cameron, O. mandibularis Cameron, O. megaceros Fauvel, O. nilgiriensis Cameron, O. subferrugineus Cameron, O. subincisus Cameron, O. sublucidus Cameron, O. subsculptus Cameron, O. antennalis Fauvel (New Caledonia, New Guinea and Australia), O. cheesmani Bernhauer (New Hebrides), O. cheesmanianus Cameron (Papua New Guinea), O. hingstoni Cameron (India), and O. tibetanus Bernhauer (China). The species O. (Anotylus) transversipennis Scheerpeltz is transferred to the genus Anotylus. Other than the new species, nine previously known species are redescribed, a key to 30 species and line drawings or color photographs of 42 species are provided. Thus, a total of 30 species are recorded from Southeast Asia. Key words: Staphylinidae, Oxytelinae, Oxytelus, Southeast Asia, Taxonomy, key, lectotypes, new synonyms, new species OXYTELUS IN SOUTHEAST ASIA

Zootaxa 3992 (1) © 2015 Magnolia Press ·

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Introduction The genus Oxytelus Gravenhorst, 1802 is the fourth most species-rich genus in the subfamily Oxytelinae (Coleoptera: Staphylinidae). It had 193 extant species in the latest catalogue of the family Staphylinidae (Herman 2001). With generic synonymies established by Makranczy (2006), the species number rises to 218 with modification by recent studies (Makranczy 2006, 2012; Schülke 2009, 2011, 2012; Lü & Zhou 2012). Members of Oxytelus live in the organic litter and other decaying habitats (Lü & Zhou 2012), and some Afrotropical taxa were also collected from or around nests of social insects. Oxytelus is distributed in all the Earth’s ecozones except Antarctica, among which the Afrotropics is, without doubt, its distribution center, harboring more than half of the whole number (139 species); Indo-Malaya (= Oriental Realm) is the second, with 38 species; Palearctic the third, with 29 species; and only a few species currently known in the Nearctic, Neotropics, Australasia, and Oceania. Southeast Asia consists of eleven countries: Brunei, Cambodia, Indonesia, Laos, Malaysia, Myanmar (Burma), Philippines, Singapore, Thailand, Timor-Leste (East Timor), and Vietnam (Fig. 1). It is one of the most important biodiversity hotspots, containing a large amount of species both widespread ones as well as those with exclusive occurrence and threatened by exceptionally serious loss of habitat (Myers et al. 2000; Sodhi et al. 2010; Koh et al. 2013). This region occupies a major part of Indo-Malaya and small parts of Australasia and Oceania and has 33 Oxytelus species as surveyed before this study. Bernhauer (1936) briefly treated the “Oxytelus” species in the Philippines, of which most are Anotylus in the current sense. Hammond (1975) reviewed the tribe Oxytelini from Ceylon (now Sri Lanka), and all of the seven treated Oxytelus species are common in Southeast Asia as well. Lü & Zhou (2012) revised the Oxytelus species known from China (21 spp.), out of which 14 species are found to cooccur in Southeast Asia. That is probably because of geographic adjacency, natural migration, or human activity. Nevertheless, after Scheerpeltz (1978), no more species have been described from this region. Different from the fauna of northern China or other many typical Palaearctic areas but similar to that of southwestern China, numerous species (e.g. O. javanus and O. ruptus; O. lividus, O. insulanus sp. nov., and O. sublividus sp. nov.; O. ferreirai, O. lucens, O. lompobatangensis sp. nov., and O. poecilopterus sp. nov.; O. lucidulus and O. subferrugineus; O. fallax, O. grandiculus sp. nov., O. subsculptus, and O. subincisus) found in Southeast Asia are exclusive to this region or smaller areas and also very closely related to sibling species that are apparently difficult to separate from each other. While our current knowledge does not allow separation of females for some species, luckily, male specimens have plenty and well diversified traits in genitalia and terminalia that distinguish congeners with such confusingly similar external appearance. There is a further batch of species (e.g. O. finitimus sp. nov., O. bellicosus, O. castaneus sp. nov.) with often strongly modified external appearance, but somewhat simplified genitalia, which is also found in the myrmecophilic species in Afrotropic (Gy. Makranczy, pers. comm.).

Material and methods During this study, a total of 776 specimens, including 203 types, collected from Southeast Asian and adjacent regions were examined. Specimens (or the parts of interest) were relaxed and cleared in hot saturated solution of potassium hydroxide (KOH) for 5 minutes (for mounted dry specimens) or 10 minutes (for alcohol-preserved specimens), and then washed in fresh water to stop further bleaching. The softened specimens were moved into glycerol and partly dissected for acquiring head capsule, mouthparts, terminalia, genitalia, and other structures for photographing or drawing. After this, all the removed parts were glued back onto the cards with Arabic gum or stored in glycerol vials for future study. This protocol is in accordance with that described in Lü & Zhou (2012, 2015). Observation, drawing, measurement, and photographing were performed using a Leica MZ APO (magnification: up to ×250) or a Zeiss Discovery V 20 (up to ×150) stereomicroscope with an ocular micrometer and mounted on Nikon D90/D300/D600 or Canon EOS 700D DSLR camera. The programs Nikon Capture NX2 (trial), digiCamControl, Canon EOS Utility, Canon Digital Photo Professional, Helicon Focus (trial), and Adobe Photoshop (trial) were used in remote control and digital post-processing of the color pictures, and the Inkscape was used to make the line-art plates. The map was drawn in Global Mapper (trial). All the calculation was performed in R 3.0.3 (R Core Team 2014).


LÜ & ZHOU

In this study as well as in our previous paper (Lü & Zhou 2012), we examined some type specimens (mainly from SDEI, BMNH, and FMNH) that were labeled as “lectotype” or “paralectotype” by P. M. Hammond (BMNH). Many of these “designations” were not published by himself, but the labels placed on the type specimens about 40 years ago could be misleading for the subsequent researchers (e.g. Gaedike 1981). Here we formally designated all those specimens involved as long as we saw Hammond’s “lectotype” specimen and followed most of his decisions unless the choice was controversial or the “lectotype” was seriously damaged and there were better candidates within the primary types. Consistent with Lü & Zhou (2015), the label data of type specimens were just copied verbatim and the “//” was used as a separator between each individual label; the data of other specimens were structured based on the information yet simply reproduced verbatim in quotation marks “ ” when the meaning was unclear. Chinese characters (including place names) were translated into English, even for those of type specimens (literally and in original order). Any notes made by us were placed in square brackets “[ ]”. In addition, a few explanatory words are necessary as regards a slight change of the style in illustration and verbal redescription compared to the preceding article (Lü & Zhou 2012). Consistency in a taxonomic article makes it easier for the readers to find relevant information. On one hand, this study lasted for five years, resulting in an obviously evolving methodology; on the other hand, changes were inflicted by necessities deriving from the conditions of type materials studied. In the earlier work, specimens were at our disposal for a complete dissection of both head and genital segments. This was not the case in several instances in the current study, for specimens with historical value and uniqueness. While dissection of the genitalia in the vast majority of depositories (museums) is currently understood as necessary for the correct establishment of taxon identity, procedures like bleaching and dissection of the head are to be avoided and considered destructive. This leads to neck and mandibular characters shown in less detail in some species here than in the earlier treatment. A further complication is caused by the fact that a previous worker, P. M. Hammond, who studied most of the type material in the former “Oxytelini” during the 1970s, habitually detached the parameres from the median lobes of the aedeagi and mounted these on the cards of the type specimens. This makes it very difficult to illustrate the types and the species known exclusively from their type materials comparable to the others. Species studied by us in earlier years were only illustrated with line drawings, as our lab was not equipped with a suitable microscopic digital photographing outfit then, but some photographs are here provided retrospectively. While the fourteen species that had already been treated in our article on the Chinese fauna (Lü & Zhou 2012) are not redescribed in this paper, additional photographs are provided. The descriptive terminology introduced by Makranczy (2006) and Lü & Zhou (2012) is followed.

The following abbreviations are used for type depositories, and museums or collections housing the material examined: BMNH: The Natural History Museum, London, UK; FMNH: Field Museum of Natural History, Chicago, Illinois, USA; HNHM: Hungarian Natural History Museum, Budapest, Hungary; IRSNB: Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium; IZ-CAS: Institute of Zoology, Chinese Academy of Sciences, Beijing, China; NHMB: Naturhistorisches Museum, Basel, Switzerland; NHMW: Naturhistorisches Museum, Wien, Austria; MSNG: Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italia; RMNH: Naturalis Biodiversity Centre [Rijksmuseum van Natuurlijke Historie], Leiden, the Netherlands; SDEI: Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany; SMNH: Swedish Museum of Natural History, Stockholm, Sweden; SMNS: Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany; TIPC: Tateo Ito’s Personal Collection, Yawata, Kyoto, Japan; ZMUC: Zoological Museum, University of Copenhagen, Copenhagen, Denmark.

OXYTELUS IN SOUTHEAST ASIA


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The following codes are used in Tab. 1 for the measurements (all measured from dorsal view): BL (body length): Approximate length from the front apex of head through the rear apex of abdomen. FBL (forebody length): Length from the front apex of head to the posterior margin of the elytra. HL (head length): Distance from the front apex of the clypeus through the level of demarcating points between temples (postocular region) and neck. HWE (head width with eyes): Maximum distance between the curves of eyes in dorsal view. HWT (head width at temples): Maximum distance between the outlines of temples in dorsal view. CL (clypeal length): distance from the anterior margin of clypeus to the posterior end of lateral portions of epistomal suture; EL (eye length): Distance between the line across the anterior apices of the eyes and the line across the posterior apices in dorsal view. TL (temple length): Distance between the line across the posterior apices of eyes and the line across the demarcating points between temple curves and neck in dorsal view. ETR (ratio of eye length at temple length): Ratio of EL at TL. PNL (pronotal length): Length of pronotum (measured along the midline). PNW (pronotal width): Width of the widest portion of pronotum. PNR (ratio of pronotal length at pronotal width): Ratio of PNL at PNW. ELA (average of elytra length): length from elytral anterolateral angle to the posterior margin, averaged between left and right; EWA (average of elytra length): width between the elytral suture and the lateral margins, averaged between left and right; ABDW (abdominal width): The width of the widest segment of abdomen.

Taxonomy Key to species from Southeast Asia 1. 2. 3. 4. 5. 6. 7. 8.

9. 10.

Antennomere 4 without basal dish (Fig. 1F in Lü & Zhou 2012). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Antennomere 4 with basal dish (Fig. 1G in Lü & Zhou 2012) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Supra-antennal ridge formed as sharp spine pointing anteriorly (Fig. 3B, C) . . . . . . . . . . . . . . . . . . . . . . . 1. O. armiger Fauvel Supra-antennal ridge not like above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Posterior margin of ♂ sternite VIII narrowly bi-ruptured near middle (as in Fig. 6D, J). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Posterior margin of ♂ sternite VIII without such ruptures (as in Figs. 11F; 21F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Elytral lateral longitudinal ridge distinct (as in Fig. 4C, D); ♂ sternite VII with posterior margin protruding in middle (as in Fig. 6C); ♀ sternite VIII with posterior margin protruding (as in Fig. 8G) or sinuate in middle (as in Fig. 9J) . . . . . . . . . . . . . 5 Elytral lateral longitudinal ridge feeble or invisible (Fig. 5D, F); ♂ sternite VII with posterior margin straight or bi-denticulate in middle (Figs. 7E; 12E in Lü & Zhou 2012); ♀ sternite VIII with posterior margin evenly curved (as in Fig. 46G). . . . . . . 10 Pronotal lateral margin crenulate in posterior half (as in Fig. 4C, D); aedeagal apico-medial hook with apex not upcurved at tip (as in Fig. 6G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Pronotal lateral margin not crenulate (or finely crenulate at posterior angles, as in Fig. 8C); aedeagal apico-medial hook with apex upcurved at tip (as in Fig. 8K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Forebody surface grainy, barely punctate (Fig. 4C, D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. O. bengalensis Erichson Forebody surface not grainy, densely punctate (as in Fig. 5B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 ♂ sternite VII with posterior margin broadly protruding in middle and truncate at apex (Fig. 13E in Lü & Zhou 2012) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. O. nigriceps Kraatz ♂ sternite VII with posterior margin protruding in middle but narrowing to the apex (Fig. 6C) . . . . . . . 4. O. javanus Cameron Pronotum and elytra variegated: pronotum brown in center and yellowish around with black margin, elytron with wide and longitudinal darker stripe in middle (Fig. 8A, B); ♂ median plate of sternite VIII with two long teeth at ends of posterior margin (Fig. 8F); ♀ sternite VIII sharply protruding in middle of posterior margin (Fig. 8G) . . . . . . . . . . . . . . 5. O. ruptus Fauvel Pronotum and elytra not variegated and without above patterns; ♂ median plate of sternite VIII without teeth on posterior margin; ♀ sternite VIII bi-sinuate in middle of posterior margin (as in Fig. 9J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 ♂ median plate of sternite VIII with tiny denticle in middle of posterior margin (Fig. 14F in Lü & Zhou 2012), paramere with highlight band near apex (as in Fig. 6H). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. O. piceus (Linné) ♂ median plate of sternite VIII with tiny notch in middle of posterior margin (Fig. 22F in Lü & Zhou 2012), paramere without highlight band near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. O. varipennis Kraatz Tergite X without incision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. O. migrator Fauvel


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11. 12. 13. 14. 15. 16. 17. 18. -

19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. -

Tergite X with deep incision at middle of posterior margin (Fig. 7H, I in Lü & Zhou 2012) . . . . . . . 9. O. incisus Motschulsky Posterior margin of ♂ sternite VIII modified by only two teeth in middle (as in Fig. 11F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Posterior margin of ♂ sternite VIII modified more complicated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Anterior angles of pronotum prominent and slightly constricted a little behind (as in Fig. 12B) . . . . . . . . . . . . . . . . . . . . . . . 13 Anterior angles of pronotum round (Fig. 11C, D). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. O. finitimus sp. nov. ♂ clypeus with two spines (Figs. 12B; 13A) or triangular protrusions (as in Fig. 13B) . . . . . . . . . . . . .11. O. bellicosus Fauvel ♂ clypeus without spines but with round protrusions (Fig. 15B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12. O. castaneus sp. nov. Eyes with coarse facets (as in Fig. 16B, D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Eyes with fine facets (as in Fig. 22C, G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Posterior margin of ♂ sternite VII with two incurved small teeth in middle and narrow and shallow emargination between teeth (Fig. 18C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. O. sublividus sp. nov. Posterior margin of ♂ sternite VII with two outcurved larger teeth near middle and gradually narrowed and deep emargination between teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Emargination on posterior margin of ♂ sternite VII with lateral edge evenly curved (Fig. 16F), aedeagus as Fig. 16H–L . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. O. lividus Motschulsky Emargination with lateral edge nearly parallel in anterior half (Fig. 18A), aedeagus as Fig. 19A–E . . 14. O. insulanus sp. nov. Lateral margin of pronotum with crenulation (at least posterior half, as in Figs. 20B, F; 22C, G) . . . . . . . . . . . . . . . . . . . . . . 18 Lateral margin of pronotum without crenulation (as in Fig. 27C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Clypeus weakly protruding, anterior margin with two pointed teeth; ♂ sternite VIII with broad triangular plate on posterior margin, furnished with tiny teeth at each side (Fig. 20C, G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. O. punctipennis Fauvel Clypeus different between sexes; ♂ clypeus longer than wide and obviously protruding, anterior margin slightly emarginate forming two blunt teeth (as in Fig. 22C, G), sternite VIII with narrow lobe protruding from middle of posterior margin (as in Fig. 23D); ♀ clypeus slightly protruding, anterior margin truncate or slightly emarginate (as in Fig. 22B, D). . . . . . . . . . . . . 19 Lateral margin of pronotum crenulate; ♂ teeth on posterior margin of sternite VII tiny (as in Fig. 21E), posterior margin of sternite VIII not emarginate beside the middle lobe (as in Fig. 21F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Lateral margin of pronotum not crenulate in front half; ♂ teeth on posterior margin of sternite VII more prominent than above (Fig. 25A, E), posterior margin of sternite VIII emarginate beside the middle lobe (Fig. 25B, F) . . . . . . . . . . . . . . . . . . . . . . 21 ♂ aedeagal paramere expanded near apex (Fig. 21J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. O. ferreirai Scheerpeltz ♂ aedeagal paramere narrowed near apex (Fig. 23B, G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. O. lucens Bernhauer Elytra with color pattern from anterior angle to posterior margin (Fig. 24B, D); ♂ aedeagal paramere narrowed near apex and with short and broad process near corner (Fig. 26D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. O. lompobatangensis sp. nov. Elytra with color pattern at posterior angle (Fig. 24G, I); ♂ aedeagal paramere expanded near apex and short and without broad process near corner (Fig. 26H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20. O. poecilopterus sp. nov. Elytra variegated (Fig. 27A–C). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. O. barbatus Fauvel Elytra not variegated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 ♂ clypeus and mandibles extremely elongate twice as long as main part of head (Fig. 28C), posterior margin of sternite VII as in Fig. 28F22. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. mandibularis Cameron ♂ clypeus and mandibles not elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 ♂ clypeus and mandibles asymmetric (Fig. 29B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. O. puncticeps Kraatz ♂ clypeus and mandibles symmetric. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 ♂ sternite VII with shallow or feeble emargination in middle (Fig. 31C), sternite VIII roundly emarginate beside median lobe, with crest-shaped patch (Fig. 10G in Lü & Zhou 2012) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. O. lucidulus Cameron ♂ sternite VII with deep, broad, arcuate emargination in middle (Fig. 32G), sternite VIII with two V-shaped emarginations beside median lobe (Fig. 32H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25. O. subferrugineus Cameron Eyes with fine facets (Fig. 35B); ♂ clypeus protruding (Fig. 34C, D) . . . . . . . . . . . . . . . . . . . . . . . . . . 26. O. megaceros Fauvel Eyes with coarse facets (as in Fig. 36B); ♂ clypeus almost truncate (as in Fig. 42B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 ♂ sternite VII with posterior margin broadly emarginate and with membranous border in emargination (Fig. 36C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. O. fallax Fauvel ♂ sternite VII with posterior margin straight (as in Fig. 40F) or slightly emarginate (Fig. 38E). . . . . . . . . . . . . . . . . . . . . . . . 28 Body larger (≥ 4.5 mm); ♂ tergite VIII depressed in center and posterior margin incrassate (Fig. 38F). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. O. grandiculus sp. nov. Body smaller (≤ 4 mm); ♂ tergite VIII normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Metatibia widest at basal 1/3 (Fig. 40E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. O. subsculptus Cameron Metatibia widest at nearly middle (Fig. 41D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. O. subincisus Cameron

1. Oxytelus armiger Fauvel, 1895 (Figs. 2 & 3) Fauvel 1895: 202 (Type locality: Bírmanie, Carin Chebà, 900–1100 m).

Type material examined. Lectotype [designated here, Fig. 3]: ♂, Syntype// Carin [=Kayah, Myanmar] Chebà, 900-1100.m., L. Fea V XII-88 [=V–XII.1888]// armiger Fvl.// Coll. et det A. Fauvel, Oxytelus armiger Fauv., OXYTELUS IN SOUTHEAST ASIA


7

R.I.Sc.N.B. 17.479 (IRSNB); Paralectotypes: 1♂, Carin [=Kayan, Myanmar] Chebà, 900–1100.m., L. Fea V XII88// Coll. et det A. Fauvel, Oxytelus armiger Fauv., R.I.Sc.N.B. 17.479 (IRSNB); 1♀, Syntype// Carin [=Kayan, Myanmar] Chebà, 900–1100.m., L. Fea V XII-88// Coll. et det A. Fauvel, Oxytelus armiger Fauv., R.I.Sc.N.B. 17.479 (IRSNB). Other material examined. 5 specs. (5♀♀). LAOS: Champasak, Bolaven Plateau, Ban Thôngvay (=Xékatam) vic.: 2♀♀, old logging road, N of village, 15º14.494'N 106º31.807'E, 1170m, selectively logged forest, dung trap (human), A. Solodovnikov, M. Thayer & A. Newton legg. (FMNH); 3♀♀, Muang Paxong, 15º14.054'N 106º31.867'E, ca.1200m, Edge of disturbed primary rainforest, near clearing, flight intercept trap, 2008.VI.8–16. A. Solodovnikov & J. Pedersen legg. (ZMUC). Diagnosis. This species can be readily recognized by the combination of the reddish color, the fine-faceted eyes (Fig. 2B, C), the spine-formed supra-antennal ridges (Fig. 2B, C), and the absence of aedeagal apico-medial hook (Fig. 2I, J). Redescription. Body (Fig. 3A) dark rufescent to blackish, and shining; head and pronotum darker and elytra a little lighter in some cases. Maxillary palpi, antennae, and legs testaceous. Length [average]: ♂, 5.9 mm; ♀, 5.5 mm. Male. Head (Figs. 2A; 3B) widest at eyes or temples. Disc as well as supra-antennal ridges densely punctate, nearly glabrous. Clypeus transverse, shorter than 1/3 head length, surface glabrous, coriaceous, scattered with a few fine punctures; anterior margin protruding and narrowed forward, furnished with two triangular teeth in near middle (in some case the teeth upward pointed). Supra-antennal ridges produced forward into long, punctate, pointed, and slightly out- and down-curved spines, beyond clypeus. Epistomal suture with lateral portions straight, extremely short, medially directed, and extending to level of anterior margin of eyes. Vertex densely punctate, concave, posterior part well-delimited; mid-longitudinal suture fine and obvious, longitudinal paralateral sutures almost invisible. Eyes with fine facets, shorter than temples. Temples broadly rounded and dilated. Occipital suture with middle portion present; nuchal ridge a little feeble and interrupted in middle, dorsal basal ridge absent. Mandible (Fig. 2C) robust and moderately curved; two denticles on inner edge, one near middle and the other at apical 1/4. Antenna (Fig. 3A) as long as head and pronotum together; antennomere 4 subglobose, without basal dish; apical antennomere as long as or a little shorter than preceding two together. Pronotum (Fig. 3B) transverse, broadest at around anterior 1/4, broader than head. Disc 5-sulcate, median sulcus and two slightly outcurved paramedial sulci deep and punctate; two paralateral sulci straight, deep, and shorter. Lateral margins evenly curved in front but slightly crenulate behind, posterior angles not prominent. Elytra (Fig. 3A) punctate and longitudinally rugose around dorsal disc, with epipleural ridge present on each piece. Abdomen coriaceous and pubescent, broadest at segment V. Sternite VII (Figs. 2E; 3D) with posterior margin broadly and shortly protruding in middle but narrowed posteriorly. Sternite VIII (Figs. 2F; 3E) with transverse subbasal ridge continuous and straight in middle, posterior margin with two deep and narrow emarginations at each side forming rectangular plate in middle, apical margin of median plate broadly rounded. Tergite X (Fig. 2H) longer than wide, parallel sided, posterior margin with pointed tooth on each side and truncate in between, with two lines of setae on pubescent posterior part. Aedeagus (Figs. 2K–N; 3G–L) with median lobe oblong-ovoid, broadest at middle, with membranous internal sac inside (no sclerites); apico-medial hook absent; dorsal membranous area covering majority of dorsum of median lobe and across whole length, in each lateral wall near apex there is membranous semicircular excision. Paramere sword-like, with furrow demarcating apical and basal arm; basal arm furnished with an upward lamella; apical arm roundly curved in lateral view, with ventrally directed seta at apical 1/3. Female. Head (Figs. 2B; 3C) smaller than male, clypeus broad, with anterior margin gradually narrowed apically and truncate or a little emarginate in middle but without teeth, supra-antennal ridges protruding into spines shorter than male, temples slightly dilated but not so much as male, vertex flat or a little convex. Mandible (Fig. 2D) shorter and slenderer than male. Abdominal sternite VII without denticles on posterior margin. Sternite VIII (Figs. 2G; 3F) with posterior margin broadly rounded but shortly protruding in middle. Tergite X (Fig. 2I) with anterior margin narrower and posterior margin with two teeth on sides and protruding a little in middle. Spermatheca (Figs. 2J; 3M) dumbbell-shaped and bent at middle, moderately inflated into sphere at both base and apex. Distribution. Laos, Myanmar. Remarks. One of the most astonishing finds when dissecting and examining a male syntype of this species is the lack of apico-medial hook on the aedeagal median lobe. But there is, instead, a weakly sclerotized lamella at the


LÜ & ZHOU

basal end of each paramere (Figs. 2K; 3K), which seems to function as the lost apico-medial hook. The other character that makes this species unique and easily recognized is the spine-like supra-antennal ridges in both male and female individuals.

2. Oxytelus bengalensis Erichson, 1840 (Figs. 4; 6J) Erichson 1840 : 789 (Type locality: Bengala). Oxytelus bicolor Walker 1859: 52 (Type locality: Ceylon); Scheerpeltz 1933: 1094 (syn. of bengalensis). Oxytelus opacifrons Sharp 1874: 93 (Type locality: Japan); Hammond 1975: 155 (syn. of bengalensis).

Material examined. 39 specs. (24♂♂, 15♀♀). VIETNAM: 9♂♂, 3♀♀, Tonkin, Hoa-Binh, A. De Cooman leg., Oxytelus bengalensis Er. det. P.M.Hammond 1980. LAOS: Vientiane Pr., Phou Khao Khouay NBCA: 2♂♂, Ban Vangheua (=Khua) school, 18º20.37'N 102º48.529'E, 775m, settlement near forests & rice fields, at light (UV & MV), 2008.V.25–30, A. Newton, M. Thayer et al. legg. (FMNH); 1♂, 18º20.369'N 102º48.523'E, 700–800m, nr. strongly distributed primary rainforests, on light, 2008.V.25–30, A. Solodovnikov & J. Pedersen legg. (ZMUC). MYANMAR: 1♀, E Shan State, Kengtung (Kyaingtong), 1997.VI.14–15, J. Rejsek leg. (SMNS). NEPAL: 1♂, 1♀, Kathmandu, Baneshwar, 1350m, 2000.VI.18–24, W. Schawaller leg. (SMNS); 2♂♂, Kaski distr., Begnas Lake, 780m, at light, 1994.X.11, G. Csorba & L. Ronkay legg. (HNHM). SRI LANKA: 1♂, 3♀♀, Ceylon E.Prov [= Eastern Prov.], Pottuvil, 1983.VII.1–12, Ole Mehl leg., Sri Lanka, Ex coll. Viggo Mahler (ZMUC); 1♂, same data as previous except: W. Prov. [= Westen Prov.]1983.VIII.2 (ZMUC); 1♀, same data as previous except: 1983.VI.17–19 (ZMUC); 1♀, E.Prov., Panama area, 1995.I.2–6, Ole Mehl legit, Sri Lanka, Ex coll. Viggo Mahler (ZMUC). THAILAND: N Khon Kaen: 1♂, 2♀♀, ad lucem [=at light], 1981.I.26, S. Saowakontha leg. (HNHM); 3♂♂, same data as previous except: 1981.II.20 (HNHM); 3♂♂, 2♀♀, same data as previous except: 1981.II.21 (HNHM); 1♀, same data as previous except: 1981.II.25 (HNHM). Diagnosis. Oxytelus bengalensis can be readily recognized by the reddish brown color (Fig. 4A, B) and the grainy surface (Fig. 4C, D). Apart from these, it can be separated from O. nigriceps by the shape of posterior margin of male sternites VII and VIII (Fig. 5E, F in Lü & Zhou 2012), and aedeagal characters (cf. Figs. 5E, F; 13E, F in Lü & Zhou 2012); can be separated from O. javanus by the shorter apex (the apical portion delimited by the seta) of aedeagal paramere (compare Fig. 6H with Fig. 5M in Lü & Zhou 2012). Distribution. Laos, Malaysia, Myanmar, Singapore, Thailand, Vietnam; Bangladesh, China, India, Japan, Nepal, Pakistan, South Korea, Sri Lanka.

3. Oxytelus nigriceps Kraatz, 1859 (Fig. 5A, B) Kraatz 1859: 171 (Type locality: India orientalis; Ceylan).

Material examined. 72 specs. (43♂♂, 29♀♀). BANGLADESH: 6♂♂, 1♀, Bengala [= Bangladesh + INDIA: West Bengal], Oxytelus nigriceps Kr., P.M. Hammond det. 1973 (ZMUC). MALAYSIA: 1♂, Malay Insel, Kraatz leg., Oxytelus nigriceps Kr. (minor) det. M. Bernhauer (FMNH); 1♂, Borneo, Sabah, Kinabalu, N.P., Poring [=Poring Hot Springs], 500m, 1996.XI.15–16, W. Schawaller leg. (SMNS). NEPAL: 2♂♂, Kaski distr., Begnas Lake, 780m, at light, 1994.X.11, G. Csorba & L. Ronkay (HNHM). PHILIPPINES: 1♂, Leyte Prov., San Jose, at light, 1945.I.30-31, E. Ray leg. (FMNH). SRI LANKA: 1♀, Tissamaharama, 1979.XII.12, Oxytelus nigriceps det. anonym, Sri Lanka Ex coll. Viggo Mahler (ZMUC). PAPUA NEW GUINEA: 3♂♂, 3♀♀, Bismarck Islands, Lavongai, Banatam, 1962.III.23, Noona Dan Exp. 61-62, caught by mercury-light (ZMUC). VIETNAM: 1♂, Prov. Nghe-An, forestière Quy-châu, 200m, à la lumière, forêt pluv. trop., semidecidue, 1963.VIII.23, T. Pócs leg. (HNHM); 2♀♀, same data as previous except: 1963.VIII.26 (HNHM); 2♂♂, Hanôi, 40m, l’hôtel, à la lumière, 1963.X.29, T. Pócs leg. (HNHM); 1♂, same data as previous except: 1963.X.30 (HNHM); 1♂, 1♀, same data as previous except: 1963.X.31 (HNHM); 2♂♂, same data as previous except: 1963.XI.1 (HNHM); 2♂♂, 1♀, same data as previous except: 1963.XI.2 (HNHM); 1♂, 3♀♀, Hanôi, 1963.IX.11–19, Manninger leg. (HNHM); 4♂♂,



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6♀♀, same data as previous except: 1963.XI.4–10 (HNHM); 1♀, same data as previous except: 1963.XI.20–30 (HNHM); 1♀, same data as previous except: 1963.XII.1–10 (HNHM); 1♂, 1♀, same data as previous except: 1963.XII.11–20 (HNHM); 2♂♂, same data as previous except: 1964.I.6–7 (HNHM); 8♂♂, 6♀♀, Hanôi, Xuan dinh, singled from under dead snake, Nr. 220, 1966.IV.26–29, Gy. Topál leg. (HNHM); 1♀, Tuong linh, near Phu ly, collected by lamp, Nr. 565, 1966.V.24–28, Gy. Topál leg. (HNHM); 1♂, River Chay, No. 248, 1971.XII.6, Gy. Topál leg. (HNHM); 2♂♂, 1♀, Thanh Hóa, 10 km N from city, at light, No. 53, 1986.I.23, Mahunka-Oláh legg. (HNHM); 1♂, Trung Trang, I. Cat Ba, 20º48'N 107º00'E, 200m, at light, No. 171, 1987.V.16, Matskási, Oláh & Topál legg. (HNHM). Diagnosis. Oxytelus nigriceps can be separated from other species in piceus group by the combination of the shape of posterior margin of male sternites VII and VIII, and the carination of apico-medial hook (Fig. 13 in Lü & Zhou 2012). It can be separated from O. bengalensis by the not grainy surface, from O. javanus and O. ruptus by the broader and truncate border of the posterior protrusion of male sternite VII, and from O. piceus by the tiny tooth at each end of posterior margin of median plate of male sternite VIII. The cephalic sexual dimorphism of O. nigriceps is very weak compared with O. bengalensis, O. javanus, O. piceus, and O. ruptus. Distribution. Indonesia, Malaysia, Myanmar, Philippines, Singapore, Thailand, Vietnam; Bangladesh, Bismarck Islands, China, India, Japan, Nepal, New Guinea, Pakistan, South Korea, Sri Lanka.

4. Oxytelus javanus Cameron, 1936 (Fig. 6A–I) Cameron 1936: 32 (subgenus Caccoporus; Type locality: Java: Tjilatjap).

Type material examined. Lectotype [designated here, Fig. 6A–I]: ♂, JAVA: F. C. Drescher. B.M. 1934-264.// Tjilatjap, Java Drescher, 4.XII.1926// O. javanus, TYPE Cam.// Oxytelus javanus Cam., P.M.Hammond det. 1970 LECTOTYPE ♂ (BMNH). Other material examined. 1♀, PHILIPPINES: Luzon, Baker, Mt. Makiling, Oxytelus javanus Cam det. P.M.Hammond 1977 (FMNH). Diagnosis. Oxytelus javanus can be separated from other species in piceus group except O. bengalensis by the combination of the shape of posterior margin of male sternites VII and VIII (Fig. 6C, D), and from O. bengalensis by the longer apex (the apical portion delimited by the seta) of aedeagal paramere (Fig. 6H). Redescription [lectotype]. Body (Fig. 6A) light and yellowish brown; head a little darker and elytra slightly infuscate. Length: 4.6 mm. Head (Fig. 6B) widest at eyes. Disc punctate posteriorly, not pubescent. Clypeus subrectangular, flat, almost 1/ 4 head length, surface impunctate; anterior margin truncate. Epistomal suture with lateral portions incurved and running posteriorly to level of anterior margin of eyes. Vertex weakly convex, definitely distinguished from neck by middle portion of occipital suture; with mid-longitudinal suture distinct. Eyes with coarse facets, convex and protruding laterally, twice as long as temples. Temples slightly dilated. Occipital suture broadly curved in dorsal view; nuchal ridge feeble in middle. Mandible slender and slightly curved. Antenna (Fig. 6A, showing only basal 7 articles) a little shorter than head and pronotum together; antennomere 4 subglobose, without basal dish; and apical antennomere as long as or a little shorter than preceding two together. Pronotum (Fig. 6B) transverse, broadest at near anterior 1/3, wider than head. Disc punctate and 3-sulcate; two paralateral depressions shallow, striate; lateral margin smooth in front and weakly crenulate behind, with posterior angles not prominent. Elytra (Fig. 6B) rugose and finely punctate, with lateral longitudinal ridge present. Abdomen coriaceous and covered with dense fine setae, broadest at segment V. Sternite VII (Fig. 6C) with posterior margin shortly protruding in middle and slightly emarginate at apex. Sternite VIII (Fig. 6D) with subbasal ridge curved; posterior margin with deeply V-shaped rupture at each 1/3, forming large and rectangular median plate, and posterior margin of which bi-emarginate, with sharp tubercle in center, with mid-longitudinal internal ridge. Aedeagus (Fig. 6E–H) with median lobe oblong in ventral view, wider in basal half; with membranous internal sac inside (no sclerites); apical orifice smaller than median lobe width; apico-medial hook U-shaped and bent at middle, with apical tip pointed, basal process well developed and with round tip; dorsal membranous area covering


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almost dorsum. Paramere with seta dwelling at 1/2 of apical arm, with highlight band nearby; apex pointed; basal arm with shallow furrow. Distribution. Indonesia, Philippines. Remarks. Similar to O. bengalensis, O. nigriceps, and O. ruptus. O. javanus differs from the other three species in color: O. javanus is lighter and prone to be yellow and without obvious dark patches; and in the parameres of aedeagus: the portion anterior to the situation of parameral seta is much longer than the other three (cf. Figs. 6E, F, H; 7K, L, N; 8I, J, L and Lü & Zhou (2012: 18 Fig. 5K–M; 37 Fig. 13K, L, N)). Additionally, the protrusion on the posterior margin of sternite VII of male O. javanus is a little shorter than that in O. bengalensis (cf. Fig. 6C and Lü & Zhou (2012: 18 Fig. 5E)), and it is slightly emarginate in apex where it is truncate in that of O. nigriceps. The male O. ruptus has two prominent processes on the posterior margin of the median plate of sternite VIII (Figs. 7F; 8F) but O. javanus does not have the homologous processes (Fig. 6D).

5. Oxytelus ruptus Fauvel, 1904 (Figs. 7; 8; 9E–G) Fauvel 1904: 100 (Type locality: Java: Monts Tengger, 1300 m; Sukabumi. Sumbawa: Mont Tambora. Celebes). Oxytelus (Caccoporus) sublucidus Cameron 1941: 433 (Type locality: Philippines: Mindanao; Momungan). Syn. nov.

Type material examined. [O. ruptus] Syntype: 1♀, Oxytelus ruptus Fvl., Sumbawa// Coll. R. I. Sc. N. B. (IRSNB). [O. sublucidus] Lectotype [designated here, Fig. 9E–G]: 1♀, Momungan Mindanao// sublucidus Brnh. Cotyp./ / O. sublucidus Cam. TYPE// M. Cameron. Bequest. B.M.1955–147// Oxytelus sublucidus Cam., P.M.Hammond det. 1970, LECTOTYPE ♀ (BMNH). Other material examined. 12 specs. (5♂♂, 7♀♀). PHILIPPINES: 2♂♂, 5♀♀, Luzon, Imugan, Boettcher leg., ruptus det. Bernhauer (FMNH); 1♂, Luzon, Los Banos, Boettcher leg., ruptus det. Bernhauer, Oxytelus ruptus Fvl det. P.M.Hammond 1975 (FMNH); 1♂, Luzon, Buranen, Boettcher leg., ruptus det. Bernhauer (FMNH); 1♂, Oxytelus ruptus Fvl det. P.M.Hammond 1977 (FMNH); 1♀, Leyte, Imugan, Boettcher leg. (FMNH); INDONESIA: 1♀, Sumatra, Fort de Kock [=Bukittinggi], 920m, 1926, E. Jacobson leg., ruptus Fauv det. M. Cameron (FMNH). Diagnosis. Oxytelus ruptus can be separated from other species in piceus group by its particular coloration (Fig. 8A–D) and the combination of the shape of posterior margin of male sternites VII and VIII (Fig. 8E, F). The female of O. ruptus can be also separated by the shapely protruding posterior margin of sternite VIII (Fig. 8G). Redescription. Body (Fig. 8A, B) yellowish brown with dark patches; head pitchy, pronotum with dark patches in center and margins, elytron with dark patch in center and reaching posterior margin; abdomen with longitudinal dark patches along center and boundary between tergites and laterosternites. Length [average] ♂, 4.0 mm; ♀, 3.7 mm. Male. Head (Figs. 7A; 8C) broader than long, widest at eyes. Disc densely punctate, less pubescent. Clypeus not protruding, shorter than 1/3 head length, anterior margin weakly up-flanging, surface impunctate and weakly coriaceous; anterior margin truncate. Supra-antennal ridges moderately elevated upward. Vertex slightly convex, definitely distinguished from neck by middle portion of occipital suture; with mid-longitudinal suture distinct. Eyes with coarse facets, strongly convex and protruding laterally, 2.5 times as long as temples. Temples broadly rounded at posterior angles. Occipital suture arcuate in dorsal view; nuchal ridge present and visible in middle, dorsal basal ridge present. Mandible (Fig. 7C) stout, slightly curved; two denticles on inner edge near middle. Antenna (Fig. 8A) a little shorter than head and pronotum together; antennomere 4 subglobose, without basal dish; apical antennomere twice longer than preceding one Pronotum (Fig. 8C) transverse, broadest at near anterior 1/4, slightly wider than or equal to head. Disc 3sulcate, and with two shallow paralateral depressions; lateral margin not crenulate, posterior angles not prominent. Elytra (Fig. 8A) punctate and rugose, and with lateral longitudinal ridge present. Abdomen pubescent on ventral side, broadest at segment V. Sternite VII (Figs. 7E; 8E) with posterior margin shortly produced in middle and weakly emarginate at apex. Sternite VIII (Figs. 7F; 8F) with subbasal ridge curved in middle, posterior margin with deeply V-shaped rupture at each 1/3, forming large and subrectangular median plate, apical margin of median plate a little produced in middle, in center of median plate with small tubercle, at



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each posterolateral corner with prominent process bearing seta, with mid-longitudinal internal ridge on inside surface. Aedeagus (Figs. 7K–N; 8I–L) with median lobe oblong-ovoid, widest at middle; with membranous internal sac inside (no sclerites); apical orifice large; apico-medial hook C-shaped in lateral view and bent at near middle, with apical tip pointed and upcurved at tip, basal process well developed and with round tip; dorsal membranous area covering most part of dorsum and across length of median lobe. Paramere arm-like, with seta on apical 1/3 of apical arm and with highlight band nearby; apex pointed; with shallow furrow on basal arm. Female. Head (Figs. 7B; 8D) smaller than male. Eyes convex and protruding laterally beyond temples, temples almost invisible in lateral view. Mandible (Fig. 7D) smaller, slenderer and shaper than male. Abdominal sternite VII with posterior margin straight and not modified. Posterior margin of sternite VIII (Figs. 7G; 8G) sharply protruding in middle. Spermatheca (Figs. 7J; 8H) slim and roundly incurved at apex. Distribution. Indonesia, Philippines. Remarks. When examining the types of O. ruptus and O. sublucidus, these were found to share the same coloration and terminalia. Thus we synonymize O. sublucidus with O. ruptus, although both of the two species were originally described based on female types.

6. Oxytelus piceus (Linné, 1767) (Fig. 9A–D) Linné 1767: 686 (Staphylinus; Type locality: Upsaliae). Oxytelus humilis Heer 1839: 204 (Type locality: Genf); Erichson 1840: 788 (syn. of piceus). Oxytelus japonicus Motschulsky 1862: 10 (Type locality: Japan); Fauvel 1902: 64 (syn. of piceus). Oxytelus mamillatus Hochhuth 1851: 53 (Type locality: Dahurien); Gusarov 1991: 4 (Oxytelus; syn. of piceus; lectotype designation). Oxytelus piceus ssp. defectivus Normand 1947: 5 (Type locality: Afrique du Nord); Herman 2001: 1450 (syn. of piceus). Oxytelus sulcatus Gebler 1848: 79 (preoccupied; Type locality: Not cited); Fauvel 1876a: 242 [= Fauvel 1876b: 57] (syn. of piceus). Staphylinus sulcatus Müller 1776: 97 (Type locality: Danicae); Olivier 1811: 614 (Oxytelus; syn. of piceus).

Material examined. 14 specs. (7♂♂, 7♀♀). LAOS: 1♂, Vientiane Pr., Phou Khao Khouay NBCA, Ban Vangheua (=Khua) school, 18º20.37'N 102º48.529'E, 775m, settlement near forests & rice fields, at light (UV & MV), 2008.V.25–30, A. Newton, M. Thayer et al. legg. (FMNH); N. KOREA: 1♀, Prov. South Pyongan [=Phyŏngannamdo], Pyonggyan [=Pyongyang], room of Hotel Te-dong, No. 309, 1975.VIII.2, J. Papp & A. Vojnits legg. (HNHM); 1♀, Mt. Pektusan [=Changbaishan =Mt. Changbai], before Sam-zi yan hotel, lake-shore, light trap, No. 379, 1977.VII.19, Dely & Draskovits legg. (HNHM); 1♂, Prov. N Hwanghae [=Hwanghaebukto], Sariwon [=Sariwŏn], 1978.IX.28, No. 425, Dr. A. Vojnits & L. Zombori legg. (HNHM); S. KOREA: 4♂♂, 5♀♀, Pusan [=Busan], on human excrement, 1945.X.30, E. Ray leg. (FMNH); MALAYSIA: 1♂, Borneo, Sabah, Tamboonan, 1984.III.29–30, G. Hangay leg. (HNHM). Diagnosis. Oxytelus piceus can be separated from other species in piceus group except O. varipennis by the pichy and more striate head and pronotum (Fig. 9A–D) and the combination of the shape of posterior margin of male sternites VII and VIII (Fig. 14E, F in Lü & Zhou 2012). The female of O. piceus can be also separated by the sinuate posterior margin of sternite VIII (Fig. 14G in Lü & Zhou 2012). It can be separated from O. varipennis by the tiny and pointed tooth in middle (cf. Figs. 14F; 22F) and by the aedeagal (cf. Figs. 14K–M; 22K–N) and spermathecal characters (cf. Figs. 14J; 22J). Distribution. Laos, Malaysia; Africa (including Madagascar, Canary Islands, and Madeira), Azerbaijan, China, Europe, Georgia, Iran, Japan, Kazakhstan, Kyrgyzstan, Mongolia, North & South Korea, Russia, Saudi Arabia, Syria, Tajikistan, Uzbekistan.

7. Oxytelus varipennis Kraatz, 1859 (Fig. 9H–J) Kraatz 1859: 172 (Type locality: Ceylan).


LÜ & ZHOU

Oxytelus varipennis ssp. pharaonum Koch 1934: 44 (Type locality: Aegypten: Pyramiden von Ghizeh; Mansourieh; Sakkarah; Ismailia).

Type material examined. Lectotype [inadvertently designated by Gaedike (1981) referring to label of Hammond placed in 1974, Fig. 9H–J]: ♀, 168.// Ceylon// Coll. Kraatz// Oxytelus varipennis Kr. P.M. Hammond det. 1974 LECTOTYPE ♀// coll. DEI Müncheberg// DEI Müncheberg, Col-03033// Oxytelus varipennis Kr. (SDEI). Paralectotypes: 1♀, Ceylon// Coll. Kraatz// Oxytelus varipennis Kr. Paralectotype P.Hammond det. 1974 ♀// coll. DEI Müncheberg// DEI Müncheberg, Col-03034 (SDEI); 1♀, Ceylon// Coll. Kraatz// Oxytelus varipennis Kr. Paralectotype P.Hammond det. 1974 ♀// coll. DEI Müncheberg// DEI Müncheberg, Col-03035 (SDEI); 1♀, Ceylon// Coll. Kraatz// Oxytelus varipennis Kr. Paralectotype P.Hammond det. 1974 ♀// coll. DEI Müncheberg// DEI Müncheberg, Col-03036 (SDEI). Other material examined. 10 specs. (3♂♂, 7♀♀). BANGLADESH: 2♀♀, Bengala [= Bangladesh + INDIA: West Bengal], Oxytelus varipennis Kr., P.M. Hammond det. 1973 (ZMUC). INDIA: 2♀♀, India occ., Maharashtra state, 40 km W of Pune, MULSHI env., F. Kantner leg. (SMNS). INDONESIA: 1♀, Madura Is., Chambaganoor, Donckier leg. (FMNH); 2♂♂, 1♀, same data as previous except: 1912 [year] (FMNH). SRI LANKA: 1♂, 1♀, Hambantota, 1979.XII.10, Sri Lanka, Ex coll. Viggo Mahler (ZMUC). Diagnosis. Oxytelus varipennis can be separated from other species in piceus group except O. piceus by the combination of the pichy and less striate head and pronotum (Fig. 9H, I) and the tiny notch in the middle of posterior margin of median plate of male sternite VIII. The female O. varipennis differs from O. piceus in spermathecal shape (cf. Figs. 14J; 22J in Lü & Liang 2012). Distribution. Indonesia, Myanmar; Bangladesh, China, India, Japan, Nepal, Pakistan, South Korea, Sri Lanka, Egypt (for O. varipennis ssp. pharaonum).

8. Oxytelus migrator Fauvel, 1904 (Fig. 5C, D) Fauvel 1904: 100 (Type locality: Siam: Bangkok; Sumatra: Palembang; Deli; Java: Batavia; Malang). Oxytelus akazawensis Bernhauer 1907: 379 (Type locality: Akazawa, 2200 ft.); Hammond 1975: 152 (syn. of migrator).

Type material examined. [O. akazawensis] Holotype [monotypy]: ♂, Akazawa, Japan·Sauter// Hans Sauter 3939, Akazawa 2200', 6–8.VIII.05. - Kamen, zur Lampe// Akazawensis Bernh. Typus// Chicago NHMus, M.Bernhauer Collection// Oxytelus akazawensis Bernh., P. M. Hammond, det. 1974, HOLOTYPE ♂// Oxytelus migrator Fvl., P. M. Hammond, det. 1974 (FMNH). Other material examined. 45 specs. (19♂♂, 26♀♀). N. KOREA: Prov. South Pyongan [=Phyŏngannamdo], Pyonggyan [=Pyongyang]: 1♀, Hotel garden, No. 143, 1971.VIII.6, S. Horvatovich & J. Papp legg. (HNHM); 1♀, same data as previous except: No. 157, 1971.VIII.10 (HNHM); 1♀, same data as previous except: No. 167, 1971.VIII.12 (HNHM); 1♂, 1♀, same data as previous except: No. 185, 1971.VIII.17 (HNHM); 2♂♂, 2♀♀, 1ex., room of Hotel Te-dong, No. 298, 1975.VII.29, J. Papp & A. Vojnits legg. (HNHM); 1♂, 3♀♀, same data as previous except: No. 302, 1975.VII.30 (HNHM); 1♀, same data as previous except: No. 309, 1975.VIII.2 (HNHM); 2♂♂, 2♀♀, same data as previous except: No. 329, 1975.VIII.9 (HNHM). INDONESIA: 1♂, Sumatra, Fort de Kock [=Bukittinggi], 920m, 1924, Jacobson leg., ferrugineus Kr. (minor) det. Bernhauer (FMNH); 1♀, Java-Preanger, Tjigembong, 1915.VI, J. B. Corporaal leg., migrator Fauvel det. anonym (FMNH). SRI LANKA: 1♀, Peradeniya, 1979.XI.29, Sri Lanka, Ex coll. Viggo Mahler, Oxytelus migrator Fvl., det. A. Solodovnikov 2013 (ZMUC). VIETNAM: 1♂, Prov. Ha-Tinh, forestière Hüöng-sön, 150m, forêt trop. pluv., à la lumière [= in light], 1963.VIII.13, T. Pócs leg. (HNHM); 2♀♀, same data as previous except: 1963.VIII.14 (HNHM); 2♂♂, same data as previous except: 1963.VIII.15 (HNHM); 1♀, baie d’Ha-long, Hông-gai, 0–10m, l’hôtel, à la lumière, 1963.IX.5, T. Pócs leg. (HNHM); 1♂, Hanôi, 40m, l’hôtel, à la lumière, 1963.VIII.7–8, T. Pócs leg. (HNHM); 1♂, 2♀♀, same data as previous except: 1963.VIII.8–9 (HNHM); 1♀, same data as previous except: 1963.IX.12, (HNHM); 1♂, same data as previous except: 1963.X.2 (HNHM); 2♂♂, same data as previous except: 1963.X.5 (HNHM); 1♂, same data as previous except: 1963.X.25 (HNHM); 1♀, same data as previous except: 1963.X.29 (HNHM); 2♀♀, same data as previous except: 1963.X.30 (HNHM); 1♂, 1♀, Hanôi, 1963.IX.11–19, Manninger leg. (HNHM); 1♂, 2♀♀, same data as previous except: 1963.XI.4–10 (HNHM); 1♂, same data as previous except: 1963.XII.11–20 (HNHM). OXYTELUS IN SOUTHEAST ASIA


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Diagnosis. Oxytelus migrator can be separated from other species in piceus group by the combination of the small size body (≤ 3 mm) and the less-modified median plate of the male sternite VIII (Fig. 12 in Lü & Zhou 2012). Distribution. Indonesia, Malaysia, Thailand, Vietnam; China, Japan, North Korea, Sri Lanka, Europe.

9. Oxytelus incisus Motschulsky, 1857 (Fig. 5E, F) Motschulsky 1857: 504 (Type locality: Indes orientales). Oxytelus bledioides Blackburn 1885: 125 (Type locality: near Honolulu); Fauvel 1901: 71 (syn. of ferrugineus). Oxytelus cordovensis Bernhauer 1910: 358 (Type locality: Mexiko: Vera-Cruz, Cordoba); Hammond 1975 (syn. of incisus). Oxytelus ferrugineus Kraatz 1859: 173 (Type locality: Ceylan); Bernhauer 1943: 96 (syn. of incisus). Oxytelus laevior Sharp 1874: 92 (Type locality: Hiogo); Fauvel 1901: 71 (syn. of ferrugineus). Oxytelus laxipennis Fairmaire 1893: 527 (Type locality: Grande Comore); Fauvel 1901: 71 (syn. of ferrugineus).

Type material examined. [O. ferrugineus] Syntype: 1♂, Kraatz Ceylan 856// ferrugineus Kr. det. Bernhauer// ferrugineus Kr. Cotypus// Chicago NHMus, M.Bernhauer Collection// Oxytelus ferrugineus Kr., P. M. Hammond, det. 1975// PARALECTOTYPE ♂// Oxytelus incisus Mots., P. M. Hammond, det. 1975// = incisus Motsch., ferrugineus Kraatz, Wiegm. Arch. 1859. 173 (FMNH). Other material examined. 113 specs. (51♂♂, 62♀♀). MYANMAR: 1♂, 1♀, Moulmein, Berlese funnel from cow dung, 1954.I–III, Lois Jones leg. (FMNH). INDIA: Uttar Pradesh, Rishikesh, 6♂♂, 1♀, 1989.VII.2–4, A. Riedel leg. (SMNS); 1♂, 1989.VIII.6, A. Riedel leg. (SMNS); 1♂, Karnataka, Gersoppa (Jog Falls), ca.600m, 1977.XI.19–24, Zool. Mus. Copenhagen Exp. (ZMUC). INDONESIA: 1♂, 1♀, Java, Bandung, Bandung Institute of Technology, ex pony manure and compost near campus, 1988.III.30, D. L. Wrensch leg. (FMNH); 2♂♂, 2♀♀, Simalur [= Aceh, Simeulue], Teluk Dalam, 1986.IX.22, Lichtfg [light-trap]. Erber leg. (SMNS); 1♂, Bali, Sanur, 1994.I.24, J. Pedersen leg. (ZMUC). LAOS: 1♀, Vientiane prov., Vientiane, Bank of river Mekong, 160m, 17º57.597'N 102º36.518'E, 2008.VI.22, in decaying debris, A. Solodovnikov & J. Pedersen legg. (ZMUC). MALAYSIA: 17♂♂, 65♀♀, Borneo, Sabah, Mt. Kinabalu Nat. Pk., 480m, Poring Hot Spgs, 1988.VIII.19, A. Smetana leg. (FMNH); 1♂, 1♀, same data as previous except: 485m (FMNH); 1♀, Terengganu, Kampung Ayre Puteh, Berlese funnel from cattle dung, 1977.IV.27, L. E. Watrous leg. (FMNH). PHILIPPINES: Mindanao Is., Davao Prov.: 1♂, Maco, Tagum, near sea level, 1946.X, H. Hoogstraal leg. (FMNH); 1♀, Todaya, E slope Mt. Apo, 2800ft, 1946.XI, H. Hoogstraal leg. (FMNH). SRI LANKA: 1♀, Hanwella, 1979.XII.20, Sri Lanka, Ex coll. Viggo Mahler (ZMUC); 1♀, Peradeniya, 1979.XI.29, Sri Lanka, Ex coll. Viggo Mahler, Oxytelus incisus Motsch. det. A. Solodovnikov 2013 (ZMUC); 1♀, Ceylon C.Prov [=Central Prov.], Nuwara Eliya, 1983.VII.2–5, Sri Lanka, Ex coll. Viggo Mahler (ZMUC). VIETNAM: 1♀, Binh Thuy, Army Post, 1971.XI.25, H. J. Harlan leg. (FMNH); 1♀, Hanôi [=Hà Nội], 40m, l’hôtel, à la lumière, 1963.VIII.10, T. Pócs leg. (HNHM); 1♂, same data as previous except: 1963.X.4 (HNHM); 1♀, Hanôi, Kim-lien, sifted litter, Nr. 10, 1966.IV.9–11, Gy. Topál leg. (HNHM); 1♂, Hanôi, 1963.IX.11–19, Manninger leg. (HNHM); 2♀♀, same data as previous except: 1963.XI.4–10 (HNHM); 1♂, same data as previous except: 1963.XII.1–10 (HNHM); 1♂, same data as previous except: 1964.I.6–7 (HNHM); 1♀, Prov. Ha-Tinh, forestière Hüöng-sön, 150m, forêt trop. pluv., à la lumière [= in light], 1963.VIII.14, T. Pócs leg. (HNHM); 1♂, same data as previous except: 1963.VIII.15 (HNHM); 1♀, Prov. Nghe-An, forestière Quy-châu, 200m, à la lumière, forêt pluv. trop., semidecidue, 1963.VIII.23, T. Pócs leg. (HNHM); 1♀, same data as previous except: 1963.VIII.24 (HNHM); 2♀♀, same data as previous except: 1963.VIII.25 (HNHM); 1♂, same data as previous except: 1963.VIII.26 (HNHM); 1♂, same data as previous except: 1963.VIII.28 (HNHM); 1♂, Lao-cai, forêt trop. second., 300m, à la lumière, 1963.IX.21, T. Pócs leg. (HNHM); 2♂♂, 2♀♀, baie d’Ha-long, Hông-gai, 0–10m, l’hôtel, à la lumière, 1963.IX.5, T. Pócs leg. (HNHM); 1♂, Lam Dong Prov., Doc Me (Maria stream), 15 km S of Bao Loc, 1988.X.23, No. 372, S. Mahunka & T. Vásárhelyi legg. (HNHM); 2♂♂, 1♀, Lam Dong Prov., Bao Loc, No. 351, 1988.X.21, Mahunka & Vásárhelyi legg. (HNHM); 3♂♂, Lam Dong Prov., Suoi Loc chau (stream), 5km E of Bao LocNo. 377, 1988.X.24, Mahunka & Vásárhelyi legg. (HNHM); 1♂, 1♀, Trung Trang, I. Cat La., 200m, 20º48'N 107º0'E, at light, No. 171, 1987.V.16, Matskasii, Oláh & Topál legg. (HNHM); 1♂, Bac Thai Prov., Quang Chu, 150m, at light, 1993.XII.2–3, Bankovics & Csorba legg. (HNHM); 1♀, Da Lat., singled from horse dungon road, No. 700, 1994.XII.5, S. Mahunka, Gy. Szirâki & L. Zombori legg. (HNHM); 1♂, Nghe An Prov., Po Phuong village, 280m, at light, 1999.X.21–22, 19º38.442'N 104º58.302'E, Ference Kassai leg. (HNHM).


LÜ & ZHOU

Diagnosis. Oxytelus incisus can be readily distinguished from other Oxytelus species by the combination of small sized body (≤ 4 mm) and the incised tergite X in both male and female (Fig. 7H, I in Lü & Zhou 2012). Distribution. Indonesia, Malaysia (Borneo), Myanmar, Philippines, Vietnam; Cosmopolitan.

10. Oxytelus finitimus sp. nov. (Figs. 10 & 11) Type material examined. Holotype [Fig. 11A, C, M]: ♂, HOLOTYPE// LAOS: Champasak Pr., Bolaven Plateau, Ban Thôngvay (=Xékatam) vic., old logging road N of village 1095m, 15º14.288'N 106º31.891'E, 8–16.vi.2008// selectively logged forest; FMHD#2008-038, carrion trap (squid), A. Solodovnikov, M. Thayer & A. Newton; ANMT site 1231 ex 95% FIELD MUS. NAT. HIST.// Oxytelus finitimus Lü & Zhou, Holotype, ♂. Det. LÜ Liang, 2014 (FMNH). Paratypes: 2♂♂, 4♀♀, same data as holotype (FMNH); 5♂♂, 2♀♀, same data as holotype except: FMHD#2008-039, dung trap (human) (FMNH); 1♂, 2♀♀, same data as holotype except: 1035m, 15º13.96'N 106º31.731'E,// FMHD#2008-035, ANMT site 1230 (FMNH); 2♂♂, 4♀♀, same data as holotype except: 995m, 15º13.761'N 106º31.749'E,// FMHD#2008-032, M. Thayer, A. Solodovnikov & A. Newton; ANMT site 1229 (FMNH); 1♀, same data as previous except: FMHD#2008-031, flight intercept trap, A. Newton, M. Thayer, & J. Pedersen (FMNH); 2♀♀, same data as holotype except: 1170m, 15º14.494'N 106º31.807'E,// FMHD#2008-041, ANMT site 1232 (FMNH). Diagnosis. This new species differs from other Oxytelus species by the combination of the reddish coloration (Fig. 11A, B), the male clypeal shapes (Figs. 10A; 11C), the integrated occipital suture and the interrupted nuchal ridge (Figs. 10A, B; 11C, D), absence of the mid-longitudinal and paralateral sutures (Figs. 10A, B; 11C, D), and the outline of the posterior margin of male sternite VIII (Figs. 10F; 11F). Description. Body (Fig. 11A, B) yellow to ferruginous to brown and shining. Posterior part of elytra and basomedial part of tergites II – VIII dark. Length [average] ♂, 4.5 mm; ♀, 4.3 mm. Male. Head (Figs. 10A; 11C) widest at eyes. Disc densely punctate, nearly glabrous. Clypeus wider than long, moderately protruding beyond anterior margin of supra-antennal ridges, as long as 1/3 head length, slightly depressed centrally, surface glabrous, coriaceous, and punctate; anterior margin narrowed anteriorly and emarginate in middle. Epistomal suture with lateral portions incurved and running backward to level of anterior margin of eyes. Vertex densely punctate, posterior part well-delimited; mid-longitudinal suture quite obsolete and only obscurely visible in few cases, paralateral sutures invisible. Eyes with fine facets, longer than temples. Temples broadly rounded and lightly dilated. Occipital suture with middle portion present; nuchal ridge interrupted in middle, dorsal basal ridge present. Mandible (Fig. 10C) with two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 11A) as long as head and pronotum together or shorter; antennomere 4 subglobose, without basal dish; apical antennomere nearly as long as two preceding together. Pronotum (Fig. 11C) transverse, broadest at around anterior 1/3, broader than head. Disc 3-sulcate, median sulcus and two slightly curved paramedial sulci deep and punctate. Lateral margins even and curved in front and straight behind, posterior angles prominent. Elytra (Fig. 11C) punctate but not rugose, with lateral longitudinal ridge absent. Abdomen (Fig. 10A) coriaceous and pubescent, broadest at segment V. Sternite VII (Figs. 10E; 11E) with posterior margin straight, without modification. Sternite VIII (Figs. 10F; 11F) with transverse subbasal ridge continuous and straight in middle, with posterior margin bi-emarginate and with two closely neighbored short triangular teeth in middle. Aedeagus (Figs. 10K–N; 11I–L). Median lobe long, slightly broadened basally, with membranous internal sac inside (no sclerites); apico-medial hook with apical part lightly upcurved, with transverse ridge (like process in lateral view) a little behind apex, basal part erect but not produced; dorsal membranous area covering about 4/5 of dorsum of median lobe and not across whole length. Paramere strongly curved in lateral view, with seta at near apex, furnished with rounded process at middle. Female. Head (Figs. 10B; 11D) a little smaller than male; clypeus protruding beyond anterior margin of supraantennal ridges but not so much as male, with anterior margin slightly emarginate; temples shorter than eyes, not dilated; occipital suture continuous in middle. Mandible (Fig. 10D) a little shorter than male. Abdominal sternite VIII (Figs. 10G; 11G) with posterior margin gradually narrowed. Tergite X (Fig. 10I) much longer than wide, with



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anterior margin narrower and posterior margin truncate. Spermatheca (Figs. 10J; 11H) υ-shaped, bent at basal 2/5, basal portion obviously inflated into triangular shape, apical portion curved and with apex rounded. Distribution. Laos. Etymology. We use a Latin adjective finitimus (neighboring) as the specific epithet for the type locality of this species is China’s neighboring country, Laos.

11. Oxytelus bellicosus Fauvel, 1895 (Figs. 12 & 13) Fauvel 1895: 202 (Type locality: Bírmanie, Carin Chebà, 900–1100 m).

Type material examined. Lectotype [designated here, Fig. 12A–D, F–I, K]: ♂, Carin [=Kayan, Myanmar] Chebà, 900–1100 m. L. Fea. V XII-88// bellicosus Fvl.// Coll. et det. A. Fauvel, Oxytelus bellicosus Fauv., R.I.Sc.N.B. 17.479// Syntype// Oxytelus bellicosus Fvl. 1895, Lectotype ♂, det. LÜ Liang, 2015 (IRSNB). Paralectotypes: 4♂♂, Carin [=Kayan, Myanmar] Chebà, 900–1100 m. L. Fea. V XII-88// Coll. et det. A. Fauvel, Oxytelus bellicosus Fauv., R.I.Sc.N.B. 17.479// Syntype// Oxytelus bellicosus Fvl. 1895, Paralectotype ♂, det. LÜ Liang, 2015 (IRSNB); 1♂, PARALECTOTYPE// Co-type// 69474// Carin Chebà, 900–1100 m. L. Fea.// Birmah// Fry Coll. 1905.100.// O. bellicosus sp. n.// Oxytelus bellicosus Fvl. Paralectotype, P. Hammond det. 1970, ♂ (BMNH); 1♀, PARALECTOTYPE// Carin Chebà, 900–1100 m. L. Fea.// M. Cameron. Bequest. B.M.1955–147.// Oxytelus bellicosus Fvl. Paralectotype, P. Hammond det. 1970, ♀ (BMNH). Other material examined. 2 specs. (1♂, 1♀). 1♂, MYANMAR: Carin [=Kayan], Asciuii Cheba, 1200– 1300m, I.1888, L. Fea leg. (FMNH); 1♀, “bellicosus Fvl. Birmania v. Krātz” det. Bernhauer, Oxytelus bellicosus Fvl ♀ det. P. Hammond 1969 (FMNH). Diagnosis. This species differs from other Oxytelus species except O. finitimus sp. nov., O. castaneus sp. nov., O. cheesmanianus, and O. validus by the combination of the relatively light and vivid coloration (Fig. 12A), the non-modified posterior margin of sternite VII (Fig. 12C), and the shape of the posterior margin of male sternite VIII (Fig. 12D). It can be separated from these four species by the spined or bi-denticulate clypeus in males (Fig. 11A–C) and the aedeagal and spermathecal characters (Fig. 12F–J). Redescription. Body (Fig. 12A) yellow to ferruginous to brown and shining; antennomeres 5 – 11 dark. Posterior part of elytra infuscate, abdomen partly dark. Length [average] ♂, 5.8 mm; ♀, 4.9 mm. Male. Head (Figs. 12B; 13A–E) widest at temples (“major” male) or eyes (“minor” male). Disc densely punctate, nearly glabrous. Clypeus protruding into a pair of short spines or triangular teeth beyond anterior margin of supra-antennal ridges, as long as 1/3 head length or longer, slightly depressed centrally, surface glabrous, and punctate or slightly striate. Epistomal suture with lateral portions incurved and running backward to level of anterior margin of eyes. Vertex densely punctate, posterior part well-delimited; mid-longitudinal and paralateral sutures absent. Eyes with fine facets, shorter than temples. Temples broadly rounded and strongly (“major” male) to moderately (“minor” male) dilated. Occipital suture with middle portion present; nuchal ridge interrupted in middle. Mandible (Figs. 12B; 13A, D) with two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 12A) shorter than head and pronotum together; antennomere 4 subglobose, without basal dish; apical antennomere shorter than two preceding together. Pronotum (Figs. 12B; 13A–E) transverse, reversely trapezoidal, broadest at around anterior 1/5, narrower, broader than or as wide as head. Disc 3-sulcate, median sulcus and two slightly curved paramedial sulci punctate. Lateral margins with slight but obvious constriction near anterior angles, straight and finely crenulate behind, anterior and posterior angles prominent. Elytra (Figs. 12A; 13C–E) punctate and rugose, with lateral longitudinal ridge absent. Abdomen (Fig. 12A) pubescent on ventral side, broadest at segment IV. Sternite VII (Fig. 12C) with posterior margin straight, without modification. Sternite VIII (Fig. 12D) with transverse subbasal ridge continuous in middle, with posterior margin bi-emarginate and with two closely neighbored short triangular teeth in middle. Aedeagus (Fig. 12F–I) with median lobe long, slightly broadened basally, with membranous internal sac inside (no sclerites); apico-medial hook not upcurved at apex, with transverse ridge (like process in lateral view) a little behind apex, basal part erect and a little curved. Paramere curved and broadened and furnished with triangular process at base of apical arm, with seta at middle of apical arm, with shallow furrow in basal arm.


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Female. Head (Fig. 13F) a little smaller and narrower than pronotum; clypeus protruding beyond anterior margin of supra-antennal ridges but not so much as male, with anterior margin slightly emarginate; temples equal to or a little shorter than eyes, weakly dilated; occipital suture continuous and straight in middle. Mandible (Fig. 13F) shorter than male. Sternite VIII (Fig. 12E) with posterior margin broadly rounded. Spermatheca (Fig. 12J) comma-shaped, basal portion strongly bulbous, apical portion curved. Distribution. Myanmar. Remarks. The males of this species vary in head, especially as regards clypeus and temples. Some males look similar (“minor” males) to the conspecific female (cf. Fig. 13D–F). It should be noted here that O. bellicosus can be representative of a group which includes O. finitimus sp. nov. (Figs. 10 & 11), O. castaneus sp. nov. (Figs. 14 & 15), O. cheesmanianus Cameron (Fig. 49), and O. validus Cameron (Fig. 55), as they share these characters: straight posterior margin of the sternite VII, the posterior margin of sternite VIII modified by two closely allied teeth in middle, and similar profile of the aedeagal parameres. Oxytelus finitimus sp. nov., O. bellicosus, and O. castaneus sp. nov. could be further clustered as a subgroup and the other two species, O. cheesmanianus and O. validus as another subgroup. The two subgroups are different in profile of the aedeagal apico-medial hook: the latter subgroup’s apico-medial hook bears a tiny process at base forming a C-shaped “hook” in lateral view (cf. Figs. 10M; 11K; 12I; 14G; 15E; 49G; 55G).

12. Oxytelus castaneus sp. nov. (Figs. 14 & 15) Type material examined. Holotype [Fig. 15]: ♂, HOLOTYPE// Vietnam, Tam dao, 27.5–2.6.1986, Vinh phu prov., Jan Horák lgt.// NHMB// Oxytelus castaneus, HOLOTYPE ♂, det. LÜ Liang, 2014 (NHMB). Paratype: 1♂, Vietnam, Tam dao, 27.5–2.6.1986, Vinh phu prov., Jan Horák lgt.// Freiwilliger Museumsverein Basel 1987// Oxytelus castaneus, PARATYPE ♂, det. LÜ Liang, 2014 (NHMB). Diagnosis. This new species is close to O. finitimus sp. nov., O. castaneus sp. nov., O. cheesmanianus, and O. validus, but can be distinguished by the round teeth on the clypeus (Figs. 14A; 15B), the finely punctate head and pronotum (Fig. 15B), and the aedeagal characters (Figs. 14E–H; 15C–G). Description. Body (Fig. 15A) yellowish brown or castaneous and shining. Elytra infuscate, abdominal tergites dark around convergent region of subbasal and basolateral ridges. Length [average] ♂, 6.8 mm. Male. Head (Figs. 14A; 15B) widest at temples. Disc punctate, nearly glabrous. Clypeus protruding into a pair of round processes beyond anterior margin of supra-antennal ridges, as long as 1/3 head length, slightly depressed centrally, surface glabrous and sparsely punctate. Epistomal suture with lateral portions incurved and running backward to level of anterior margin of eyes. Vertex densely punctate; mid-longitudinal and paralateral sutures absent. Eyes with fine facets, shorter than temples. Temples broadly rounded and dilated. Occipital suture and nuchal ridge discontinuous in middle. Mandible (Figs. 14B; 15B) with two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 15A) nearly equal to head and pronotum together; antennomere 4 subglobose, without basal dish; apical antennomere shorter than two preceding together. Pronotum (Fig. 15B) transverse, reversely trapezoidal, broadest at around anterior 1/5, as wide as head. Disc with three punctate sulci, two paramedial sulci shallow and slightly curved. Lateral margins feebly crenulate, with slight but obvious constriction near anterior angles, straight behind constriction, anterior and posterior angles prominent. Elytra (Fig. 15B) punctate and rugose, with lateral longitudinal ridge absent. Abdomen (Fig. 15A) pubescent on ventral side, broadest at segment IV. Sternite VII (Fig. 14C) with posterior margin almost straight, with feeble modification near middle. Sternite VIII (Fig. 14D) with transverse subbasal ridge continuous in middle, with posterior margin bi-emarginate and with two closely neighbored short triangular teeth in middle. Aedeagus (Figs. 14E–H; 15C–G) with median lobe long, slightly broadened basally, with membranous internal sac inside (no sclerites); apico-medial hook not upcurved at apex, with transverse ridge (like process in lateral view) a little behind apex, basal part erect and a little curved. Paramere broadened and furnished with triangular process at base of apical arm, with seta at middle of apical arm, with shallow furrow in basal arm. Female. Unknown. Distribution. Vietnam. Etymology. The specific epithet is from the color, castaneus. This Latin adjective means castaneous or chestnut colored. OXYTELUS IN SOUTHEAST ASIA


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Remarks. The new species seems to be close to O. bellicosus and O. finitimus sp. nov. (see above). O. bellicosus has the most exaggerated clypeal modification, but this trait is variable even among type specimens. Nevertheless, O. castaneus sp. nov. differs from O. bellicosus in the following two traits (only males): 1) the round processes of clypeal anterior margin (Figs. 14A; 15B) beyong the intraspecific variation in that of O. bellicosus (Figs. 12B; 13A–E); 2) the pattern of the aedeagal internal sac in dorsal view (cf. Figs. 12G; 14F; 15D).

13. Oxytelus lividus Motschulsky, 1857 (Fig. 16) Motschulsky 1857: 503 (Type locality: Indes orientales). Oxytelus gigantulus Fauvel 1904: 99 (Type locality: Siam: Bangkok); Hammond 1975: 156 (syn. of lividus).

Type material examined. [O. gigantulus] Lectotype [designated here, Fig. 16A, B, E–I, M]: ♂, Bangkok// 192. Bangkok.// Coll. et det. A. Fauvel, Oxytelus gigantulus Fauv. R.I.Sc.N.B. 17.479// Syntype (IRSNB). Paralectotypes: 1♂, Java// gigantulus Fvl.// Syntype// Coll. et det. A. Fauvel, Oxytelus gigantulus Fauv. R.I.Sc.N.B. 17.479// Syntype (IRSNB); 1♀, Java// Syntype// Coll. et det. A. Fauvel, Oxytelus gigantulus Fauv. R.I.Sc.N.B. 17.479// Syntype (IRSNB); 1♀, Deli Sumatra// Syntype// Coll. et det. A. Fauvel, Oxytelus gigantulus Fauv. R.I.Sc.N.B. 17.479// Syntype (IRSNB); 1♀, Dili, May, 1892. W. Doherty.// Dili = Deli probablement// frontalis Epp. in litt.// Coll. et det. A. Fauvel, Oxytelus gigantulus Fauv. R.I.Sc.N.B. 17.479 (IRSNB). [Note: The last specimen (from “Dili”) was treated as syntype by Hammond (1975: 156), but this location did not appear in the original description (“Siam: Bangkok; Java; Sumatra: Deli”, see Fauvel 1904: 100). The name “Dili” in present use refers to the capital of Timor-Leste far from Sumatra, where there used to be two “Deli”: Sultanate of Deli and Deli Serdang Regency. No more material has been found supporting “Dili = Deli”, which seems to be speculated by Fauvel (“probablement”). We follow Hammond’s (1975) decision, however, as this female specimen does not conflict with the identities of other type specimens examined here.] Other material examined. 32 specs. (27♂♂, 5♀♀). CAMBODIA: 1♀, Mondolkiri Pr., Seima Biodiv. Cons. Area, road between Seima and O’Rang, 12º12'12''N 107º01'09''E, 300m, at light, 2006.I.30, G. Csorba, L. Duval & G. Ronkay legg. (HNHM); INDIA: 1♂, Goa, Molem [=Mollem], from cattle-dung singled, 1980.XI.17, Gy. Topál leg. (HNHM); INDONESIA: 5♂♂, SE-Sulawesi, Rawa Aopa Nat. Park, Aopa vill., 1994.II.8–10, M. Strba & I. Jenis leg. (NHMW); 1♂, same data as previous except: Kendari Airport, 30 km W of Kendari, 1994.II.11–14 (NHMW); 2♂♂, C-Sulawesi, 45 km SE Palu, 01º11'S 120º08'E, 1994, Haft leg. (NHMW); 1♂, NORDSUMATRA, Pematang-Siantar, 1986.II–III, Diehl leg. (SMNS); LAOS: 1♂, Phungsoli [=Phongsali], Gnoi-ou, 1700m, 2007.III.23/25, Jing-Ke Li leg. (FMNH); 1♂, Louangphrabang pr., Khan riv., 19º53'N 102º09'E, 300m, 1999.IV.21, Vít Kubáň leg. Oxytelus lividus Motschulsky det. Makranczy, 2000 (NHMB); 4♂♂, Louang Phrabang prov., Muang Ngoy, 20º43'N 102º41'E, 500m, 1999.IV.22, Vít Kubáň leg. Oxytelus lividus Motschulsky det. Makranczy, 2000 (NHMB); PHILIPPINES: 1♂, Negros Is., Mambucal, 1994.II.18, Seyfert leg. (NHMW); 1♂, N Palawan, Bahile, 50m, 1992.XII.22, Bolm leg. (NHMB); THAILAND: 1♀, Nan prov., Ban Huay Kon env., 2002.V.27–VI.10, P. Průdek & M. Obořil leg. (HNHM); VIETNAM: 1♀, Cao Bang, Ba Bể National Park, 1997.III.28–IV.7, A. Monastyrskii leg. (FMNH); 1♂, Prov. Nghe-An, forestière Quy-châu, 200m, à la lumière, forêt pluv. trop., semidecid., 1963.VIII.25, T. Pócs leg. (HNHM); 8♂♂, N Vietnam 100 km S from Banoi, CUCPHUONG nat. park, 1991.V.2–12, E. Jendek leg. Oxytelus lividus Motschulsky det. Makranczy, 2000 (NHMB); 1♀, Cuc Phuong NP, 20º15.586'N 105º42.320'E, 147m, 2005.IV.30–V.1, A. Kun leg. (HNHM); 1♀, Tuyen Quang Prov., 3km SE of Pac Ban Village, Na Hang Nature Reserve, 22.20ºN 105.25ºE, 380m, at light, 1997.II.22–26, G. Csorba leg. (HNHM). Diagnosis. Oxytelus lividus can be readily recognized by the combination of the large size (≥ 5 mm), the khaki colored and finely punctate body (Fig. 16A, C), the serrate anterior margin of male clypeus (Fig. 16B), the dark medial stripes on the abdominal tergites (Fig. 16C), and the deeply and sharply emarginate male sternite VII (Fig. 16F). It differs from the subsequent two new species in the sexually dimorphic and aedeagal characters (see the relevant species accounts below). Distribution. Cambodia, Indonesia, Laos, Malaysia, Myanmar, Philippines, Thailand, Vietnam; China (southern part), Sri Lanka, India.


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14. Oxytelus insulanus sp. nov. (Figs. 17A–E; 18A, B; 19A–F) Type material examined. Holotype [Fig. 17A, B, E]: ♂, HOLOTYPE// BORNEO: SABAH, S Keningau, 350m, 20.–22.III.2007, leg. W. SCHAWALLER// Oxytelus insulanus Holotype ♂ det. LÜ Liang, 2014 (SMNS). Paratypes: 1♂, same data as holotype except identification label (SMNS); 1♀, PARATYEP// BORNEO: SABAH, Tambunan, 530m, 2007.III.14–15, W. Schawaller leg. (SMNS). 1♂, PARATYPE// C Sulawesi, 17 km E Pendolo, 800m, 120.45.49E, 2.06.33S 4–9 Jul 1999, Bolm lgt.// Oxytelus insulanus Paratype ♂ det. LÜ Liang, 2014 (SMNS); 6♀♀, same data as previous except identification label (SMNS); 1♂, PARATYPE// C Sulawesi, 50m, 1999, 6 km E Tambarana, 120.28.06E, 1.11.15S, Bolm lgt. 9.–11.July// Oxytelus insulanus Paratype ♂ det. LÜ Liang, 2014 (SMNS). Diagnosis. This new species similar to O. lividus and the following new species O. sublividus but can be distinguished by the depth, sharpness, and curvature of the emargination on the posterior margin of male sternite VII (Fig. 18A). The males also differ in aedeagal characters (cf. Figs. 16H–L; 19) but the females are difficult to separate. Description. Body (Fig. 17A, C) yellowish brown or khaki colored, flattened, cuticle rough. Lateral parts of pronotum, posterolateral margins of elytra yellow, extreme margins of pronotum black. Mandibles reddish brown. Abdomen along middle with longitudinal dark stripe. Length [average] ♂, 7.5 mm; ♀, 7.1 mm. Male. Head (Figs. 17B) subtrapezoidal and roughly coriaceous, widest at temples or eyes. Disc rugose on basal half, not pubescent. Clypeus reversed trapezoidal and forward barely beyond anterior margin of supra-antennal ridges, as long as near 1/4 head length, slightly depressed in basal part, surface glabrous, impunctate and coriaceous; anterior margin shallowly tri-emarginate, with four denticles. Epistomal suture with lateral portions straight but slightly incurved at posterior end, running posteromedially to level of anterior margin of eyes. Vertex slightly convex but flat anteriorly, posterior part not well-delimited and continuous with neck; mid-longitudinal suture obvious. Eyes with coarse facets, as long as temples, convex. Temples dilated posterolaterally. Occipital suture with middle portion absent; nuchal ridge present but interrupted in middle and extending to lateral plate, dorsal basal ridge present. Mandible (Fig. 17B) stout, falciform, strongly incurved; two denticles on inner edge, one near middle and one near apex. Antenna (Fig. 17A) nearly as long as head and pronotum together; antennomere 4 subglobose, without basal dish; with apical antennomere elongated as long as two preceding together. Pronotum (Fig. 17B) transverse, broadest at near anterior 1/4, as wide as head or a little narrower. Disc rugose and 3-sulcate, two paramedial sulci deep and curved; two paralateral depressions shallow and striate; lateral margin finely crenulate across length. Elytra (Fig. 17B) rough and rugose, without lateral longitudinal ridge. Abdomen (Fig. 17A) coriaceous and pubescent on ventral side, broadest at segment V. Sternite VII (Fig. 18A) with no sclerotized area anterior to basal ridge; posterior margin with deep and acute emargination in middle, on each side with process. Sternite VIII (Fig. 18B) strongly sclerotized in area anterior to basal ridge, with subbasal ridge feeble or discontinuous near middle; posterior margin deeply bi-emarginate forming three lobes; median lobe dark and with mid-longitudinal internal ridge, narrowed posteriorly and attached by finger-formed projection at each side of base and furnished with a pair of setae near apex, apex weakly emarginate and behind with transverse carina; lateral lobes triangular covered with long setae. Aedeagus (Figs. 19A–E) with median lobe oblong-ovoid, weakly constricted at apical 1/4; with membranous internal sac and one semicirluar and two V-shaped sclerites in apical part of median lobe; apico-medial hook bent at near middle, with apex alatiform, on downside with two transverse carinae (like two denticles in lateral view); dorsal membranous area not covering all dorsum and not across length of median lobe. Parameres arm-like, with hand-like apex enlarged and trifurcate (with three finger-formed projections), on ventral projection situated with long seta, with small process on lateral surface. Female. Head (Fig. 17D) subtriangular, smaller than male; clypeus subquadrate, as long as 1/3 head length, flat and not depressed in basal part; anterior margin broadly rounded but truncate in middle; epistomal suture with lateral portions incurved; vertex flat, even with clypeus, posterior part well-delimited and separated from neck by occipital suture; eyes over 4 times length of temples, convex and protruding laterally beyond temples, covering most of lateral surface of head; occipital suture with median portion present and obvious. Mandible (Fig. 17D) slender, incurved but not so strongly as in male. Abdominal sternite VII with posterior margin straight. Sternite



19

VIII with posterior margin broadly rounded, but protruding in middle. Spermatheca (Fig. 19F) comma-shaped, basal portion strongly bulbous, constricted at middle, and sharply slimmed near apex. Distribution. Indonesia (Sulawesi), Malaysia (Borneo). Etymology. The specific epithet is derived from a Latin word insulanus, which has two meanings: “pertaining to an island” (adjective) or “a person living on an island” (noun). Here we use the first meaning, as this species has hitherto been found only on islands (Borneo or Sulawesi). Remarks. This species differs from O. lividus in the shape of the posterior margin of sternites VII – VIII (Figs. 16E–G; 18A, B) and the shape of the apex of parameres (Figs. 16K–L; 19D, E).

15. Oxytelus sublividus sp. nov. (Figs. 17F–J; 18C, D; 19G–K) Type material examined. Holotype [Fig. 17F, G, J]: ♂, HOLOTYPE// LAOS-N, 21.iv.1999, Louangphrabang pr., 19º53'N 102º09'E, KHAN riv., 300m, Vít Kubáň leg.// NHMB// Oxytelus sublividus sp.n. det. Makranczy, 2001// Oxytelus sublividus, ♂, sp.n. Holotype, det. Liang Lü, 2014// Oxytelus sublividus HOLOTYPE ♂ det. LÜ Liang, 2014 (NHMB). Paratypes: 1♂, VIET NAM, Quang Chu Prov. Bac Thai, 500 m, 22º00'N 105º50'E// singling, No.236, 25.V.1987, Matskási-Oláh-Topál// Oxytelus lividus Motschulsky, det. Makranczy, 2000// Oxytelus sublividus PARATYPE ♂ det. LÜ Liang, 2014 (HNHM); 1♂, VIET NAM, Lam Dong Prov. Suoi Loc chau (stream), 5 km E of Bao Loc// No. 377, 24.X.1988, leg. Mahunka S. & Vásárhelyi T.// Oxytelus sublividus sp.n. det. Makranczy, 2001// Oxytelus sublividus PARATYPE ♂ det. LÜ Liang, 2014 (HNHM); 2♀♀, VIET NAM, Lam Dong Prov. Suoi Loc Chau (stream), 5 km E of Bao Loc// 24.X.1988, No. 377, leg. S. Mahunka & T.Vásárhelyi// Oxytelus sublividus PARATYPE ♀ det. LÜ Liang, 2014 (HNHM); 1♂, S-VIETNAM, Nam Cat Tien Nat. Park, 1.15.5.1994, Pacholatko & Dembicky// NHMW// Oxytelus sublividus PARATYPE ♂ det. LÜ Liang, 2014 (NHMW); 2♂♂, S-VIETNAM, 40km NW An Khe Buon Luoi, 620-750m// 14º10'N 108º30'E, 28.3.-12.4.1995, Pacholatko & Dembicky// NHMW// Oxytelus sublividus PARATYPE ♂ det. LÜ Liang, 2014 (NHMW); 1♂, LAOS: 18º16'N 103º10'E, 70km NE Vientiane, 150m, Ban Phabat, 27.IV.-1.V.1997, leg. Strba & Hergovits// NHMW// Oxytelus sublividus PARATYPE ♂ det. LÜ Liang, 2014 (NHMW); 1♂, LAOS-N, 22.iv.1999, Louang Phrabang prov., 20º43'N 102º41'E, MUANG NGOY, 500m, Vít Kubáň leg.// Oxytelus lividus Motschulsky det. Makranczy, 2000// Oxytelus sublividus PARATYPE ♂ det. LÜ Liang, 2014 (NHMB). Diagnosis. This new species is similar to the preceding two species O. lividus and O. sublividus sp. nov. but can be distinguished by the small notch and teeth in the middle of the posterior margin of male sternite VII (Fig. 18C), the shorter median plate of male sterntie VIII (Fig. 18D), and aedeagal characters (Fig. 19G–K). Description. Body (Fig. 17F, H) yellowish brown or khaki colored, flattened, cuticle rough. Lateral parts of pronotum, posterolateral margins of elytra yellow, extreme margins of pronotum black. Mandibles reddish brown. Abdomen along middle with longitudinal dark stripe. Length [average] ♂, 6.7 mm; ♀, 6.2 mm. Male. This species is so similar to O. lividus and O. insulanus sp. nov. that it prevents any attempt to distinguish them on the basis of external/dorsal traits. Oxytelus sublividus sp. nov. differs from the other two species in the following characters: 1) processes on posterior margin of sternite VII smaller than both O. lividus and O. insulanus sp. nov., and emargination between processes shallower than O. lividus and O. insulanus sp. nov. (Figs. 16F; 18A, C); 2) difference in median lobe of sternite VIII (Figs. 16G; 18B, D); 3) aedeagal parameres, especially shape of apex and size of process on lateral surface of anterior arm (Figs. 16H, I, K, L; 19A, B, G, H, D, E, J, K). Female. Distinct from male (see the sexual differences described in O. insulanus sp. nov.). Indeed the females of O. lividus, O. insulanus sp. nov., and O. sublividus sp. nov. can hardly be separated, the spermatheca is slightly different but seems not definite and is based on examination of a small sample. Here we judge the identity of femals by the location they were collected (in company with conspecific males). Distribution. Laos, Vietnam. Etymology. The specific epithet is derived from the name of O. lividus, as it resembles the latter species.


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16. Oxytelus punctipennis Fauvel, 1905 (Fig. 20) Fauvel 1905: 113 (Type locality: Sikkim: Darjeeling). Oxytelus discalis Cameron 1930: 222 (subgenus Tanycraerus; Type locality: Simla Hills: Kotgarh; Fagu. Mussoorie district: Kolhu Khet Gad, alt. 6000 to 8000 feet); Lü & Zhou 2012: 48 (syn. of punctipennis).

Material examined. [O. punctipennis] Holotype [monotypy, Fig. 20A–D]: ♂, Darjeeling, Sikkim// punctipennis Fvl.// TYPE// Coll. et det. A. Fauvel, Oxytelus punctipennis Fauv. R I.Sc.N.B. 17.479 (IRSNB). [O. discalis] Lectotype [designated here, Fig. 20E–G]: 1♂, Kotgarh 7000', Simla Hills.// Dr. Cameron. IX. 1921.// O. discalis Cam. TYPE// M. Cameron. Bequest. B.M. 1955–147.// Oxytelus discalis Cam. P.M.Hammond det.1970, LECTOTYPE ♂ (BMNH). Paralectotypes: 1♂, Kotgarh 7000', Simla Hills.// Dr. Cameron. 11.IX. 1921/ / Cotypus don. Cameron// discalis Cam CO-TYPE// Chicago NHMus, M.Bernhauer Collection// Oxytelus discalis Cam, Paralectotype, P.Hammond det. 1970, ♂ (FMNH); 1♂, Kotgarh 7000', Simla Hills.// Dr. Cameron. 19.IX. 1921// Chicago NHMus, M.Bernhauer Collection// Oxytelus discalis Cam, Paralectotype, P.Hammond det. 1970, ♀ (FMNH). Other material examined: 3 specs. (2♂♂, 1♀). INDIA: 1♂, West Bengal, Ghum district, 1931.V–VII, M. Cameron leg. (NHM); 1♀, Uttar Pradesh, Mussorie [=Mussoorie in Uttarakhand since 2000], Dhanolt [=Dhanaulti], 2250m, 1989.VII.11, A. Riedel leg. (SMNS); 1♂, Darjeeling Indien, 1977.VII.26, O. Mehl leg. (ZMUC). Diagnosis. This species can be distinguished by the combination of the bi-denticulate clypeus in both males and females (Fig. 20B, F) and the particular shape of the posterior margin of male sternite VII: triangularly protruding with two sharp teeth at base (Fig. 20C, G). Distribution. Myanmar; Bangladesh, China (central part and Xizang), India.

17. Oxytelus ferreirai Scheerpeltz, 1978 (Fig. 21) Scheerpeltz 1978: 187 (subgenus Tanycraerus; Type locality: Timor: Maubisse).

Type material examined. Holotype [by original designation, Fig. 21A, C, E–K]: ♂, LAQUECO (Maubisse)// Voando ao cair da tarde iunte a uma mata de cafe', 1965.III.3, R.N.F// ex coll. Scheerpeltz// TYPUS, Oxytelus Ferreirai. Scheerplatz// Oxytelus lucens Bernh., P.M.Hammond, det. 1987// Oxytelus ferreirai Scheer. Holotype ♂, det. LÜ Liang, 2014// Oxytelus ferreirai valid species, det. LÜ Liang, 2014 (NHMW). Paratypes: 1♀, LAQUECO (Maubisse)// Voando ao cair da tarde iunte a uma mata de cafe', 1965.III.3, R.N.F// ex coll. Scheerpeltz// TYPUS, Oxytelus Ferreirai. Scheerplatz// Oxytelus ferreirai Scheer. Paratype ♀, det. LÜ Liang, 2014 (NHMW); 1♂, LAQUECO (Maubisse)// Voando ao cair da tarde iunte a uma mata de cafe', 1965.III.3, R.N.F// ex coll. Scheerpeltz// COTYPUS, Oxytelus Ferreirai. Scheerplatz// Oxytelus ferreirai Scheer. Paratype ♂, det. LÜ Liang, 2014 (NHMW); 1♀, LAQUECO (Maubisse)// Voando ao cair da tarde iunte a uma mata de cafe', 1965.III.3, R.N.F// ex coll. Scheerpeltz// COTYPUS, Oxytelus Ferreirai. Scheerplatz// Oxytelus ferreirai Scheer. Paratype ♀, det. LÜ Liang, 2014 (NHMW). Diagnosis. This species can be distinguished from other Oxytelus species except O. lucens, O. lompobatangensis sp. nov., and O. poecilopterus sp. nov. by the combination of the fine-faceted eyes, the crenulate lateral margin of pronotum (Fig. 21C, D), the reddish-infuscated elytra (Fig. 21C, D), and the shape of the posterior margin of male sternites VII and VIII (Fig. 21E, F). It can be separated from O. lucens by the non-incrassate clypeal anterior margin in males (cf. Figs. 21C; 22C, G), the distinct teeth on posterior margin of male sternite VII (cf. Figs. 21E; 23C), and aedeagal characters (Fig. 21G–J), from O. lompobatangensis sp. nov. and O. poecilopterus sp. nov. by the elytral coloration (Fig. 21C, D), the short protrusion of sternite VIII (cf. Figs. 21F; 25B, F), and the aedeagal characters (Fig. 21G–J). Redescription [holotype]. Body (Fig. 21A) blackish, pitchy, or dark testaceous; elytra dark but reddish at apex. Mandibles, maxillary palpi, first 4 antennomeres, and legs testaceous. Length: 5.9 mm.



21

Head (Fig. 21C) with widest at temples. Disc densely punctate, nearly glabrous. Clypeus protruding largely beyond anterior margin of supra-antennal ridges, as long as 1/3 head length, depressed in basal part, surface glabrous and coriaceous in depressed region, scattered with large punctures (same as on vertex) in anterior part; anterior margin emarginate and slightly thickened. Epistomal suture with lateral portions broadly incurved and running backward and medially to level of anterior margin of eyes. Vertex densely punctate, slightly convex, and even somewhat depressed near clypeus, posterior part poorly delimited; mid-longitudinal suture absent, two punctate paralateral sutures running anteriorly to level of posterior margin of eyes. Eyes with fine facets, shorter than temples. Temples dilated and broadly rounded. Occipital suture with middle portion absent; nuchal ridge interrupted in middle. Mandible (Fig. 21C) robust and strongly curved; two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 21A) shorter than head and pronotum together; antennomere 4 subglobose, without basal dish; with apical antennomere shorter than two preceding together. Pronotum (Fig. 21C) transverse, broadest at about anterior 1/4, as broad as head. Disc 5-sulcate, two paramedial sulci curved, deep, and punctate; two paralateral sulci deep, straight, and shorter, each with punctate depression on lateral side. Lateral margins evenly broad and crenulate from anterolateral to posterolateral angles. Elytra (Fig. 21C) punctate and striate but slightly rugose, without lateral longitudinal ridge. Abdomen (Fig. 21A) coriaceous and pubescent sparsely on dorsal side but densely on ventral side, broadest at segment V. Sternite VII (Fig. 21E) with two denticles near middle of posterior margin, between which shallowly emarginate. Sternite VIII (Fig. 21F) with subbasal ridge continuous and straight in middle, with posterior margin protruding and gradually narrowed in middle, apex slightly emarginate and with short and fine transverse carina a little behind. Aedeagus (Fig. 21G–J). Median lobe oblong-ovoid, with membranous internal sac formed as hanging strip; apico-medial hook with apical part short, apex beveled, a little behind with transverse ridge (like process in lateral view). Paramere arm-like, obviously broadened near apex, where furnished with seta; basal arm with shallow furrow. Distribution. Timor-Leste. Remarks. Similar to O. lucens, but can be definitely distinguished by the shape of the posterior margin of sternite VII and the shape of parameres (more broadened near apex compared with Fig. 23B). The internal sac is also different, there are a pair of hook-like sclerites in O. lucens (cf. Lü & Zhou (2012: Fig. 9L, M)) but present as a single pliable ribbon in O. ferreirai (Fig. 21G–I).

18. Oxytelus lucens Bernhauer, 1903 (Figs. 22 & 23) Bernhauer 1903: 34 (Type locality: Sumatra). Oxytelus (Tanycraerus) malaisei Scheerpeltz 1965: 155 (Type locality: Tenasserim, Sukl, 75 km E. of Moulmein, 600 m). Syn. nov.

Type material examined. [O. lucens] Lectotype [designated here, Figs. 22A, C, E; 23A, B]: ♂, Dohrn, Sumatra, Sinabong// lucens Bernh. ded. Dohrn// lucens Bernh. Type// Chicago NHMus, M.Bernhauer Collection// Oxytelus lucens Bernh., P. M. Hammond, det. 1970, LECTOTYPE ♂ (FMNH). Paralectotype: 1♀, Dohrn, Sumatra, Sinabong// Chicago NHMus, M.Bernhauer Collection// Oxytelus lucens Bernh., Paralectotype, P. M. Hammond, det. 1970 ♂ (FMNH). [O. malaisei] Holotype [by original designation and monotypy, Figs. 22F–H; 23C–G]: ♂, TENASSERIM Sukli, 75 km. E. of Moulmein. 600 m, 27-31.X.34 Malaise// HOLOTYPUS// TYPUS Oxytelus (Tanycraerus) Malaisei O. Scheerpeltz// 8134 E91 +// Oxytelus (Tanycraerus) Malaisei n. sp. det. Scheerpeltz, 1940// Oxytelus lucens Bernh. = O. malaisei Schrpltz. det. LÜ Liang, 2013// NHRS-JONI 000000209 (SMNH). Other material examined. 22 specs. (11♂♂, 11♀♀). INDONESIA: 3♂♂, 1♀, Java orient., Mt. Tengger, 4000ft, 1890, H. Fruhstorfer leg., lucens Brh. det. Bernhauer (FMNH); 2♀♀, East Java, Nongkodjadjar, 1934, I. P. A. Kalis leg., lucens Brh. det. Bernhauer (FMNH). LAOS: 6♂♂, 5♀♀, Champasak prov., Bolaven Plateau, Muang Paxong, Ban Thôngvay (=Xékatam) vic., 15º14.054'N 106º31.867'E, 1200m, Edge of distributed primary rainforest, near clearing, flight intercept trap, 2008.VI.8–16, A. Solodovnikov & J. Pedersen legg. (ZMUC). PHILIPPINES: 1♂, Mindanao, Davao, East slope Mt. McKinley, 3000ft, in rotten stem of succulent plant beside


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strm. orig. for., H. Hoogstraal leg. (FMNH); 1♀, same data as previous except: 3300ft, flight at dusk, F. G. Werner leg. (FMNH); 1♀, Luzon, Immugan, COTYPUS [note: Luzon was not included in the original type locality], (FMNH-A. Bierig colln.). VIETNAM: 1♂, 1♀, Cuc Phuong, Ninh binh, 1966.V.5–18, Nr. 385, from trap in soil, Gy. Topál leg. (HNHM). Diagnosis. Oxytelus lucens can be distinguished from other Oxytelus species except O. ferreirai, O. lompobatangensis sp. nov., and O. poecilopterus sp. nov. by the combination of the fine-faceted eyes, the crenulate lateral margin of pronotum (Fig. 22C, G), and the shape of the posterior margin of male sternites VII and VIII (Fig. 23C, D). It can be separated from the three sibling species by the incrassate clypeal anterior margin in males (Fig. 22C), the obsolete teeth on posterior margin of male sternite VII (Fig. 23C), and aedeagal characters (Fig. 23B, E– G). Distribution. Indonesia, Laos, Malaysia, Myanmar, Philippines, Vietnam; Bangladesh, China (southern part), India. Remarks. We propose a synonymy of O. lucens with O. malaisei according to comparison of the shape of the posterior margin of sternite VIII (with obsolete denticles near middle), the shape of the apex of parameres (Fig. 23B, G), and the internal sac configuration (compare Fig. 23E–G and Lü & Zhou (2012: Fig. 9 K–M)).

19. Oxytelus lompobatangensis sp. nov. (Figs. 24A–E; 25A–D; 26A–D) Type material examined. Holotype [Fig. 24A, B, E]: ♂, HOLOTYPE// S Sulawesi, 7 kmS Mamasa, Gn. Lompobatang, 950m, 119.20.32.E 2.56.13.S, 29.Jul 1999, Bolm lgt.// Oxytelus lompobatangensis, HOLOTYPE ♂, det. LÜ Liang, 2015 (SMNS). Paratypes: 1♂, 2♀♀, PARATYPE// S SULAWESI, Malino, Gn. Lompobatang, 1800m, 119.53.31.E 5.17.50.S, 13.-14.Jul 2001, Bolm lgt. (SMNS). Diagnosis. This new species has no one feature that immediately separates it from all of its congeners. However, it is obviously close to O. lucens and O. ferreirai, but can be distinguished from the two species by the combination of the lateral margin of pronotum increnulate in front half (Fig. 24B, D), the longer posterior protrusion on the sternite VIII (Fig. 25B), the shape of parameral apex, and the internal sac configuration (Fig. 26A–C). Description. Body (Fig. 24A, C) blackish; pronotum lighter and brownish, elytra with broad reddish stripe slightly oblique from anterior angle to posterior margin. Maxillary palpi, first 4 antennomeres, and legs testaceous. Length [average] ♂, 5.9 mm; ♀, 5.6 mm. Male. Head (Fig. 24B) with widest at temples. Disc sparsely punctate, nearly glabrous. Clypeus broad, and protruding beyond anterior margin of supra-antennal ridges, a little shorter than 1/3 head length, slightly depressed in basal part, surface glabrous and coriaceous in depressed region, with scattered fine punctures (same as those on vertex) in anterior part; anterior margin slightly emarginate in middle. Epistomal suture with lateral portions broadly incurved and running backward and medially to level of anterior margin of eyes. Vertex sparsely punctate, slightly convex, and depressed near clypeus (about same level with latter), posterior part poorly delimited; midlongitudinal suture fine and short, two punctate paralateral sutures running anteriorly to level of posterior margin of eyes. Eyes with fine facets, shorter than temples. Temples dilated. Occipital suture with middle portion absent; nuchal ridge interrupted in middle, dorsal basal ridge present. Neck scattered with punctures larger than those on vertex. Mandible (Fig. 24B) robust and strongly curved; two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 24A) shorter than head and pronotum together; antennomere 4 subglobose, without basal dish; with apical antennomere shorter than two preceding together. Pronotum (Fig. 24B) transverse, broadest at about anterior 1/4, as broad as (but in some cases wider or narrower than) head. Disc 5-sulcate, two paramedial sulci curved, deep, and punctate; two paralateral sulci deep, straight, and shorter, each with punctate depression on lateral side. Lateral margins crenulate in posterior half. Elytra (Fig. 24B) punctate, striate, weakly rugose, without lateral longitudinal ridge. Abdomen (Fig. 24A) coriaceous and pubescent on ventral side, broadest at segment IV or V. Sternite VII (Fig. 25A) with two sharp denticles near middle of posterior margin, between which roundly emarginate. Sternite VIII (Fig. 25B) with subbasal ridge continuous and straight in middle, with two obsolete tubercles in center, with posterior margin protruding into reverse trapezoidal lobe in middle, apex truncate and with short and fine transverse carina a little behind, emarginate at each side of lobe. OXYTELUS IN SOUTHEAST ASIA


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Aedeagus (Figs. 26A–D) with median lobe oblong-ovoid, with membranous internal sac and hook-like sclerites inside; apico-medial hook with apical part short, apex blunt, with transverse ridge (like process in lateral view). Paramere arm-like, with seta at near apex, slightly constricted near apex, processed at near corner between apical and basal arm. Female. Head (Fig. 24D) smaller than male, narrower than pronotum; clypeus flat and less protruding, with anterior margin truncate; temples almost as long as eyes, not dilated, with mid-longitudinal suture absent, with occipital suture present in middle. Mandible (Fig. 24D) smaller than male. Abdominal sternite VII without denticles on posterior margin. Sternite VIII (Fig. 25C) with posterior margin broadly rounded and weakly protruding in middle. Spermatheca (Fig. 25D) υ-shaped, basal portion not bulbous, apex rounded and slightly curved. Distribution. Indonesia. Etymology. The specific epithet is derived from the type locality “Lompobatang”.

20. Oxytelus poecilopterus sp. nov. Figs. 24F–J; 25E–H; 26D–H) Type material examined. Holotype [Fig. 24F, G, J]: ♂, HOLOTYPE// S Sulawesi, 8 km W Mamasa, 950m, 119.20.32.E 2.56.13.S, 18.-21.Jul 1999, Bolm lgt. // Oxytelus poecilopterus ♂, HOLOTYPE, det. LÜ Liang, 2015 (SMNS). Paratypes: 6♂♂, 4♀♀, PARATYPE// S Sulawesi, 8 km W Mamasa, 950m, 119.20.32.E 2.56.13.S, 18.21.Jul 1999, Bolm lgt. (SMNS). Diagnosis. Same as O. lompobatangensis sp. nov., the new species O. poecilopterus is also close to O. lucens and O. ferreirai, and similarly can be distinguished from the two species by the combination of the lateral margin of pronotum increnulate in front half (Fig. 24G, I), the longer posterior protrusion on the sternite VIII (Fig. 25F), the shape of parameral apex, and the internal sac configuration (Fig. 26E–G). It can be readily separated from O. lompobatangensis sp. nov. by the pale or yellowish posterolateral angle of elytra (cf. Fig. 24B, D, G, I) and the profiles of the aedeagal apico-medial hook and parameres (cf. Fig. 26C, D, G, H). Description. Body (Fig. 24F, H) black or brownish black. Mouthparts, antennae, legs lighter; Posterolateral angle of elytra yellow or pale. Length [average] ♂, 4.8 mm; ♀, 4.3 mm. Male. Head (Fig. 24G) sub-triangular. Disc finely punctate, nearly glabrous. Clypeus moderately protruding beyond anterior margin of supra-antennal ridges, as long as 1/4 head length, slightly depressed centrally, surface glabrous, slightly coriaceous, and sparsely punctate; anterior margin narrowed anteriorly and truncate or slightly emarginate in middle. Epistomal suture with lateral portions incurved and running backward to level of anterior margin of eyes. Vertex sparsely punctate, slightly convex, and depressed near clypeus (about same level with latter), posterior part poorly delimited; mid-longitudinal suture invisible, two punctate paralateral sutures running anteriorly to level of posterior margin of eyes. Eyes with fine facets. Temples dilated, longer than twice eye length (“major” male). Occipital suture with middle portion absent; nuchal ridge interrupted in middle. Neck scattered with punctures similar to those on vertex. Mandible (Fig. 24G) robust and strongly curved; two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 24F) shorter than head and pronotum together; antennomere 4 subglobose, without basal dish; with apical antennomere a little longer than penultimate one. Pronotum (Fig. 24G) transverse, broadest at about anterior 1/4, narrower than (typical) or as broad as head. Disc 5-sulcate, two paramedial sulci curved, deep, and sparsely punctate; two paralateral sulci deep, straight, and shorter, each with punctate depression on lateral side. Lateral margins crenulate in posterior half. Elytra (Fig. 24G) punctate, striate, weakly rugose, without lateral longitudinal ridge. Abdomen (Fig. 24F) pubescent on ventral side, broadest at segment IV or V. Sternite VII (Fig. 25E) with two sharp denticles near middle of posterior margin, between which roundly emarginate. Sternite VIII (Fig. 25F) with subbasal ridge continuous and straight in middle, with two obsolete tubercles in center, with posterior margin protruding into reverse trapezoidal lobe in middle, apex truncate and with fine transverse carina a little behind, emarginate at each side of lobe. Aedeagus (Fig. 26E–H) with median lobe oblong-ovoid, abruptly constricted and narrowing anteriorly, with membranous internal sac and sword-shaped sclerites inside; apico-medial hook with apical part short, apex beveled, with transverse ridge (like process in lateral view). Paramere arm-like, broadened near apex, with seta at apical 1/3 of apical arm.


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Female. Head (Fig. 24I) smaller than male, narrower than pronotum; clypeus flat and less protruding, with anterior margin truncate; temples almost as long as eyes, not dilated, with mid-longitudinal suture absent, paralateral sutures feeble, with occipital suture continuous in middle. Mandible (Fig. 24I) smaller than male. Abdominal sternite VII without denticles on posterior margin. Sternite VIII (Fig. 25G) with posterior margin broadly rounded and a little protruding in middle. Spermatheca (Fig. 24H) υ-shaped, basal portion not bulbous, apex rounded and slightly curved. Distribution. Indonesia. Etymology. The specific epithet is composed of two Greek words: “ποικιλ-” (“poecil-” = variegated) and “πτερόν” (“pteron” = wing), for the elytra with a yellowish speckle on black background.

21. Oxytelus barbatus Fauvel, 1905 (Fig. 27) Fauvel 1905: 113 (Type locality: Java).

Type material examined. Holotype [monotypy; Fig. 27A, C, F, G, J–N]: ♂, Java//barbartus Fvl.// TYPE// Coll. et det. A. Fauvel, Oxytelus barbatus Fauv., R.I.Sc.N.B. 17.479 (IRSNB). Other material examined. 4 specs. (2♂, 2♀). 1♂, INDONESIA: Sumatra, Fort de Kock [=Bukittinggi], 920m, 1926, E. Jacobson leg., barbatus Fvl det. Cameron, barbatus Fauv. don. Cameron det. Bernhauer, Oxytelus barbatus Fvl ♀ det. P. Hammond 1969 (FMNH); 1♂, same data as previous except: Oxytelus barbatus ♂ Fauvel. det. M. Cameron 1927 (BMNH); 1♀, same data as previous except: barbatus Fauv. don. Jacobson det. M. Cameron 1928, Oxytelus barbatus Fvl ♀ det. P. Hammond 1969 (FMNH); 1♀, Sumatra, Fort de Kock [=Bukittinggi], 920m, 1925, E. Jacobson leg., Oxytelus barbatus ♀ Fauvel. det. M. Cameron 1927 (BMNH). Diagnosis. The male of O. barbatus can be distinguished from the males of other Oxytelus species by the pale patch in center of elytra (Fig. 27C) and the shape of the posterior margin of sternites VII and VIII (Fig. 27F, G). The females are akin to female O. ferreirai, but they differ in forebody punctation and elytral coloration (cf. Figs. 21B, D; 27B, E). Redescription. Body (Fig. 27A, B) dark brown; mouthparts, antennae, legs testaceous; elytra with pale pattern in center. Length ♂ [holotype], 4.3 mm; ♀, 5.0 mm. Male. Head (Fig. 27C) widest at eyes or temples. Disc punctate posteriorly, not pubescent. Clypeus flat, almost 1/4 head length, surface sparsely punctate; anterior margin slightly protruding and with two tiny teeth in middle. Epistomal suture with lateral portions incurved and running posteriorly to level of anterior margin of eyes. Vertex weakly convex; with mid-longitudinal suture feeble, two punctate paralateral sutures running anteriorly to level of posterior margin of eyes. Eyes with fine facets, convex and protruding laterally, shorter than temples. Temples obviously dilated. Occipital suture with middle portion absent; nuchal ridge interrupted in middle. Mandible (Fig. 27C) robust and strongly curved; two denticles on inner edge, one near middle and one at apical 1/4. Antenna (Fig. 27A) as long as head and pronotum together; antennomere 4 subglobose, without basal dish; with apical antennomere equal to two preceding together. Pronotum (Fig. 27C) transverse, broadest at about anterior 1/4, narrower than head. Disc 5-sulcate, midlongitudinal sulcus and two paramedial sulci deep, and punctate; two paralateral sulci straight, vague and shorter, each with punctate depression on lateral side. Lateral margins not crenulate. Elytra (Fig. 27C) punctate, striate, rugose, without lateral longitudinal ridge. Abdomen (Fig. 27A) pubescent, broadest at segment V. Sternite VII (Fig. 27F) with posterior margin shortly protruding and densely pubescent in middle. Sternite VIII (Fig. 27G) with subbasal ridge continuous in middle, with posterior margin broadly emarginate and shortly protruding in middle of emargination, apex of protrusion with two denticles. Aedeagus (Fig. 27J–M) with median lobe oblong-ovoid, anterior part narrower, with membranous internal sac inside (no sclerites); apico-medial hook with apical part short, apex blunt, with transverse ridge (like process in lateral view). Paramere arm-like, apical arm slender, and curved, with seta at apical 1/3; basal arm with shallow furrow. Female. Head (Fig. 27E) smaller than male, narrower than pronotum; clypeus flat and less protruding, with anterior margin truncate; temples shorter than eyes, not dilated, paralateral sutures absent, with occipital suture OXYTELUS IN SOUTHEAST ASIA


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continuous in middle. Mandible (Fig. 27E) smaller than male. Abdominal sternite VII without modification on posterior margin. Sternite VIII (Fig. 27H) with posterior margin broadly rounded and a little protruding in middle. Spermatheca (Fig. 27I) υ-shaped, basal portion not bulbous, apex rounded and slightly curved. Distribution. Indonesia. Remarks. The males of this species vary in head outline and even presence of the middle portion of occipital suture (e.g. Fig. 27D).

22. Oxytelus mandibularis Cameron, 1929 (Fig. 28) Cameron 1929: 442 (subgenus Tanycraerus; Type locality: Selangor: The Gap).

Type material examined. Lectotype [designated here, Fig. 28A–C, F–G, J–N]: ♂, LECTO-TYPE// Type// The Gap, Selangor, F.M.S. Dr. Cameron.// Dung// O. mandibularis Cam. TYPE// M. Cameron. Bequest. B.M.1955– 147.// Oxytelus mandibularis Cam. P.M.Hammond, det. 1970, LECTOTYPE ♂ (BMNH). Paralectotype: 1♀, PARA-LECTO-TYPE// The Gap, Selangor, F.M.S. Dr. Cameron.// M. Cameron. Bequest. B.M.1955–147.// Oxytelus mandibularis Cam. Paralectotype, P.Hammond, det. 1970, ♀ (BMNH). Diagnosis. The males of O. mandibularis can be readily recognized by the extremely elongated clypeus and mandibles (Fig. 28A, C). The females do not have these features but can be separated by the combination of the small size (≤ 4 mm), the fine-faceted eyes, and the appearance of pronotum (Fig. 28E). Redescription. Body (Fig. 28A, B, D) brown; elytra pale; mouthparts, antennomeres 1 – 4, legs testaceous. Length: ♂, 4.2 mm; ♀, 2.68 mm. Male. Head (Fig. 28C) widest at eyes. Disc punctate posteriorly, not pubescent. Clypeus surface sparsely punctate, with anterior margin extremely protruding into long and slender beak, longer than rest part of head. Epistomal suture with lateral portions incurved and running posteriorly to level of anterior margin of eyes. Vertex depressed anteriorly; with mid-longitudinal and paralateral sutures suture absent. Eyes with fine facets, convex and protruded laterally. Temples shorter than eyes. Occipital suture and nuchal ridge discontinuous in middle. Mandible (Fig. 28C) extremely elongated, as long as clypeus, and curved near apex; two denticles at basal 1/3 of inner edge and closely situated. Antenna (Fig. 28A) almost equal to head (excluding elongated part of clypeus) and pronotum together; antennomere 4 subglobose, without basal dish; with apical antennomere longer than penultimate one. Pronotum (Fig. 28A) transverse, broadest at near anterior 1/3, wider than head. Disc punctate and 3-sulcate; two paralateral depressions shallow and vague, striate; lateral margin not crenulate, with posterolateral angles not prominent. Elytra (Fig. 28A) punctate and rugose, with lateral longitudinal ridge absent. Abdomen (Fig. 28B) weakly coriaceous and covered with dense fine setae on ventral side, broadest at segment V. Sternite VII (Fig. 28D) with posterior margin slightly protruding in middle. Sternite VIII (Fig. 28E) with subbasal ridge discontinuous in middle but forming two small processes, with posterior margin protruding in middle and forming subtriangular lobe but truncate at apex, at base moderately and evenly emarginate and equipped with two tiny teeth a little behind. Aedeagus (Fig. 28F–I) with median lobe oblong in ventral view, wider in basal half; with membranous internal sac inside (no sclerites); apical orifice smaller than median lobe width; apico-medial hook reduced and appressed to median lobe, with apical tip pointed, carinate a little behind tip. Paramere with seta at 1/3 of apical arm, with triangular process near basal 1/3 of apical arm, and articulated by small sclerite; basal arm with shallow furrow. Female. Head (Fig. 28E) smaller than male, narrower than pronotum; clypeus flat and not protruding, with anterior margin truncate; temples shorter than eyes, slightly dilated, with occipital suture continuous in middle. Mandible (Fig. 28E) not elongated. Abdominal sternite VII without modification on posterior margin. Sternite VIII (Fig. 28H) with posterior margin broadly rounded. Spermatheca (Fig. 28I) slender and bent twice. Distribution. Malaysia. Remarks. Close to O. puncticeps. They are small in size (disregarding the hugely extended clypeus and mandibles of O. mandibularis); they share similar pronotal appearance (Figs. 28A; 29A–C; 30A–C); and they both bear extraordinary modification in clypeus. Otherwise they differ remarkably, though.


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23. Oxytelus puncticeps Kraatz, 1859 (Figs. 29 & 30) Kraatz 1859: 176 (Type locality: Ceylan). Oxytelus (Anotylus) micantoides Scheerpeltz 1978: 193 (Type locality: Dadim-Bere, Same, Timor). Syn. nov. Oxytelus (Anotylus) monoceros Cameron 1919: 226 (subgenus; Type locality: Ceylon: Colombo, Belligam, Nuwera Eliya); Cameron 1930: 228 (subgenus Tanycraerus, syn. of micans); Hammond 1975: 150 (syn. of puncticeps).

Type material examined. [O. micantoides] Holotype [by original designation, Fig. 30A, D, L]: ♂, ♂// TIMOR (Same) sobre folhias mortes// e excremento de cavalo [= on dead leaves and horse excrement], 12-I-66 R. N. F.// TYPUS Oxytelus micantoides O. Scheerpeltz// Oxytelus (Anotylus) micantoides Scheerp.// ex coll. Scheerpeltz// Oxytelus puncticeps Kr. P. M. Hammond, det. 1987// Oxytelus puncticeps = O. micantoides det. LÜ Liang, 2014 (NHMW). Paratype: 1♀, ♀// TIMOR (Same)// sobre folhias mortes// e excremento de cavalo, 12-I-66 R. N. F.// TYPUS Oxytelus micantoides O. Scheerpeltz// Oxytelus (Anotylus) micantoides Scheerp.// ex coll. Scheerpeltz// Oxytelus puncticeps = O. micantoides det. LÜ Liang, 2014 (NHMW). Other material examined. 14 specs. (7♂♂, 7♀♀). INDIA: 7♂♂, 5♀♀, Uttar Pradesh, Mussorie [=Mussoorie, governed by the state Uttarakhand since 2000], Rabbit-Farm, 1300m, 1989.VII.10, A. Riedel leg. (SMNS). INDONESIA: 1♀, N-Sumatra, ob. Ambarita, Samosir, 1200m, Weide-Strauch-Veget. [Willow shrub], in Rinderdung [in cattle dung], 1991.VIII.5, Erber leg. (SMNS). SRI LANKA: 1♀, Ceylon C. Prov [= Central Prov.], Knucles 1600m, 1983.VI.28, Ole Mehl leg., Sri Lanka, Ex coll. Viggo Mahler (ZMUC). VIETNAM: Tonkin, HoaBinh, 1939.VII, A. de Cooman leg., Oxytelus puncticeps Kr. det. P.M.Hammond 1980 (IZ-CAS). Diagnosis. The males of O. puncticeps can be readily recognized by the asymmetrical clypeal anterior margin and mandibles (Fig. 29B), although there are a few of exceptional individuals (Fig. 29C, D). The females have symmetrical clypeal anterior margin and mandibles (Fig. 30C) but can be separated by the combination of the small size (≤ 3 mm), the fine-faceted eyes, and the appearance of pronotum (Fig. 30C). The females of O. puncticeps and O. mandibularis can be divided by the striation of the pronotum (cf. Figs. 28E, 30C). Distribution. Borneo, Indonesia, Philippines, Vietnam; China (southern part), India, Japan, Sri Lanka, Nigeria, Cameroon, Congo, Uganda, Kenya, Tanzania, South Africa, Comoros, Madagascar. Remarks. The clypeal anterior margin is commonly useful for identifying O. puncticeps without dissection, but among SMNS specimens we found a male (Fig. 29C, D) with symmetrical clypeus (bearing two tiny teeth) but share other trait with normal O. puncticeps specimens.

24. Oxytelus lucidulus Cameron, 1929 (Fig. 31) Cameron 1929: 442 (subgenus Tanycraerus; Type locality: Selangor: Bukit Kutu). Oxytelus ginyuenensis Bernhauer 1935: 5 (Type locality: Tempelberg, 900 m, Ginyün b. Bébé, C. China); Lü & Zhou 2012: 28 (syn. of lucidulus). Oxytelus takahashii Ito 1994: 44 (subgenus Tanycraerus; Type locality: Japan: Nara Pref.: Nara Park); Lü & Zhou 2012: 28 (syn. of lucidulus).

Type material examined. [O. lucidulus] Lectotype [designated here, Fig. 31A–D]: ♂, Rotting Fruit.// Bukit Kutu, Selangor, F.M.S. Dr. Cameron.// Oxytelus lucidulus Cam TYPE// M.Cameron. Bequest. B.M.1955–147.// Oxytelus lucidulus Cam. P.M.Hammond det.1970, LECTOTYPE, ♂ (BMNH). [O. ginyuenensis] Lectotype [designated here, Fig. 31E, G]: ♂, Tempelberg, 900m// Ginyün b. Bébé C. China, Rttr. [=Chongqing, Beibei, Jinyunshan]// Ginyünensis Brnh. Typ.// ginyünensis Brnh. Typus, Oxytelus// Chicago NHMus, M.Bernhauer Collection// Oxytelus lucidulus Cam., P.Hammond, det.1969// Oxytelus ginyunensis Bernh., P.M.Hammond, det.1970, LECTOTYPE ♂ (FMNH). Paralectotype: 1♀, Tempelberg, 900m// Ginyün b. Bébé C. China, Rttr.// ginyünanensis Brnh. Cotypus, don. Reitter Jr.// Chicago NHMus, M.Bernhauer Collection// Oxytelus lucidulus Cam., P.Hammond, det.1969// Oxytelus ginyunensis Bernh., Paralectotype, P.M.Hammond, det.1970, ♂ (FMNH). [O. takahashii] Paratype: 1♂, KASUGA, NARA, 15.IX.1988, S. TAKAHASHI// PARATYPE, Oxytelus (Tanycraerus) takahashii T. ITO sp.nov. (TIPC-IZCAS). OXYTELUS IN SOUTHEAST ASIA


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Other material examined: 92 specs. (34♂♂, 58♀♀). LAOS: Vientiane Pr., 1♂, 1♀, Phou Khao Khouay NBCA, E of Ban Vangheua (=Khua), 18º20.055'N 102º48.949'E, 800m, bamboo thicket w/some large trees, dung trap (human), 2008.V.26–31, A. Newton & M. Thayer legg. (FMNH); LAOS: Champasak Pr., Bolaven Plateau, Ban Thôngvay (=Xékatam) vic.: 6♂♂, 2♀♀, old logging road N of village, 15º13.761'N 106º31.749'E, 995m, selectively logged forest, carrion trap (squid), 2008.VI.8–16, M. Thayer, A. Solodovnikov & A. Newton legg. (FMNH); 1♀, same data as previous except: 15º13.96'N 106º31.731'E, 1035m, A. Solodovnikov, M. Thayer, & A. Newton legg. (FMNH); 1♂, 3♀♀, same data as previous except: 15º14.288'N 106º31.891'E, 1095m, A. Solodovnikov, M. Thayer, & A. Newton legg. (FMNH); 4♂♂, 22♀♀, same data as previous except: 15º14.494'N 106º31.807'E, 1170m, A. Solodovnikov, M. Thayer, & A. Newton legg. (FMNH); 6♂♂, 8♀♀, same data as previous except: old logging road N of village, 15º13.761'N 106º31.749'E, 995m, selectively logged forest, dung trap (human), 2008.VI.8–16, M. Thayer, A. Solodovnikov & A. Newton legg. (FMNH); 1♂, 6♀♀, same data as previous except: 15º13.96'N 106º31.731'E, 1035m, A. Solodovnikov, M. Thayer, & A. Newton legg. (FMNH); 5♂♂, 3♀♀, same data as previous except: 15º14.288'N 106º31.891'E, 1095m, A. Solodovnikov, M. Thayer, & A. Newton legg. (FMNH); 7♂♂, 6♀♀, same data as previous except: 15º14.494'N 106º31.807'E, 1170m, A. Solodovnikov, M. Thayer, & A. Newton legg. (FMNH); 1♂, Champasak prov., Bolaven Plateau, Muang Paxong, Ban Thôngvay, 15º14.741'N 106º31.916'E, 1300m, 2008.VI.9–16, anonym leg. (ZMUC). MALAYSIA: 1♂, 2 ♀♀, Kuala Lumpur, ex manure, 25.IX.1965, D. H., A. C., & A. H. Kistner legg., Oxytelus lucidulus Cam. det. P.M.Hammond 1974 (FMNH). PHILIPPINES: 1♀, Palawan, Bacuit, Boettcher leg., megaceros Fvl. det. Bernhauer (FMNH). VIETNAM: 1♂, 3♀♀, Tonkin, Hoa-Binh, A. de Cooman leg., Oxytelus lucidulus Cam. det. P.M.Hammond 1981. Diagnosis. Oxytelus lucidulus is close to O. subferrugineus, but differs in the shape of the posterior margin of male sternite VII: O. lucidulus has two tiny teeth on the margin and a shallow emargination between the teeth (Fig. 10F in Lü & Zhou 2012), but O. subferrugineus has two large triangular processes and a large semicircular emargination on the relevant places (Figs. 32G; 33E). Oxytelus lucidulus is also similar to O. pallidipennis (from Pakistan, India, and Nepal) that they look almost the same by appearance (Lü & Zhou 2012). Yet, O. lucidulus can be distinguished definitely by the combination of the obvious teeth on the posterior margin of male sternite VII, the different shape of apico-medial hook on aedeagus, and the ventral process of paramere without a tiny curved projection (compare Fig. 31C and Lü & Zhou (2012: Fig. 10F, K–M) with Fig. 51G, K–R). Distribution. Laos, Malaysia, Philippines, Vietnam; China (southern part), Japan, Korea.

25. Oxytelus subferrugineus Cameron, 1929 (Figs. 32 & 33) Cameron 1929: 443 (subgenus Tanycraerus; Type locality: Pahang: Bentong). Oxytelus (Tanycraerus) nilgiriensis Cameron 1930: 227 (Type locality: Nilgiri Hills); Lü & Zhou 2012: 49 (syn. of subferrugineus). Oxytelus (Tanycraerus) kedirianus Cameron 1938: 147 (Type locality: E. Java: Res. Kediri Pandan). Syn. nov.

Type material examined. [O. subferrugineus] Lectotype [designated here, Fig. 32C–E]: ♀, Bentong, Pahang, F.M.S. Dr. Cameron.// O. subferrugineus Cam. TYPE// M. Cameron. Bequest. B.M.1955–147// Oxytelus subferrugineus Cam., P.M.Hammond det. 1970, LECTOTYPE ♀ (BMNH). [O. nilgiriensis] Lectotype [designated here, Fig. 32F–K]: ♂, Nilgiri Hills, A.K.Weld Downing// a 551// M. Cameron. Bequest. B.M.1955–147// O. nilgiriensis Cam. TYPE// Oxytelus nilgiriensis Cam., P.M.Hammond det. 1970, LECTOTYPE ♂ (BMNH). [O. kedirianus] Lectotype [designated here, Fig. 33A, B, E–H, J]: 1 ♂, PANDANARDEM. RES. KEDIRI. E. JAVA.// O. Kedirianus Cam. TYPE// M. Cameron. Bequest. B.M.1955–147.// Oxytelus kedirianus Cam. P.M.Hammond det.1970 LECTOTYPE ♂ (BMNH). Other material examined. 8 specs. (3♂♂, 5♀♀). MALAYSIA: 1♂, 2♀♀, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 495m, 1988.VIII.27, A. Smetana leg. (FMNH); VIETNAM: Tonkin, Hoa-Binh: 1♀, 1939.VII, A. de Cooman leg., Anotylus glareosus (Woll.) det. P. M. Hammond 1980; 1♂, 1♀, 1930, Clermont leg., nilgiriensis Cam. det. Bernhauer, Oxytelus subferrugineus Cam. = nilgiriensis Cam. det. P. M. Hammond 1970 (FMNH); 1♂, Ha Son Binh, Than Loc, at light, No. 85, 1986.I.30, Mahunka & Oláh legg. (HNHM); 1♀, Minhxuan, 1971.XII.2, No. 213, Gy. Topál leg. (HNHM).


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Diagnosis. Oxytelus subferrugineus can be readily recognized by the two large triangular processes and a large semicircular emargination on the posterior margin of male sternite VII (Figs. 32G; 33E). Distribution. Indonesia, Malaysia, Vietnam; China (Guangxi), India. Remarks. We examined a male O. kedirianus type specimen (Fig. 33A, B, E–H, J) labeled as “lectotype” by P. M. Hammond that is indeed in Cameron’s type series, but it actually belongs to Oxytelus subferrugineus. Another type (labeled as “paralectotype” by P. M. Hammond) that we know is a female deposited in RMNH (C. J. Louwerens collection, see Fig. 33C, D, I, K). Judged by the photos provided by the curator, Dr. Hans Huijbregts, this female specimen is also O. subferrugineus. There could be other syntypes, but their whereabouts are beyond our current knowledge. What is more, the author, Cameron (1938) described the male sexual traits as “6th ventral segment with a median subtriangular lobe, furnished with a tuft of yellow setae at the apex, on each side broadly arcuately emarginate; 7th produced as a triangular plate with rounded apex” (p. 147–148). In Cameron’s terminology, the “6th ventral segment” should be sternite VII and “7th” should refer to sternite VIII. One can easily find discrepancy between the type specimen (see Fig. 33E, F) and the description, according to which this species is not O. subferrugineus. It is, however, suspicious that the described “6th ventral segment” is so complicated that it is more likely to be a sternite VIII than sternite VII, and, actually, it just agrees with sternite VIII of the male type specimen. So it is reasonable to question the accuracy of Cameron’s description of male sexual traits. Therefore, O. kedirianus is here placed in synonymy on the basis of the lectotype specimen.

26. Oxytelus megaceros Fauvel, 1895 (Figs. 34 & 35) Fauvel 1895: 201 (Type locality: Bírmanie, Carin Chebà, 900–1100 m; Carin Asciuii Ghécu, 1400–1500 m; Malacca, Perak; Sumatra, tabacs). Oxytelus kalisi Bernhauer 1934a: 172 (Type locality: Ost-Java, Nongkodjadjar, 4000 f). Syn. nov. Oxytelus megaceros var. flavicollis Bernhauer 1912: 247 (Type locality: Philippinen).

Type material examined. [O. megaceros] Lectotype [designated here, Fig. 34A, C, E–K]: ♂, Carin [=Kayan, Myanmar], Asciuii Ghécu, 1400–1500 m. L. Fea. III-IV.88// megaceros Fvl.// Coll. et det. A. Fauvel, Oxytelus megaceros Fauv., R.I.Sc.N.B. 17.479// Syntype// Oxytelus megaceros Fvl. 1895, Lectotype ♂, det. LÜ Liang, 2015 (IRSNB). Paralectotype: 1♀, Carin [=Kayan, Myanmar] Chebà, 900–1100 m. L. Fea. V XII-88// Buitenzorg 2-3 (Java)// Pasuruan Java est// Coll. et det. A. Fauvel, Oxytelus megaceros Fauv., R.I.Sc.N.B. 17.479// Syntype// Oxytelus megaceros Fvl. 1895, Paralectotype ♀, det. LÜ Liang, 2015 (IRSNB). [O. kalisi] Lectotype [designated here, Fig. 35]: ♀, NONGKODJADJAR, EAST IAVA(4000 F.), I.P.A.KALIS,1934// Oxytelus Kalisi Brnh, Typus// Kalisi Brnh Typus Oxytelus// Chicago NHMus M. Bernhauer Collection// Oxytelus kalisi Bernh. P.M.Hammond det.1970, LECTOTYPE, ♀ (FMNH). Additional material examined. 38 specs. (26♂♂, 12♀♀). CHINA: 1♂, Kouy-Tcheou [= Guizhou or KweiTschou or Kueitschou], Reg. de Pin-fa, 1906, Père Cavalerie [collector], Coll. et det. A. Fauvel Oxytelus megaceros Fauv. R.I.Sc.N.B. 17.479 (IRSNB). INDONESIA: 1♀, Java, Bandung, Bandung Institute of Technology, ex leaf compost, 1988.III.30, D. L. Wrensch leg. (FMNH); 1♂, Java, Jockaboemi, Mt. Suiczou, Malang, Coll. et det. A. Fauvel Oxytelus megaceros Fauv. R.I.Sc.N.B. 17.479 (IRSNB); 6♂♂, 3♀♀, Lombok, Sesaot, 500m, 1994.II.1, Bolm leg. (SMNS); 1♂, 1♀, Lombok Is., Senaro [=Senaru village], N slope of Rinjani, 1994.II.2–5, 1100m, Bolm leg. (SMNS); 3♂♂, Bali, Danau Buyan, 1300m, 1994.II.19–21, Bolm leg. (SMNS); 1♂, Bali, Lake Bratan, ca.1200m, 1994.I.27, J. Pedersen leg. (ZMUC); 1♂, 1♀, Kebun Raya, ca.1600m, 1994.II.4, J. Pedersen leg. (ZMUC); 1♂, Sumatra, Ajer Mantcior, Agosto, 1878, O. Beccari [collector], “Perak Malacca”, “Birmanie Helfer”, “Basakumbo Sumatra”, Coll. et det. A. Fauvel Oxytelus megaceros Fauv. R.I.Sc.N.B. 17.479 (IRSNB); 1♂, Sumatra, Mte Singalang, Luglio, 1878, O. Beccari [collector], “Basakumbo Sumatra”, Coll. et det. A. Fauvel Oxytelus megaceros Fauv. R.I.Sc.N.B. 17.479 (IRSNB); 1♀, N-Sumatra, ob. Ambarita, Samosir, 1200m, WeideStrauch-Veget. [Willow shrub], in Rinderdung [in cattle dung], 1991.VIII.5, Erber leg. (SMNS); 1♀, S Sulawesi, 8 km W Mamasa, 2º56'13''S 119º20'32''E, 950m, 1999.VII.18–21, Bolm leg. (SMNS). LAOS: 4♂♂, 1♀, Champasak prov., Bolaven Plateau, Muang Paxong, Ban Thôngvay (=Xékatam) vic., 15º14.054'N 106º31.867'E, 1200m, Edge of distributed primary rainforest, near clearing, flight intercept trap, 2008.VI.8–16, A. Solodovnikov & J. Pedersen legg. (ZMUC). PHILIPPINES: 1♀, Luzon, Subuagrn (FMNH); 1♂, 1♀, Luzon, Subuagrn, Boettcher leg. (FMNH); 1♂, same data as previous except: Oxytelus megaceros var. flavicollis Bnh. det. P.M. Hammond 1974



29

(FMNH); 1♂, same data as previous except: flavicollis Brnh. det. Bernhauer (FMNH); 1♀, Mindanao, Davao, Sitio Taglawiq, Maco, Tagum, nr. sea level, original dipterocarp for., 1946.X, H. Hoogstraal & D. Heyneman legg. (FMNH); THAILAND: 1♂, Phatthalung prov., around Wildbreeding Center, secondary forest, swept and singled, No. 48, 2003.XII.2, A. Orosz & Gy. Sziráki legg (HNHM); VIETNAM: 1♂, Lam Dong Prov., Duc Me (Maria stream), 15 km S of BaoLoc, No. 372, 1988.X.23, S. Mahunka & T. Vásárhelyi legg. (HNHM); 1♂, Luc yen [Yen Bai prov., Luc Yen Distr.], No. 239, 1971.XII.5, Gy. Topál leg. (HNHM). Diagnosis. Oxytelus megaceros can be separated from other Oxytelus species by the combination of the finefaceted eyes (Fig. 34C, D), the antenna with basal dish on the article 4 (Fig. 35C), and the shape of the posterior margin of male sternites VII and VIII (Fig. 34E, F). Distribution. Borneo, Indonesia, Malaysia, Myanmar, Philippines, Thailand, Vietnam; China (southern part), India, Pakistan. Remarks. When describing O. kalisi, Bernhauer (1934a) compared it with O. megaceros. We investigated both the text and the type specimen, finding that the difference in size claimed by the author is probably the outcome of a situation that the abdomen was (and it remains) abnormally extended (the forebody lengths are almost equal according to our measurement). And we also conclude that the differences in punctation and shape are trifling and that in coloration does not exceed intraspecific variation. The ultimate evidence is that both the antennae of the types of O. megaceros and O. kalisi have a basal dish on the antennomere 4, which we have found only in O. megaceros (apart from the sculptus species-group [Epomotylus]).

27. Oxytelus fallax Fauvel, 1878 (Fig. 36) Fauvel 1878: 215 (Type locality: Ternate); Bernhauer 1936: 81 (Oxytelus; misidentification, see subincisus; Philippines).

Type material examined. Holotype [monotypy, Fig. 36]: ♂, Ternate X, Beccari 1875// TYPUS// Museo Civ. Genova// Oxyt. fallax Fvl.// Oxytelus fallax Typus! Fauv.// HOLOTYPUS ♂ Oxytelus fallax Fauvel, 1878// fallax Fvl. (MSNG). Diagnosis. Oxytelus fallax can be easily distinguished from O. sculptus, O. subsculptus, O. grandiculus sp. nov., and O. subincisus (sculptus species-group [Epomotylus]) by the broad emargination and membranous border of the posterior margin of sternite VII (male only). However, O. antennalis (= O. cheesmani syn. nov., from Papua New Guinea, Australia, and Pacific islands) also shares this trait (Fig. 46E, H). Oxytelus fallax is different from O. antennalis in aedeagus, especially in the profiles of the apico-medial hook of median lobe (there is no tiny denticle at near the corner, cf Figs. 36G; 46M) and parameres (shorter dorsal process, cf. Figs. 36H; 46N). Redescription [holotype]. Body (Fig. 36A) brown; head a little darker. Length: 3.45 mm. Head (Fig. 36B) widest at eyes. Disc punctate, not pubescent. Clypeus subrectangular, flat, about 1/4 head length, surface sparsely punctate; anterior margin truncate and hardly beyond anterior margin of supra-antennal ridges. Supra-antennal ridges elevated upward. Epistomal suture with lateral portions weakly incurved and running to level of anterior margin of eyes. Vertex weakly convex, definitely distinguished from neck by occipital suture; with mid-longitudinal suture distinct, paralateral sutures absent. Eyes with coarse facets, extremely convex and protruding laterally, covering almost all lateral area. Temples reduced. Occipital suture continuous in middle; nuchal ridge almost invisible. Mandible (Fig. 36B) curved; two denticles on inner edge. Antenna (Fig. 36A) stout, longer than head and pronotum together; antennomere 4 subglobose, with dish at base; ultimate antennomere shorter than preceding two together. Pronotum (Fig. 36B) transverse, broadest at near anterior 1/3, wider than head. Disc punctate and 3-sulcate; two paralateral depressions shallow, weakly strigose; lateral margin not crenulate. Elytra (Fig. 36B) punctate and rugose, with lateral longitudinal ridge on each elyton. Abdomen coriaceous and densely setose on ventral side, broadest at segment V. Sternite VII (Fig. 36C) with posterior margin transparent and broadly emarginate. Sternite VIII (Fig. 36D) with transverse subbasal ridge continuous and straight in middle; posterior margin with two deep and narrow ruptures near middle, forming subrectangular median plate, apical margin of median plate truncate, slightly emarginate in middle, with tiny transverse carina a little behind, and with mid-longitudinal internal ridge.


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Aedeagus (Fig. 36E–H) with median lobe oblong-ovoid, widest at base; with two membranous strips visible inside but no sclerites; apical orifice large; apico-medial hook broad in ventral view and bent at middle in lateral view, with pointed apical tip. Paramere short, arm-like, apical arm slightly projected ventromedially at near corner between apical and basal arms, with dorsal process present, with seta on near apex, with apex rounded; basal arm with shallow furrow. Distribution. Indonesia; New Guinea. Remarks. We consider O. antennalis, O. fallax, O. grandiculus sp. nov., O. meinanderi Scheerpeltz (cf. Makranczy 2014), O. sculptus Gravenhorst (Fig. 52), O. subincisus, and O. subsculptus as “sculptus speciesgroup”, which is almost equal to the subgenus Epomotylus, based on resemblance that they are similar in head shape, the stout antennomere 1, the basal dish of antennomere 4, the obviously elevated supra-antennal ridges, the extreme coverage and convexity of eyes, the deeply bi-emarginate posterior margin of male sternite VIII, and the male and female genitalia. The O. megaceros has similar antennae (with basal dish on antennomere 4), but differs from this species-group in other characters. Bernhauer (1936) reported this species from the Philippines. We examined (at least a majority of) these specimens (FMNH) and found all the male specimens to be O. subincisus, so we remove the Philippines from the distribution of O. fallax.

28. Oxytelus grandiculus sp. nov. (Figs. 37 & 38) Type material examined. Holotype [Fig. 38A, C, N]: ♂, HOLOTYPE// BORNEO, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Spgs, 495m, 23.VIII.88, A. Smetana “[B140]”// FIELD MUSEUM ex collection of H. G. Nelson// Oxytelus grandiculus. ♂, sp. n. Holotype, det. Liang LÜ, 2014// Oxytelus grandiculus, HOLOTYPE ♂, det. LÜ Liang, 2014 (FMNH). Paratypes: 16♂♂, 11♀♀, same data as holotype (FMNH); 1♀, same data as holotype except: 21.VIII.88, “[B136]” (FMNH); 6♂♂, 4♀♀, same data as holotype except: 24.VIII.88, “[B144]” (FMNH); 1♂, BORNEO: SABAH, Kinabalu N.P.: Poring 500m, 15.-16.XI.1996, leg. W.SCHAWALLER// Oxytelus grandiculus ♂, PARATYPE, det. LÜ Liang, 2014 (SMNS); 1♂, BORNEO: SABAH, Crocker Range N.P., NW Keningau, 900-1200m, at light; 18.XI.1996, leg. D.Grimm// Oxytelus grandiculus ♂, PARATYPE, det. LÜ Liang, 2014 (SMNS). 1♀, NORD-SUMATRA: ob. Ambarita, Samosirm 5.8.1991, ERBER// 1200 m, Weide-StrauchVeget. In Rinderdung// Oxytelus grandiculus ♀, PARATYPE, det. LÜ Liang, 2014 (SMNS). 4♀♀, THAILAND, Doi Phuka, National Park, headquarters, 26-27.XI.2003,// UV light, leg. L. Peregovits, M. Földvári, Á. Kőrösi, A. Szappanos & B. Maklári-Kis, No.18// Oxytelus grandiculus ♀, PARATYPE, det. LÜ Liang, 2014 (HNHM). Diagnosis. This new species can be easily distinguished from other species in sculptus group by the combination of the large body size (≥ 4 mm) and the male tergite VIII with a round depression in center and with incrassate posterior margin (Figs. 37D; 38F). Description. Body (Fig. 38A, B) shiny, brownish; head, antennae, and pronotum a little darker. Length [average] ♂, 5.2 mm; ♀, 4.6 mm. Male. Head (Figs. 37A; 38C) subpentagonal, widest at eyes. Disc punctate, not pubescent. Clypeus subrectangular, flat, protruding beyond anterior margin of supra-antennal ridges, longer than 1/3 head length, surface sparsely punctate and coriaceous; anterior margin truncate and slightly emarginate. Supra-antennal ridges elevated upward. Epistomal suture with lateral portions incurved and running posteriorly a little beyond level of anterior margin of eyes. Vertex weakly convex, definitely distinguished from neck by middle portion of occipital suture; with mid-longitudinal suture distinct and long, nearly anteriorly touching clypeus, paralateral sutures absent. Eyes with coarse facets, extremely convex and almost hemispherical, covering almost entire lateral area. Temples reduced. Occipital suture straight in dorsal view; nuchal ridge obscure, dorsal basal ridge absent. Mandible (Fig. 37B) curved; two denticles on inner edge near middle and near apex each. Antenna (Fig. 38A) stout, longer than head and pronotum together; antennomere 4 subglobose, with dish at base; apical antennomere shorter than two preceding together. Pronotum (Fig. 38C) transverse, broadest at near anterior 1/3, wider than head. Disc punctate and extremely rugose, with 3 longitudinal sulci; two paralateral depressions shallow; lateral margin not crenulate and posterior angles quite prominent. Elytra (Fig. 38C) punctate and rugose, with lateral longitudinal ridge on each elytron. Legs slender, similar to O. subincisus but larger, not broadened (as Fig. 41D). OXYTELUS IN SOUTHEAST ASIA


31

Abdomen (Fig. 38A) coriaceous and covered with dense fine setae, broadest at segment V. Tergite VII (Fig. 37C) with sclerotized and bi-lobed area anterior to basal ridge. Sternite VII (Figs. 37E; 38E) with posterior margin straight shallowly emarginate in middle. Tergite VIII (Figs. 37D; 38F) with large but shallow round depression in center and posterior to subbasal ridge, at either side with weak carina, and with posterior margin broadly rounded and thickened. Sternite VIII (Figs. 37F; 38G) with transverse subbasal ridge straight, with mid-longitudinal internal ridge from middle to posterior margin; posterior margin broad with two deep and narrow ruptures near middle, forming subrectangular median plate, apical margin of median plate slightly produced in middle, behind which with short transverse carina. Tergite X (Fig. 37H) gradually narrowed posteriorly, with posterior margin truncate or feebly serrate, with two lines of setae on pubescent posterior part. Aedeagus (Figs. 37K–N; 38J–M) with median lobe oblong-ovoid, widest at middle; with two membranous strips and other structures visible inside but no sclerites; apical orifice large; apico-medial hook broad in ventral view and bent at middle, with apical part short and a little pointed, basal process well developed; dorsal membranous area not covering all dorsum but across length of median lobe. Paramere arm-like, apical arm projected at near corner, with dorsal process present, with seta on near apex of apical arm, with apex rounded; basal arm with shallow furrow. Female. Abdominal sternite VII with posterior margin straight and not emarginate in middle. Tergite VIII without depression or carina, with posterior margin less thickened. Sternite VIII (Figs. 37G; 38H) with posterior margin parabolically rounded. Tergite X (Fig. 37I) with anterior margin narrower. Spermatheca (Figs. 37J; 38I) dumbbell-shaped, inflated into sphere at both base and apex. Distribution. Indonesia (Sumatra), Malaysia (Borneo), Thailand. Etymology. The specific epithet is a Latin adjective. We use this name because this species is “rather large” in the sculptus group. Remarks. Oxytelus grandiculus sp. nov. differs from other species in the sculptus group by combination of larger body size, the slender metatibiae, and the male tergite VIII shallowly depressed and with the posterior margin relatively thickened.

29. Oxytelus subsculptus Cameron, 1928 (Figs. 39 & 40) Cameron 1928: 101 (subgenus Epomotylus; Type locality: Sumatra: Fort de Kock).

Type material examined. Lectotype [designated here, Fig. 40C, D, N]: ♀, Fort de Kock, Sumatra. E. Jacobson. B.M.1928–247.// Fort de Kock (Sumatra) 920 M// Oxytelus subsculptus Cam TYPE// Oxytelus subsculptus Cam. P.M. Hammond det. 1972 LECTOTYPE ♀ (BMNH). Other material examined. 11 specs (4♂♂, 7♀♀), MALAYSIA: Borneo, Sabah, Mt. Kinabalu Nat. Pk., 480m, Poring Hot Spgs, 1988.VIII.19, A. Smetana “[B126]”, FIELD MUSEUM ex collection of H. G. Nelson (FMNH). Diagnosis. This species can be easily distinguished from other species in sculptus group by the laterally expanate metatibiae (Figs. 39C; 40E, compared with Fig. 41D). Redescription. Body (Fig. 40A, C) brown; head, antennae a little darker. Length [average] ♂, 3.4 mm; ♀, 3.8 mm. Male. Head (Figs. 39A; 40B) widest at eyes. Disc punctate, not pubescent. Clypeus subrectangular, flat, about 1/4 head length, surface sparsely punctate and coriaceous; anterior margin truncate and hardly beyond anterior margin of supra-antennal ridges. Supra-antennal ridges elevated upward. Epistomal suture with lateral portions weakly incurved and running to level of anterior margin of eyes. Vertex weakly convex, definitely distinguished from neck by occipital suture; with mid-longitudinal suture distinct and long, paralateral sutures absent. Eyes with coarse facets, extremely convex and protruding laterally, covering almost all lateral area. Temples reduced. Occipital suture continuous in middle; nuchal ridge almost invisible, dorsal basal ridge absent. Mandible (Fig. 39B) curved; two denticles on inner edge near middle and near apex each. Antenna (Fig. 40A) stout, longer than head and pronotum together; antennomere 4 subglobose, with dish at base; ultimate antennomere shorter than preceding two together.


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Pronotum (Fig. 40B) transverse, broadest at near anterior 1/3, wider than head. Disc punctate and 3-sulcate; two paralateral depressions shallow, weakly strigose; lateral margin even in front and slightly crenulate behind and posterior angles prominent. Elytra (Fig. 40B) punctate and rugose, with lateral longitudinal ridge on each elytron. Legs similar to O. subincisus Cameron, but metatibia laterally expanate at basal 1/3 (Figs. 39C; 40E). Abdomen (Fig. 40A) coriaceous and covered with dense fine setae, broadest at segment V. Sternite VII (Figs. 39D; 40F) with posterior margin straight. Sternite VIII (Figs. 39E; 40G) with transverse subbasal ridge continuous and straight in middle; posterior margin with two deep and narrow ruptures near middle, forming subrectangular median plate, apical margin of median plate truncate with tiny transverse carina a little behind, and with midlongitudinal internal ridge. Tergite X (Fig. 39G) longer than wide, sharply narrowed posteriorly, posterior margin truncate, with two lines of setae on pubescent posterior part. Aedeagus (Figs. 39J–M; 40J–M) with median lobe much elongate, widest at base, and constricted near apical orifice; with two membranous strips visible inside but no sclerites; apical orifice large; apico-medial hook broad in ventral view and bent at middle in lateral view, with pointed apical tip and basal process well developed in middle; dorsal membranous area not covering all dorsum but across whole length of median lobe. Paramere arm-like, apical arm slightly projected ventromedially at near corner, with dorsal process present, with seta on near apex, with apex rounded; basal arm with shallow furrow. Female. Similar to male. Abdominal sternite VIII (Figs. 39F; 40H) with posterior margin parabolically rounded and slightly protruding in middle. Tergite X (Fig. 39H) similar to male but posterior margin a little broader. Spermatheca (Figs. 39I; 40I) dumbbell-shaped, inflated into sphere at both base and apex. Distribution. Indonesia, Malaysia (Borneo). Remarks. Among the species in sculptus group, the laterally expanate metatibiae (Figs. 39C; 40E) of O. subsculptus is such a notable character that we can use it to prove the identity of the males (all the types are females).

30. Oxytelus subincisus Cameron, 1941 (Figs. 41 & 42) Cameron 1941: 433 (subgenus Caccoporus; Type locality: Philippines: Los Banos: Mt. Banahoa); Bernhauer 1936: 81 (Oxytelus; misidentified as fallax; Philippines). Oxytelus (Caccoporus) fruhstorferi Cameron 1945: 143 (Type locality: E. Java: Pengalengan, altitude 4000 feet). Syn. nov.

Type material examined. [O. subincisus] Lectotype [designated here, Fig. 41A–C, N]: ♀, 12// Los Banos Luzon/ / O. subincisus Cam. TYPE// M. Cameron. Bequest. B.M.1955–147// Oxytelus subincisus Cam., P.M. Hammond det. 1972, LECTOTYPE ♀ (BMNH). [O. fruhstorferi] Holotype [monotypy, Fig. 42]: ♀, Java occident. Pengalengan. 4000' 1893 H. Fruhstorfer.// O. fruhstorferi Cam. TYPE// M.Cameron Bequest. B.M.1955–147. Other material examined. 23 specs. (14♂♂, 9♀♀). CHINA: 1♂, Yunnan, Kunming, Guandu Dt., Huiliu Village, in grass heap, 1950m, 2006.VII.25, X.-Y. Li leg. (IZ-CAS); 1♀, Guangxi, Napo, Defu, 1350m, 2000.VI.18, W.-Z. Li leg. (IZ-CAS). LAOS: 1♂, Vientiane Pr., Phou Khao Khouay NBCA, Ban Vangheua (=Khua) school, 18º20.37'N 102º48.529'E, 775m, settlement near forests & rice fields, at light (UV & MV), 25– 30.V.2008, A. Newton, M. Thayer et al. legg. (FMNH); PHILIPPINES: Luzon: 2♂♂, Laguna, Mt. Banahao, Boettcher leg., fallax Fvl. det. Bernhauer (FMNH); 1♂, 2♀♀, Laguna, Los Banos, Boettcher leg., fallax Fvl. det. Bernhauer, Oxytelus subincisus Cam. det. P. M. Hammond (FMNH); 7♂♂, 3♀♀, Laguna, Mt. Makiling, 4km SE Los Banos, Berlese funnel from rotten figs, 7.V.1977, L. E. Watrous leg. (FMNH); 2♂♂, 1♀, same data as previous except: in rotting figs (FMNH); 2♀♀, Ilocos Norte, Bangui, Boettcher leg., “luzonicus Brnh. Typ.”, fallax Fvl. det. Bernhauer (FMNH). Diagnosis. Oxytelus subincisus can be separated from other Southeast Asian species in sculptus group except O. subsculptus by the straight posterior margin of male sternite VII, and can be distinguished from O. subsculptus by the slender metatibia (not explanate, Fig. 41D). Distribution. Laos, Philippines, Indonesia, China (Guangxi, Yunnan).



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Remarks. The unique type of O. fruhstorferi apparently looks the same as O. subincisus in our current treatment and there is no reason not to treat them as synonyms, although the unique type of O. fruhstorferi and the types of O. subincisus are all females, so this decision has to be taken with some caution.

Species excluded from Oxytelus Anotylus transversipennis (Scheerpeltz, 1978) comb. nov. (Fig. 43) Scheerpeltz 1978: 191 (Oxytelus, subgenus Anotylus; Type locality: Betano, Alfandeça, Fomenta, Timor); Herman 2001: 1461 (Oxytelus; catalog).

Type material examined. Holotype [by original designation, Fig. 43A, B, E–K]: ♂, ♂// BETANO (Alfandeça)// Vozudo ao caù da tarde, 14-II-66, R.N.F.// Oxytelus (Anotylus) transversipennis Scheerp.// TYPUS, Oxytelus transversipennis O. Scheerpeltz// ex coll. Scheerpeltz// Anotylus transversipennis (Scheerp.) P.M.Hammond det. 1987 (NHMW). Paratype: 1♀, ♀// BETANO (Alfandeça)// Vozudo ao caù da tarde, 14-II-66, R.N.F.// Oxytelus (Anotylus) transversipennis Scheerp.// TYPUS, Oxytelus transversipennis O. Scheerpeltz// ex coll. Scheerpeltz (NHMW). Distribution. Indonesia. Remarks. This species was described as a species in the subgenus Anotylus (Scheerpeltz 1978), even though the Anotylus had been re-elevated as a genus by Herman (1970), but neither Herman (1970) nor Hammond (1976) transferred this species to Anotylus. In Herman (2001), it remained in the genus Oxytelus. Here by examining the types of Oxytelus (Anotylus) transversipennis (NHMW), we confirm that it is an Anotylus species and remove it from Oxytelus.

Some species from adjacent regions The following 11 species are outside the study region. But Southeast Asia is encompassed by India, China, Australia, and Papua New Guinea. It is, therefore, necessary to examine and comment these species here, though they are currently not recorded from the region in question. The species that co-occurr in both Southeast Asia and China were revised in Lü & Zhou (2012) and also commented in the above text, but the four species described in Lü & Zhou (2012), O. abiturus, O. ailaoshanicus, O. bajiei, and O. solus, were not illustrated by color figures in that paper. These species are distributed in China (Yunnan, Sichuan, and other provinces), and O. ailaoshanicus are even found in Nepal (see below). So the color illustrations provided here may be helpful to the readers who would be interested in the species in the boundary areas. For the five designations here, we follow the authors’ suggestion (the specimen labeled as “type/typus” but “cotype/cotypus”, though the full data were not present in the original description) and the preivous researcher’s results (unpublished), and we compared the lectotype specimen with the original description in order to make sure that the designated specimen is consistent with the description. And additionally, we propose a new synonym (O. antennalis = O. cheesmani) here. *1. O. abiturus Lü & Zhou (Fig. 44). Holotype [by original designation, Fig. 44A, B]: ♂, HOLOTYPE// CHINA, Yunnan, Lijiang, Yongshengxian, 2180m, by heap trap, 2000.VIII.2, H.-Z. Zhou & X.-D. Yu legg. (IZCAS). Paratypes: 3♂♂, 9♀♀, same data as holotype (IZ-CAS); CHINA, Sichuan: 1♂, Huilixian, Yimen, 1974.VII.29, G.-Y. Han leg. (IZ-CAS); CHINA, Yunnan: 1♂, Chuxiong, Lufeng, Guangtong, from weed heap in crop field, 2006.VII.28, X.-Y. Li leg. (IZ-CAS); 1♀, same data as previous except: K. Shi leg. (IZ-CAS). Other material. 2♂♂, 4♀♀, same data as holotype (IZ-CAS). *2. O. ailaoshanicus Lü & Zhou (Fig. 45). Holotype [by original designation, Fig. 45A, B]: ♂, HOLOTYPE// CHINA, Yunnan, Jingdong, Ailaoshan [=Mt. Ailao], South of the Ailaoshan station, pitfall trap, 2010.IX.18, L. Lü and X. Zhang legg.// N. 24º32.36', E. 101º01.67', Alt. 2475–2480m (IZ-CAS). Paratypes: 1♂, PARATYPE// CHINA, Yunnan, Jingdong, Ailaoshan [=Mt. Ailao], South of the Ailaoshan station, flight interception trap, 2010.IX.19, Y.-L. Zhou & X. Zhang legg.// N. 24º32.682', E. 101º01.668', Alt. 2513m (IZ-CAS) [Note: this specimen was misidentified as female in Lü & Zhou (2012)]; 1♀, PARATYPE// CHINA, Yunnan, Lijiangxian,


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Maoniuping, 3115m, Abies forest, in fungi, 2000.VIII.3, H.-Z. Zhou leg. (IZ-CAS). Other material. NEPAL: 1♂, “Nepal424” Lalitpur Distr., Mt.Phulchoki, 1800–2000m, 1995.IV.25, Martens & Schawaller legg., Oxytelus ailaoshanicus Lü & Zhou, ♂, det. LÜ Liang, 2013 (SMNS); 2♂♂, “Nepal426” Kaski Distr., above Pothana, 2000m, 1995.IV.27–29, Martens & Schawaller legg., Oxytelus ailaoshanicus Lü & Zhou, ♂, det. LÜ Liang, 2013 (SMNS). *3. O. antennalis Fauvel, 1889 = O. cheesmani Bernhauer, 1934b syn. nov. (Figs. 46 & 47). Type material. [O. antennalis Fauvel (Fig. 46)] Lectotype [designated here, Figs. 46A, B, E, F, J–O]: ♂, LECTO-TYPE// PARALECTO-TYPE// Tonglioué “ghe” [cramped handwriting]// Yahoué février// novembre// antennalis Fvl.// Coll. et det. A. Fauvel, Oxytelus antennalis Fauv., R.I.Sc.N.B. 17.479// Syntype// Oxytelus antennalis Fvl. P.M. Hammond det. 1972, LECTOTYPE ♂ & 3 PARALECTOTYPES (♀) (IRSNB). Paralectotypes: 3♀♀, same data and pinned together with Lectotype. [O. cheesmani Bernhauer (Fig. 47)]. Lectotype [designated here, Fig. 47A, B, E, F, H, J– O]: ♂, LECTO-TYPE// Type// New Hebrides: Santo. Viii-ix.1929. L.E.Cheesman. B. M. 1929-537.// S. 125.// Oxytelus Cheesmani Brnh. Typus// Oxytelus cheesmani Bernh. P.M.Hammond det. 1970 LECTOTYPE ♂// Oxytelus antennalis Fvl. P.Hammond det. 1969 (BMNH). Paralectotype: 1♀, PARA-LECTO-TYPE// New Hebrides: Santo. Viii-ix.1929. L.E.Cheesman. B. M. 1929-537.// S. 131.// Oxytelus cheesmani Bernh., Paralectotype, P.Hammond det. 1970 ♀ (BMNH). Other material. 1♀, AUSTRALIA: Queensland, Mulgrave River, Oxytelus antennalis Fvl. det. P.Hammond (BMNH); 1♂, PAPUA NEW GUINEA: Finschhafen, 1.IV.1944, Oxytelus antennalis Fvl. det. P.Hammond 1969 (BMNH). Remarks. We examined both the types of O. antennalis and O. cheesmani and we suppose that they are synonymous species, as they are same in head (Figs. 46B, D; 47B, D), male sternites VII–VIII (Figs. 46E, F; 47E, F), aedeagus (Figs. 46J–N; 47J–N), and spermatheca (Figs. 46I; 47I). When describing the O. cheesmani, Bernhauer (1934b) compared the new species with O. fallax. For the differences between O. fallax and O. antennalis (= O. cheesmani), please see the “Diagnosis” of O. fallax. *4. O. bajiei Lü & Zhou (Fig. 48). Holotype [by original designation, Fig. 48A, B]: ♂, HOLOTYPE// CHINA, Hunan, Bamianshan Natural Reserve, 774m, light trap, 9–12.VII.2008, Z. Yang leg. (IZ-CAS). Paratypes: 2♂♂, 1♀, same data as holotype (IZ-CAS). *5. O. cheesmanianus Cameron (Fig. 49). Lectotype [designated here, Fig. 49]: ♂, LECTO-TYPE// TYPE// PAPUA: Kokoda. 1,200ft. viii.1933. L.E.Cheesman. B. M. 1933-577.// O. Cheesmanianus Cam. TYPE (BMNH). Remarks. This species can be readily separated by the reddish color (Fig. 49A), truncate anterior margin of clypeus (Fig. 49B), and the two round and fine teeth at the posterior angle of pronotum (Fig. 49B). The aedeagal median lobe had been partly damaged before our examination, though still informative. There is no P. M. Hammond’s “lectotype” label on the pin, but this specimen was labeled as “TYPE” by Cameron (like other Cameron’s types), so we choose it as the lectotype. *6. O. hingstoni Cameron (Fig. 50). Lectotype [designated here, Fig. 50]: ♀, LECTO-TYPE// Type// Sikhim: Lachen. 10,000ft. 26.iv.1924. Maj.R.W.G.Hingston.// Everest Exp. Brit. Mus. 1924—386.// O. hingstoni TYPE Cam.// Oxytelus hingstoni Cam. P.M.Hammond det. 1970 LECTOTYPE ♀ (BMNH). Remarks. This species is akin to O. solus, but they can be separated by the color of elytra (Figs. 50B; 53A), though the types of the two species are heterosexual. We also saw some specimens in Nepal like O. hingstoni, but they differ in female body size and other characters, so the males of O. hingstoni are still unknown. *7. O. pallidipennis Cameron (Fig. 51). Syntypes: 1♂, PARA-LECTO-TYPE// Dehra Dun. Dr. Cameron, 30.9.1922// pallidipennis COTYPE Cam.// Cotypus don. Cameron// Chicago NHMus, M. Bernhauer Collection// Oxytelus pallidipennis Cam, Paralectotype, P. Hammond, det. 1970 ♂ (FMNH); 1♂, Dehra Dun India// Dehra Dun. Dr. Cameron, 30.9.1922// COTYPUS// Field Mus. Nat. Hist. 1966 A. Bierig Colln. Acc. Z-13812// O. (Tanycr.) pallidipennis Cam. (FMNH); 1♀, PARA-LECTO-TYPE// Dehra Dun. Dr. Cameron, 30.9.1922// Cotypus don. Cameron// Chicago NHMus, M. Bernhauer Collection// Oxytelus pallidipennis Cam, Paralectotype, P. Hammond, det. 1970 ♀ (FMNH). [Note: see the “Diagnosis” of O. lucidulus and Lü & Zhou (2012: 31) for the differentiation between O. pallidipennis and O. lucidulus.] *8. O. sculptus Gravenhorst (Fig. 52). 1♂, USA: California, Marin Co., Bolinas Mesa, compost, 1962.VIII.2– 4, P. M. Hammond leg. (BMNH); 1♀, NEW ZEALAND: MC, Christchurch, Halswell, garden, compost, 1984.I.19, P. M. Hammond leg., Oxytelus sculptus Grav. det. P. M. Hammond, 1986 (BMNH). Remarks. Oxytelus sculptus is the type species of the subgenus Epomotylus, and it has the notable baso-dished antennomere 4, so we use it to name the species group whose members have the coarce-faceted eyes and the basal



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dish on antennomere 4. And this species used to be thought as a “cosmopolitan” species (Herman 2001), but we have not found any specimens either from China or from Southeast Asia. *9. O. solus Lü & Zhou (Fig. 53). Holotype [by original designation and monotypy, Fig. 53]: ♂, HOLOTYPE/ / CHINA: Sichuan, Wolong, Wuyipeng, Yuancaodi, 2450m, in untouched birch forest, pitfall trap, 1–4.V.2004, X.D. Yu leg. (IZ-CAS). *10. O. tibetanus Bernhauer (Fig. 54). Lectotype [designated here, Fig. 54]: ♂, Grenze Thibet Ost: Tatsien-lu [=CHINA: Sichuan, Ganzi (or Garzê Tibetan Autonomous Prefecture), Kangding (=Dardo)], Em. Reitter.// Oxytelus tibetanus Brnh., Typ. un.// tibetanus Bernh., Typus unic.// Chicago NHMus, M.Bernhauer Collection// Oxytelus tibetanus Bernh., P. M. Hammond, det. 1970, LECTOTYPE ♂ (FMNH). [Note: Please see the details of this species in Lü & Zhou (2012).] *11. O. validus Cameron (Fig. 55). Paratype: 1♂, Para-type// M. Cameron. Bequest. B.M. 1955–147.// Neth. Ind.-American New Guinea Exped. Sigi Camp 1500 m, 17 ii 1939, L. J. Toxopeus// M. Cameron det., 19 O. (Tanycraerus) validus Cam. COTYPE (BMNH). Remarks. This species can be apparently distinguished from O. cheesmanianus by the darker color (Fig. 55A) and two tiny teeth on the anterior margin of clypeus (Fig. 55B).

Acknowledgements We thank Dr R. G. Booth (BMNH), Mr J. H. Boone (FMNH), Dr Gy. Makranczy (HNHM), Mr Y. Gérard & Dr W. Dekoninck (IRSNB), Dr H. Schillhammer (NHMW), Dr H. Huijbregts (RMNH), Dr S. M. Blank & Mr L. Behne (SDEI), Dr W. Schawaller (SMNS), Mr T. Ito for sending specimens or pictures for this study. And we thank Dr M. K. Thayer (FMNH), Dr Gy. Makranczy, and Dr T.-H. Luo (IZ-CAS) for many kinds of help during our study. Dr Gy. Makranczy and one anonymous reviewer reviewed earlier versions of this manuscript and gave a lot of constructive suggestions, for which we are very grateful. This study was supported by the National Natural Science Foundation of China (NSFC-31472036, NSFC-31272358, NSFC-J1210002), and a grant from the Key Laboratory of the Zoological Systematics and Evolution of CAS (No. Y229YX5105).

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(1851) Beitraege zur naeheren Kenntniss der Staphylinen Russlands. Enthaltend Beschreibung neuer Genera und Arten, nebst Erläuterungen noch nicht hinlänglich bekannter Staphylinen des russischen Reichs. Bulletin de la Société Impériale des Naturalistes de Moscou, 24 (pt. 2), 3–58. Ito, T. (1994) Notes on the species of Staphylinidae (Coleoptera) from Japan 3. Descriptions of two new species of the genus Oxytelus Gravenhorst. Natural History Research, 3, 41–46. Koch, C. (1934) Wissenschaftliche ergebnisse der Entomologische Expeditionen seiner Durchlaucht des Fuersten Alexandro C. della Torre e Tasso nach Aegypten und auf die Halbinsel Sinai. IV. Staphylinidae (Coleoptera). Bulletin de la Société Royale Entomologique d’Égypte, 1934, 33–91.



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Koh, L.P., Kettle, C.J., Sheil, D., Lee, T.M., Giam, X., Gibson, L. & Clements, G.R. (2013) Biodiversity State and Trends in Southeast Asia. In: Levin, S.A. (Ed.), Encyclopedia of Biodiversity. 2nd Edition. Vol. 1. Academic Press, Waltham, pp. 509– 527. Kraatz, G. (1859) Die Staphylinen-Fauna von Ostindien, insbesondere der Insel Ceylan. Archiv für Naturgeschichte Berlin, Abteilung A., 25, 1–196. http://dx.doi.org/10.5962/bhl.title.66002 Linné, C. von (1767) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Duodecima Reformata. Vol. 1. pt. 2. Laurentii Salvii, Holmiae, pp. 533– 1327 + xxxvii. Lü, L. & Zhou, H.-Z. (2012) Taxonomy of the genus Oxytelus Gravenhorst (Coleoptera: Staphylinidae: Oxytelinae) from China. Zootaxa, 3576, 1–63. Lü, L. & Zhou, H.-Z. (2015) Review of the genus Platystethus Mannerheim (Coleoptera: Staphylinidae: Oxytelinae) in China. Zootaxa, 3915 (2), 151–205. http://dx.doi.org/10.11646/zootaxa.3915.2.1 Makranczy, G. (2006) Systematics and phylogenetic relationships of the genera in the Carpelimus group (Coleoptera: Staphylinidae: Oxytelinae). Annales Historico-Naturales Musei Nationalis Hungarici, 98, 29–119. Makranczy, G. (2012) Nomenclatural changes removing secondary homonymy in the genus Oxytelus (Coleoptera: Staphylinidae: Oxytelinae). Acta Entomologica Musei Nationalis Pragae, 52, 193–205. Makranczy, G. (2014) Oxytelus meinanderi Scheerpeltz, 1974 and O. tuberculifrons Eichelbaum, 1913, two Afrotropical species with problematic type material (Coleoptera: Staphylinidae: Oxytelinae). Acta Zoologica Academiae Scientiarum Hungaricae, 60, 1–12. Motschulsky, V. de (1857) Énumération des nouvelles espèces de Coléoptères rapportés de ses voyages. Bulletin de la Société Impériale des Naturalistes de Moscou, 30, 490–517. Motschulsky, V. de (1862) Entomologie spéciale, Insectes du Japan. I. Coleopteres. Etudes Entomologiques. Helsingfors, 9, 4– 39. [1860] Müller, O.F. (1776) Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. Hallager, Havniae, xxxii + 282 pp. http://dx.doi.org/10.5962/bhl.title.13268 Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858. http://dx.doi.org/10.1038/35002501 Normand, H. (1947) Contribution au catalogue des Coléoptères de la Tunisie. Bulletin Société de Géographie et d’Archélogie de la Province d’Oran. Troisième supplément, fascicule 2, 66/67 (1945–1946), 1–16. Olivier, A.G. (1811) Encyclopédie Méthodique. Histoire naturelle. Insectes. vol. 8(2). H. Agasse, Paris, 362 pp. [pp. 361–722] R Core Team (2014) R: A language and environment for statistical computing. ver. 3.0.3. R Foundation for Statistical Computing, Vienna, Austria, Available from: http://www.R-project.org/ (accessed 9 June 2015) Scheerpeltz, O. (1965) Wissenschaftliche Ergebnisse der Schwedischen Expedition 1934 nach Indien und Burma. Coleoptera Staphylinidae (except. Megalopsidiinae et Steninae). (5. Beitrag zur Kenntnis der orientalischen Staphyliniden.). Arkiv för Zoologi Uppsala, 17, 93–371. Scheerpeltz, O. (1978) Wissenschaftliche Ergebnisse der von Dr. Raul do Nascimento Ferreira—Coimbra, Portugal—während seines Aufenthaltes vom Ende des Jahres 1964 bis zum Anfang des Jahres 1966 auf der Insel Timor, der östlichsten der Kleinen Sundainseln, durchgeführten Aufsammlungen von Staphyliniden (Coleoptera). (16. Beitrag zur Kenntnis der orientalischen Staphyliniden und 7. Beitrag zur Kenntnis der australisch-polynesischen Staphyliniden). EOS: revista española de entomología, 52, 185–233. Schülke, M. (2009) Zur Taxonomie und Faunistik westpalaearktischer Staphylinidae (Coleoptera: Staphylinidae: Omaliinae, Oxytelinae et Tachyporinae). [On the taxonomy and zoogeography of western Palaearctic Staphylinidae (Coleoptera: Staphylinidae: Omaliinae, Oxytelinae et Tachyporinae)]. Linzer Biologische Beiträge, 41, 803–844. Schülke, M. (2011) Zur Kenntnis der Verwandtschaft von Bledius (Hesperophilus) atricapillus (Germar) (Coleoptera, Staphylinidae: Oxytelinae). Linzer Biologische Beiträge, 43, 1595–1608. Schülke, M. (2012) Vier neue paläarktische Oxytelini (Coleoptera, Staphylinidae, Oxytelinae). Linzer Biologische Beiträge, 44, 1641–1666. Sharp, D.S. (1874) The Staphylinidae of Japan. Transactions of the Entomological Society of London, 1874, 1–103. http://dx.doi.org/10.1111/j.1365-2311.1874.tb00159.x Sodhi, N.S., Posa, M.R.C., Lee, T.M., Bickford, D., Koh, L.P. & Brook, B.W. (2010) The state and conservation of Southeast Asian biodiversity. Biodiversity and Conservation, 19, 317–328. http://dx.doi.org/10.1007/s10531-009-9607-5 Walker, F. (1859) Characters of some apparently undescribed Ceylon insects. Annals and Magazine of Natural History, 3, 50– 56.


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FIGURE 1. Map of Southeast Asia (area filled in red).



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FIGURE 2. Oxytelus armiger Fauvel, 1895 (1): A, head ♂; B, head ♀; C, right mandible ♂; D, right mandible ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, tergite X ♂; I, tergite X ♀; J, spermatheca; K, aedeagus (ventral view); L, aedeagus (dorsal view); M, median lobe of aedeagus (lateral view); N, right paramere (in situ) of aedeagus (lateral view). Scales: J = 0.1 mm; rest, 0.3 mm.


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FIGURE 3. Oxytelus armiger Fauvel, 1895 (2) (B, N—lectotype in IRSNB; A, C–M—paralectotypes in IRSNB): A, habitus ♂; B, head and pronotum ♂; C, forebody ♀; D, sternite VII ♂; E, sternite VIII ♂; F, sternite VIII ♀; G, aedeagus (ventral view); H, aedeagus (dorsal view); I, medianlobe of aedeagus (ventral view); J, medianlobe of aedeagus (lateral view); K, parameres (ventral view); L, right paramere (in situ) of aedeagus (lateral view); M, spermatheca, N, labels of the lectotype. Scales: A = 1 mm; B, C = 0.5 mm; M = 0.1 mm; rest, 0.3 mm.



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FIGURE 4. Oxytelus bengalensis Erichson, 1840: A, habitus ♂; B, habitus ♀; C, forebody ♂; D, forebody ♀. Scales: A, B = 1 mm; C, D = 0.5 mm.


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FIGURE 5. Oxytelus nigriceps Kraatz, 1859 (A, B), O. migrator Fauvel, 1904 (C, D), and O. incisus Motschulsky, 1857 (E, F): A, C, E, habitus ♂; B, D, F, head and pronotum ♂. Scales: 0.5 mm.



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FIGURE 6. Oxytelus javanus Cameron, 1936 (A–I, lectotype in BMNH) and O. bengalensis Erichson, 1840 (J): A, habitus ♂; B, forebody ♂; C, sternite VII ♂; D, J, sternite VIII ♂; E–G, aedeagus with right paramere removed (E—ventral view, F— dorsal view, G—lateral view); H, right paramere (in situ) of aedeagus (lateral view); I, labels of lectotype. Scales: A = 1 mm; B = 0.5 mm; rest, 0.3 mm.


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FIGURE 7. Oxytelus ruptus Fauvel, 1904 (1): A, head ♂; B, head ♀; C, right mandible ♂; D, right mandible ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, tergite X ♂; I, tergite X ♀; J, spermatheca with basal portion of duct; K, aedeagus (ventral view); L, aedeagus (dorsal view); M, median lobe of aedeagus (lateral view); N, right paramere (in situ) of aedeagus (lateral view). Scales: J = 0.1 mm; rest, 0.3 mm.



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FIGURE 8. Oxytelus ruptus Fauvel, 1904 (2): A, habitus ♂; B, habitus ♀; C, head and pronotum ♂; D, head and pronotum ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, spermatheca with duct; I, aedeagus (ventral view); J, aedeagus (dorsal view); K, median lobe of aedeagus (lateral view); L, right paramere (in situ) of aedeagus (lateral view). Scales: A, B = 1 mm; C, D = 0.5 mm; H = 0.1 mm; rest, 0.2 mm.


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FIGURE 9. Oxytelus piceus (Linné, 1767) (A–D), O. sublucidus Cameron, 1941 (syn. of O. ruptus, E–G, lectotype in BMNH), and O. varipennis Kraatz, 1859 (H–J, lectotype in SDEI): A, habitus ♂; B, E, H, habitus ♀; C, forebody ♂; D, F, I, forebody ♀; G, labels of lectotype; J, terminalia ♀ (ventral view). Scales: A, B, E, H = 1 mm; C, D, F, I = 0.5 mm; J = 0.2 mm.



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FIGURE 10. Oxytelus finitimus sp. nov. (1): A, head ♂; B, head ♀; C, right mandible ♂; D, right mandible ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, tergite X ♂; I, tergite X ♂; J, spermatheca; K, aedeagus (ventral view); L, aedeagus (dorsal view); M, median lobe of aedeagus (lateral view); N, right paramere (in situ) of aedeagus (lateral view). Scales: J = 0.1 mm; rest, 0.3 mm.


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FIGURE 11. Oxytelus finitimus sp. nov. (2) (A, C, M—holotype; B, D–L—paratypes): A, habitus ♂; B, habitus ♀; C, forebody ♂; D, forebody ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, spermatheca; I, aedeagus (ventral view); J, aedeagus (dorsal view); K, medianlobe of aedeagus (lateral view); L, right paramere (in situ) of aedeagus (lateral view); M, labels of holotype. Scales: A–D = 1 mm; H = 0.05 mm; rest, 0.2 mm.



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FIGURE 12. Oxytelus bellicosus Fauvel, 1895 (1) (A–D, F–I, K—lectotype in IRSNB; E, J—paralectotype in BMNH): A, habitus ♂; B, forebody ♂; C, sternite VII ♂; D, sternite VIII ♂; E, sternite VIII ♀; F–I, aedeagus (F—ventral view; G—dorsal view; H—ventro-lateral view; I—dorso-lateral view); J, spermatheca; K, labels of lectotype. Scales: A, B = 1 mm; J = 0.1 mm; rest, 0.3 mm.


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FIGURE 13. Oxytelus bellicosus Fauvel, 1895 (2) (sexual dimorphism and variation on head; B–E, paralectotypes in IRSNB; F, paralectotype in BMNH): A, B, head and pronotum ♂; C–E, forebody ♂; F, head and pronotum ♀. Scales: 1 mm.



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FIGURE 14. Oxytelus castaneus sp. nov. (1): A, head ♂; B, right mandible ♂; C, sternite VII ♂; D, sternite VIII ♂; E, aedeagus (ventral view); F, aedeagus (dorsal view); G, median lobe of aedeagus (lateral view); H, right paramere (in situ) of aedeagus (lateral view). Scales: A, B = 0.5 mm; rest, 0.3 mm.


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FIGURE 15. Oxytelus castaneus sp. nov. (2) (Holotype, ♂): A, habitus; B, forebody; C, aedeagus (ventral view); D, aedeagus (dorsal view); E, medianlobe of aedeagus (lateral view); F–G, right paramere (in situ) of aedeagus (F—lateral view, G— ventro-lateral view); H, labels. Scales: A, B = 2 mm; rest, 0.3 mm. OXYTELUS IN SOUTHEAST ASIA


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FIGURE 16. Oxytelus lividus Motschulsky, 1857 (A–I, M—lectotype and paralectotype of O. gigantulus Fauvel in IRSNB; J– L—aedeagus of O. lividus): A, habitus ♂ (lectotype); B, forebody ♂ (lectorype); C, habitus ♀ (paralectotype); D, head and pronotum ♀ (paralectotype); E, terminalia ♂ (ventral view, lectotype); F, sternite VII ♂ (lectotype); G, sternite VII ♂ (lectotype); H, aedeagus (ventral view, lectotype); I, aedeagus (dorsal view, lectotype); J, median lobe of aedeagus (lateral view); K, right paramere (in situ) of aedeagus (lateral view); L, left paramere (in situ) of aedeagus (lateral view); M, labels of lectotype. Scales: A–D = 2 mm; rest, 0.3 mm.


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FIGURE 17. Oxytelus insulanus sp. nov. (A–E) and O. sublividus sp. nov. (F–J) (1): A, F, habitus ♂ (holotype); B, G, forebody ♂ (holotype); C, H, habitus ♀ (paratype); D, I, forebody ♀ (paratype); E, J, labels of holotype. Scales: 1 mm.



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FIGURE 18. Oxytelus insulanus sp. nov. (A, B) and O. sublividus sp. nov. (C, D) (2): A, C, sternite VII ♂ (paratype); B, D, sternite VIII ♂ (paratype). Scales: 0.3 mm.


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FIGURE 19. Oxytelus insulanus sp. nov. (A–F) and O. sublividus sp. nov. (G–K) (3): A, G, aedeagus (ventral view, paratype); B, H, aedeagus (dorsal view, paratype); C, I, median lobe of aedeagus (lateral view, paratype); D, J, left paramere (in situ) of aedeagus (lateral view, paratype); E, K, right paramere (in situ) of aedeagus (lateral view, paratype); F, spermatheca. Scales: F = 0.05 mm; rest, 0.3 mm. OXYTELUS IN SOUTHEAST ASIA


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FIGURE 20. Oxytelus punctipennis Fauvel, 1905 (A–D—holotype in IRSNB) and O. discalis Cameron 1930 (E–G— lectotype in BMNH): A, E, habitus ♂; B, F, head and pronotum ♂; C, G, terminalia ♂ (ventral view); D, H, type labels. Scales: A, E = 1 mm; B, F = 0.5 mm; C, G = 0.2 mm.


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FIGURE 21. Oxytelus ferreirai Scheerpeltz, 1978 (A, C, E–K—holotype in NHMW; B, D—paratype in NHMW): A, habitus ♂; B, habitus ♀; C, forebody ♂; D, forebody ♀; E, sternite VII ♂; F, sternite VIII ♂; G, aedeagus (ventral view); H, aedeagus (dorsal view); I, medianlobe of aedeagus (lateral view); J, right paramere (in situ) of aedeagus (lateral view); K, labels of holotype. Scales: A, B = 1 mm; C, D = 0.5 mm; rest, 0.3 mm.



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FIGURE 22. Oxytelus lucens Bernhauer, 1903 (1) (A, C, E—lectotype in FMNH; B, D—paralectotype in FMNH; F–H— holotype of O. malaisei Scheerpeltz in SMNH): A, habitus ♂; B, habitus ♀; C, head and pronotum ♂; D, head and pronotum ♀; E, labels of lectotype; F, habitus ♂; G, forebody ♂; H, labels of holotype. Scales: A, B, F = 1 mm; C, D, G = 0.5 mm.


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FIGURE 23. Oxytelus lucens Bernhauer, 1903 (2) (A, B—lectotype in FMNH; C–G—holotype of O. malaisei Scheerpeltz in SMNH): A, sternite VIII ♂; B, aedeagus; C, sternite VII ♂; D, sternite VIII ♂; E, aedeagus (ventral view); F, aedeagus (dorsal view); G, aedeagus (lateral view). Scales: 0.3 mm.



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FIGURE 24. Oxytelus lompobatangensis sp. nov. (A–E) and O. poecilopterus sp. nov. (F–J) (1): A, F, habitus ♂ (holotype); B, G, forebody ♂ (holotype); C, H, habitus ♀ (paratype); D, I, forebody ♀ (paratype); E, J, labels of holotype. Scales: A, C, F, H = 1 mm; B, D, G, I = 0.5 mm.


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FIGURE 25. Oxytelus lompobatangensis sp. nov. (A–D, paratypes) and O. poecilopterus sp. nov. (E–H, paratypes) (2): A, E, sternite VII ♂; B, F, sternite VIII ♂; C, G, sternite VIII ♀; D, H, spermatheca. Scales: D, H = 0.1 mm; rest, 0.3 mm.



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FIGURE 26. Oxytelus lompobatangensis sp. nov. (A–D, paratype) and O. poecilopterus sp. nov. (E–H, paratype) (3): A, E, aedeagus (ventral view); B, F, aedeagus (dorsal view); C, G, median lobe of aedeagus (lateral view); D, H, right paramere (in situ) of aedeagus (lateral view). Scales: 0.3 mm.


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FIGURE 27. Oxytelus barbatus Fauvel, 1905 (A, C, F, G, J–N—holotype in IRSNB): A, habitus ♂; B, habitus ♀; C, forebody ♂; D, head and pronotum ♂ (“minor”); E, head and pronotum ♀; F, sternite VII ♂; G, sternite VIII ♂; H, sternite VIII ♀; I, spermatheca; J–M, aedeagus (J—ventral view, K—dorsal view, L—ventro-lateral view, M—dorso-lateral view); N, labels of holotype. Scales: A–E = 0.5 mm; I = 0.05mm; rest, 0.2 mm.



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FIGURE 28. Oxytelus mandibularis Cameron, 1929 (A–C, F–G, J–N—lectotype in BMNH; D, E, H, I—paralectotype in BMNH): A, forebody ♂; B, abdomen ♂; C, head ♂; D, habitus ♀; E, forebody ♀; F, sternite VII ♂; G, sternite VIII ♂; H, sternite VIII ♀; I, spermatheca; J–L, median lobe of aedeagus (J—ventral view, K—dorsal view, L—lateral view); M, right paramere (in situ) of aedeagus (lateral view); N, labels of lectotype. Scales: A, B, D = 0.5 mm; I = 0.05 mm; rest, 0.2 mm.


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FIGURE 29. Oxytelus puncticeps Kraatz, 1859 (1): A, habitus ♂; B, C, head and pronotum ♂ (B, normal male; C, “minor” male); D, ventral terminalia of the “minor” male. Scales: D = 0.2 mm; rest, 0.5 mm.



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FIGURE 30. Oxytelus puncticeps Kraatz, 1859 (2) (A, D–L—holotype of O. micantoides Scheerpeltz in NHMW; B, C, M— paratype of O. micantoides Scheerpeltz in NHMW): A, forebody ♂; B, habitus ♀; C, forebody ♀; D, abdominal segment VII ♂ (showing posterior margin of sternite VII); E, sternite VII ♂; F, sternite VIII ♂; G, aedeagus (ventral view); H, aedeagus (dorsal view); I, median lobe of aedeagus (ventro-lateral view); J, median lobe of aedeagus (lateral view); K, right paramere (in situ) of aedeagus (lateral view); L, M, type labels. Scales: A–C = 0.5 mm; rest, 0.2 mm.


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FIGURE 31. Oxytelus lucidulus Cameron, 1929 (A–D—lectotype in BMNH; E, G—lectotype of O. ginyuenensis Bernhauer in FMNH; F—paralectotype of O. ginyuenensis Bernhauer in FMNH): A, habitus ♂; B, head and pronotum ♂; C, terminalia ♂; D, G, labels of lectotype; E, forebody ♂; F, head and pronotum ♀. Scales: C = 0.2 mm; rest, 0.5 mm.



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FIGURE 32. Oxytelus subferrugineus Cameron, 1929 (1) (C–E—lectotype of O. subferrugineus in BMNH; F–K—lectotype of O. nilgiriensis Cameron in BMNH): A, habitus ♂; B, F, forebody ♂; C, habitus ♀; D, forebody ♀; E, K, labels of lectotype; G, sternite VII ♂; H, sternite VIII ♂; I, median lobe of aedeagus (ventro-lateral view); J, parameres of aedeagus (ventral view). Scales: G–J = 0.2 mm; rest, 0.5 mm.


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FIGURE 33. Oxytelus subferrugineus Cameron, 1929 (2) (A, B, E–H, J—lectotype of O. kedirianus Cameron in BMNH; C, D, I, K—paralectotype of O. kedirianus Cameron in RMNH): A, habitus ♂; B, forebody ♂; C, habitus ♀; D, head and pronotum ♀; E, abdominal segment VII ♂ (showing posterior margin of sternite VII); F, sternite VIII ♂; G, median lobe of aedeagus (lateral view); H, parameres of aedeagus (ventral view); I, metathorax and abdomen ♀ (ventral view); J, K, type labels. Scales: A, B = 0.5 mm; rest, 0.2 mm. [Photos C, D, I, K © RMNH, Dr. Hans Huijbregts, used by permission]



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FIGURE 34. Oxytelus megaceros Fauvel, 1895 (1) (A, C, E–K—lectotype in IRSNB; B, D—paralectotype in IRSNB): A, habitus ♂; B, habitus ♀; C, forebody ♂; D, forebody ♀; E, sternite VII ♂; F, sternite VIII ♂; G–J, aedeagus (G—ventral view, H—dorsal view, I—ventro-lateral view, J—dorso-lateral view); K, labels of lectotype. Scales: A–D = 0.5 mm; rest, 0.2 mm.


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FIGURE 35. Oxytelus megaceros Fauvel, 1895 (2) (lectotype of O. kalisi Bernhauer in FMNH): A, habitus ♀; B, forebody ♀; C, right antenna ♀; D, labels of lectotype. Scales: 1 mm.



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FIGURE 36. Oxytelus fallax Fauvel, 1878 (holotype in MSNG): A, habitus ♂; B, forebody ♂; C, sternite VII ♂; D, sternite VIII ♂; E, aedeagus (ventral view); F, aedeagus (dorsal view); G, median lobe of aedeagus (lateral view); H, right paramere (in situ) of aedeagus (lateral view); I, type labels. Scales: A, B = 0.5 mm; rest, 0.2 mm.


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FIGURE 37. Oxytelus grandiculus sp. nov. (1): A, head ♂; B, right mandible ♂; C, tergite VII ♂; D, tergite VIII ♂; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♂; H, tergite X ♂; I, tergite X ♀; J, spermatheca; K, aedeagus (ventral view); L, aedeagus (dorsal view); M, median lobe of aedeagus (lateral view); N, right paramere (in situ) of aedeagus (lateral view). Scales: J = 0.1 mm; rest, 0.3 mm.



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FIGURE 38. Oxytelus grandiculus sp. nov. (2) (A, C, N—holotype; B, D–M—paratypes): A, habitus ♂ (holotype); B, habitus ♀; C, forebody ♂ (holotype); D, forebody ♀; E, sternite VII ♂; F, tergite VIII ♂; G, sternite VIII ♂; H, sternite VIII ♀; I, spermatheca; J, aedeagus (ventral view); K, aedeagus (dorsal view); L, medianlobe of aedeagus (lateral view); M, right paramere (in situ) of aedeagus (lateral view); N, labels of holotype. Scales: A–D = 1 mm; I = 0.05 mm; rest, 0.3 mm.


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FIGURE 39. Oxytelus subsculptus Cameron, 1928 (1): A, head ♂; B, right mandible ♂; C, right metatibia (ventral view); D, sternite VII ♂; E, sternite VIII ♂; F, sternite VIII ♀; G, tergite X ♂; H, tergite X ♀; I, spermatheca; J, aedeagus (ventral view); K, aedeagus (dorsal view); L, median lobe of aedeagus (lateral view); M, right paramere (in situ) of aedeagus (lateral view). Scales: I = 0.1 mm; rest, 0.2 mm.



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FIGURE 40. Oxytelus subsculptus Cameron, 1928 (2) (C, D, N—lectotype in BMNH): A, habitus ♂; B, forebody ♂; C, habitus ♀; D, forebody ♀; E, right hind leg ♂; F, sternite VII ♂; G, sternite VIII ♂; H, sternite VIII ♀; I, spermatheca; J, aedeagus (ventral view, left paramere removed); K, aedeagus (dorsal view, left paramere removed); L, median lobe of aedeagus (lateral view); M, right paramere (in situ) of aedeagus (lateral view). Scales: A–D = 0.5 mm; I = 0.05 mm; rest, 0.2 mm.


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FIGURE 41. Oxytelus subincisus Cameron, 1941 (1) (A–C, N—lectotype in BMNH): A, habitus ♀; B, forebody ♀; C, terminalia ♀; D, right hind leg ♂; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, spermatheca; I, median lobe of aedeagus (ventral view); J, median lobe of aedeagus (dorsal view); K, median lobe of aedeagus (lateral view); L, parameres of aedeagus (ventral view); M, right paramere (in situ) of aedeagus (lateral view); N, labels of lectotype. Scales: A, B = 0.5 mm; H = 0.05 mm; rest, 0.2 mm.



79

FIGURE 42. Oxytelus subincisus Cameron, 1941 (2) (holotype of Oxytelus fruhstorferi Cameron, BMNH): A, habitus ♀; B, forebody ♀; C, labels of holotype. Scales: 0.5 mm.


LÜ & ZHOU

FIGURE 43. Anotylus transversipennis (Scheerpeltz, 1978) (A, B, E–K—holotype in NHMW; C, D, L—paratype in NHMW): A, habitus ♂; B, forebody ♂; C, habitus ♂; D, forebody ♂; E, sternite VII ♂; F, sternite VIII ♂; G–J, aedeagus (G— ventral view, H—dorsal view, I—ventro-lateral view, J—dorso-lateral view); K, L, type labels. Scales: A–D = 0.5 mm; rest, 0.2 mm. OXYTELUS IN SOUTHEAST ASIA


81

FIGURE 44. Oxytelus abiturus Lü & Zhou, 2012 (A, B—holotype in IZ-CAS; C–N—paratypes in IZ-CAS): A, habitus ♂; B, head and pronotum ♂; C, habitus ♀; D, head and pronotum ♀; E, right mandible ♂; F, right mandible ♀; G, sternite VII ♂; H, sternite VIII ♂; I, sternite VIII ♀; J, spermatheca; K, aedeagus (ventral view); L, aedeagus (dorsal view); M, median lobe of aedeagus (lateral view); N, right paramere (in situ) of aedeagus (lateral view). Scales: A–D = 0.5 mm; J = 0.05 mm; rest, 0.2 mm.


LÜ & ZHOU

FIGURE 45. Oxytelus ailaoshanicus Lü & Zhou, 2012 (A, B—holotype in IZ-CAS; C–J—paratypes in IZ-CAS): A, habitus ♂; B, head and pronotum ♂; C, habitus ♀; D, head and pronotum ♀; E, aedeagus (ventral view); F, aedeagus (dorsal view); G, median lobe of aedeagus (ventral view); H, median lobe of aedeagus (lateral view); I, left paramere (in situ) of aedeagus (lateral view), J, right paramere (in situ) of aedeagus (lateral view). Scales: A–D = 0.5 mm; E–J, 0.2 mm.



83

FIGURE 46. Oxytelus antennalis Fauvel, 1889 (1) (A, B, E, F, J–O—lectotype in IRSNB; C, D, G, I—paralectotype in IRSNB): A, habitus ♂; B, forebody ♂; C, habitus ♀; D, forebody; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, terminalia (♂, ventral view); I, spermatheca; J, aedeagus (ventral view); K, aedeagus (dorsal view); L, medianlobe of aedeagus (ventro-lateral view); M, medianlobe of aedeagus (lateral view); N, right paramere (in situ) of aedeagus (lateral view); O, labels of lectotype. Scales: A–D = 0.5 mm; I = 0.05 mm; rest, 0.2 mm.


LÜ & ZHOU

FIGURE 47. Oxytelus antennalis Fauvel, 1889 (2) (A, B, E, F, H, J–O—lectotype of O. cheesmani Bernhauer in BMNH; C, D, G, I—paralectotype of O. cheesmani Bernhauer in BMNH): A, habitus ♂; B, forebody ♂; C, habitus ♀; D, forebody ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, terminalia ♂ (ventral view); I, spermatheca; J–M, aedeagus (J— ventral view, K—dorsal view, L—ventro-lateral view, M—lateral view); N, right paramere (in situ) of aedeagus; O, labels of lectotype. Scales: A–D = 0.5 mm; I = 0.05 mm; rest, 0.2 mm.



85

FIGURE 48. Oxytelus bajiei Lü & Zhou, 2012 (paratypes in IZ-CAS): A, habitus ♂; B, forebody ♂; C, habitus ♀; D, forebody ♀; E, terminalia (ventral view) ♂; F, terminalia (ventral view) ♀; G, aedeagus (ventral view); H, aedeagus (dorsal view); I, median lobe of aedeagus (lateral view); J, right paramere (in situ) of aedeagus (lateral view). Scales: A–D = 1 mm; E–J = 0.5 mm.


LÜ & ZHOU

FIGURE 49. Oxytelus cheesmanianus Cameron, 1937 (lectotype in BMNH): A, habitus ♂ (with abdomen reversely glued and segments VIII–X removed); B, forebody ♂; C, abdominal segment VII ♂; D, sternite VIII ♂; E–G, median lobe of aedeagus (E—ventral view, F—dorsal view, G—lateral view); H, right paramere (in situ) of aedeagus (lateral view); I, labels of lectotype. Scales: A, B = 1 mm; rest, 0.3 mm.



87

FIGURE 50. Oxytelus hingstoni Cameron, 1928 (lectotype in BMNH): A, habitus ♀; B, forebody ♀; C, sternite VIII ♀; D, spermatheca; E, labels of lectotype. Scales: A, B = 0.5 mm; C = 0.2 mm; D = 0.1 mm.


LÜ & ZHOU

FIGURE 51. Oxytelus pallidipennis Cameron, 1930 (syntypes in FMNH): A, habitus ♂; B, habitus ♀; C, head and pronotum ♂; D, head and pronotum ♀; E, right mandible ♂; F, right mandible ♀; G, sternite VII ♂; H, sternite VIII ♂; I, sternite VIII ♀; J, spermatheca; K, aedeagus (ventral view); L, aedeagus (dorsal view); M–P, median lobe of aedeagus (M, P—lateral view, N—ventral view, O—dorsal view); Q, parameres of aedeagus (ventral view); R, right paramere (in situ) of aedeagus (lateral view). Scales: A–D = 0.5 mm; J = 0.05 mm; rest, 0.2 mm.



89

FIGURE 52. Oxytelus sculptus Gravenhorst, 1806: A, habitus ♂; B, habitus ♀; C, head and pronotum ♂; D, head and pronotum ♀; E, sternite VII ♂; F, sternite VIII ♂; G, sternite VIII ♀; H, spermatheca; I–K, aedeagus (I—ventral view, J— dorsal view, K—ventro-lateral view); L, median lobe of aedeagus (lateral view); M, right paramere (in situ) of aedeagus. Scales: A–D = 0.5 mm; H = 0.05 mm; rest, 0.2 mm.


LÜ & ZHOU

FIGURE 53. Oxytelus solus Lü & Zhou, 2012 (holotype ♂, in IZ-CAS): A, habitus; B, head and pronotum; C, right mandible; D, terminalia (ventral view); E, sternite VII; F, sternite VIII ♂; G–L, aedeagus (G—ventral view, H—dorsal view); I–K, median lobe of aedeagus (I—apical view, J—lateral view, K—ventro-lateral view); L, right paramere (in situ) of aedeagus (lateral view). Scales: A, B = 1 mm; C, D = 0.5 mm; rest, 0.3 mm.



91

FIGURE 54. Oxytelus tibetanus Bernhauer, 1933 (lectotype in FMNH): A, forebody ♂; B, sternite VII ♂; C, sternite VIII ♂; D, median lobe of aedeagus (dorsal view); E, left paramere (in situ) of aedeagus (medial side in lateral view); F, labels of lectotype. Scales: A = 0.5 mm; rest, 0.2 mm.


LÜ & ZHOU

FIGURE 55. Oxytelus validus Cameron, 1952 (paratype in BMNH): A, habitus ♂ (with abdominal segments VIII–X removed); B, forebody ♂; C, abdominal segment VII ♂; D, sternite VIII ♂; E–G, median lobe of aedeagus (E—ventral view, F—dorsal view, G—lateral view); H, right paramere (in situ) of aedeagus (lateral view). Scales: A, B = 1 mm; rest, 0.3 mm.



93


LÜ & ZHOU

0.81 (1)

1.03 (1) 0.80 (1)

1.05 (1) 0.69 (1)

0.25 (1) 0.20 (1)

0.27 (1) 0.25 (1)

0.35 (1) 0.15 (1)

0.77 (1) 1.67 (1)

2.53 (1)

0.73 (1)

1.02 (1)

0.90 (1)

0.29 (1)

0.28 (1)

0.16 (1)

1.75 (1)

1.68 (1)

0.44 (1)

0.68 (1)

0.49 (1)

0.12 (1)

0.29 (1)

0.03 (1)

11.00 (1)

1.01 (1) 0.87 (1)

0.76 (1) 0.78 (1)

0.96 (1) 0.87 (1)

0.63 (1) 0.59 (1)

1.20 (1) 1.13 (1)

1.15 (1)

0.70 (1)

1.03 (1)

0.76 (1)

1.30 (1)

0.72 (1)

0.74 (1)

0.69 (1)

0.46 (1)

0.87 (1)

0.70 (1)

0.97 (1)

0.86 (1)

0.17 (1)

0.37 (1)

0.13 (1)

2.75 (1)

4.60 (1)

♂

2.20 (1)

7.11±0.68(7) 3.35±0.11(7) 0.96±0.04(7) 1.32±0.05(7) 1.11±0.04(7) 0.32±0.01(7) 0.47±0.02(7) 0.13±0.01(7) 3.59 ± 0.20(7)

♀

♂

4.16 (1)

2.48 (1)

1.48 (1)

1.31 (1) 0.37 (1)

0.69 (1) 0.52 (1)

0.67 (1) 0.46 (1)

0.98 (1) 0.13 (1)

0.21 (1) 0.18 (1)

0.12 (1) 0.07 (1)

1.75 (1)

0.57 (1)

1.08 (1)

0.75 (1) 0.63 (1)

0.77 (1) 0.78 (1)

0.65 (1) 0.60 (1)

0.44 (1) 0.38 (1)

0.72 (1) 0.61 (1)

3.74±0.2(6)

6.73±0.57(8) 3.50±0.24(8) 1.22±0.16(8) 1.52±0.16(8) 1.46±0.21(8) 0.34±0.03(8) 0.51±0.05(8) 0.31±0.07(8) 1.72 ± 0.46(8)

6.19±0.09(2) 3.38±0.00(2) 0.98±0.00(2) 1.32±0.00(2) 1.10±0.00(2) 0.32±0.00(2) 0.46±0.01(2) 0.15±0.01(2) 3.21 ± 0.11(2)

3.44±0.15(4) 1.75±0.05(4) 0.43±0.02(4) 0.64±0.01(4) 0.43±0.01(4) 0.13±0.01(4) 0.30±0.01(4) 0.01±0.00(4) 53.00 ±29.42(4) 0.50±0.01(4) 0.67±0.01(4) 0.75±0.02(4) 0.69±0.01(4) 0.43±0.01(4) 0.80±0.02(4)

3.77±0.10(5) 1.86±0.06(5) 0.44±0.01(5) 0.68±0.02(5) 0.41±0.01(5) 0.14±0.01(5) 0.30±0.01(5) 0.00 (5)

♂

♀

♂

♀

---

0.53±0.01(5) 0.71±0.01(5) 0.75±0.02(5) 0.72±0.02(5) 0.46±0.01(5) 0.91±0.03(5)

1.03±0.01(2) 1.49±0.02(2) 0.69±0.00(2) 1.16±0.31(2) 1.15±0.28(2) 1.75±0.00(2)

1.05±0.06(8) 1.57±0.13(8) 0.67±0.03(8) 1.34±0.08(8) 0.92±0.04(8) 1.81±0.09(8)

1.89±0.09(6) 0.47±0.03(6) 0.76±0.04(6) 0.57±0.06(6) 0.15±0.01(6) 0.30±0.02(6) 0.06±0.09(6) 11.65 ± 6.27(6) 0.62±0.03(6) 0.86±0.04(6) 0.72±0.01(6) 0.79±0.03(6) 0.53±0.02(6) 0.99±0.05(6)

0.63±0.03(6) 0.92±0.06(6) 0.69±0.01(6) 0.77±0.07(6) 0.54±0.03(6) 0.95±0.06(6)

4.00±0.29(6) 2.08±0.11(6) 0.54±0.08(6) 0.83±0.09(6) 0.71±0.14(6) 0.17±0.01(6) 0.31±0.02(6) 0.08±0.05(6) 6.07 ± 4.07(6)

♀

0.72±0.03(4) 0.97±0.05(4) 0.74±0.02(4) 0.90±0.05(4) 0.67±0.09(4) 1.19±0.13(4)

0.79±0.08(6) 1.13±0.11(6) 0.70±0.02(6) 0.97±0.09(6) 0.72±0.13(6) 1.17±0.14(6)

0.49 (1)

0.58 (1)

0.90±0.00(2) 1.22±0.01(2) 0.74±0.01(2) 1.17±0.02(2) 0.83±0.01(2) 1.52±0.05(2)

♂

0.73 ± 0.10(6)

2.57 (1)

0.88 (1)

4.34±0.25(4) 2.33±0.14(4) 0.63±0.04(4) 0.83±0.04(4) 0.74±0.06(4) 0.20±0.00(4) 0.25±0.01(4) 0.15±0.01(4) 1.68 ± 0.18(4)

4.77±0.69(6) 2.88±0.30(6) 1.01±0.18(6) 1.08±0.12(6) 1.15±0.19(6) 0.34±0.06(6) 0.28±0.04(6) 0.39±0.1(6)

2.68 (1)

0.68 (1)

♀

♂

♀

1.00 (1)

0.93±0.05(2) 1.38±0.06(2) 0.68±0.01(2) 1.23±0.08(2) 0.88±0.04(2) 1.48±0.07(2)

0.68 (1)

1.04±0.03(7) 1.47±0.05(7) 0.71±0.01(7) 1.15±0.24(7) 1.09±0.20(7) 1.78±0.04(7)

1.04±0.09(4) 1.63±0.15(4) 0.64±0.02(4) 1.32±0.11(4) 0.94±0.07(4) 1.77±0.13(4)

4.60±0.40(6) 2.21±0.08(6) 0.50±0.02(6) 0.78±0.02(6) 0.54±0.02(6) 0.19±0.01(6) 0.32±0.01(6) 0.02±0.00(6) 19.78 ± 3.86(6) 0.66±0.02(6) 0.86±0.03(6) 0.77±0.01(6) 0.90±0.02(6) 0.58±0.02(6) 1.12±0.04(6)

7.54±0.78(4) 3.62±0.41(4) 1.24±0.24(4) 1.57±0.25(4) 1.54±0.34(4) 0.34±0.03(4) 0.52±0.06(4) 0.34±0.13(4) 1.75 ± 0.78(4)

♀

0.69±0.01(6) 0.98±0.02(6) 0.71±0.02(6) 0.93±0.02(6) 0.63±0.01(6) 1.19±0.02(6)

0.71±0.02(6) 1.06±0.04(6) 0.67±0.01(6) 0.95±0.02(6) 0.64±0.03(6) 1.18±0.07(6)

0.54 (1)

0.92±0.04(2) 1.44±0.06(2) 0.64±0.00(2) 1.19±0.05(2) 0.83±0.06(2) 1.56±0.05(2)

0.80 (1)

0.86±0.12(5) 1.30±0.23(5) 0.66±0.03(5) 1.09±0.14(5) 0.83±0.18(5) 1.45±0.12(5)

0.68 (1)

0.77 (1)

0.87±0.07(3) 1.15±0.08(3) 0.76±0.01(3) 0.96±0.07(3) 0.67±0.05(3) 1.23±0.09(3)

Abbreviation: BL—body length; FBL—forebody length; HL—head length; HWE—head width at eyes; HWT—head width at temples; CL—clypeal length; EL—eye length; TL—temple length; ETR—Ratio of EL at TL; PNL—pronotal length; PNW—pronotal width; PNR—Ratio of PNL at PNW; ELA—elytra length (average); EWA—elytra length (average); ABDW—abdominal width.

subsculptus

sublividus

ABDW

♂

5.62±0.44(2) 2.91±0.13(2) 0.75±0.01(2) 1.01±0.01(2) 0.92±0.00(2) 0.27±0.00(2) 0.29±0.01(2) 0.17±0.00(2) 1.65 ± 0.07(2)

ruptus

EWA

5.19±0.67(6) 2.32±0.12(6) 0.50±0.01(6) 0.81±0.03(6) 0.56±0.01(6) 0.18±0.01(6) 0.31±0.06(6) 0.02±0.01(6) 18.19 ± 4.05(6) 0.70±0.03(6) 0.90±0.04(6) 0.78±0.01(6) 0.93±0.04(6) 0.61±0.03(6) 1.11±0.09(6)

5.88±0.71(2) 3.38±0.09(2) 1.18±0.05(2) 1.30±0.07(2) 1.35±0.05(2) 0.39±0.01(2) 0.33±0.01(2) 0.44±0.04(2) 0.77 ± 0.04(2)

poecilopterus

ELA

4.26±0.21(6) 2.28±0.03(6) 0.58±0.01(6) 0.88±0.01(6) 0.75±0.02(6) 0.21±0.01(6) 0.30±0.00(6) 0.09±0.00(6) 3.30 ± 0.14(6)

♀

mandibularis

PNR

♂

4.53±0.37(6) 2.48±0.12(6) 0.72±0.06(6) 1.00±0.05(6) 0.91±0.06(6) 0.27±0.02(6) 0.31±0.01(6) 0.18±0.04(6) 1.75 ± 0.31(6)

3.45 (1)

6.83±0.14(2) 3.38±0.16(2) 1.20±0.07(2) 1.41±0.10(2) 1.48±0.11(2) 0.46±0.01(2) 0.33±0.01(2) 0.43±0.03(2) 0.77 ± 0.02(2)

4.93 (1)

5.83±0.89(5) 2.97±0.47(5) 1.07±0.29(5) 1.24±0.25(5) 1.26±0.33(5) 0.41±0.14(5) 0.31±0.03(5) 0.37±0.15(5) 0.92 ± 0.28(5)

0.60 (1)

PNW

0.96±0.06(2) 1.36±0.11(2) 0.71±0.01(2) 1.06±0.01(2) 0.76±0.04(2) 1.34±0.04(2)

PNL

♀

♂

♂

♂

♀

♂

2.10 (1)

2.60 (1)

4.30 (1)

ETR

5.00 (1)

TL

♂

EL

♀

CL

5.48±0.65(3) 2.69±0.21(3) 0.82±0.07(3) 0.98±0.08(3) 0.92±0.08(3) 0.22±0.02(3) 0.24±0.02(3) 0.22±0.02(3) 1.12 ± 0.03(3)

HWT

5.94±0.24(2) 3.09±0.00(2) 1.11±0.05(2) 1.21±0.06(2) 1.22±0.10(2) 0.27±0.01(2) 0.27±0.01(2) 0.39±0.03(2) 0.70 ± 0.03(2)

HWE

♀

HL

♂

FBL

lompobatangensis ♂

javanus

insulanus

grandiculus

finitimus

falax

castaneus

bellicosus

barbatus

armiger

BL

TABLE 1. Measurements (in millimeters) of adults of some Oxytelus species from Southeast Asia (Mean ± Standard deviation (Sample size))