Revision of the Asian Tribe Megarthropsini (Coleoptera: Staphylinidae: Tachyporinae)

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3430, 72 pp., 211 ®gures, 2 tables February 27, 2004

LEE H. HERMAN1

CONTENTS Abstract ...... 2 Introduction ...... 2 Abbreviations ...... 2 Megarthropsini ...... 3 Key to Genera of the Megarthropsini ...... 7 Megarthropsis ...... 7 Key to Species of Megarthropsis ...... 8 Description of Species ...... 10 Nepaliodes ...... 23 Key to Species of Nepaliodes ...... 28 Description of Species ...... 29 Peitawopsis ...... 30 Key to Species of Peitawopsis ...... 34 Description of Species ...... 34 Lacvietina, new genus ...... 39 Key to Species of Lacvietina ...... 43 Description of Species ...... 44 Discussion ...... 54 Acknowledgments ...... 67 References ...... 67 Appendix: Paratachinus laticollis Cameron ...... 69 Addendum ...... 72

1 Division of Invertebrate Zoology American Museum of Natural History. e-mail: [email protected]

ABSTRACT The species and genera of the Megarthropsini are revised. Descriptions and illustrations of the genera and species and keys for identi®cation are presented. Lacvietina, a new genus, with four new species is described. Three of the new species, L. aurora, L. copiosa, and L. cuprina, are from Vietnam, and the fourth, L. paricosta, is from Malaysia and Thailand. Lacvietina punctatissima (Hayashi), from Taiwan, is a new combination and was transferred from Tach- inus. A new species of Nepaliodes, N. solangelae from Thailand and China, is described. Seven new species of Megarthropsis are described; six of them, M. deverra, M. durga, M. empusa, M. frazerensis, M. parca, and M. smetanai, are from Malaysia; the seventh, M. djawaensis, is from Indonesia. A phylogenetic analysis supports Megarthropsini as a monophyletic tribe with Deropini as its sister group. Within Megarthropsini, Peitawopsis is the most basal genus to the remaining three genera, and Lacvietina is the sister group to Megarthropsis and Nepaliodes. The tribe is most readily distinguished from other taxa of the Tachyporinae by the densely and coarsely punctate body. Finally, the type species of Paratachinus, P. laticollis Cameron, is redescribed and discussed. Lacvietina takashii is a new combination.

INTRODUCTION tribe in the Tachyporinae. In the same article Smetana extended the known range of Me- This revision was prompted by the col- garthropsis from Singapore to Borneo, and lection of some odd specimens of unknown extended that of Nepaliodes from central subfamilial assignment from the mountains Nepal (Bagmati Province) to Uttar Pradesh of western Vietnam in 1998 and 1999. and West Bengal in India. The geographic They turned out to be three undescribed range of the tribe was extended and the an- species of a new genus of the rather anom- atomical diversity increased when Smetana alous Asian tribe of the Tachyporinae, the (1992) added the new monotypic, Taiwa- Megarthropsini. Understanding the position nese genus Peitawopsis; in the same article of the Vietnamese species required exami- Smetana further illustrated Megarthropsis. nation of the other genera of the tribe. This More recently a second (Smetana, 1995a) process led to the discovery of nine other and third species (Herman and Smetana, unnamed species distributed among the 2002) of Peitawopsis, both from Taiwan, other three genera, which until now includ- were named. ed a total of only four species in three gen- The present article describes new taxa, era. presents new distributional data and char- Megarthropsini was proposed by Cam- acters, and discusses the relationships of the eron (1919: 231), who, while providing no Megarthropsini and its components. The characters speci®c to the tribe, presented a treatment of Peitawopsis herein is an expan- detailed description of his new monotypic sion of a review by Herman and Smetana genus from Singapore, Megarthropsis. (2002). The descriptions of Peitawopsis and When the tribe was published it was not P. watanabei, the locality data, and the key assigned to a subfamily, but Cameron to species are included so that this revision- (1919: 232) stated that it appeared to be ary study stands alone and presents a com- related to the ``Trichophyini'' and ``Tach- prehensive and coherent treatment of the yporini''. Later it was placed in the Tach- tribe and its species. yporinae (Cameron, 1921: 349). The tribe included only Megarthropsis, ABBREVIATIONS and neither the tribe nor genus was cited again except in catalogs (Scheerpeltz, 1933: AMNH American Museum of Natural History, 1478; Blackwelder, 1952: 236) until 1983. New York, New York ASC A. Smetana collection, Ottawa, Ontar- Smetana (1983a) moved the monotypic io, Canada Nepalese genus Nepaliodes Coiffait to the CNC Canadian National Collection of In- Megarthropsini, provided detailed descrip- sects, Ottawa, Ontario, Canada tions and numerous illustrations for both GRC G. de Rougemont collection, Londini- genera, and discussed the position of the eÁres, France 2004 HERMAN: MEGARTHROPSINI 3

IEBR Institute of Ecology and Biological Re- sources, Hanoi, Vietnam MHNG MuseÂum d'Histoire Naturelle, GeneÁve, Switzerland MSC Michael SchuÈlke collection, Berlin, Germany NHMW Naturhistorisches Museum Wien, Wien, Austria

MEGARTHROPSINI Megarthropsini Cameron, 1919: 231. Type genus: Megarthropsis, ®xed by monotypy. ÐScheerpeltz, 1933: 1478 (world catalog supple- ment).ÐSmetana, 1983a: 142 (characters; discussion; key to genera).ÐSmetana, 1983b: 274 (tribe of Tachyporinae; key to tribes; characters).ÐNewton and Thayer, 1992: 66 (tribe of Tachyporinae).ÐHerman, 2001a: 678 (catalog).

DIAGNOSIS: Megarthropsini is distinguished from all tribes of the Tachyporinae by the dense, coarse punctation of the head, pronotum, and elytra (®gs. 1, 171, 183, 188), the re¯exed, explanate, anterolateral cephalic margin (®gs. 171, 183, 189, 194), and the ventrobasal groove on the median lobe of the aedeagus (®gs. 48, 80, 97, 152). It is separated from other tribes except the Vatesini by the emarginate posterior elytral margin (®gs. 25, 69, 91, 114, 131, 143), and from other tribes but the Deropini by the presence of a neck (®g. 194). DESCRIPTION: Length 2.7±4.9 mm; width 1.1±1.5 mm. Color dark to pale reddish brown and with or without yellowish brown lateral margins of pronotum and elytra. Head with vertex coarsely punctate (®gs. 1, 171, 183, 188); punctures setate. Head with anterolateral margin explanate and weakly to strongly re¯exed from antenna to anterior margin of clypeus (®gs. 171, 183, 189, 194); clypeus with (®g. 184) or without (®gs. 189, 194) re¯exed anterior margin. Epistomal suture present and complete (®g. 189) or incomplete (®g. 184); suture, when complete, angulate at middle; midcranial suture (®gs. 1, 189) present, well developed or rudimentary, and extending posteriorly from middle of epistomal suture Fig. 1. Lacvietina cuprina. or absent. Postocular lateral margin of head with carina (®gs. 172, 184) or rounded ridge (®gs. 189, 194) extending medially from posterior margin of eye to neck. Neck present and broad (®g. 194). Gular sutures 4 AMERICAN MUSEUM NOVITATES NO. 3430 moderately to widely separated and diver- short, basal, median carina. Tergite VII gent from about middle. Submentum with with palisade fringe of posterior margin coarse punctation or impunctate. Mentum well developed, reduced, or absent. Ter- coarsely punctate. Labial palp three-seg- gites IX divided middorsally by tergum X mented; glossa emarginate medially. Max- (®gs. 13, 73, 119, 144). illary palp four-segmented; fourth segment MALE: Sternites V±VII and sternum VIII long, stout, and fusiform (®gs. 184, 189); variously modi®ed; V±VII with large to galea with dense cluster of apical setae and small rounded lobe on internal surface of an- with row of setae on lateral margin. Man- terior margin (®gs. 3, 4, 11, 87, 95, 108). dible short, broad, and edentate; mola well Sternite VII with (®gs. 4, 77) or without (®g. developed. Antenna long or short and 40) peg setae. Sternum VIII (®gs. 6, 51, 88, reaching to about middle of elytra or ex- 111) with deep, apically wide, basally ta- tending beyond posterior margin; antennal pered, median emargination; narrow basal insertion beneath re¯exed margin of clyp- portion of emargination with tiny setae along eus and frons; antennomere 2 sparsely pu- margins; apical margin with four lobes, each bescent; antennomeres 3±11 with ®ne, dense pubescence. with long, stout, dark, apical seta; lateral lobe Pronotum (®gs. 1, 66, 79, 130, 188) wider small, paramedial lobe large. than long; surface with coarse, setate punc- Aedeagus with medial surfaces of para- tation; lateral region explanate and strongly meres contiguous (®gs. 5, 74, 93, 121); me- or feebly punctate; anterior and posterior an- dian lobe with shallow to deep longitudinal gles angulate (®gs. 79, 183) or broadly groove on ventral surface of bulbous base rounded (®gs. 1, 188). Tergosternal suture (®gs. 48, 80, 97, 152). present. Hypomeron strongly in¯exed and FEMALE: Sternites V±VI unmodi®ed. Ster- not visible in lateral view; postprocoxal lobe nite VII with truncate or rounded posterior present and visible in lateral view. Furcas- margin. Sternum VIII with ®ve (®gs. 68, 82) ternum medially carinate. or six (®gs. 7, 27, 141) apical lobes; lateral Elytra with coarse, setate punctation (®gs. and paralateral apical lobes paired; median 1, 171, 183, 188); lateral margin moderately region with one or two apical lobes; median to strongly explanate and forming ``epipleur- lobe(s) ¯attened and with apical, fanlike ar- al gutter''; epipleural ridge with row of rangement of setae of variable number. Ter- spinelike setae (®g. 1); spinelike setae in- gum VIII with four (®gs. 9, 16), ®ve (®gs. creasingly short posteriorly; epipleuron visi- 28, 140), or six (®g. 81) apical lobes; lateral ble in lateral view; posterior margin emar- lobe with (®gs. 9, 81, 140) or without (®g. ginate (®gs. 1, 25, 69, 91, 114); posterolat- 117) small secondary lobe on lateral edge; eral angle strongly (®g. 91) to slightly (®g. lateral and paralateral lobes paired, long and 143) produced. Mesospiracular peritreme slender; median region with one or two lightly sclerotized, triangular, moderately lobes. Valvifers paired, broad, and curved long, tapered medially; peritremes separated (®gs. 14, 72, 118, 145). Coxite long, slender, by small, interperitremal sclerite. Mesoster- cylindrical, and pubescent (®g. 14); stylus num with or without short midbasal carina; long, slender, and pubescent (®g. 14) or paramedial carina present or absent; median surface with curved carina near base of me- short. sosternal process; mesosternal process Spermatheca as in ®gures 8, 92, 99, 142. grooved (®g. 146). DISCUSSION: Cameron (1919) established Tarsal formula 5±5±5. the tribe with one genus, Megarthropsis. Abdomen strongly tapered posteriorly Little was known or written about the (®g. 1). Segment II with one pair of para- group until Smetana (1983a, 1992) rede- tergites. Segment III with two pairs of par- scribed and discussed the tribe and its atergites. Segments IV±VII with one pair of placement, added two genera (Nepaliodes paratergites. Sternum I present as narrow and Peitawopsis), and published illustra- sclerite anterior to II. Sternite II fused to tions of characters and a habitus for the III and with median carina. Sternite III with genera. 2004 HERMAN: MEGARTHROPSINI 5

Figs. 2±9. Megarthropsis decorata. 2. Aedeagus, lateral. 3. Sternite VI, male. 4. Sternite VII, male. 5. Aedeagus, ventral. 6. Sternum VIII, male. 7. Sternum VIII, female. 8. Spermatheca. 9. Tergum VIII, female. 6 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 10±15. Megarthropsis deverra. 10. Aedeagus, ventral. 11. Sternite VII, male. 12. Aedeagus, lateral. 13. Tergites IX and tergum X, female. 14. Segment IX, ventral, female. 15. Sternum VIII, male (broken lines represent completion of missing portion of segment; apical seta on right side cut off).

Apically expanded setae on the dorsal sur- Asia. At present, species are reported from face of the head, pronotum, and elytra are Nepal, India, Thailand, Malaysia, Singapore, uniquely characteristic of the tribe but are too Vietnam, China, and Taiwan. It is probable dif®cult to view without the high magni®ca- that the tribe occurs throughout the region, tion of a compound microscope or scanning including other sites in China and Indonesia electron microscope to be useful as a practical and other countries of Southeast Asia and diagnostic feature of the tribe. that many more species will be found. All DISTRIBUTION: The species of this tribe are the species have been collected from moist all known from the southeastern quadrant of to wet leaf litter, adjacent to or in stream 2004 HERMAN: MEGARTHROPSINI 7

Figs. 16±19. Megarthropsis deverra. 16. Tergum VIII, female. 17, 18. Spermatheca. 19. Sternum VIII, female. beds, or on the forest ¯oor. Collections have ¯exed (®g. 184); anterior pronotal angle been made at elevations of 220±2900 m. angulate (®g. 79) . . Nepaliodes Coiffait 3(1). Elytral surface with lateral third concave, median portion convex (®g. 188); meta- KEY TO GENERA OF THE MEGARTHROPSINI sternum without pit adjacent to apex of Here, in the other keys, and in the mesosternal process ...... descriptions care must be taken when ...... Peitawopsis Smetana searching for peg setae on the abdominal Ð Elytral surface convex to near margin (®g. sternites. Often their small size make them 1); metasternum with deep pit adjacent to dif®cult to see with a stereoscopic apex of mesosternal process (®gs. 146, 196, 197) ...... Lacvietina, new genus microscope and they are more easily and reliably viewed with the higher magni®cation of a compound microscope. Megarthropsis Cameron Figures 2±73, 171±182 1. Basal angles of pronotum angulate (®gs. 66, Megarthropsis Cameron, 1919: 231. Type spe- 79); head with well-developed laterobasal cies: Megarthropsis decorata Cameron, ®xed postocular carina (®gs. 172, 184) ...... 2 by monotypy. Ð Basal angles of pronotum broadly rounded ÐCameron, 1921: 355, 406 (characters; check- (®gs. 1, 96); head with rounded laterobasal list).ÐScheerpeltz, 1933: 1478 (catalog).Ð postocular ridge (®gs. 189, 194) ...... 3 Blackwelder, 1952: 236 (type species).ÐSme- 2(1). Anterior margin of clypeus not re¯exed but tana, 1983a: 144 (characters; illustrations).Ð with weak ridge; anterior pronotal angle Smetana, 1992: 204 (characters; illustrations; rounded (®gs. 66, 172) ...... habitus).ÐHerman, 2001a: 678 (catalog)...... Megarthropsis Cameron Ð Anterior margin of clypeus strongly re- DIAGNOSIS: Megarthropsis can be separat- 8 AMERICAN MUSEUM NOVITATES NO. 3430 ed from Nepaliodes by the angulate postocu- al angle strongly angulate; median three- lar carina (®g. 172), the rounded anterior an- quarters of notum coarsely and moderately gle of the pronotum (®gs. 66, 172), and, ex- densely punctate and strongly convex; lat- cept for M. djawaensis, the narrowly explan- eral eighth explanate and with coarse, mod- ate lateral margin of the elytra (®g. 171). The erately dense punctation. ®rst antennal segment of most species of Me- Elytra convex (®g. 171), slightly convex garthropsis is parallel-sided to slightly ta- medially and strongly convex laterally; sur- pered (®g. 171), but in one species, M. dja- face evenly, densely, and coarsely punctate; waensis, the scape is moderately strongly ta- lateral sixteenth to ®fth explanate and with pered apically and similar to that of Nepali- margin re¯exed; lateral margin with row of odes. Megarthropsis is distinguished from short spinelike setae extending from humer- Peitawopsis and Lacvietina by the distinct al angle to near basal angle; setae of epi- postocular carina (®g. 172) and the angulate pleural ridge increasingly short posteriorly; posterior margin (®gs. 25, 69) slightly, mod- basal angle of the pronotum (®g. 66); the lat- erately, or strongly emarginate laterally; ter two genera lack both features. posterolateral angle slightly, moderately, or DESCRIPTION: Length 3.8±4.2 mm; width strongly produced (®gs. 25, 69). Mesoster- 1.1±1.4 mm. num with short, thick, midlongitudinal ca- Color reddish brown with yellowish rina extending posteriorly from basal mar- brown pronotal lateral margins; head usually gin; paramedial carina rudimentary, present dark reddish brown to nearly black. only basally. Metasternum with shallow de- Head (®g. 171) with dorsum densely and pression adjacent to apex of mesosternal coarsely punctate. Clypeal surface largely process; circum-mesocoxal ridge enlarged impunctate, but with a few punctures pre- medially. sent. Head (®g. 171) with lateral margin Procoxa without carina on medial surface. strongly re¯exed from antenna to anterior MALE: Sternite VII (®gs. 4, 11, 21) with margin of clypeus; anterior margin of clyp- wide emargination of posterior margin; sur- eus with feeble ridge. Epistomal suture face with deep to shallow median depres- complete and angulate at middle; midcranial sion and with or without median peg setae. suture rudimentary, minute, and poorly de- Tergum VIII (®gs. 26, 36, 39, 67) with four veloped (best seen in lightly pigmented or apical lobes. cleared specimens). Dorsum of head without Aedeagus (®gs. 5, 10, 49, 59) with para- midlongitudinal groove; surface with or meres straight or slightly curved; parameres without broad, shallow, median depression. of nearly equal length and width; median Postocular lateral margin of head with ca- lobe with deep groove adjacent to midline rina extending posteromedially from eye; of ventral surface of base (®g. 48). carina angulate near eye (®g. 172); posto- FEMALE: Sternum VIII with ®ve (®g. 68) cular vertical carina present and extending or six lobes (®gs. 7, 19); median lobe or pair ventrally from lateral carina. Gular sutures of lobes with one fanlike cluster of setae. widely separated. Submentum impunctate. Tergum VIII with four (®gs. 9, 16) or ®ve Antennae long, slender, and reaching be- (®g. 28) apical lobes; lateral lobe with small secondary lobe on lateral edge. yond posterior margin of elytra; scape more Spermatheca (®gs. 8, 57) with apex of or less parallel-sided from near base to apex capsule enlarged and globose, then abruptly (see M. decorata, M. deverra, M. frazeren- constricted and tapered proximally to sper- sis) or slightly (see M. durga, M. empusa, mathecal duct; juncture of capsule with sper- M. parca, M. smetanai) to moderately (see mathecal duct not enlarged. M. djawaensis) tapered apically; dorsal sur- DISCUSSION: Cameron (1919: 231) de- face of scape with short, moderately dense scribed the genus in detail; Smetana (1983a: pubescence and ventral surface sparsely pu- 144; 1992: 204±206) supplemented it with bescent. new characters and numerous illustrations. Pronotum (®gs. 66, 172) with anterior angle broadly rounded and extending beyond KEY TO SPECIES OF MEGARTHROPSIS median portion of anterior margin; lateral 1. Sternum VIII (®g. 6) with posterior margin margin unevenly curved to basal angle; bas- emarginate (males) ...... 2 2004 HERMAN: MEGARTHROPSINI 9

Figs. 20±25. Megarthropsis djawaensis. 20. Sternum VIII, male. 21. Sternite VII, male. 22. Aedea- gus, ventral. 23. Aedeagus, lateral. 24. Spermatheca. 25. Elytron, apical margin, left (setae omitted).

Ð Sternum VIII with posterior margin entire, not dense mat of pubescence in median de- emarginate (females) ...... 9 pression .... M. smetanai, new species 2(1). Sternite VII without peg setae (®gs. 21, 40) Ð Sternites V and VI without dense median ...... 3 mat of pubescence ...... 5 Ð Sternite VII with peg setae (®gs. 4, 61) . . 5(4). Sternite V (®g. 45) with small peg setae me- ...... 4 dially ..... M. frazerensis, new species 3(2). Antennal scape tapered anteriorly from Ð Sternite V without peg setae ...... 6 base; cephalic postocular vertical carina 6(5). Sternite VII (®g. 11) with broad, deep emar- obliquely oriented ...... gination; parameres (®g. 10) obliquely ...... M. djawaensis, new species truncate apically (in ventral or dorsal Ð Antennal scape of more or less equal di- view) ...... M. deverra, new species ameter throughout; cephalic postocular Ð Sternite VII (®gs. 4, 33, 53) with broad, vertical carina dorsoventrally oriented . . moderately deep emargination; parameres ...... M. empusa, new species acute or subacute apically, not obliquely 4(3). Sternites V (®g. 63) and VI (®g. 65) with truncate (in ventral or dorsal view) . . 7 10 AMERICAN MUSEUM NOVITATES NO. 3430

7(6). Sternite VII (®g. 53) without peg setae on the evenly curved emargination of sternite disc ...... M. parca, new species VII (®g. 4), and from M. djawaensis, M. em- Ð Sternite VII (®gs. 4, 33) with peg setae on pusa, M. parca, and M. durga by the large disc ...... 8 cluster of discal peg setae on sternite VII (®g. 8(7). Sternite VII (®g. 33) with sinuate posterior margin ...... M. durga, new species 4). Ð Sternite VII (®g. 4) with evenly curved pos- Females of M. decorata have two median terior margin . . . M. decorata Cameron lobes on sternum VIII (®g. 7) and a pair of 9(1). Sternum VIII (®g. 68) with one median lobe straight, basally broad median lobes on ter- ...... M. smetanai, new species gum VIII (®g. 9). Tergum VIII is straight Ð Sternum VIII (®g. 7) with two median lobes from the base to the apex; the median region ...... 10 is not tumid. 10(9). Tergum VIII (®g. 28) with ®ve lobes on DESCRIPTION: Length 3.2±3.8 mm; width apical margin ...... M. djawaensis, new species 1.1±1.3 mm. Color reddish brown and yel- Ð Tergum VIII (®gs. 9, 34, 71) with four lowish brown. Head dark reddish brown, lobes on apical margin ...... 11 nearly black. Pronotum with convex median 11(10). Tergum VIII (®g. 55) tumescent at base region reddish brown, darker anteriorly than of median lobes ...... posteriorly; lateral, explanate region yellow- ...... M. parca, new species ish brown. Elytra dark reddish brown. Ab- Ð Tergum VIII without median tumescence, domen reddish brown. Antennae and legs surface (in lateral view) straight from pale reddish brown. base to apex ...... 12 Head with deep, moderately large punc- 12(11). Tergum VIII (®g. 16) with pair of short, broad, widely separated median lobes tation dorsally; postocular vertical carina ...... M. deverra, new species dorsoventrally oriented. Antennal scape more Ð Tergum VIII (®gs. 9, 34) with pair of or less parallel-sided. long narrowly separated median lobes Pronotum with deep, moderately large ... M. decorata Cameron, M. durga, punctation. new species Elytra with deep moderately large punctation; lateroapical emargination moderately DESCRIPTION OF SPECIES deep; epipleural gutter moderately wide. Megarthropsis decorata Cameron Wings fully developed. Tergite VII with Figures 2±9 well-developed palisade fringe on posterior margin. Megarthropsis decorata Cameron, 1919: 232. MALE: Sternite V with shallow median im- Type locality: Singapore: Mandai. Lectotype, pression; surface without peg setae or dense male, designated by Smetana (1983a: 146) in The Natural History Museum (London); not ex- mat of setae medially; posterior margin with amined. broad, shallow, indistinct emargination. ÐCameron, 1921: 379, 406 (characters; Singa- Sternite VI (®g. 3) with shallow median pore).ÐScheerpeltz, 1933: 1478 (catalog).Ð impression; impression with dense cluster of Smetana, 1983a: 146 (lectotype designation; setae near posterior margin; posterior margin characters; ®gures 1±7 [only, the other illustra- with broad, shallow emargination. tions refer to M. parca]; bionomics; Singapore). Sternite VII (®g. 4) with broad, shallow DIAGNOSIS: Megarthropsis decorata has no median depression; posterior margin with one feature that immediately separates it broad, deep, evenly curved, median emargi- from all of its congeners; various characters nation; emargination without median notch; must be used to distinguish it. The males of apical region of disc with two medially sep- M. decorata can be separated from M. sme- arated clusters of peg setae that also extend tanai by the absence of a dense mat of setae to and laterally along posterior margin; peg on sternites V and VI, from M. frazerensis setae along posterior margin arranged in by the absence of peg setae on sternite V, even or slightly sinuate row. from M. deverra by the shallower emargi- Sternum VIII (®g. 6) with deep, broad, nation of sternite VII (®g. 4) and the more median emargination; emargination strongly slender parameres (®g. 5), from M. durga by constricted near middle and with basal por- 2004 HERMAN: MEGARTHROPSINI 11

Figs. 26±28. Megarthropsis djawaensis. 26. Tergum VIII, male. 27. Sternum VIII, female. 28. Ter- gum VIII, female. tion of emargination strongly narrowed; mar- more specimens were collected from the Nee gins of emargination strongly sinuate. Soon swamp forest, which is in the Mandai Aedeagus (®gs. 2, 5) with parameres region. straight, slender, and tapered to subacute A male collected on Mt. Kinabalu, Bor- apex. neo, is indistinguishable from the specimens FEMALE: Tergum VIII (®g. 9) with median from Singapore. Two females were collected pair of lobes straight, tapered apically, and with the male from Borneo. This disjunction broad basally; surface without tumescence at of the distribution of M. decorata is surpris- base of middle lobe. Sternum VIII (®g. 7) ing given the diversity of species in the Kin- with three pairs of apical lobes. abalu region. However, because most species Spermatheca as in ®gure 8. discussed herein are known from only one DISCUSSION: The illustrations cited in 1992 collecting site or are from different eleva- as M. decorata (Smetana, 1992: ®gs. 30±42) tions on the same mountain, little can be said are of M. smetanai and in 1983 the illustra- about the distribution of the species. tions of the female abdominal segments Specimens of the type series were collect- (Smetana, 1983a: ®gs. 8±10) are of M. parca. ed in wet rotting leaves on the edge of a jun- Illustrations of the male of M. decorata gle stream (Cameron, 1919: 233) and from from Singapore were published by Smetana ¯ood debris (Smetana, 1983a: 146). Smetana (1983a: 1±7). The ®gures of M. decorata in (op. cit.) concluded that the species lives in the present article were taken from a speci- leaf litter and other debris in very wet habi- men from Borneo. tats. Although no elevational data were in- DISTRIBUTION AND HABITAT: The species cluded on the locality labels, the Mandai re- was described from Singapore. The type se- gion is at low elevation (the highest point in ries was collected from Mandai; later two Singapore is about 175 m). The specimens 12 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 29±33. Megarthropsis durga. 29. Aedeagus, ventral. 30. Aedeagus, lateral. 31. Sternum VIII, male. 32. Spermatheca. 33. Sternite VII, male.

from Mt. Kinabalu were collected from for- Megarthropsis deverra, new species est ¯oor litter near a stream in the vicinity of Figures 10±19 the headquarters of the National Park at an HOLOTYPE: Male. [Malaysia:] ``SABAH: Poring elevation of about 1500 m. Hot Springs, Langanan Falls, 900±950 m MATERIAL EXAMINED: Five specimens: 3 12.V.1987 BurckhardtÐLoÈbl''. Deposited in males, 2 females. Singapore: Nee Soon the MuseÂum d'Histoire Naturelle, GeneÁve, Swamp forest, 24-V-68, R.W. Taylor (1 male, Switzerland (MHNG). MHNG; 1 male, NHMW). Malaysia: Sabah: PARATYPES: One male, 1 female. Same data as ho- near Kinabalu N.P. headquarters, forest ¯oor lotype (1 female, MHNG). Malaysia: Sabah: litter near Liwagu Stream, 29-VII-82, G.-M. Crocker Range, Mawar Waterfall env. (9c), 17.6.1996, vegetation debris and forest litter de Rougemont (1 male, 2 females, GRC). around fallen trees (1 male, NHMW). Three males and one female were dissected for genitalic and abdominal features. DIAGNOSIS: The males of M. deverra can 2004 HERMAN: MEGARTHROPSINI 13

Figs. 34±36. Megarthropsis durga. 34. Tergum VIII, female. 35. Sternum VIII, female. 36. Tergum VIII, apex, male. be distinguished from males of all other spe- Wings fully developed. Tergite VII with cies by the broad deep emargination of ster- well-developed palisade fringe on posterior nite VII (®g. 11) and the short, broad, margin. obliquely truncate parameres of the aedeagus MALE: Sternite V with feeble, shallow me- (®g. 10). dian impression; surface without peg setae or The females can be distinguished from dense mat of setae; posterior margin feebly those of other species by the short, widely emarginate. separated median pair of lobes of tergum Sternite VI with broad, shallow depres- VIII (®g. 16). sion; surface with cluster of setae on poste- DESCRIPTION: Length 3.7±4.2 mm; width rior margin near lateral margin of impres- 1.2 mm. Color dark reddish brown, reddish sion, but without dense discal mat of setae; brown, and yellowish brown. Head dark red- posterior margin with broad moderately deep dish brown, nearly black. Pronotum with ex- emargination. planate lateral region yellowish brown; con- Sternite VII (®g. 11) with broad, moder- vex median region dark reddish brown, near- ately deep, median depression; posterior ly black anteriorly and paler reddish brown margin with broad, deep emargination; emar- posteriorly. Elytra dark reddish brown, near- gination truncate or slightly rounded medi- ly black. Abdomen reddish brown. Antennae ally and without median notch; apical region reddish brown. Legs pale reddish brown. of disc with two medially separated clusters Head with deep, moderately large punc- of numerous peg setae; peg setae along pos- tation dorsally; postocular vertical carina terior margin arranged in evenly curved row. dorsoventrally oriented. Antennal scape more Sternum VIII (®g. 15) with deep, broad, or less parallel-sided. median emargination; emargination strongly Pronotum with deep, moderately large constricted near middle and with basal por- punctation. tion strongly narrowed; margins of emargi- Elytra with deep moderately large punc- nation strongly sinuate. tation; lateroapical emargination moderately Aedeagus (®gs. 10, 12) with parameres deep; epipleural gutter moderately wide. slightly bent to right (in ventral view); par- 14 AMERICAN MUSEUM NOVITATES NO. 3430 ameres short, broad medially, and obliquely reddish brown posteriorly; lateral, explanate truncate apically. region reddish brown. Elytra dark reddish FEMALE: Tergum VIII with median pair of brown. Abdomen reddish brown with apical lobes broad, short, and widely separated (®g. segments darker reddish brown. Antennae 16). Sternum VIII with three pairs of apical and legs reddish brown. lobes (®g. 19). Head with large, deep punctures on dorsal Spermatheca as in ®gures 17, 18. surface. Postocular vertical carina obliquely, ETYMOLOGY: The name of this species is dorsoposteriorly to ventroanteriorly, orient- from the Latin for the Roman guardian of ed. Antenna with ®rst segment moderately newborn children (Deverra) and is symbol- tapered from base to apex. ized by a broom for sweeping away evil in- Pronotum with large, deep punctures. ¯uences (Jordan, 1993); used in apposition. Elytra with large, deep punctation; latero- DISTRIBUTION AND HABITAT: This species is apical emargination deep (®g. 25); epipleural known from two localities, one in the Mt. gutter wide. Kinabalu National Park at about 900 m, the Wings fully developed. Tergite VII with other in the nearby Crocker Range. well-developed palisade fringe on posterior MATERIAL EXAMINED: Three specimens: margin. holotype male and male and female para- MALE: Sternite V without median impres- types. sion, peg setae, or dense mat of setae medi- All specimens were dissected for genitalic ally; posterior margin without emargination. and abdominal characters. Sternite VI with feeble median impression; surface without special cluster of setae; pos- Megarthropsis djawaensis, new species terior margin without emargination. Figures 20±28 Sternite VII (®g. 21) with broad, shallow, HOLOTYPE: Male. ``INDONESIA: W Java `Ranca median impression; impression without setae Upas' ca. 1300m 10 km S Ciwidey lg. Schuh medially, but with short setae along lateral 9.8.1994''. Deposited in the Naturhistorisches margins; posterior margin with moderately Museum Wien, Wien, Austria (NHMW). deep, broadly V-shaped emargination; pos- PARATYPES: Two males, 3 females. Same data as holotype but collected from ``forest litter'' (1 terior margin evenly curved, not sinuate, to male, 2 females, NHMW). Indonesia: W. Java, median emargination; surface without peg Situ Lembang, ca 1500 m, 15 km N. Bandung, setae. lg. Schuh, 7.8.1994 (1 male, 1 female, Sternum VIII (®g. 20) with deep, broad, NHMW). median emargination; emargination more strongly narrowed near basal quarter; base of DIAGNOSIS: This species is separated from all others of the genus by the apically tapered emargination narrowly rounded; margins of antennal scape, the oblique orientation of the emargination weakly sinuate. postocular vertical carina, and the strongly Aedeagus (®gs. 22, 23) with parameres produced posterolateral angle of the elytra straight, slender, tapered, and narrowly (®g. 25). The punctation of the pronotum and rounded apically. dorsal surface of the head and elytra is coars- FEMALE: Tergum VIII (®g. 28) with para- er and deeper and the punctation of the pro- medial lobes separated by smaller median notal disc is more sparse than for any of the lobe. Sternum VIII (®g. 27) with three pairs other species of the genus. Only this species of apical lobes. and M. empusa lack peg setae on sternite VII Spermatheca as in ®gure 24. (®g. 21); these two species can be separated ETYMOLOGY: The name for this species is by the features listed in the preceding two taken from a variant of the Indonesian name sentences. (Djawa) for Java. DESCRIPTION: Length 3.3±3.8 mm; width DISTRIBUTION AND HABITAT: The species is 1.3±1.4 mm. Color reddish brown and yel- known only from western Java where spec- lowish brown. Head dark reddish brown to imens were collected from forest litter at nearly black. Pronotum with convex median 1300 m and 1500 m in August. region dark reddish brown anteriorly and MATERIAL EXAMINED: Six specimens: ho- 2004 HERMAN: MEGARTHROPSINI 15 lotype male, 2 paratype males, and 3 para- Wings fully developed. Tergite VII with type females. well-developed palisade fringe on posterior Two males and one female were dissected margin. for characters of the abdomen and genitalia. MALE: Sternite V with feeble median impression; surface without peg setae or dense Megarthropsis durga, new species mat of setae; posterior margin without emar- Figures 29±36 gination. HOLOTYPE: Male. [Malaysia]: ``SABAH Mt. Kin- Sternite VI with feeble median impression; abalu 1580 m, 27.IV.1987 BurckhardtÐLoÈbl''. impression with slightly denser row of setae Deposited in the MuseÂum d'Histoire Naturelle, along posterior margin, but without dense GeneÁve, Switzerland (MHNG). discal mat of setae; posterior margin with PARATYPES: Two males. Same data as holotype (1 broad, shallow emargination. male, MHNG). [Malaysia]: Sabah: Poring Hot Sternite VII (®g. 33) with broad, shallow Springs, Langanan Falls, 900±950 m, 12-V- 1987, Burckhardt and LoÈbl (1 male, MHNG). median impression; posterior margin with broad, moderately deep, sinuate emargina- DIAGNOSIS: Males of M. durga can be sep- tion; emargination without median notch; arated from other species by the broad, mod- surface with small, median cluster of peg se- erately deep, sinuate posterior margin of ster- tae on disc and with row of peg setae along nite VII (®g. 33) and the sinuate row of peg posterior margin; peg setae of posterior mar- setae near the margin. The disc of sternite gin arranged in sinuate row and row narrow- VII has only a few peg setae medially. ly divided medially. Females are separated from other species, Sternum VIII (®g. 31) with deep, broad, except M. decorata and M. smetanai,bythe median emargination; emargination slightly pair of median lobes of tergum VIII (®g. 34) more strongly constricted from about apical that are straight and broad at the base; the quarter and gradually convergent to base; surface adjacent to the base of the median margins of emargination moderately sinuate. lobes is not tumescent. This pair of median Aedeagus (®gs. 29, 30) with parameres lobes is broader basally in M. durga than in slightly curved to right (in ventral view), M. decorata (®g. 9). Sternum VIII (®g. 35) long, slender, and tapered to acute apices. of M. durga has a pair of median lobes in FEMALE: Tergum VIII (®g. 34) with me- contrast to the single median lobe of M. sme- dian pair of lobes straight, tapered apically, tanai. Denser cephalic, pronotal, and elytral punctation separate M. durga from M. dja- and broad basally; surface without tumes- waensis. cence near base of median lobes. Sternum VIII (®g. 35) with three pairs of apical lobes. DESCRIPTION: Length 3.4±3.8 mm; width 1.4 mm. Color reddish brown and yellowish Spermatheca as in ®gure 32. brown. Head reddish brown to dark reddish ETYMOLOGY: Named for the many armed brown to nearly black. Pronotum with con- Hindu goddess Durga, who is depicted as a vex median region reddish brown to dark woman riding on a lion or tiger and in whose reddish brown; lateral explanate region yel- hands are weapons. In later Hindu tradition, low brown. Elytra and abdomen reddish particularly in northern India, Durga is linked brown. Antennae and legs pale reddish with the fertility of crops and is celebrated brown. at harvest time in the Durga Puja (Jordan, Head with deep, moderately large punc- 1993); used in apposition. tation dorsally; postocular vertical carina DISCUSSION: The females identi®ed as M. dorsoventrally oriented. Antennal scape more durga are not designated as paratypes be- or less parallel-sided to slightly tapered api- cause they cannot be distinguished reliably cally. from M. decorata. Pronotum with deep, moderately large DISTRIBUTION AND HABITAT: This species is punctation. known only from Mt. Kinabalu in Borneo Elytra with deep moderately large punc- where it was collected at 900±1580 m. tation; lateroapical emargination moderately MATERIAL EXAMINED: Six specimens: ho- deep; epipleural gutter moderately wide. lotype male, 2 paratype males, 3 females. 16 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 37±41. Megarthropsis empusa. 37. Aedeagus, ventral. 38. Aedeagus, lateral. 39. Tergum VIII, male (broken lines represent completion of missing portion of segment; long apical seta on left side of illustration missing). 40. Sternite VII, male. 41. Sternum VIII, male (long lateroapical setae missing).

The females are from the type locality and an, 18.V.87 BurckhardtÐLoÈbl''. Deposited in were collected with the holotype. the MuseÂum d'Histoire Naturelle, GeneÁve, Three males and one female were dissect- Switzerland (MHNG). ed for genitalic and abdominal characters. DIAGNOSIS: The absence of peg setae on sternite VII (®g. 40) will separate the males Megarthropsis empusa, new species of this species from the others of the genus Figures 37±41 except M. djawaensis. This species is distin- HOLOTYPE: Male. [Malaysia]: ``SABAH: Crocker guished from M. djawaensis by the denser Ra. 1600m, km 51 rte Kota Kinabalu-Tambun- cephalic and pronotal punctation, the nearly 2004 HERMAN: MEGARTHROPSINI 17 cylindrical ®rst antennomere, the dorsoven- portion of emargination strongly narrowed; tral orientation of the vertical postocular ca- margin of emargination strongly sinuate. rina that extends ventrally from the dorsolat- Aedeagus (®gs. 37, 38) with paramere eral postocular carina, the less strongly pro- straight, moderately broad, and tapered to duced posterolateral elytral angle, and the subacute apex. narrower epipleural gutter. The emargination FEMALE: Unknown. of sternite VII of the male of M. empusa (®g. ETYMOLOGY: The name is from the Greek 40) is deeper than in M. djawaensis (®g. 21). (Empousa) for hobgoblin or a mischievious The female is unknown. goblin, and is used for the species found hid- DESCRIPTION: Length 3.4 mm; width 1.2 den in a long series of M. smetanai. mm. Color reddish brown and yellowish DISTRIBUTION AND HABITAT: This species is brown. Head dark reddish brown. Pronotum known only from the Crocker Range in with convex median region dark reddish northern Borneo where it was collected at brown anteriorly and reddish brown to yel- 1600 m with M. smetanai. lowish brown posteriorly; explanate lateral MATERIAL EXAMINED: One specimen: ho- region yellowish brown. Elytra dark reddish lotype male. brown. Abdomen reddish brown with darker apical segments. Antennae and legs reddish Megarthropsis frazerensis, new species brown. Figures 42±48 Head with moderately large punctation on HOLOTYPE: Male. [Malaysia]: ``PAHANG Frazer's dorsal surface; postocular ventral carina dor- Hill 11.III.90 Rougemont''// ``Megarthropsis soventrally oriented. Antennal scape more or decorata Cam. det Rougemont'' [labels hand- less parallel-sided to slightly tapered apical- written]. In collection of G.-M. de Rougemont ly. (GRC) to be deposited in Natural History Mu- Pronotum with moderately large puncta- seum, London. tion. PARATYPE: One male. Malaysia: Prov. Johor, Gg. Elytra with moderately large punctation; Ledang (14), Hutan Lipur, 200 m, 4.2.1992, leg. posterolateral emargination moderately deep; Schillhammer (1 male, NHMW). epipleural gutter narrow. DIAGNOSIS: The male of Megarthropsis Wings fully developed. Tergite VII with frazerensis can be separated from the males well-developed palisade fringe on posterior of all other species of the genus by the margin. unique presence of a cluster of peg setae MALE: Sternite V slightly ¯attened medi- on the middle of the disc of sternite V (®g. ally; surface without peg setae or dense mat 45). The emargination of the posterior mar- of setae medially; posterior margin with gin of sternite VII is notched medially and moderately wide feeble median emargina- the peg setae are divided into two clusters tion. (®g. 44). Sternite VI with wide, feeble median im- The female is unknown. pression; impression with slightly denser pu- DESCRIPTION: Length 3.4 mm; width 1.2 bescence near posterior margin; posterior mm. Color reddish brown and yellowish margin with wide, shallow median emargi- brown. Head dark reddish brown, nearly nation. black. Pronotum with explanate lateral re- Sternite VII (®g. 40) with broad, shallow, gion yellowish brown; convex median region median impression; impression without setae reddish brown, darker anteriorly and paler medially near posterior margin, but with posteriorly. Elytra and abdomen reddish short setae along lateral margin; posterior brown. Antennae and legs pale reddish margin with broad, sinuate emargination; brown. emargination deeper and more or less broad- Head with deep, moderately large punc- ly V-shaped medially; surface without peg tation dorsally; postocular vertical carina setae. dorsoventrally oriented. Antennal scape more Sternum VIII (®g. 41) with broad, deep, or less parallel-sided. median emargination; emargination strongly Pronotum with deep, moderately large constricted from about middle and with basal punctation. 18 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 42±48. Megarthropsis frazerensis. 42. Aedeagus, ventral. 43. Aedeagus, lateral. 44. Sternite VII, male. 45. Sternite V, male. 46. Sternum VIII, male (lateroapical setae cut off). 47. Sternite VI, male. 48. Aedeagus, anterior end, cross section.

Elytra with deep moderately large punc- disc with scattered cluster of small peg setae tation; lateroapical emargination shallow; in feeble impression. epipleural gutter moderately wide. Sternite VI (®g. 47) with broad, shallow, Wings fully developed. Tergite VII with median impression; posterolateral edge of well-developed palisade fringe on posterior impression with small dense cluster of setae; margin. posterior margin with moderately deep, MALE: Sternite V (®g. 45) with broad, rounded median emargination. shallow emargination of posterior margin; Sternite VII (®g. 44) with broad, shallow 2004 HERMAN: MEGARTHROPSINI 19 median depression; posterior margin with deagus; this con®guration is unique among broad, deep median emargination; emargi- the known species. nation with small median notch; median re- Females of M. parca can be separated gion of disc with peg setae arranged in two from other species by the tumid apical third dense, medially separated clusters; peg setae of tergum VIII (®g. 55). Each lobe of the of anterior region scattered; peg setae along median pair of lobes of tergum VIII curves posterior margin aligned in more or less laterad from the midlongitudinal line and the even, medially separated rows. lobes are slender (®g. 54). Sternum VIII (®g. 46) with deep, broad, DESCRIPTION: Length 3.7±4.1 mm; width median emargination; emargination wide about 1.4 mm. Color reddish brown; head, apically and strongly constricted near middle convex median region of pronotum, elytra, and with basal portion of emargination and abdomen with darker infusions; head strongly narrowed; margin of emargination darker than remainder of body; pronotum strongly sinuate. with explanate lateral portion paler reddish Aedeagus (®gs. 42, 43) with parameres brown; antennae and legs pale reddish feebly bent to left (in ventral view) and ta- brown. pered to subacute apex. Head with deep, moderately large punc- FEMALE: Unknown. tation dorsally; postocular vertical carina ETYMOLOGY: The name for this species is dorsoventrally oriented. Antennal scape more taken from the type locality, Frazer's Hill in or less parallel-sided to slightly tapered api- West Malaysia. cally. DISTRIBUTION AND HABITAT: This species is Pronotum with deep, moderately large known by only two Malaysian specimens punctation. and both lack further collecting data. Elytra with deep moderately large punc- MATERIAL EXAMINED: Two specimens: ho- tation; lateroapical emargination moderately lotype and paratype males. deep; epipleural gutter moderately wide. Both specimens were dissected for geni- Wings fully developed. Tergite VII with talic and abdominal characters. well-developed palisade fringe on posterior margin. Megarthropsis parca, new species MALE: Sternite V with feeble median im- Figures 49±58 pression; surface without peg setae or dense setal mat; posterior margin with broad feeble HOLOTYPE: Male. [Malaysia]: ``SABAH Mt. Kin- emargination. abalu 1900 m, 26.IV.1987 BurckhardtÐLoÈbl''. Sternite VI (®g. 52) with broad, shallow Deposited in the MuseÂum d'Histoire Naturelle, GeneÁve, Switzerland (MHNG). median depression; impression with moder- PARATYPES: Three males, 5 females. Same data as ately dense cluster of setae near posterior holotype (2 males, 3 females, MHNG). [Malay- margin, but without dense setal mat; poste- sia]: Sabah: Mt. Kinabalu, 6200 ft, 22± rior margin with broad, shallow emargina- 26.VI.68, R.W. Taylor (1 male, MHNG). [Ma- tion. laysia]: Borneo: Sabah: Mt. Kinabalu N.P., Sternite VII (®g. 53) with broad, shallow Summit Trail, 1890 m, [26.IV.86], A. Smetana median depression; posterior margin with (1 female, ASC). [Malaysia]: Sabah Mt. Kina- broad, deep, evenly curved, median emar- balu 1580 m, 27.IV.1987 BurckhardtÐLoÈbl (1 gination; emargination without median female, MHNG). notch; disc without peg setae; peg setae ar- ÐSmetana, 1983a: 146 (®gs. 8±10; misidenti®ed as M. decorata). ranged in curved row adjacent to posterior margin; row continuous or slightly separat- DIAGNOSIS: The evenly curved posterior ed medially. margin of sternite VII (®g. 53), the single Sternum VIII (®g. 51) with deep, broad, row of peg setae near the posterior margin, median emargination; emargination moder- and absence of discal peg setae will separate ately constricted near middle and gradually the males of M. parca from other species. convergent to narrow base; margins of emar- The ventral surface of each paramere is gination moderately strongly sinuate. oblique to the midsagittal plane of the ae- Aedeagus (®gs. 49, 50). Parameres more 20 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 49±53. Megarthropsis parca. 49. Aedeagus, ventral. 50. Aedeagus, lateral. 51. Sternum VIII, male. 52. Sternite VI, male. 53. Sternite VII, male.

or less straight (in ventral view), moderately Spermatheca as in ®gures 57 and 58. long, moderately broad, and rounded apical- ETYMOLOGY: The name of this species is ly; ventral surface oriented obliquely to mid- from the Latin (Parca) for the goddess of sagittal plane; apex with small point on ven- destiny and birth (Leach, 1992); used in ap- tral surface. position. FEMALE: Tergum VIII (®g. 54) with me- DISCUSSION: Megarthropsis parca was re- dian pair of lobes slightly curved laterally; ported as M. decorata from East Malaysia, median margin curved; lobes slender and Sabah, Mt. Kinabalu (Smetana, 1983a: 146 moderately tapered apically; surface near and ®gs. 8±10). base of median lobes strongly tumescent (®g. DISTRIBUTION AND HABITAT: This species is 55). Sternum VIII (®g. 56) with three pairs known only from Mt. Kinabalu in Borneo of lobes. where it was collected at 1580 to 1900 m. 2004 HERMAN: MEGARTHROPSINI 21

Figs. 54±58. Megarthropsis parca. 54. Tergum VIII, female. 55. Tergum VIII, lateral, female. 56. Sternum VIII, female. 57, 58. Spermatheca.

MATERIAL EXAMINED: Nine specimens: ho- males, 3 females, MHNG); 1550±1650 m, 24- lotype male and three male and ®ve female IV-1987 (1 male, 2 females, MHNG; 1 male, paratypes. MSC); 1750 m, 27-IV-1987 (1 female, Three males and three females were dis- MHNG). Borneo: Sabah: Mt. Kinabalu Nation- sected for genitalic and abdominal features. al Park HQ, Liwagu Riv. Tr., 1520 m, 12-VIII- 88, A. Smetana [B106] (1 male, AMNH); 11- VIII-88 [B100] (1 female, AMNH). Borneo: Megarthropsis smetanai, new species Sabah Mt., Kinabalu National Park HQ, Silau- Figures 59±73, 171±182 Silau Tr., 1550 m, 4-IX-88, A. Smetana [B177] (1 male, AMNH); 2-IX-88 [B171] (1 male, HOLOTYPE: Male. [Malaysia]: ``SABAH Crocker Ra. 1600 m, km 51 rte Kota Kinabalu-Tambun- AMNH). an, 18.V.87 BurckhardtÐLoÈbl''. Deposited in ÐSmetana, 1992: 204±206 (®gs. 30±42; misiden- the MuseÂum d'Histoire Naturelle, GeneÁve, ti®ed as M. decorata). Switzerland (MHNG). DIAGNOSIS: Males of M. smetanai can be PARATYPES: Thirty-two males, 16 females. [Ma- distinguished from their congeners by the laysia]: Same data as holotype (15 males, 5 fe- dense mat of pubescence in the large median males, MHNG). Sabah: Mt. Kinabalu, 1430 m, 22-V-1987, Burckhardt and LoÈbl (3 males, depression of sternites V and VI (®gs. 63, MHNG); 1500 m, 30-IV-1987 (1 male, 65). Sternite VII (®g. 61) has a deep, rounded MHNG); 1500 m, 21-V-1987 (1 male, 2 fe- median depression, the peg setae are ar- males, MHNG); 1550 m, 28-IV-1987 (2 males, ranged in a dense cluster along the posterior 2 females, MHNG); 1550 m, 29-IV-1987 (5 margin, and the posterior margin is sinuate 22 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 59±66. Megarthropsis smetanai. 59. Aedeagus, ventral. 60. Aedeagus, lateral. 61. Sternite VII, male. 62. Sternite VII, apical margin, male (setae omitted). 63. Sternite V, male. 64. Sternum VIII, male. 65. Sternite VI, male. 66. Pronotum (setae omitted). 2004 HERMAN: MEGARTHROPSINI 23 with a small, rounded median notch (®g. 62); base; margins of emargination moderately this con®guration is unique to M. smetanai. sinuate. The females of M. smetanai are distin- Aedeagus. Parameres (®gs. 59, 60) bent guished from those of all other species by the moderately to left (in ventral view) and ta- presence of only ®ve lobes on sternum VIII, pered to subacute apex. the median lobe of which is broad (®g. 68). FEMALE: Tergum VIII (®g. 71) with me- DESCRIPTION: Length 3.3±4.6 mm; width dian pair of lobes straight and broad basally. 1.3±1.4 mm. Color reddish brown and yel- Sternum VIII (®g. 68) with ®ve lobes, one lowish brown. Head dark reddish brown to lateral and one paralateral pair of lobes and nearly black. Pronotum convex, median re- one broad median lobe. gion dark reddish brown to reddish brown, Spermatheca as in ®gure 70. posterior portion usually paler; explanate lat- ETYMOLOGY: This species is named to hon- eral region pale reddish brown to yellowish or my friend and colleagueAlesÏ Smetana for brown. Elytra and abdomen reddish brown. his many sterling contributions to our under- Antennae and legs pale reddish brown. standing of the Staphylinidae and who, over Head with deep, moderately large punc- the years, I have come to admire for the qual- tation dorsally; postocular vertical carina ity of his work and depth of his knowledge. dorsoventrally oriented. Antennal scape more In large part his studies and collections of the or less parallel-sided to slightly tapered api- Megarthropsini made the present work pos- cally (®g. 171). sible. Pronotum with deep, moderately large DISCUSSION: Figures 30±42 published by punctation. Smetana (1992) and identi®ed as M. decor- Elytra with deep moderately large punc- ata are illustrations of M. smetanai. tation; lateroapical emargination moderately Sternum VIII (®g. 68) of the females has deep; epipleural gutter moderately wide. one broad median lobe, but other members Wings fully developed. Tergite VII with of the tribe, except the species of Nepaliodes, well-developed palisade fringe on posterior have a median pair of lobes. The broad lobe margin. in M. smetanai is clearly the result of a fu- MALE: Sternite V (®g. 63) with large, sion of this pair of median lobes. moderately deep, median depression; depres- DISTRIBUTION AND HABITAT: This species is sion with dense mat of setae and depression known only from Mt. Kinabalu and the near- nearly as long as segment; surface without by Crocker Range where it was collected at peg setae; posterior margin with broad, shal- elevations of 1430±1750 m. According to low median emargination. ®eld notes for specimens collected by A. Sternite VI (®g. 65) with large, deep, me- Smetana this species has been collected from dian depression; depression nearly as long ¯eshy mushrooms and soft polyporus-type segment and surface covered with dense mat mushrooms growing on a fallen rotting tree, of setae; posterior margin with broad, mod- and from fresh and rotting fruits of the ``ker- erately deep, median emargination. osene tree''. Sternite VII (®g. 61) with large, deep, MATERIAL EXAMINED: Forty-nine speci- rounded, median depression; depression pol- mens: holotype male, 32 male and 16 female ished medially and without setae in middle, paratypes. but with short setae laterally and basally; sur- Six males and one female were dissected face with dense cluster of peg setae along for genitalic and abdominal characters; one posterior margin; cluster of peg setae slightly male was disarticulated. separated medially; posterior margin with broad sinuate emargination; emargination Nepaliodes Coiffait with small, rounded median notch (®g. 62). Figures 74±90, 183±187 Sternum VIII (®g. 64) with broad, deep Nepaliodes Coiffait, 1977: 272. Type species: Ne- median emargination; emargination moder- paliodes variolosus Coiffait, ®xed by original ately strongly constricted at about apical designation and monotypy. third then gradually convergent to narrow ÐSmetana, 1983a: 150 (characters; illustrations; 24 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 67±73. Megarthropsis smetanai. 67. Tergum VIII, male. 68. Sternum VIII, female. 69. Elytron, apical margin, left (setae omitted). 70. Spermatheca. 71. Tergum VIII, female. 72. Segment IX, ventral, female. 73. Tergites IX and tergum X, female. 2004 HERMAN: MEGARTHROPSINI 25

Figs. 74±80. Nepaliodes variolosus. 74. Aedeagus, ventral. 75. Aedeagus, lateral. 76. Sternites V and VI, male. 77. Sternite VII, male. 78. Sternum VIII, male (long apical setae missing). 79. Pronotum (setae omitted). 80. Aedeagus, anterior end, cross section.

habitus).ÐSmetana, 1992: 204 (characters; il- lateral third or fourth of the elytra (®g. 183). lustrations).ÐHerman, 2001a: 679 (catalog). The parameres (in ventral view) of Nepali- DIAGNOSIS: Nepaliodes is separated from odes are curved to the right, and the right other genera of the tribe by the apically ta- paramere is about twice as long as the left pered antennal scape (®g. 184), the angulate (®gs. 74, 84) and is more strongly curved. anterior and posterior angles of the pronotum Females have ®ve lobes on the posterior mar- (®g. 79), and the explanate, sparsely punctate gin of sternum VIII (®gs. 82, 89), and tergum 26 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 81±83. Nepaliodes variolosus. 81. Tergum VIII, female. 82. Sternum VIII, female. 83. Sper- matheca.

VIII has six slender lobes, the median pair dorsal surface of scape densely pubescent, of which is more slender than the others. The ventral surface sparsely pubescent. spermathecal capsule which tapers proximal- Pronotum (®gs. 79, 183) with anterior an- ly from the apex is abruptly expanded before gle angulate, narrowly rounded, and strongly attaching to the spermathecal duct (®gs. 83, produced beyond median portion of anterior 92). margin; lateral margin broadly and unevenly DESCRIPTION: Length 3.7±5.0 mm; width rounded; basal angle angulate; median three- 1.7±2.0 mm. Color evenly reddish brown to ®fths coarsely and densely punctate and dark reddish brown. strongly convex; lateral ®fth strongly explan- Head (®g. 184) with dorsum coarsely and ate and surface sparsely and feebly punctate. densely punctate. Clypeus sparsely punctate. Elytra (®g. 183) with median two-thirds or Head with lateral margin strongly re¯exed three-quarters convex and densely and from antenna to anterior margin of clypeus coarsely punctate; lateral third to quarter (®g. 184); anterior margin of clypeus strong- concave, sparsely and feebly punctate, ly re¯exed (®g. 184). Epistomal suture (®g. strongly explanate, and with lateral margin 184) present laterally on re¯exed portion of re¯exed; lateral margin with row of short, clypeus just anterior to antennal insertion, spinelike setae from humeral angle to near absent from median three-®fths; midcranial middle of lateral margin; spinelike setae in- suture absent. Dorsum of head without mid- creasingly short posteriorly; posterior margin longitudinal groove; surface with or without deeply emarginate laterally (®gs. 91, 183); broad, shallow median depression. Postocu- posterolateral angle strongly produced and lar lateral margin of head with carina ex- acute (®g. 91). Mesosternum with midlon- tending medially from eye (®g. 184); carina gitudinal carina extending from basal margin straight; vertical postocular carina absent. to near base of mesosternal process; para- Gular sutures moderately widely separated. medial carina rudimentary. Metasternum Submentum coarsely punctate. Antenna long with shallow depression adjacent to apex of and slender, reaching nearly to apical margin mesosternal process; circum-mesocoxal ridge of elytra; scape tapered apically (®g. 184); enlarged medially. 2004 HERMAN: MEGARTHROPSINI 27

Figs. 84±88. Nepaliodes solangelae. 84. Aedeagus, ventral. 85. Sternites V and VI, male. 86. Ae- daegus, lateral. 87. Sternite VII, male. 88. Sternum VIII, male.

Procoxa without carina on median surface. curved to right; right paramere thicker, more MALE: Sternite VII (®gs. 77, 87) with wide strongly curved, and about twice as long as emargination of posterior margin; surface left; median lobe with deep longitudinal with broad median depression and with me- groove near middle of ventral base (®g. 80). dian peg setae. Tergum VIII (®gs. 78, 88) FEMALE: Sternum VIII (®gs. 82, 89) with with three apical lobes; median lobe with ®ve apical lobes; median lobe broad and with shallow emargination. two small, apical, fanlike clusters of setae, Aedeagus (®gs. 74, 75, 84, 86), in ventral one on each side of midline; median lobe view, with parameres closely appressed and with midapical region more lightly pigment- 28 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 89±92. Nepaliodes solangelae. 89. Sternum VIII, apical portion, female. 90. Tergum VIII, female. 91. Elytron, apical margin, left (setae omitted). 92. Spermatheca. ed (and dif®cult to see) than lateroapical re- stance that can be removed after soaking gion. Tergum VIII (®gs. 81, 90) with six them in ammonia. slender apical lobes; median lobes more slen- Nepaliodes includes two species, N. vari- der and shorter than lateral and paramedial olosus, from Nepal and India and N. solan- lobes; lateral lobe without secondary lobe on gelae from Thailand and China. lateral edge. Spermathecal (®gs. 83, 92) capsule curved KEY TO SPECIES OF NEPALIODES and gradually tapered from apex to spermathecal duct; capsule expanded just before 1. Sternite VII (®g. 77) with posterior margin juncture with spermathecal duct. deeply emarginate (males) ...... 2 DISCUSSION: The detailed description of Ð Sternite VII with posterior margin truncate or Nepaliodes published by Coiffait (1977: 272) slightly emarginate (females) ...... 3 was ampli®ed by Smetana (1983a: 150; 2(1). Sternite VII (®g. 77) with broad, deep emar- 1992: 204), who also presented numerous il- gination; sternite V with setae posteriorly lustrations. directed; sternite VI with setae of median Females are identi®ed by their collection region directed slightly to middle (®g. 76) in association with the males and by distri- ...... N. variolosus Coiffait bution. Characters have not been found that Ð Sternite VII (®g. 87) with broad, moderately permit species identi®cation of the females. deep emargination; sternites V and VI Illustrations of segment VIII (®gs. 81, 82, 89, (®g. 85) with setae of median region me- 90) of the two species suggest that females dioposteriorly directed ...... can be separated by features of the terga and ...... N. solangelae, new species sterna, but those variations are found in spec- 3(1). Present in India and Nepal ...... imens of both species...... N. variolosus Coiffait Specimens of both species are usually Ð Present in Thailand and China ...... coated with a ®ne-grained, mudlike sub- ...... N. solangelae, new species 2004 HERMAN: MEGARTHROPSINI 29

DESCRIPTION OF SPECIES examined by Smetana, is Coiffait's determi- nation label for the species; it reads ''Ne- Nepaliodes variolosus Coiffait paliodes variolosa''. Figures 74±83, 183±187 The labels record some specimens from Nepaliodes variolosus Coiffait, 1977: 272. Type ``Dobate Ridge NE Barahbise''. According locality: Weg v. Pokhara z. Goropani, Zentral- to Smetana (1988: 176), that locality should Nepal. Holotype, male, deposited in Naturhis- read ``Pokhare NE Barahbise''; it is east of torisches Museum Wien; examined. Kathmandu (Smetana, 1988: 172, site 25). ÐSmetana, 1983a: 154 (characters; habitus; ae- DISTRIBUTION: Nepaliodes variolosus is deagus; body parts; bionomics; India; Nepal). known from Uttar Pradesh and West Bengal DIAGNOSIS: The deeper emargination of in India and Bagmati Province in Nepal at sternite VII (®g. 77), the ®ner, less medially elevations of 1700, 1900, and 2700 m, re- directed pubescence of sternites V and VI spectively. In Nepal the species was collected (®g. 76), and the less strongly tapered, more from accumulations of deep, wet leaf litter in apically blunt right paramere (®gs. 74, 75) depressions and small gullies in a deciduous will separate the males of N. variolosus from forest with dense undergrowth (Smetana, N. solangelae. The females of the two spe- 1983a: 155). cies can be separated by locality. MATERIAL EXAMINED: Twenty-seven spec- DESCRIPTION: Length 3.7±5.0 mm; width imens: 13 males, 14 females. Nepal: holo- 1.2±1.8 mm. Color reddish brown to dark type, locality cited above, September±Octo- reddish brown; explanate margins of head, ber 1971, collected by H. Franz (1 male, pronotum and elytra paler; head darker. NHMW); (Prov. Bagmati), Dobate Ridge Wings fully developed. [Pokhare: see Discussion above] NE Barah- Tergite VII with palisade fringe present. bise, 2700 m, 2.V.81, LoÈbl and Smetana (1 MALE: Sternite V (®g. 76) with pubes- male, 2 females, MHNG; 1 male, 1 female, cence of median region ®ne and more or less AMNH); Prov. Bagmati, Dobate Ridge [Pok- posteriorly directed. Sternite VI (®g. 76) with hare; see remarks under Discussion above] pubescence of median region slightly direct- NE Barahbise, 2800 m, 2.V.81, LoÈbl and ed to middle. Sternite VII (®g. 77) with Smetana (1 female, MHNG); Lalitpur Distr., broad, deep emargination of posterior mar- 2 km S Godavari, 1700 m, 19-X-83, Smetana gin; disc with dense cluster of peg setae and LoÈbl (1 male, 2 females). Inde: Garhwal along posterior margin and in shallow me- (UP), Mussoorie, No. 15, 1700 m, 19-X-79, dian depression of apical half. Sternum VIII I. LoÈbl (6 males, 2 females, MHNG); Ku- (®g. 78) with deep median emargination; maon (UP), Rangarh, No. 6B, 2000 m, emargination broad apically and narrow ba- 9.X.79, I. LoÈbl (4 males, 4 females, MHNG); sally; margins of emargination strongly sin- Uttar Pr., Kumaon, Rangarh, No. 6B, 2250 uate and convergent to narrow base. m, 9.X.79, I. LoÈbl (1 female, MHNG). India: Aedeagus (®gs. 74, 75) with right para- W. Bengal, Darjeeling dist., Algarah-Labha, mere about half as long as left; left paramere 1900 m, 11-X-78, Besuchet and LoÈbl(1fe- stout, strongly curved to left and tapered api- male, MHNG). cally to moderately broad, rounded apex (in Dissections of three males and two fe- dorsal or ventral view). males were examined for features of the ab- FEMALE: Tergum VIII (®g. 81), sternum domen and for the male and female genitalia VIII (®g. 82), and spermatheca (®g. 83). Spe- and genital segments. cies distinguished from N. solangelae by as- No specimens were disarticulated. sociation with male and distribution. DISCUSSION: Two spellings of the name Nepaliodes solangelae, new species were used by Coiffait (1977: 244, 272, 273) Figures 84±90 in the original publication, namely Nepali- HOLOTYPE: Male. ``THAILAND: Chiang Mai Do'i odes variolasa and N. variolosa. As ®rst re- Suthep, 1550 m 4.XI.1985, Burckhardt-LoÈbl''. viser Smetana (1983a: 154) selected and cor- Holotype deposited in the MuseÂum d'Histoire rected the name to N. variolosus. Attached Naturelle, GeneÁve, Switzerland (MHNG). to the pin with the holotype, which was not PARATYPES: Six males. Thailand: Chiang Mai, 30 AMERICAN MUSEUM NOVITATES NO. 3430

Doi Inthanon, 1650 m, 7.XI.1985, Burckhardt- cies distinguished from N. variolosus by as- LoÈbl (2 males, MHNG); Chiang Mai distr., Doi sociation with male and distribution. Suthep-Pui NP, 1080 m, 17.2.1995, leg P. ETYMOLOGY: This species is named for So- Schwendinger, evergr. hill forest (1 male, langel Casabona who has worked with me as MSC); ``Doi Phu Nan Province, 500 m, Kha a preparator for 25 years. Nat Park, 1700 m, Schwendinger, 6.10.91'' (1 male, MHNG) [according to information pro- DISCUSSION: Females were not selected as vided by I. LoÈbl, this label should read: ``N. paratypes because they can only be identi®ed Thailand, Nan Province, Doi Phu Kha National by distribution and by their collection in as- Park, 1700 m, 6.10.91, P. Schwendiger'']; Tak- sociation with males. I have been unable to sin Maharat N.P., 1000 m, 9.2.1993, Schwen- ®nd characters of the females that distinguish dinger (1 male, MHNG). China: Guangxi: Da- the species. wangling, 1190 m, 6.VIII.1999, J.R. Fellowes, DISTRIBUTION: This species is known from 15 m closed canopy, broadleaf forest ¯oor litter northern Thailand where it was collected at (1 male, GRC). elevations of 1000±1780 m, and Guangxi Province, China, where it was collected at DIAGNOSIS: The males of this species are 1190 m. separated from those of N. variolosus by the MATERIAL EXAMINED: Seven males and 9 shallower emargination of the posterior mar- females: holotype male, 6 male paratypes, gin of sternite VII (®g. 87), the more strongly and 9 females. The localities from which the tapered, more apically acute right paramere females were collected are the following. (in ventral view; ®g. 84), and the slightly Thailand: Chiang Mai, Doi Inthanon, 1650 coarser, more medially directed median setae m, 7.XI.1985, Burckhardt-LoÈbl (6 females, of sternites V and VI (®g. 85). The females MHNG); Chiang Mai, Doi Inthanon, 1780 are identi®ed by association with the male m, 17.XII.86, P. Schwendinger (1 female, and by locality. MHNG); Chiang Mai, Doi Suthep, 1400 m, ESCRIPTION: Length 4.0±4.7 mm; width D 5.XI.1985, Burckhardt-LoÈbl (1 female, 1.8±2.0 mm. Color reddish brown to dark MHNG); Prov. Chiang Mai, Doi Suthep, reddish brown; explanate margins of head, 1320 m, 27.XII.1986, P. Schwendinger (1 fe- pronotum, and elytra paler; head darker. male, MHNG). Wings fully developed. Dissections of ®ve males and three fe- Tergite VII with palisade fringe present. males were examined for features of the ab- MALE: Sternite V (®g. 85) with setae of domen and characters of the male and female median region slightly coarse and mediopos- genitalia and genital segments. teriorly directed. Sternite VI (®g. 85) with No specimens were disarticulated. pubescence of median region slightly coarse and strongly medioposteriorly directed; pos- Peitawopsis Smetana terior margin with small median lobe. Ster- Figures 93±120, 188±193 nite VII (®g. 87) with broad, moderately deep emargination of posterior margin; disc Peitawopsis Smetana, 1992: 199. Type species: with dense cluster of peg setae in broad, shal- Peitawopsis monticola Smetana, ®xed by orig- low, median depression of apical half and inal designation and monotypy. along posterior margin. Sternum VIII (®g. ÐHerman, 2001a: 679 (catalog).ÐHerman and 88) with deep median emargination; emar- Smetana, 2002: 198 (diagnosis; key to species). gination broad apically and narrow basally; DIAGNOSIS: The broadly rounded posterior margin of emargination strongly sinuate to pronotal angle will separate Peitawopsis narrow base. (®gs. 96, 188) from Nepaliodes (®g. 79) and Aedeagus (®gs. 84, 86) with left paramere Megarthropsis (®g. 66) which have strongly (in ventral view) moderately curved and angulate posterior angles. The lateral third of about half as long as right; right paramere the elytra of Peitawopsis is concave (®g. stout, strongly curved and tapered apically to 188), the elytra of Megarthropsis (®g. 171) acute apex. and Lacvietina (®g. 1) are convex to the nar- FEMALE: Tergum VIII (®g. 89), sternum rowly re¯exed epipleural ridge, and the ely- VIII (®g. 90), and spermatheca (®g. 92). Spe- tra of Nepaliodes (®g. 183) are convex to 2004 HERMAN: MEGARTHROPSINI 31

Figs. 93±100. Peitawopsis monticola. 93. Aedeagus, ventral. 94. Aedeagus, lateral. 95. Sternite VII, male. 96. Pronotum (setae omitted). 97. Aedeagus, anterior end, cross section. 98. Spermatheca (coiled cluster omitted). 99. Spermatheca. 100. Sternum VIII, male. about the lateral third or fourth, which is con- lateroapical angle to the elytral suture, but is cave, explanate, and sparsely punctate. The emarginate laterally in the other genera. The posterior margin of the elytra of Peitawopsis distinct postocular cephalic carina of Nepa- is broadly and evenly emarginate from the liodes (®g. 184) and Megarthropsis (®g. 172) 32 AMERICAN MUSEUM NOVITATES NO. 3430 is modi®ed to a rounded ridge in Peitawopsis to humeral angle; posterior margin with (®g. 189). The antennal scape of Peitawopsis broad emargination; posterolateral angle (®g. 189) is nearly parallel-sided from near strongly produced. Mesosternum without the base to the apex, whereas the scape of midlongitudinal basal ridge or carina; para- Nepaliodes (®g. 184) is tapered apically from medial carina present basally and moderately the base. long. Metasternum without depression or pit DESCRIPTION: Length 3.3±4.6 mm; width adjacent to apex of mesosternal process; cir- 1.3±1.7 mm. cum-mesocoxal ridge moderately developed Color pale to dark reddish brown; head medially. dark reddish brown, nearly black; anterior Procoxa with carina on median surface. and lateral margins of pronotum and elytra MALE: Sternite VII (®gs. 95, 108, 110) pale reddish brown to yellowish brown. with broad emargination of posterior margin; Head (®g. 189) with dorsum moderately surface with peg setae and broad depression densely and moderately coarsely punctate. medially. Sternum VIII (®gs. 100, 101, 111) Clypeal punctation ®ne and sparse or absent. with deep anteriorly tapered emargination. Head with lateral margin moderately strongly Tergum VIII with three apical lobes; median re¯exed from antenna to anterior margin of lobe with broad, shallow apical emargina- clypeus (®g. 189); anterior margin of clypeus tion. Aedeagus (®gs. 93, 103, 112) with par- not re¯exed and with ®ne transverse micro- ameres tapered apically, straight, and of sculpturing. Epistomal suture (®g. 189) pres- about equal length and width; median lobe ent, complete, and angulate at middle; mid- with shallow groove (or depression) near cranial suture short (®g. 189). Dorsum of middle of ventral surface of base (®g. 97). head with midlongitudinal groove extending FEMALE: Sternum VIII (®g. 104) with three posteriorly from epistomal suture to near pairs of lobes on posterior margin; median neck and well-developed (®g. 189) to feeble. pair of lobes with fanlike cluster of setae on Postocular lateral margin of head with round- each lobe. Tergum VIII (®gs. 117, 120) with ed ridge extending medially from eye (®g. four apical lobes; lateral lobe without sec- 189); postocular vertical carina absent. Gular ondary lobe on lateral edge. sutures widely separated. Submentum mod- Spermathecal capsule enlarged at apex erately coarsely punctate. Antenna short, and strongly constricted then tapered prox- reaching to near middle of elytra; scape (®g. 189) more or less parallel-sided from near imally (®gs. 105, 116) or gradually tapered base to apex, and with scattered setae dor- from apex to spermathecal duct (®gs. 98, sally and ventrally. 99); juncture of capsule with spermathecal Pronotum (®gs. 96, 188) with anterior and duct not enlarged; spermathecal duct with posterior angles broadly rounded; anterior one or more loops near capsule. angles produced beyond median portion of DISCUSSION: Peitawopsis, with three spe- anterior margin; lateral margin broadly and cies, is known only from the Chung-Yang evenly curved; median and lateral surfaces Shan of Taiwan. The species have reduced densely and coarsely punctate and with some elytra and wings, are ¯ightless, and are anastomosing punctures; punctation slightly found at elevations of 1500±2900 m. Two denser anteriorly and laterally than medially species, P. monticola and P. watanabei, live and posteriorly. on Peitawushan where both were collected Elytra (®g. 188) reduced, combined width from moist litter on the forest ¯oor. The two greater than length as measured from scutel- species were collected at one locality where lar apex to line across lateroapical angles of P. watanabei was found in litter in depres- elytra; surface with dense punctation. Elytra sions and gullies of the forest ¯oor, and P. (®g. 188) with median two-thirds of dorsal monticola was found in litter and debris surface shallowly convex and lateral third along a trail. Although the two species over- shallowly concave; surface evenly, densely, lap in their elevational distribution, P. wa- and moderately strongly punctate; lateral tanabei is found at higher elevations to margin explanate and moderately re¯exed; about 2900 m. The third species (P. inexsp- lateral margin with spinelike setae restricted ectata) is from Kuanshan and Peinantashan 2004 HERMAN: MEGARTHROPSINI 33

Figs. 101±108. Peitawopsis inexspectata. 101. Sternum VIII, male. 102. Aedeagus, lateral. 103. Aedeagus, ventral. 104. Sternum VIII, female. 105±107. Spermatheca, variation. 108. Sternite VII, male. 34 AMERICAN MUSEUM NOVITATES NO. 3430 where it also was collected from moist, for- P. monticola are about as long as the median est ¯oor litter. lobe (®g. 93) in contrast to the longer parameres of P. inexspectata (®g. 103). KEY TO SPECIES OF Peitawopsis The spermatheca of the females of P. mon- 1. Abdominal sternite VII (®gs. 95, 110) with ticola has the apical end gradually enlarged emarginate posterior margin (males) .... 2 (®gs. 98, 99) in contrast to the abruptly en- Ð Abdominal sternite VII with rounded posterior larged apical end of the other two species margin (females) ...... 4 (®gs. 106, 115). The spermathecal duct is 2(1). Sternite VII (®g. 110) with uninterrupted densely coiled near the spermathecal capsule row of peg setae near posterior margin in P. monticola (®g. 99), but is only slightly ... P. watanabei Herman and Smetana coiled in P. inexspectata (®g. 106) and P. Ð Sternite VII (®gs. 95, 108) with two medi- watanabei (®g. 116). ally separated rows of peg setae near posterior margin ...... 3 DESCRIPTION: Length 3.4±4.6 mm; width 3(2). Sternite VII (®g. 95) with two rows of peg 1.4±1.6 mm. setae along posterior margin and with two Color pale to dark reddish brown, head small groups of peg setae on disc ..... usually dark reddish brown to nearly black...... P. monticola Smetana Pronotum with central region reddish brown Ð Sternite VII (®g. 108) with two rows of peg to dark reddish brown to nearly black and setae beginning at posterior margin and with lateral, anterior, and posterior margins extending onto disc and without separate pale reddish brown to yellowish brown. El- small group of peg setae on disc ...... ytra with central region reddish brown to ...... P. inexspectata Smetana 4(1). Spermathecal capsule (®g. 99) gradually en- dark reddish brown to nearly black and with larged at one end; spermathecal duct with lateral and anterior margins pale reddish multiple loops near capsule ...... brown to yellowish brown. Antennae reddish ...... P. monticola Smetana brown to pale reddish brown. Legs pale red- Ð Spermathecal capsule (®gs. 105, 116) dish brown. abruptly expanded at one end; spermathe- Wings reduced to minute pads. cal duct with a few loops near capule . . Tergite VII with minute, irregularly inter- ...... 5 rupted palisade fringe on posterior margin. 5(4). Present on Peitawushan and its foothills . . ... P. watanabei Herman and Smetana MALE: Sternite V with feeble, median im- Ð Present on Kuanshan and Peinantashan . . . pression on apical half; impression without ...... P. inexspectata Smetana pubescence near apical margin, but present on remainder. Sternite VI with shallow, median depres- DESCRIPTION OF SPECIES sion extending for most of length of segment; Peitawopsis monticola Smetana depression without pubescence on apical Figures 93±100, 188±193 ®fth, remainder sparsely pubescent. Sternite VII (®g. 95) with moderately Peitawopsis monticola Smetana, 1992: 204. Type locality: Taiwan: Pingtung Hsien, Peitawushan broad, moderately deep, arcuate median Trail at 2000 m. Holotype in the Smetana Col- emargination of posterior margin; sternite lection (ASC), to be deposited in the MuseÂum with medially separated row of peg setae on d'Histoire Naturelle, GeneÁve, Switzerland; ex- posterior margin and with two, small, medi- amined. ally separated clusters of peg setae on disc ÐHerman and Smetana, 2002: 199 (characters; near apical half; row of peg setae with longer distribution). spinelike seta near lateral end; sternite with DIAGNOSIS: The posterior margin of ster- 14±19 peg setae; posterior row with 10±14 nite VII of the males of P. monticola has a peg setae and discal clusters each with one medially separated row of peg setae and the to three, but usually with two peg setae; disc disc has a small cluster of peg setae on each with broad, shallow, slightly tapered depres- side of the midline (®g. 95); this con®gura- sion from posterior margin for most of tion of peg setae will separate P. monticola length; depression without punctation or pu- from the other two species. The parameres of bescence on most of surface, but with ®ne 2004 HERMAN: MEGARTHROPSINI 35

Figs. 109±116. Peitawopsis watanabei. 109. Aedeagus, lateral. 110. Sternite VII, male. 111. Sternum VIII, male. 112. Aedeagus, ventral. 113. Sternite VII, middle of posterior margin, male (setae omitted from median portion of disc). 114. Elytron, apical margin, left (setae omitted). 115, 116. Spermatheca, two views. punctation and ®ne setae on basal third and curved medially to about basal third then along lateral margin. abruptly and narrowly separated; basal third Sternum VIII (®g. 100) with broad, deep, of emargination narrow and tapered to nar- median emargination; margins of emargina- rowly rounded basal margin. tion widely separated apically and gradually Aedeagus with parameres about as long as 36 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 117±120. Peitawopsis watanabei. 117. Tergum VIII, female. 118. Segment IX, ventral, female. 119. Tergites IX and tergum X, female. 120. Tergum VIII, female, setae omitted. median lobe (®gs. 93, 94), as measured from description because the sclerotized structures apical margin of basal ori®ce to apex of par- of the internal sac of the other two species ameres. are much smaller and indescript. FEMALE: Spermathecal capsule gradually DISTRIBUTION AND HABITAT: This species is and moderately enlarged at apex and tapered known only from Peitawushan (see Smetana, from there toward spermathecal duct (®gs. 1995b: 6, site 10) in Pingtung Hsien where 98, 99); spermathecal duct with dense cluster it was collected at 1500±2325 m. Specimens of multiple loops near capsule (®g. 99). were collected in mature broadleaf forests by DISCUSSION: The genitalic structures of this sifting leaves, humus, and other debris from species differ signi®cantly from the other the forest ¯oor. species. The spermathecal duct is densely MATERIAL EXAMINED: Fifty-one speci- coiled near the spermathecal capsule, where- mens: 34 males, 17 females. as the spermathecal duct of the other species Taiwan: Pingtung Hsien: Peitawushan have a few loops near the spermathecal cap- trail at 2000 m, May 23, 1991, A. Smetana, sule. sifting fallen leaves, various other debris and The aedeagus of P. monticola has a large, humus accumulated along large fallen trees strongly sclerotized capsule within the ae- [T91] (holotype male, allotype, paratypes: 1 deagus that is presumably part of the internal male, 2 females, ASC). Peitawushan, Kuai- sac. This structure was not included in the Ku Hut, 2325 m, May 22, 1991, A. Smetana, 2004 HERMAN: MEGARTHROPSINI 37 sifting fallen leaves and other debris along Color pale to dark reddish brown with trail in mature broadleaf evergreen forest dark reddish brown to nearly black head. [T90] (paratypes: 11 males, 3 females, ASC); Pronotum with central region reddish brown 2125 m, April 27, 1992, A. Smetana, sifting to dark reddish brown and lateral, anterior, fallen leaves, dead vegetation, and various and posterior margins pale reddish brown to other moist debris in mature broadleaf ever- yellowish brown. Elytra mostly pale reddish green forest [T102] (6 males, 2 females, brown with darker central region. Antennae ASC); 2130 m, April 27, 1992, A. Smetana, mostly dark reddish brown, paler basally. sifting fallen leaves, dead vegetation, and Legs reddish brown. various other moist debris in mature broad- Wings reduced to minute pads. leaf evergreen forest [T101] (4 males, 4 fe- Tergite VII with minute, irregularly inter- males, ASC; 2 males, 1 female, AMNH); rupted palisade fringe on posterior margin. 2135 m, April 30, 1992, A. Smetana, sifting MALE: Sternite V with shallow median de- fallen leaves, various other debris and humus pression on apical two-thirds; pubescence accumulated along large fallen trees [T108] present on basal half of depression; apical (5 males, 3 females, ASC). Peitawushan Trail half of depression without pubescence or at 1500 m, May 1, 1992, A. Smetana, sifting punctation. fallen leaves, various other debris, and hu- Sternite VI with shallow median depres- mus accumulated along large fallen trees sion extending for most of length of segment; [T110] (4 males, 1 female, ASC). depression with punctation and pubescence Five males and two females were dissect- on basal third and without punctation or pu- ed for characters of the genital segments and bescence on midapical two-thirds. genitalia, and one female was disarticulated. Sternite VII (®g. 108) with moderately deep, arcuate, median emargination of pos- Peitawopsis inexspectata Smetana terior margin; sternite with row of peg setae Figures 101±108 on each side of midline of posterior region; Peitawopsis inexspectata Smetana, 1995a: 131. rows begin at posterior margin and curve an- Type locality: Taiwan: Kaohsiung Hsien, Kuan- teromedially onto posteromedial portion of shan trail above Kaunshanchi River, 2550 m. disc; posterior margin with long, spinelike Holotype in the Smetana Collection (ASC), to seta near posterolateral end of row of peg be deposited in the MuseÂum d'Histoire Natu- setae; sternite with 8±12 peg setae; median relle, GeneÁve, Switzerland; examined. surface of sternite with shallow depression ÐHerman and Smetana, 2002: 200 (characters; reaching from posterior margin to near base; distribution). depression slightly tapered anteriorly and DIAGNOSIS: Sternite VII (®g. 108) of the with scattered ®ne punctation and minute se- males of P. inexspectata has a row of peg tae around basal and lateral margins, but re- setae on each side of the midline that begins gion anterior to, between, and posterior to at the posterior margin and curves medioan- rows of peg setae devoid of punctation or teriorly onto the disc; the disc lacks separate ®ne pubescence. clusters of peg setae. This arrangement of Sternum VIII (®g. 101) with broad, deep, peg setae is unique among the three species median emargination; margins of emargina- of the genus. tion widely separated apically and gradually The females of P. inexspectata are sepa- curved medially to about basal third, then rated from P. monticola by the abruptly ex- strongly convergent and narrowly separated panded apical end of the spermatheca of the to base; basal third of emargination tapered former species (®g. 107) in contrast to the to rounded basal margin. gradually expanded portion near the sper- Aedeagus with parameres longer than me- mathecal capsule (®g. 99). Females of P. dian lobe (®gs. 102, 103) as measured from inexspectata and P. watanabei are similar apical margin of basal ori®ce to apex of par- and identi®ed by distribution and association ameres. with the male. FEMALE: Spermathecal capsule with apical DESCRIPTION: Length 3.2±4.4 mm; width end globular and abruptly enlarged and from 1.3±1.7 mm. there tapered toward spermathecal duct; sper- 38 AMERICAN MUSEUM NOVITATES NO. 3430 mathecal duct with several loops near cap- to be deposited in the MuseÂum d'Histoire Na- sule (®gs. 105±107). turelle, GeneÁve, Switzerland. DISTRIBUTION AND HABITAT: This species is DIAGNOSIS: Males of Peitawopsis watana- known only from Kuanshan and Peinanta- bei are separated from those of P. inexspec- shan (see Smetana, 1995b: 6, sites 18 and 48) tata and P. monticola by the uninterrupted in Kaohsiung Hsien, Taiwan, where it was row of peg setae near the posterior margin collected at 2080±2550 m. Specimens were found in litter and debris on the ¯oor of ma- of sternite VII (®g. 110). ture broadleaf evergreen, mixed broadleaf The females of P. watanabei are separated evergreen, and coniferous forests. from those of P. monticola by the abruptly enlarged apical end of the spermatheca and MATERIAL EXAMINED: Thirty-®ve specimens: 19 males, 16 females. Taiwan: Kao- by the slightly coiled end of the spermathecal hsiung Hsien: Kuanshan trail above Kaun- duct near the capsule (®gs. 115, 116). Fe- shanchi River, 2550 m, April 21, 1992, A. males of P. watanabei and P. inexspectata Smetana, sifting fallen leaves, twigs, and hu- are similar and separated by the site of col- mus from ¯oor of mature broadleaf ever- lection. green forest [T96] (holotype male, allotype, DESCRIPTION: Length 3.3±3.7 mm; width paratypes: 3 males, 3 females, ASC; 1 fe- 1.3±1.4 mm. male, AMNH); July 22, 1993, sifting leaves, Color pale to dark reddish brown. Head twigs, and debris from ¯oor of mature mixed dark reddish brown to nearly black. Prono- broadleaf evergreen and coniferous (Abies) tum with central region dark reddish brown; forest [T160] (1 male, paratypes: 2 males, lateral, anterior, and posterior margins pale ASC; 1 male, AMNH). Peinantashan trail, reddish brown to yellowish brown. Elytra 2500 m, July 4, 1993, A. Smetana, sifting with central region dark reddish brown and fallen leaves and other debris from under de- with lateral and anterior margins pale reddish ciduous bushes along trail [T136] (3 males, brown to yellowish brown. Antennae reddish 1 female, ASC); 2390±2490 m, July 4, 1993, brown to dark reddish brown. Legs pale red- sifting fallen leaves and other debris from dish brown to yellowish brown. under deciduous bushes along trail [T138] (3 Wings reduced to minute pads. males, 4 females, ASC); 2080 m, July 6, Tergite VII with minute, irregularly inter- 1993, sifting layers of moist fallen leaves and rupted palisade fringe on posterior margin. other debris accumulated along wall of old MALE: Sternite V with slight impression forest road in mature broadleaf evergreen on midapical half; impression with microse- forest [T141] (4 males, 3 females, ASC); tae and micropunctures. 2450 m, May 2, 1995, sifting debris from a Sternite VI with feeble, anteriorly taper- few moist spots on ¯oor of very dry mature ing, median depression; depression nearly as mixed forest (Quercus, Pinus, Chamaecypar- long as sternite; depression with microsetae is) [T170] (1 male, 3 females, ASC). and micropunctures basally and laterally and One specimen lacking abdominal seg- without punctures or setae midapically. ments VI±X was collected at 2500 m along Sternite VII (®g. 110) with moderately the Peinantashan Trail and is almost certainly broad, moderately deep, arcuate median emar- P. inexspectata. gination of posterior margin; surface adjacent Five males and a female were dissected for to emargination with uninterrupted row of peg features of the genital segments and genitalia setae; gap between row of peg setae and pos- and one female was disarticulated. terior margin of sternite variable; sternite with 9±16 peg setae (®gs. 110, 113); spinelike seta Peitawopsis watanabei Herman and present near lateral end of row of peg setae; Smetana sternite without cluster of peg setae on disc Figures 109±120 separated from posterior row; surface with Peitawopsis watanabei Herman and Smetana, shallow, anteriorly tapering, median depres- 2002: 202. Type locality: Taiwan: Pingtung sion extending anteriorly from posterior mar- Hsien, Peitawushan, above Kuai-Ku Hut, 2680 gin; depression without setae or punctures on m. Holotype in the Smetana Collection (ASC), midposterior surface, but with micropunctures 2004 HERMAN: MEGARTHROPSINI 39 and microsetae laterally, basally, and on most tion, Km 16±17, 1700±1800 m, April 28, of median region. 1998, A. Smetana, sifting moist to wet fallen Sternum VIII (®g. 111) with broad, deep, leaves and other debris accumulated along anteriorly tapering emargination; margin of rock wall at edge of forest road in mature emargination broadly and shallowly sinuate broadleaf evergreen forest [T190] (paratypes: and medially convergent. 1 male, 2 females, ASC). Aedeagus with parameres about as long as Six males and three females were dissect- median lobe (®gs. 109, 112), as measured ed for features of the genital segments and from apical margin of basal ori®ce to apex genitalia; a male and female were disarticu- of parameres. lated. FEMALE: Spermathecal capsule with apical end globular and abruptly enlarged then ta- Lacvietina, new genus pered to spermathecal duct; spermathecal Figures 121±170, 194±203 duct with several loops near capsule (®gs. 115, 116). Type species: Lacvietina cuprina, new species, DISTRIBUTION AND HABITAT: This species is ®xed here by original designation. known in Pingtung Hsien from Peitawushan (see Smetana, 1995b: 6, site 10) and its foot- DIAGNOSIS: Lacvietina can be separated hills that extend south into northern Kao- from all other genera of the tribe by the deep hsiung Hsien (about 25 km south of site 10 metasternal pit adjacent to the apex of the in Smetana, 1995b: 6), where it was collected mesosternal process (®gs. 146, 196, 197). at elevations of 1700 to 2910 m. Specimens The presence of a rounded ridge (®g. 194) in were found in moist leaf litter and debris and place of a postocular carina on the lateral moss on the ¯oor of broadleaf evergreen and side of the head and the rounded anterior and Abies forests. Unlike P. monticola, this spe- posterior angles of the pronotum (®g. 130) cies ascends to the main ridge of Peitawu- separate it from Megarthropsis and Nepali- shan and occurs there in the Abies forest at odes. Lacvietina is distinguished from Pei- close to 3000 m. tawopsis by the convex elytra (®g. 1) and MATERIAL EXAMINED: Thirty specimens: 18 absence of a midlongitudinal groove on the males, 12 females. Taiwan: Pingtung Hsien: vertex of the head (®g. 194). Taiwan: Pingtung Hsien, Peitawushan, above DESCRIPTION: Length 2.7±3.7 mm; width Kuai-Ku Hut 2680 m, 29.IV.1992, A. Sme- 1.1±1.4 mm. Color dark to pale reddish tana, sifting moist fallen leaves, moss, and brown. other debris from small seepage in mature Head (®gs. 1, 194) with dorsum densely Abies forest with lush undergrowth [T106] and coarsely punctate. Clypeus impunctate or (holotype male; paratypes: 9 males and 6 fe- with a few scattered punctures. Head with males, ASC; 3 males, 2 females, AMNH). lateral margin moderately re¯exed from an- Peitawushan, Kuai-Ku Hut, 2325 m, May 21, tenna to near anterior margin of clypeus (®g. 1991, A. Smetana, sifting litter in forest ¯oor 194); anterior margin of clypeus not re¯exed. depression and in small gullies in mature Epistomal suture (®g. 1) present, entire, and broadleaf evergreen forest [T88] (paratype: 1 angulate at middle; midcranial suture short male, ASC). Peitawushan, above Kuai-Ku and moderately to feebly developed and only Hut, 2750 m, April 29, 1992, A. Smetana, visible in some specimens when cleared. sifting fallen leaves, old vegetation, moss and Dorsum without median impression or mid- various other debris in forest ¯oor depressions longitudinal groove (®g. 194). Postocular lat- and small gullies in mature broadleaf forest eral margin of head with rounded ridge ex- intermixed with conifers [T107] (paratype: 1 tending posteromedially from eye; vertical male, ASC). Peitawushan ridge, 2800±2910, postocular carina absent. Gular sutures wide- April 28, 1992, A. Smetana, sifting of moss, ly separated. Submentum coarsely punctate. twigs, and various other debris on shaded ar- Antenna long, slender, and extending from eas below groups of low Abies trees [T105] about middle of elytra to near posterior mar- (paratypes: 2 males, 2 females, ASC). Kao- gin; scape (®g. 194) more or less parallel- hsiung Hsien: Road above Tona Forest Sta- sided from near base to apex, not tapered api- 40 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 121±133. Lacvietina cuprina. 121. Aedeagus, ventral. 122. Sternite VI, male. 123. Sternite VII, male. 124±126. Sternite VII, median posterior region, variation, male. 127. Aedeagus, lateral. 128. Sternum VIII, male. 129. Tergum VIII, female. 130. Pronotum (setae omitted). 131. Elytron, apical margin, left (setae omitted). 132, 133. Tergum VIII, apex, male (setae omitted). 2004 HERMAN: MEGARTHROPSINI 41

Figs. 134±139. Lacvietina aurora. 134. Aedeagus, ventral. 135. Aedeagus, lateral. 136. Tergum VIII, apex, male (setae omitted). 137. Sternum VIII, male. 138. Sternite VII, male. 139. Sternite VI, male. cally, and with scattered pubescence on dor- lateral margin narrowly re¯exed and explan- sal and ventral surfaces. ate; lateral margin with short, spinelike setae Pronotum with anterior and posterior an- extending posteriorly from humeral angle; gles broadly rounded (®gs. 1, 130); anterior posterior margin moderately (®gs. 1, 131) to angles produced beyond median portion of slightly (®g. 143) emarginate laterally; fe- anterior margin; lateral margin evenly male of one unnamed species without emar- curved; median and lateral surfaces coarsely, gination; posterolateral angle moderately to densely, and evenly punctate; punctures dis- slightly produced (®gs. 131, 143). Mesoster- tinct, not anastomosing; at least one species num (®g. 146) without midlongitudinal basal with moderately coarse, moderately dense carina; paramedial basal carina present and punctation laterally and across base and with short. Metasternum with deep pit adjacent to ®ne punctation medially and anteriorly apex of mesosternal process (®gs. 146, 196, (based on undescribed female [sp. A]). 197); circum-mesocoxal ridge moderately Elytra (®g. 1) convex, moderately convex developed medially. medially and strongly convex laterally; sur- Procoxa without carina on medial surface. face evenly, densely, and coarsely punctate; MALE: Sternite VII (®gs. 123, 138, 150) 42 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 140±146. Lacvietina aurora. 140. Tergum VIII, female. 141. Sternum VIII, female. 142. Sper- matheca. 143. Elytron, apical margin, left (setae omitted). 144. Tergites IX and tergum X, female. 145. Segment IX, ventral, female. 146. Mesosternum and metasternum. with wide emargination of posterior margin, left paramere (in ventral view) shorter and broad median depression, and median cluster wider than right; median lobe with shallow of peg setae. Tergum VIII with four apical groove (or depression) near middle of ventral lobes (®gs. 132, 133, 136, 153). surface of base (®g. 152). Aedeagus (®gs. 121, 134, 147) with par- FEMALE: Sternum VIII (®gs. 141, 168) ameres more or less straight in ventral view; with three pairs of apical lobes; median pair 2004 HERMAN: MEGARTHROPSINI 43 of lobes with apical, fanlike cluster of setae Cameron, 1926, T. (T.) debilis Horn, 1877, on each lobe. Tergum VIII with four (®gs. T. (T.) himalayicus Cameron, 1926, and T. 129, 170) or ®ve (®gs. 140, 151) apical (T.) monticola Cameron, 1926; all have ®ne lobes; median lobe or median pair of lobes pronotal punctation and lack the metasternal moderately wide; lateral lobe with small sec- pit. However, a species from Sumatra, for ondary lobe on lateral edge. which I have seen only one female, has mod- Spermatheca (®gs. 142, 170) globose api- erately coarse pronotal punctation basally cally, abruptly constricted, then tapered api- and laterally and ®ne punctation medially cally to spermathecal duct; juncture of cap- and anteriorly and belongs in Lacvietina. sule with spermathecal duct not enlarged. Based on this specimen and on a habitus il- DISCUSSION: Five species from Thailand, lustration by Pace (1986: ®g. 10) it is likely Malaysia, Vietnam, and Taiwan are known that Tachinus (Tachinoderus) rougemonti and characterized herein. A sixth species, Pace, 1986: 49, from Thailand, should be from Indonesia, was examined but is known transferred to Lacvietina; that transfer must from only one female. It is likely that a sev- await study of the type or accurately identi- enth species (see below) from Thailand ®ed specimens. Examination of other species should be included. in Tachinus (Tachinoderus) may result in a The species were collected from montane few additional reassignments. regions at relatively low elevations. They ETYMOLOGY: The genus is named after the were all collected from wet to very wet litter Lac Viet, an ancient group of people in Vi- and debris. Without doubt other species will etnam. The gender of the name is feminine. be found in southern China, Southeast Asia, and perhaps Indonesia. KEY TO SPECIES OF LACVIETINA Tachinus punctatissimus (Hayashi) is transferred herein to Lacvietina. Hayashi 1. Sternite VII (®g. 123) emarginate (males) 2 (1991) described the species in Paratachinus Ð Sternite VII truncate (females) ...... 6 2(1). Sternite VII (®g. 150) with large cluster (ca. Blackwelder, 1952, a genus group name that 25±30) of peg setae on median surface of was ®rst proposed and characterized by disc ...... L. copiosa, new species Cameron (1932: 396) who, however, did not Ð Sternite VII (®gs. 123, 138) with smaller designate a type species, so the name was cluster (ca. 3±13) of peg setae on median unavailable. Blackwelder (1952: 293) made surface of disc ...... 3 the name available when he designated the 3(2). Sternite V (®g. 160) with pair of posteriorly type species (Herman, 2001b: 11). Paratach- diverging carinae medially ...... inus, originally established with two species, ...... L. paricosta, new species Paratachinus monticola Cameron and Par- Ð Sternite V without carinae ...... 4 atachinus laticollis Cameron (a secondary 4(3). Sternite VII (®g. 162) with posterior row of peg setae separated medially ...... homonym replaced by Tachinus oblongo- ...... L. punctatissima (Hayashi) punctatus Ullrich), is currently cited as a ju- Ð Sternite VII (®gs. 123, 138) with posterior nior synonym of Tachinus (Tachinoderus) row of peg setae continuous, not inter- (Ullrich, 1975: 291; Campbell, 1988: 276; rupted medially ...... 5 Li, 1995b: 207), but see Coiffait (1982: 39, 5(4). Sternite VII (®g. 123) with row of peg setae 116; 1984: 117, 138) and Hayashi (1991: 48) evenly arranged along posterior margin who cited it as valid...... L. cuprina, new species Tachinus (Tachinoderus) currently in- Ð Sternite VII (®g. 138) with row of peg setae cludes 42 species (extracted from Herman, sinuously arranged along posterior mar- 2001a). Based on the original descriptions of gin...... L. aurora, new species 6(1). the pronotal punctation of the species in Tergum VIII (®gs. 140, 151) with ®ve apical lobes ...... 7 Tachinoderus, only P. punctatissimus Hay- Ð Tergum VIII (®gs. 129, 171) with four api- ashi is reported to have the pronotum cov- cal lobes ...... 8 ered with coarse punctation, a characteristic 7(6). Tergite VII with strong ground sculpturing feature of species in Lacvietina. The few spe- between punctures; surface between cies of the subgenus that were examined in- punctures shining dully ...... clude Tachinus (Tachinoderus) brunneicollis ...... L. copiosa, new species 44 AMERICAN MUSEUM NOVITATES NO. 3430

Ð Tergite VII without or with feeble ground interrupted row of peg setae on the posterior sculpturing between punctures, surface margin of sternite VII (®gs. 123±126), the polished and strongly shining ...... shallow, dully shining depression of sternite ...... L. aurora, new species VI, and the absence of carinae on sternite V. 8(6). Elytral epipleuron with two rows of punc- The base of the emargination of sternum VIII tures, one along ventral margin and another along middle (®g. 195) ...... (®g. 128) is rounded and narrower than in ...... L. cuprina, new species the other species (®gs. 137, 154, 157, 161). Ð Elytral epipleuron with one row of punc- Females of L. cuprina are separated from tures along ventral margin ...... those of the other species by the presence of ...... L. punctatissima (Hayashi) a row of punctures along the ventral margin of the elytral epipleuron and another along the length of the middle (®g. 195); the other DESCRIPTION OF SPECIES species have a row of punctures adjacent to Lacvietina cuprina, new species the ventral margin only. Tergum VIII of L. Figures 121±133, 194±203 cuprina is comprised of four apical lobes, whereas both L. aurora and L. copiosa have HOLOTYPE: Male. ``VIETNAM: Hatinh Pr., RaoÁ ®ve (®gs. 140, 151); L. punctatissima also An 13 km W Huang Son, rt. 8 7 km S NuoÁc has four apical lobes (®g. 171). SoÃt village 18Њ21ЈN, 105Њ15ЈE, 350 m V-3± 1998, L. Herman, litter nr. drying forest DESCRIPTION: Length 2.7±3.4 mm; width stream''. On long-term loan to the American 1.1±1.3 mm. Color pale to dark reddish Museum of Natural History (AMNH), to be de- brown. Head darker than remainder of body, posited in the Institute of Ecology and Biolog- elytra darker than pronotum and abdomen. ical Resources, Hanoi, Vietnam (IEBR). Pronotum with dark disc and paler margins. PARATYPES: Sixty-eight males, 46 females. Same Abdomen with segments VII and VIII darker data as holotype (20 males, 10 females, than or concolorous with preceding seg- AMNH; 15 males, 12 females, IEBR; 5 males, ments. Antennae and legs reddish brown. 4 females, MHNG); same locality and collector Pronotal punctation extending to anterior as holotype: V-2±1998, 450 m, litter near seep margin and without lateroapical impunctate, along river (3 males, 2 females, AMNH; 3 males, 1 female, IEBR); IV-21±1998, 220 m, polished spot mesad of anterolateral angle. decaying litter near stream (3 females, AMNH); Elytra with posterolateral angle moderate- IV-26±1998, 220 m, litter near stream (2 fe- ly to slightly produced (®g. 131); epipleuron males, AMNH); IV-30±1998, 270 m, litter near (®g. 195) with two rows of punctures, one stream (1 female, AMNH); IV-25±1998, 1150 near vental margin and second on midlon- m, litter near drying stream (2 males, 2 females, gitudinal surface. Wings present and fully IEBR), IV-28±1998 (4 males, 4 females, developed. AMNH), IV-20±1998 (2 males, 1 female, Tergite VII with moderately strong ground AMNH). Vietnam: Quang Binh Pr., SW of sculpturing between punctures; surface dully Chalo, nr. Laos Bdr., 17Њ40ЈN, 105Њ46ЈE, 400 shining. m, IV-14, 15±1998, L. Herman (1 male, AMNH). Vietnam: Quang Nam Prov., 25 km MALE: Sternite V with shallow median de- (by air) S.W. Tra My, 940 m, IV-12±1999, pression on apical half; depression tapered 15Њ39ЈN, 108Њ02ЈE, ¯ood debris, L. Herman (4 basally; surface of depression with ground males, 1 female, AMNH, 3 males, IEBR); sculpturing and with setae and punctation ba- 15Њ12Ј14ЉN, 108Њ02Ј13ЉE, 850 m, IV-14±1999 sally and laterally but absent midapically; (3 males, 2 females, AMNH; 3 males, 1 female, posterior margin with broad, shallow median IEBR). Paratypes followed by the abbreviation emargination. ``IEBR'' are on long-term to the American Mu- Sternite VI (®g. 122) with shallow median seum (AMNH) and will eventually be deposited depression; depression nearly as long as ster- in the Institute of Ecology and Biological Re- nite; surface of depression with ground sources (IEBR). sculpturing and without punctures or setae DIAGNOSIS: The males of Lacvietina cu- except basally and laterally; posterior margin prina can be separated from their congeners of sternite with broad shallow median emar- by the small cluster of 3±7 peg setae on the gination. disc of sternite VII, the evenly curved, un- Sternite VII (®g. 123) with shallow me- 2004 HERMAN: MEGARTHROPSINI 45

Figs. 147±154. Lacvietina copiosa. 147. Aedeagus, ventral. 148. Aedeagus, lateral. 149. Sternite VI, male. 150. Sternite VII, male. 151. Tergum VIII, female. 152. Aedeagus, anterior end, cross section. 153. Tergum VIII, apex, male (setae omitted). 154. Sternum VIII, male.

dian depression on apical three-®fths; de- curved row of peg setae bordered laterally pression elliptical and longer than wide; de- by long, stout, darkly pigmented seta. pression with ground sculpturing and with Sternum VIII (®g. 128) with broad, deep small group of 3±7 peg setae (®gs. 123±126) median emargination; margins of emargina- on disc; posterior margin with broad, evenly tion moderately sinuate and converging to curved, moderately deep emargination; sur- base and with surface adjacent to basal two- face adjacent to emargination with evenly ®fths beveled; basal margin narrowly round- 46 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 155±160. Lacvietina paricosta. 155. Sternite VII, male. 156. Tergum VIII, male (setae omitted). 157. Sternum VIII, male (apical setae missing). 158. Aedeagus, ventral. 159. Aedeagus, lateral. 160. Sternite V, male. ed; sternum with short, curved, medially known from the northern Vietnamese prov- notched ridge on inner surface adjacent to inces of Hatinh, Quang Binh, and Quang base of sternal emargination. Nam. It was collected at elevations of 220± Tergum VIII (®gs. 132, 133) with shallow 1150 m from wet leaf litter near streams in to moderately deep, rounded emargination Hatinh Province and from ¯ood debris in between median pair of apical lobes; emar- Quang Nam Province. gination wider than deep. ETYMOLOGY: The name of this species is Aedeagus asymmetrical (®gs. 121, 127); from the Latin for coppery (cuprinus). parameres with lateral margins slightly sin- MATERIAL EXAMINED: One hundred ®fteen uate, right (in ventral view) more sinuate specimens. The holotype and 68 male and 46 than left; left paramere (in ventral view) female paratypes. Six males were dissected shorter and slightly wider than right; left par- for the genitalic features and a male and fe- amere slightly curved medially and slightly male were disarticulated. curved ventrally at apex; right paramere more strongly curved medially and ventrally Lacvietina aurora, new species near apex. Figures 134±146 FEMALE: Tergum VIII (®g. 129) with four HOLOTYPE: Male. ``VIETNAM: Hatinh Pr., RaoÁ apical lobes. An 13 km W Huang Son, rt. 8 7 km S NuoÁc DISTRIBUTION AND HABITAT: This species is SoÃt village 18Њ21ЈN, 105Њ15ЈE, 1150 m IV-25± 2004 HERMAN: MEGARTHROPSINI 47

Figs. 161±167. Lacvietina punctatissima. 161. Sternum VIII, male. 162. Sternite VII, male. 163. Sternite V, median region, male. 164. Sternite VI, male. 165. Sternum VIII, male. 166. Aedeagus, lateral. 167. Aedeagus, ventral.

1998, L. Herman, litter nr. drying stream''. On holotype (2 males, 1 female, AMNH; 2 males, long-term loan to the American Museum of 1 female, IEBR; 1 male, MHNG); same locality Natural History (AMNH), to be deposited in the and collector as holotype: IV-20±1998 (1 male, Institute of Ecology and Biological Resources, AMNH; 1 male, IEBR); IV-28±1998 (2 fe- Hanoi, Vietnam (IEBR). males, AMNH; 1 female, IEBR). Paratypes fol- PARATYPES: Seven males, 5 females. Same data as lowed by the abbreviation ``IEBR'' are on long- 48 AMERICAN MUSEUM NOVITATES NO. 3430

term to the American Museum (AMNH) and tion, or ground sculpturing; depression abrupt- will eventually be deposited in the Institute of ly demarcated basally; lateral margins of de- Ecology and Biological Resources (IEBR). pression slightly curved but approximately DIAGNOSIS: The males of Lacvietina au- parallel to one another; posterior margin of rora are distinguished from those of the other sternite with broad, shallow emargination. two species of the genus by the sinuous row Sternite VII (®g. 138) with moderately of peg setae near the posterior margin of ster- deep median depression on apical two-thirds; nite VII (®g. 138), the evenly curved emar- depression broad, slightly longer than wide gination of sternite VII (®g. 138), the small and ovoid; surface of depression shining, cluster of 7±9 peg setae on the midbasal por- with patch of minute setae and punctures on tion of sternite VII, the polished median de- lateral portion, and with small basal cluster pression of sternite VI, and the absence of of 7±9 peg setae; posterior margin of sternite carinae on sternite V. with broad, deep, evenly curved emargina- Females of L. aurora are separated from tion; emargination bordered by sinuous row L. cuprina by the single row of punctures on of peg setae; row of peg setae bordered lat- the elytral epipleuron and the ®ve apical erally by long, stout darkly pigmented seta. lobes of tergum VIII (®g. 140), from L. co- Sternum VIII (®g. 137) with broad, deep piosa and L. punctatissima (and probably L. emargination; lateral margins of emargina- paricosta) by the strongly shining interpunc- tion sinuate and converging to base and with tational surface of tergite VII, and from L. surface adjacent to basal third beveled; basal punctatissima by the presence of ®ve apical margin of emargination truncate, beveled, lobes of tergum VIII (®g. 140). and with small median notch. DESCRIPTION: Length 3.0±3.5 mm; width Tergum VIII (®g. 136) with moderately 1.1±1.4 mm. Body pale to dark reddish deep, rounded emargination between median brown. Head darker than remainder of body, pair of apical lobes; emargination slightly elytra darker than pronotum and abdomen. wider than deep. Pronotum with dark disc and paler margins. Aedeagus (®gs. 134, 135) asymmetrical; Antennae and legs reddish brown. parameres with lateral margins sinuously Pronotal punctation extending to anterior curved; left paramere (in ventral view) wider margin and without lateroapical impunctate and shorter than right and with apical portion spot mesad of anterolateral angle, but some straight; apex of right slightly curved medi- individuals with slightly less dense puncta- ally and with apex slightly bent ventrally. tion on small lateroapical spot. FEMALE: Tergum VIII (®g. 140) with ®ve Elytra with posterolateral angle moderate- lobes on posterior margin; median lobe ta- ly to slightly (®g. 143) produced; epipleuron with row of punctures near ventral edge and pered apically and with small seta at apex. without row of punctures in median region. DISTRIBUTION AND HABITAT: The species is Wings present and fully developed. known from Hatinh Province in northern Vi- Tergite VII without ground sculpturing be- etnam at 1150 m. It was collected near the tween punctures or feeble if present; surface end of the dry season from leaf litter near a polished. slow ¯owing, drying stream. MALE: Sternite V with shallow median de- ETYMOLOGY: The name of this species is pression on apical ®fth; depression increas- from the Latin for dawn (aurora) and refers ingly deep and wide apically; basal and lat- to the fact that to reach the site of collection eral margins of depression with cluster of I had to leave camp before sunrise to have a short setae; midapical surface of depression suf®cient amount of time to collect. polished; posterior margin of sternite with MATERIAL EXAMINED: Thirteen specimens. shallow median emargination. Male holotype and 7 male and 5 female para- Sternite VI (®g. 139) with moderately deep types. median depression on apical three-fourths; Three males and one female were dissect- depression broad, about as wide as long; de- ed for genitalic features. One male was dis- pression polished and without setae, puncta- articulated. 2004 HERMAN: MEGARTHROPSINI 49

Lacvietina copiosa, new species lateroapical impunctate, polished spot mesad Figures 147±154 of anterolateral angle; spots separated or connected by narrow to feebly developed, im- HOLOTYPE: Male. ``VIETNAM: Hatinh Pr., RaoÁ An 13 km W Huang Son, rt. 8 7 km S NuoÁc punctate band. SoÃt village 18Њ21ЈN, 105Њ15ЈE, 220 m IV-26± Elytra with posterolateral angle moderate- 1998, litter near stream, L. Herman collector''. ly to slightly produced; epipleuron with row On long-term loan to the American Museum of of punctures near ventral margin and without Natural History (AMNH), to be deposited in the row of punctures in median region. Wings Institute of Ecology and Biological Resources, present and fully developed. Hanoi, Vietnam (IEBR). Tergite VII with strong ground sculpturing PARATYPES: Thirty-six males, 39 females. Same between punctures; surface shining dully. data as holotype (2 males, 3 females, AMNH). MALE: Sternite V with moderately deep, mi- Same locality and collector as holotype: 350 m, litter near drying forest stream, V-3±1998 (5 dapical depression; depression with cluster of males, 6 females, AMNH; 4 males, 5 females, short dense setae medially and basally, but pol- IEBR; 2 males, 2 females, MHNG); 300±350 ished and without setae midapically; posterior m, litter near forest stream, V-1±1998 (1 male, margin with shallow median emargination. 2 females, AMNH; 1 male, 3 females, IEBR); Sternite VI (®g. 149) with broad, deep, 450 m, litter near seep along river, V-2±1998 (1 polished, median depression; depression male, 1 female, AMNH; 1 male, IEBR); 1150 without setae, punctures, or ground sculptur- m, IV-28±1998 (1 male, 1 female, AMNH; 2 ing; depression abruptly demarcated basally; males, IEBR); IV-25±1998, 1150 m, litter nr. width of depression about twice length; lat- drying stream (1 male, AMNH). Vietnam: eral margins of depression diverging poste- Quang Nam Prov.: 25 km (by air) S.W. Tra My, 940 m, IV-12±1999, 15Њ39ЈN, 108Њ02ЈE, ¯ood riorly; posterior margin broadly and shallow- debris, L. Herman (4 males, 1 females, AMNH; ly emarginate. 5 males, 2 females, IEBR; 2 males, 2 females, Sternite VII (®g. 150) with broad, mod- MHNG); 15Њ12Ј14ЉN, 108Њ02Ј13ЉE, 850 m, IV- erately deep, median depression; depression 14±1999 (2 males, 6 females, AMNH; 2 males, slightly longer than wide; surface of depres- 5 females, MHNG). Paratypes followed by the sion glabrous midapically and with remain- abbreviation ``IEBR'' are on long-term to the der shining dully; depression with dense American Museum (AMNH) and will eventu- cluster of 26±30 peg setae; cluster of peg se- ally be deposited in the Institute of Ecology and tae denser basally than apically; cluster of Biological Resources (IEBR). peg setae on posteriorly sloping plateau; ster- DIAGNOSIS: The males of Lacvietina copio- nite with irregular row of peg setae adjacent sa can be separated from those of the other to posterior margin; apical row of peg setae four species by the large cluster of 26±30 peg separated medially and bordered laterally by setae on the disc of sternite VII (®g. 150), long, darkly pigmented spinelike seta; pos- the medially interrupted row of peg setae terior margin with broad, moderately deep, along the posterior margin of sternite VII sinuate emargination; emargination as wide (®g. 150), and the wide parameres (®g. 147). as median depression and with middle more Females can be separated from those of deeply emarginate; median third of posterior the other species by the single row of punc- margin without setae, polished, and more tures on the elytral epipleuron and the strong lightly sclerotized. interpunctational ground sculpturing of ter- Sternum VIII (®g. 154) with broad, deep gum VII; the ®ve apical lobes of tergum VIII median emargination; lateral margin of emar- (®g. 151) will separate it from L. cuprina gination sinuously curved; emargination nar- (®g. 129) and L. punctatissima (®g. 171). rower basally than apically; surface of ster- DESCRIPTION: Length 3.1±3.7 mm; width num adjacent to basal half of emargination 1.3±1.4 mm. Body dark to pale reddish slightly beveled and shallowly impressed; brown. Head darker than remainder of body, base of emargination moderately beveled, elytra darker than pronotum and abdomen. shallowly curved, and with pair of setae on Pronotum with dark disc and paler margins. each side of midline; base of emargination Antennae and legs reddish brown. more or less truncate, and with small, median Pronotum with small to moderately large notch. 50 AMERICAN MUSEUM NOVITATES NO. 3430

Tergum VIII (®g. 153) with moderately DIAGNOSIS: The males of this species can deep, rounded emargination between median be separated easily from the others of the ge- pair of apical lobes; emargination wider than nus by the pair of posteriorly diverging, me- deep. dial carinae on sternite V (®g. 160) and the Aedeagus asymmetrical (®gs. 147, 148); deep incision between the median pair of parameres each with small, subapical process lobes of tergum VIII (®g. 156). The para- on ventral surface (viewed laterally); lateral meres of L. paricosta (®g. 158) are more margins sinuously curved; left paramere (in slender than are those of L. aurora (®g. 134) ventral view) wider and shorter than right; and there are fewer peg setae along the pos- apex of left paramere nearly straight; apex of terior margin of sternite VII (®g. 155) than right paramere curved medially. for L. aurora (®g. 138) and L. cuprina (®g. FEMALE: Tergum VIII (®g. 151) with ®ve 123) and fewer medial peg setae in L. pari- lobes on posterior margin; median process costa (®g. 155) than L. copiosa (®g. 150). short or long, tapered apically, and with DESCRIPTION: Length 3.5 mm; width 1.2 small seta at apex. mm. Color reddish brown. Head darker than DISTRIBUTION AND HABITAT: This species is remainder of body. Pronotum with darker known from the northern Vietnamese prov- reddish brown median region and pale red- inces of Hatinh and, farther to the south, dish brown to yellowish brown lateral and Quang Nam. It was collected at elevations of basal margins. Abdominal segments darker 220±940 m from wet leaf litter near streams basally than apically. in Hatinh Province and from ¯ood debris in Pronotum with moderately large, impunc- Quang Nam Province. tate, polished spot adjacent to anterior mar- ETYMOLOGY: The name of this species is gin and mesad of anterolateral angle; im- from the Latin for furnished abundantly (co- punctate spots connected by similarly im- piosus) and refers to the large number of peg punctate and polished narrow band along an- setae on sternite VII of the males. terior margin. DISCUSSION: Some males may have some- Elytra with posterolateral angle moderate- what fewer or somewhat more peg setae on ly strongly to weakly produced; epipleuron sternite VII than indicated in the description. with one row of punctures adjacent to ventral The numbers in the description are based on edge. Wings fully developed. Tergum with a sampling of the males; no attempt was moderately strong ground sculpturing be- made to count the peg setae on all the males tween punctures; surface dully shining. examined. The point is that L. copiosa has MALE: Sternite V (®g. 160) with medial signi®cantly more peg setae than the other pair of posteriorly diverging carinae; carinae species. larger apically than basally; median region MATERIAL EXAMINED: Seventy-six speci- without punctures or setae, but with cluster mens. Male holotype and 36 male and 39 of minute, stout, cuticular processes between female paratypes. and at base of carinae (not shown in ®gure Four males and one female were dissected and visible only at high magni®cation of for genitalic features and a male and female compound microscope; visible when viewed were disarticulated. in glycerine with stereo microscope as pale spots); posterior margin feebly emarginate. Lacvietina paricosta, new species Sternite VI with broad, shallow median de- Figures 155±160 pression; depression nearly as long as segment and about a quarter of width; depression HOLOTYPE: Male. ``Thailand, Chiangmai Doi Yu- without punctures or setae on most of surface thep [ϭ Suthep?], 1500 m 13.11.1995 leg. P. but with one stout seta near lateroapical mar- Wunderle''//``Paratachinus aurora Herm. det. gin; posterior margin feebly emarginate. M. SchuÈlke, 2002''. Specimen missing right antenna. Deposited in collection of Michael Sternite VII (®g. 155) with broad, shallow, SchuÈlke (MSC). indistinctly margined, median depression; PARATYPE: 1 male. Malaysia: Cameron Highl. Gn. depression with weak ground sculpturing and Jasar, 1400±1600 m, 6.IV.1990, leg. A. Riedel small median cluster of 11±12 peg setae; (1 male, MSC). posterior margin with broad, evenly curved, 2004 HERMAN: MEGARTHROPSINI 51 moderately deep emargination; emargination the disc of sternite VII (®g. 150) and those with row of peg setae medially; peg setae of of L. punctatissima have a small cluster (®g. posterior margin separated, not contiguous; 162). Other salient features of the males of peg seta of posterior margin bordered later- L. punctatissima include the even, medially ally by row of short setae; lateral edge of row separated marginal row of peg of sternite VII of setae and peg setae marked by long, stout, (®g. 162), and the broad, moderately deep, darkly pigmented seta. polished depression on sternite VI. Lacvieti- Sternum VIII (®g. 156) with broad, deep na punctatissima, with one row of punctures median emargination; margins of emargina- near the ventral edge of the elytral epipleu- tion strongly sinuate and converging to base; ron, is separated from L. cuprina, which has surface adjacent to basal ®fth of emargina- two rows, one near the ventral margin and tion narrowly beveled; basal margin broad, one along the length of the middle (®g. 195). truncate, beveled, and medially notched. The females of L. punctatissima have four Tergum VIII (®g. 156) with deep emargi- apical lobes on sternite VIII (®g. 170) and nation between median pair of lobes; emar- those of L. copiosa (®g. 151) and L. aurora gination much deeper than wide. (®g. 140) have ®ve. Aedeagus (®gs. 158, 159) asymmetrical; DESCRIPTION: Length 3.3±3.6 mm; width parameres with lateral margins moderately 1.3 mm. Color reddish brown to dark reddish sinuate; right paramere (in ventral view) less brown. Head darker than remainder of body, strongly sinuate than left; left paramere (in elytra darker than abdomen and about as dark ventral view) shorter and wider than right. as median portion of pronotum. Pronotum FEMALE: Unknown. with dark disc and paler margins. Abdomen DISTRIBUTION AND HABITAT: The species is evenly pigmented. known from Cameron Highlands in West Ma- Pronotum with moderately large, impunc- laysia and the Chiang Mai region of Thailand. tate, polished spot adjacent to anterior mar- ETYMOLOGY: The name of this species is gin and mesad of anterolateral angle; im- from the Latin for equal (par) and rib (costa) punctate spots connected by similarly im- and refers to the pair of ridges or carinae on punctate and polished narrow band along an- sternite V of the males. terior margin. MATERIAL EXAMINED: Two specimens. Elytra with posterolateral angle moderate- Male holotype and paratype. ly strongly to weakly produced; epipleuron Both males were dissected for abdominal with one row of punctures adjacent to ventral and genitalic features. No specimens were edge. Wings fully developed. disarticulated. Tergum VII with moderately strong ground sculpturing between punctures; sur- Lacvietina punctatissima (Hayashi), new face dully shining. combination MALE: Sternite V (®g. 163) with arc- Figures 161±170 shaped median depression; depression about half of length and about one-quarter of width Paratachinus punctatissimus Hayashi, 1991: 48. of sternite; basal and medial portion of de- Type locality: Taiwan: Taipei Hsien, Mt. pression with dense cluster of small, spini- Yangming. Holotype, female, deposited in col- form setae; apical edge of depression strong- lection of T. Shibata; not examined. Tachinus punctatissimus (Hayashi): Herman, ly shining and without setae; posterior mar- 2001a: 978. gin with broad, shallow emargination. Sternite VI (®g. 164) with broad, curved DIAGNOSIS: The pronotum of L. punctatis- median depression; depression about two- sima has a moderately large, polished, im- thirds of length and about one-third of width punctate spot adjacent to the anterior margin of sternite; depression with minute setae and near the anterolateral angle. Only L. copiosa punctures along perimeter but absent medi- has a similar, but smaller spot; the pronotal ally and with long seta near lateroapical mar- punctation of L. cuprina and L. aurora gin; posterior margin with broad, shallow reaches the anterior margin. The males of L. emargination. copiosa have a large cluster of peg setae on Sternite VII (®g. 162) with broad, shallow, 52 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 168±170. Lacvietina punctatissima. 168. Sternum VIII, female. 169. Spermatheca. 170. Ter- gum VIII, female. indistinctly margined, median depression; description of this species on one female and pression with weak ground sculpturing, scat- placed it in Paratachinus Blackwelder, 1952. tered minute setae, and small, median cluster I did not examine the type, but the characters of 7±13 peg setae; posterior margin with published by Hayashi (1991), including the broad, slightly sinuate, moderately deep emar- glabrous spots on the anterior margin of the gination; posterior margin with row of medi- pronotum and features of the tergum and ally separated peg setae in emargination; lat- sternum VIII, are shared by the females stud- eral edge of cluster of peg setae marked by ied in the present work. The males examined long, stout, darkly pigmented seta. herein comprise one species and the males Sternum VIII (®g. 161) with broad, deep and females share features of the distribution median emargination; margins of emargina- of the pronotal punctation, so I conclude they tion strongly sinuate and converging to base are conspeci®c. In addition to the coarse ce- and with surface adjacent to basal two-thirds phalic, pronotal, and elytral punctation, spec- beveled; basal margin broad, truncate, bev- imens of P. punctatissimus have a weakly eled, and medially notched. re¯exed anterolateral margin of the head and, Tergum VIII (®g. 165) with shallow emar- most importantly, a deep metasternal pit (see gination between median pair of apical lobes; ®g. 146); these features support its placement emargination wider than deep. in Lacvietina, and it is so transferred herein. Aedeagus (®gs. 166, 167) asymmetrical; The postocular ridge is obsolete to poorly de- parameres with lateral margins strongly sin- veloped and evident as a sharper curve in the uate; right paramere (in ventral view) more even curvature from the dorsal surface ven- strongly sinuate than left; left paramere (in trally. ventral view) shorter and wider than right. DISTRIBUTION AND HABITAT: Lacvietina FEMALE: Tergum VIII (®g. 170) with four punctatissima (Hayashi) is restricted to Tai- apical lobes. wan, where it has been collected in ®ve DISCUSSION: Hayashi (1991) based his de- Hsiens, four in the north (Taitung, Ilan, Nan- 2004 HERMAN: MEGARTHROPSINI 53 tou, and Taipei) and one in the south (Kao- Smetana, sifting moist plant debris in forest hsiung). The species has been collected at el- ¯oor depressions, subtropical, mostly broad- evations of 550±1800 m. Most (perhaps all) leaf evergreen forest [T184] (1 male, ASC). of the specimens were collected from plant Dissections of eight males and ®ve fe- debris in wet to very wet habitats, both near males were examined for features of the ab- ¯owing water and from moist spots on the domen, genital segments, and genitalia. One forest ¯oor. The species has been collected disarticulated male and female were exam- along with species of such wet-habitat genera ined. as Stenus, Dianous, Derops, and Atanygna- The holotype, not examined for this study, thus. is from Taipei Hsien. MATERIAL EXAMINED: Thirty-®ve specimens: 15 males, 20 females. Taiwan: Ilan Lacvietina sp. A Hsien: Shen-Mi Lake, 24Њ22Ј43ЉN, 121Њ44Ј 12ЉE, 1110 m, 10.V.1995, A. Smetana, sift- A sixth species was examined that differs ing lea¯itter and other moist debris under from the preceding ®ve. It is represented by shrubs/trees near lake [T177] (7 females, only one female and so is not named because ASC). [Hsien cited as ``Illan'']: Near Chih- it is often impossible to identify the females tuan, July 23, 1978, Y. Shibata leg. (1 male, or to associate them with males by means CNC); July 24, 1979, Y. Shibata leg. (1 fe- other than collecting site. The species is dis- male, CNC). Kaohsiung [Hsien]: Tengshih, tinguished from all the others in the genus 1400 m, 21±23.VII.2000, leg. H. Sugaya by the absence of medial pronotal punctation. (1 male, 1 female, MSC); Tengchih, 1400 m, Tergum VIII (of female) has ®ve apical 21±23.VII.2000, leg. H. Sugaya (1 male, lobes, as do L. aurora and L. copiosa. MSC). Nantou [Hsien]: Chosen-lintao, 1800 DESCRIPTION: Female. Length 4.0 mm; m, 11.X.1998, leg. H. Sugaya (2 males, 1 width 1.5 mm. Color reddish brown. Head female, MSC); Tehuashe, 900 m, 25.X.2001, nearly black and darker than remainder of H. Sugaya leg. (1 male, MSC). Taitung body. Pronotum with pale reddish brown lat- Hsien: Hsinkangshan above Chengkang, 750 eral and basal margins. m, 18.IV.1998, A. Smetana, sifting moist Head with a few scattered ®ne punctures plant debris along edges of abandoned forest and with scattered punctulation; surface pol- road in subtropical, mostly broadleaf ever- ished. Antenna reaching nearly to posterior green forest [T182] (2 males, 5 females, margin of elytra. ASC); 800 m, 17.IV.1998, A. Smetana, Lise Pronotum with coarse, moderately dense Robillard, sifting of wet plant debris at base punctation on lateral third and along basal of rock wall over which thin layer of water third; median third and anterior margin with was running [T180] (1 male, 1 female, ASC); scattered punctulation and ®ne punctures and 550±600 m, 22.IV.1998, A. Smetana, sifting surface strongly shining to polished. of soaking wet plant debris accumulated at Elytra with posterolateral angle not pro- base of dripping wet rock wall and collected duced; epipleuron with one row of punctures together with Atanygnathus sp. [T187] (2 adjacent to ventral edge. males, 1 female, ASC); 750 m, 21.IV.1998, Tergum VIII with ®ve apical lobes. A. Smetana, sifting of moist plant debris DISCUSSION: In the key to species for La- along edges of abandoned forest road in sub- cvietina, this species will run to couplet 7, tropical, mostly broadleaf evergreen forest where it can be separated from both L. au- [T185] (1 female, ASC); 800 m, 26.IV.95, A. rora and L. copiosa by the impunctate me- Smetana, sifting moist to wet debris along dian third of the pronotum. edges of small creek [T167] (1 male, 1 fe- Although the posterolateral elytral margin male, ASC); 800 m, 27.IV.95, A. Smetana, is rounded, not emarginate as is the case for sifting wet plant debris at base of rock wall others of the Megarthropsini, the lateral mar- over which thin layer of water was running gin of the head is re¯exed from above the and collected together with Derops lisae and antennal insertion anteriorly, the postocular species of Stenus and Dianous [T168] (2 ridge is modestly developed, the neck is males, 1 female, ASC); 900 m, 19.IV.98, A. present, and the cephalic, pronotal, and ely- 54 AMERICAN MUSEUM NOVITATES NO. 3430 tral punctation is coarse. The species is in- tana, 1995). The tribe with only three genera cluded in Lacvietina because of the presence and four species was still minuscule. of the intermesocoxal metasternal pit. In 1998 and 1999, while collecting in Vi- DISTRIBUTION: Indonesia: N. Sumatra, Ko- etnam, I found many small, short and wide, tacane, Gn. Sinabung, 2000 m, 7±8.X.1990, densely punctate, odd, omaliine-like speci- leg. A. Riedel (1 female, MSC). mens that turned out to represent three new MATERIAL EXAMINED: One female. species of an unknown genus of the Megar- thropsini and that are herein placed in a new DISCUSSION genus, Lacvietina. To describe the new genus and understand its association with others of The Megarthropsini was ®rst proposed as the tribe I requested specimens of the named a ``n. group'' (but without characters speci®c Megarthropsini from colleagues who had to it) for one monotypic genus, Megarthrop- collected in southern and eastern Asia from sis, from Singapore (Cameron, 1919: 231). India and Nepal to Taiwan. As a result, in Megarthropsis was stated to be ``undoubt- addition to the 3 new Vietnamese species, 10 edly . . . closely related to the Tachyporini'' others belonging to the new genus or to the and further on, ``related to the Trichophyini three previously described genera were dis- and Tachyporini'' (Cameron, 1919: 231± covered. The new species of Nepaliodes ex- 232). Later, Cameron (1921: 349, 355, 406) tends the range of the genus from Nepal and speci®cally included the tribe in the Tachy- India to Thailand and China. Peitawopsis, porinae and provided a few characters to sep- now with three species, is con®ned to the arate it from the Tachyporini. That subfamil- mountains of Taiwan (Herman and Smetana, ial assignment was followed by Scheerpeltz 2002). The collections of Megarthropsis re- (1933: 1478), but there was no further dis- sulted in a minor explosion of species. The cussion or characterization of the genus (or genus grew from one species to eight and the tribe) or addition of other genera or species geographical range now reaches Java. Be- until 1983, nor were there any published il- cause six of the seven new species of Me- lustrations of characters or the body form. garthropsis were collected on Mt. Kinabalu The genus and the tribe remained somewhat and in the Crocker Range, it seems likely that a mystery group. many more will be found as other areas are Coiffait (1977: 272±275) described Ne- explored. Since the species of Megarthrop- paliodes with one species from Nepal and sini all live in leaf litter, the likelihood is illustrated the antenna, maxillary palp, la- great of ®nding additional taxa of the Me- brum, sternum VIII (with parts of VII and garthropsini throughout the region from In- IX) of the male, the aedeagus, and outline of dia and China to Taiwan and through the In- the body. Although Coiffait assigned the ge- domalay peninsula and Indonesia, particular- nus to the Tachyporinae, he wrote nothing ly Sumatra, Java, and Borneo. about its tribal af®liation. Although the Megarthropsini is in the Tach- Smetana (1983a) published a de®nition of yporinae, the relationships of the tribe to the the Megarthropsini, moved Nepaliodes into it, others is unclear, as are the relationships of and added new characters for both Nepaliodes the genera within the tribe. Currently six and Megarthropsis. He also introduced nu- tribes are included in the subfamily (Cordo- merous and detailed line illustrations and banini, Deropini, Megarthropsini, Mycetopor- scanning electron microscope photographs of ini, Tachyporini, and Vatesini), but no derived salient features of both genera, included a larval (Ashe and Newton, 1993) or adult fea- habitus for both species, and extended the tures have been found that clearly de®ned it. range of Nepaliodes to India and Megarthrop- A seventh tribe, the Symmixini, is a synonym sis to Borneo. Smetana's work permitted rec- of the Tachyporini (SchuÈlke, 2003). ognition of the tribe without reference to types Smetana (1983a: 143), in a detailed dis- of the genera for the ®rst time. cussion of the subfamilial assignment of the A third monotypic genus, Peitawopsis, Megarthropsini, discussed its relationship to from Taiwan was added (Smetana, 1992) and the seven subfamilies of the Tachyporine later a second species of it was named (Sme- group (see Herman, 2001a: v, for list) and 2004 HERMAN: MEGARTHROPSINI 55 concluded that derived features of abdominal TABLE 1 segments VII±X and the aedeagus supported Character Matrix its inclusion in the Tachyporinae. The relationship of the Megarthropsini within the subfamily was not addressed. Smetana (1983a: 143) argued that Megarthropsis and Nepaliodes should be combined in the Me- garthropsini because both have the antenna inserted under the raised and explanate lateral margin of the frons and possess exocrine glands and a dorsal, postocular, cephalic carina. These features are unique in the Tach- yporinae. To examine the phylogenetic relationships of the genera of Megarthropsini and assess the tribe's position among the tribes of the Tachyporinae, examples of genera of the Deropini (Derops divalis [Sanderson]), Me- garthropsini (all genera and species), Myce- toporini (Lordithon fungicola Campbell) Tachyporini (Coproporus ventriculus (Say), Sepedophilus castaneus (Horn), Tachinus ®mbriatus Gravenhorst, Tachyporus califor- nicus Horn), and Vatesini (Vatesus clypeatus (Wasmann) and V. splendidus [Wasmann]) were dissected. No specimens of the Cordo- banini were available for study. Cordobanus, from Mexico and about which little has been written, is monotypic. The following characters were employed in the analysis, and the distribution of the character states is summarized in the matrix (table 1). Character 0. Surface punctation (head, pronotum, elytra): 0 (®ne); 1 (medium); 2 (coarse). CI/RI ϭ 100/100. The cephalic, ples include Mycetoporus bipunctatus Camp- pronotal, and elytral dorsal surfaces of most bell and Mycetoporus lucidulus LeConte species of the Tachyporinae are smooth and (Campbell, 1991) with more evenly distrib- polished to shining dully with weak to strong uted, sparse pronotal punctation, and Boli- microsculpturing. The surface has sparse to topunctus punctatissimus Campbell (Camp- dense, ®ne micropunctation to larger, mod- bell, 1993a: 31, 64) with dense, minute pro- erately coarse punctation; less commonly the notal punctulation. Species of Parabolitobius punctation is large and coarse. The pronotal and Bolitopunctus have moderately dense punctation may be more or less uniform or and coarse elytral punctation in addition to arranged marginally, submarginally, or dis- one or more serial rows (Campbell, 1993a: cally and is often in rows. The elytral punc- 29±34, 65; Li et al., 2000b). By contrast, tation may be uniform and sparse to dense, Derops (Deropini) has dense, moderately arranged in serial rows, or both. For many coarse punctation, and the genera of the Me- Mycetoporini, the surface is smooth and pol- garthropsini (®gs. 1, 171, 183, 188, 194) all ished and the few punctures are arranged in have prominent, dense, coarse, deep puncta- serial rows, but some species have more uni- tion. Some species currently included in form and denser pronotal and/or elytral punc- Tachinus (Tachinoderus) have dense, coars- tation (M. SchuÈlke, correspondence). Exam- er, cephalic, pronotal, and elytral punctation, 56 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 171±176. Megarthropsis smetanai. 171. Head, pronotum, and elytra. 172. Head and pronotum, left lateral side. 173. Head, dorsal, seta in deep puncture. 174. Head, dorsal, seta in shallow depression. 175. Pronotum, seta in deep puncture. 176. Pronotum, setae in shallow depression. 2004 HERMAN: MEGARTHROPSINI 57

Figs. 177±182. Megarthropsis smetanai. 177. Pronotum, seta in shallow depression. 178. Pronotum, seta in shallow depression. 179. Elytral seta in deep puncture. 180. Elytral seta in deep puncture. 181. Elytral seta in shallow depression. 182. Abdominal seta, dorsal. 58 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 183±187. Nepaliodes variolosus. 183. Head, pronotum, and elytra. 184. Head. 185. Head, dorsal setae in deep punctures. 186. Pronotal seta in deep puncture. 187. Elytral seta in deep puncture. 2004 HERMAN: MEGARTHROPSINI 59

Figs. 188±193. Peitawopsis monticola. 188. Head, pronotum, and elytra. 189. Head. 190. Head, dorsal seta in deep puncture. 191±193. Pronotal setae in deep punctures. 60 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 194±199. Lacvietina cuprina. 194. Head. 195. Elytron, left, lateral view. 196, 197. Pterothorax, ventral view. 198, 199. Head, dorsal setae in deep punctures. 2004 HERMAN: MEGARTHROPSINI 61

Figs. 200±203. Lacvietina cuprina. 200, 201. Pronotum, setae in deep punctures. 202. Elytral seta in deep puncture. 203. Elytral seta, enlarged view of ®gure 202. for example, Tachinus (Tachinoderus) rou- Uniquely, Derops and the Megarthropsini gemonti (Pace, 1986: 49); some other species (®g. 194) have a broad, distinct neck. in the subgenus have dense, moderately Character 2. Postocular laterodorsal sur- coarse elytral punctation, but most have ®ne, face: 0 (unmodi®ed); 1 (ridged); 2 (carinate). sparse to moderately dense punctation. The CI/RI ϭ 66/90. The postocular laterodorsal coarse, dense punctation in the Megarthrop- region of the head of most Tachyporinae sini is visually the most distinctive feature lacks a ridge or carina. Vatesus (Vatesini) shared by these tachyporines. Associated and Coproporus (Tachyporini) both have a with the punctures are apically expanded se- feebly developed, rounded ridge that gradu- tae that are uniquely present in all Megar- ally curves posteriorly from the eye. In the thropsini (®gs. 173, 175, 179, 180, 185±187, Megarthropsini the laterodorsal postocular 190±193, 198±203). region has a rounded ridge (®gs. 189, 194) Character 1. Neck: 0 (absent); 1 (pre- or strongly developed carina (®gs. 172, 184) sent). CI/RI ϭ 100/100. Most genera of the extending posteriorly from the eye. The pos- Tachyporinae lack a neck; the postocular por- tocular ridge of Lacvietina is modestly de- tion of the head simply tapers gradually from veloped (®g. 194), and on some individuals the eyes to the posterior margin of the head. of L. punctatissimus the ridge is obsolete and 62 AMERICAN MUSEUM NOVITATES NO. 3430 evident only as a break in the even curvature (®g. 188) of the Megarthropsini. The poste- from the dorsal to posteroventral surface. rior angle is acute in Megarthropsis (®g. 66), Character 3. Postocular laterodorsal ca- and both the anterior and posterior angles are rina/ridge: 0 (absent); 1 (straight); 2 (angu- acute in Nepaliodes (®g. 79). In Derops and late). CI/RI ϭ 66/88. The angulate postocular Tachinomorphus the posterior angle is acute, carina (®g. 172) is found only in Megar- and in some species of Coproporus and Va- thropsis.InNepaliodes the carina (®g. 184) tesus the anterior angle is acute. is straight and extends diagonally postero- Character 9. Postprocoxal lobe: 0 (pres- medially from the eye, and the postocular ent); 1 (absent). CI/RI ϭ 100/100. The post- ridge of Peitawopsis (®g. 189) and Lacvie- procoxal lobe is absent in Vatesus (Vatesini), tina (®g. 194) is similarly straight. The feeble Coproporus (Tachyporini), and the Myceto- ridge of Coproporus and Vatesus simply fol- porini and is present in other tachyporines. lows the curvature of the head and is coded Character 10. Elytral posterior margin: 0 herein as straight. (truncate or rounded); 1 (emarginate). CI/RI Characters 4 and 5: ϭ 50/75. In most species of the Tachyporinae 4. Clypeal lateral margin: 0 (unmodi®ed); the posterior margin of the elytra is rounded 1 (re¯exed). CI/RI ϭ 100/100. or truncate. In the Megarthropsini and in 5. Clypeal anterior margin: 0 (unmodi- some species of Vatesus (for example V. cly- ®ed); 1 (re¯exed). CI/RI ϭ 100/100. peatus, also see Seevers, 1958) it is emargin- In the Megarthropsini (®gs. 171, 184, 189, ate. The posterior margin of Peitawopsis 194) the lateral margin of the head is re¯exed (®gs. 114, 188) is broadly and evenly emar- beginning at the base of the antenna and ex- ginate and the posterolateral angle large and tending onto the clypeus. The re¯exed lateral acute. The posterior margin of Lacvietina, margin is strongly developed in Megarthrop- Megarthropsis, and Nepaliodes (®gs. 25, 69, sis (®g. 171) and Nepaliodes (®g. 184), and 91, 131) is emarginate laterally. The postero- in the latter genus even the anterior clypeal lateral emargination and acute angle are large margin is re¯exed (®g. 184). In Peitawopsis in Nepaliodes (®g. 91) and Megarthropsis (®g. 189) and Lacvietina (®g. 194) the lateral djawaensis (®g. 91), moderately large in Pei- margin is less strongly re¯exed and is only tawopsis (®g. 114), moderately large to small modestly developed in Lacvietina. Such re- in Lacvietina (®g. 131, 143) and the other ¯exed margins of the head do not occur in species of Megarthropsis (®g. 69), and are other tachyporines. Only Nepaliodes has a particularly small in some individuals of M. re¯exed anterior clypeal margin. decorata, L. cuprina, and L. copiosa. A sim- Character 6. Epistomal suture: 0 (present); ilarly emarginate elytral margin occurs in 1 (absent). CI/RI ϭ 50/50. The epistomal su- many Aleocharinae (Hammond, 1975: 169 ture (®gs. 1, 189) is well developed in most [character 12], 170). A female of an un- tachyporines, but is incomplete in Nepaliodes named species of Lacvietina (sp. A) lacks the (®g. 184) and absent in some species of Va- lateroapical emargination. tesus (Vatesini). Most genera of the Tachy- Character 11. Elytral (dorsolateral quarter) porini and Megarthropsini have a midcranial surface: 0 (convex); 1 (concave). CI/RI ϭ suture extending posteriorly from the middle 100/100. The elytra of Peitawopsis differ sig- of the epistomal suture (®gs. 1, 189). It is ni®cantly from those of the other species of lacking in Nepaliodes and short to rudimen- the Megarthropsini and other Tachyporinae. tary in the other three genera. The midcranial In Peitawopsis (®g. 188) the dorsal elytral suture was not coded in the matrix. surface is depressed and broadly concave be- Characters 7 and 8: ginning at the lateral margin and extending to 7. Pronotal posterolateral angle: 0 (round- about the lateral third or quarter; the remained); 1 (angulate). CI/RI ϭ 50/90. der of the surface is shallowly convex. In oth- 8. Pronotal anterolateral angle: 0 (round- er tachyporines the elytra are convex; they are ed); 1 (angulate). CI/RI ϭ 50/66. shallowly convex from the elytral suture to The anterior and posterior angles of the just beyond the lateral third and then more pronotum are rounded in most tachyporines strongly convex to the epipleural ridge. In and in Lacvietina (®g. 130) and Peitawopsis some species the surface adjacent to the ely- 2004 HERMAN: MEGARTHROPSINI 63 tral suture is more or less ¯attened. In La- in Sepedophilus and Euconosoma (Campbell, cvietina (®g. 1) and Megarthropsis (®g. 171) 1976a, 1976b) and Tachyporus (Campbell, the elytra are convex from the sutural margin 1979); in the latter two genera the parameres laterally. The lateral elytral margin of Lacvie- are narrowly separated or touching in some tina (®g. 1) and Megarthropsis (®g. 171) is species. The parameres are separated in Va- modi®ed with a narrow to moderately wide tesus (Vatesini), Coproporus (Tachyporini), epipleural gutter; the convexity of the elytra and Mycetoporini (for example Campbell, extends medially from this gutter. In Nepali- 1991, 1993a). Unique to Nepaliodes (®gs. odes (®g. 183) the epipleural gutter is wide, 74, 84), the parameres are so tightly contig- prominent, and causes the lateral margin of uous as to appear to be fused. Unique to the the elytra to be concave and strongly explan- Megarthropsini is a ventrobasal groove of the ate. The epipleural gutter of Megarthropsis median lobe. This groove is longitudinally djawaensis is wider and more distinctive and oriented and is deep in Nepaliodes (®g. 80) the lateral margin of the elytra more explanate and in most Megarthropsis (®g. 48), but is than for other species of the genus. shallow in Megarthropsis djawaensis, Pei- Character 12. Metasternal intercoxal pit: tawopsis (®g. 97), and Lacvietina (®g. 152). 0 (absent); 1 (present). CI/RI ϭ 100/100. Character 16. Sternite VII (male)Ðinter- Unique to Lacvietina is a metasternal pit ad- nal marginal lobe: 0 (absent); 1 (present). CI/ jacent to the apex of the mesosternal process RI ϭ 50/50. Sternite VII of the males of the (®gs. 146, 196, 197). In Megarthropsis the Megarthropsini (®gs. 4, 77, 95) has a large ridge surrounding the mesocoxal acetabulum to small lobe on the internal, lateral portion is large and well developed medially where of the basal margin. The basal margin of VI it is suf®ciently enlarged as to suggest a pit (®g. 3) and even V (®g. 63) is adorned with (see Smetana, 1983a: 206, ®g. 41). However, such lobes. A similar lobe is found in Tach- in Lacvietina the pit is an invagination of the inus ®mbriatus. body wall, but not in Megarthropsis. Character 17. Sternite VII (male)Ðpeg Character 13. Procoxal carina: 0 (absent); setae: 0 (absent); 1 (present). CI/RI ϭ 50/80. 1 (present). CI/RI ϭ 50/66. The medial sur- Peg setae on sternite VII of males are found face of the procoxa of both Peitawopsis and in Olophrinus (Campbell, 1993c), in many Derops (Deropini) is transversely carinate. species of Tachinus, in some species of Ni- Other tachyporines lack this carina, but it is tidotachinus (Campbell, 1993b), in the Der- present and more strongly developed in some opini, and in most Megarthropsini (®g. 4). genera of the Osoriinae and Piestinae. Within the Megarthropsini two species, M. Characters 14 and 15: djawaensis (®g. 21) and M. empusa (®g. 40), 14. ParameresÐmedial surface: 0 (sepa- lack peg setae, and in one species, M. fra- rated); 1 (contiguous); 2 (``fused''). CI/RI ϭ zerensis (®g. 45), the males also have peg 100/100. setae on sternite V. They are found in a few 15. Median lobeÐventrobasal groove: 0 species of the Mycetoporini (M. SchuÈlke, (absent); 1 (present). CI/RI ϭ 100/100. correspondence). They are lacking in most In most genera the parameres are narrowly other genera of the subfamily (various pub- to widely separated, but are medially contig- lished articles and M. SchuÈlke, correspon- uous in the Megarthropsini and Deropini, dence). and in some genera of the Tachyporini (M. Character 18. Tergum VIII (female)Ð SchuÈlke, correspondence) such as Tachinus secondary lobe of lateral lobe: 0 (absent); 1 (but are slightly separated in some species). (short and small); 2 (long and large). CI/RI The parameres are fused in Tachinomorphus ϭ 66/85. Tergum VIII of the females has two (Campbell, 1973b), separated basally (often) pairs of lobes or processes in most species of and contiguous or fused apically in Nitido- the subfamily. Typically the median pair of tachinus (Campbell, 1993b), contiguous to lobes has one or more moderately stout, pale narrowly separated in Coprotachinus (Camp- setae and the lateral pair has a long, stout, bell, 1994a), narrowly separated in Cileopo- darkly pigmented (usually nearly black) seta. rus (Campbell, 1994b) and in Coproporus On the lateral margin and situated more an- and Cilea (Campbell, 1975), and separated teriorly is another long, stout, ``black'' seta 64 AMERICAN MUSEUM NOVITATES NO. 3430 which extends from an unmodi®ed lateral (2.1), concave dorsolateral elytral surface margin, a ``bump'', a short lobe, or a long (11.1), and procoxal carina (13.1). Peitawop- lobe. Species of Tachinus either lack or have sis is the sister of the other three genera, a a bump (see ®gures in Campbell, 1973a, and clade supported by the presence of a second- Li et al., 2000a), and some have a small lobe ary lobe on the lateral surface of the lateral (for example Tachinus ®mbriatus; also see Li lobe of tergum VIII of the females (18.1, et al., 2002). Derops has the long, black se- 18.2). Lacvietina, with ®ve species, is mono- tae, but lacks the lobe or bump on the lateral phyletic based on the presence of a median margin of the segment (see Zheng, 2002: metasternal pit (12.1) and is the sister of Me- 193). Species of the Megarthropsini either garthropsis and Nepaliodes which together lack the lobe (®gs. 117) or have one that is form a clade supported by the presence of a short (®gs. 34, 129, 140) or long (®g. 81). postocular, dorsolateral cephalic carina (2.2) The preceding characters (0±18) were cod- and an angulate posterolateral pronotal angle ed (table 1) for all the species of the Megar- (7.1). Megarthropsis, with eight species, is thropsini along with one species for the My- monophyletic based on the shared presence cetoporini (Lordithon fungicola) and of the angulate postocular cephalic carina Deropini (Derops divalis), and two for the (3.2). The two species of Nepaliodes share Vatesini (Vatesus clypeatus and Vatesus the largest number of unique characters; splendidus) and Tachyporini (Tachinus ®m- these include the re¯exed anterior clypeal briatus and Coproporus ventriculus). No ef- margin (5.1), the absence of an epistomal su- fort was made to specify the relationships of ture (6.1), the angulate anterolateral pronotal the species within the genera of the Megar- margin (8.1), the ``fused'' parameres (14.2) thropsini, rather the relationship among the that are asymmetrical and strongly curved, genera of the Megarthropsini was the focus and the modi®cation of the secondary lobe of this study. of tergum VIII of the females into a long One most parsimonious tree (steps ϭ 34; slender process (18.2). The diagnosis of each CI ϭ 70; RI ϭ 89) resulted from applying clade and the distribution of the homopla- the implicit enumeration (IE) command of sious states are listed in table 2. A female of the Hennig86 phylogenetic program to the an unnamed species of Lacvietina lacks the matrix (table 1). The resulting hypothesis lateroapical elytral emargination, but the sig- (®g. 204; see table 2 for diagnosis of clades) ni®cance of this is unknown. Since only one suggests that the Deropini, Megarthropsini, such specimen has been examined, it is un- and part of the Tachyporini form a clade sup- clear if this is unique to the individual or ported by the contiguous parameres (14.1) characteristic of the species. and presence of peg setae on sternite VII Although there was no intention to examine (17.1) in the males, and that the Deropini and the tribal relationships within the Tachypori- Megarthropsini appear to be sister groups be- nae, additional conclusions can be drawn cause they share dense, moderately coarse to from the cladogram (®g. 204) and these hy- coarse punctation (0.1) and the presence of a potheses tested with more taxa that include broad neck (1.1). examples of all genera of the subfamily and The analysis supports the following hy- a representative cross section of the large gen- potheses. The Megarthropsini is monophy- era, more characters, and, importantly, one or letic; the genera share dense, coarse puncta- more outgroups selected from among repre- tion (0.2), the presence of a postocular later- sentatives of the other subfamilies of the odorsal cephalic ridge (2.1) or carina (2.2), a Tachyporine group (Herman, 2001a) along re¯exed lateral clypeal margin (4.1), an with Habrocerus magnus LeConte, an an- emarginate posterior elytral margin (10.1), a omolous species of uncertain relationship or ventrobasal groove on the median lobe generic assignment (Assing and Wunderle, (15.1), and a pair of lobes on the internal, 1995: 341). Genera such as Tachyporus and anterior margin of sternite VII of the males Sepedophilus must be included in any tribal- (16.1). Peitawopsis, with three species, is level analysis; the former is the type genus of monophyletic; the species share the presence the Tachyporini, the latter has a number of of a postocular, dorsolateral cephalic ridge unique features. The Mycetoporini is likely a 2004 HERMAN: MEGARTHROPSINI 65

Fig. 204. Hypothesis of relationships of genera in the Megarthropsini. Single most parsimonious tree (19 characters; CI ϭ 70; RI ϭ 89; steps ϭ 34). Numbers on clades correspond to diagnoses given in table 1 and discussed in text. monophyletic group, but its placement, se- adapt the Vatesus adults for life with army lected for purposes of this study to root the ants, then Coproporus and Vatesus look much cladogram, requires further study. It is notable alike. They share the derived loss of the post- that Coproporus and Vatesus fall into the procoxal lobe (9.1), the strongly infolded ely- same clade. The species of Vatesus are myr- tral epipleuron, and the weak postocular ce- mecophilus inquilines that live with army ants phalic ridge that follows the curvature of the (Ecitonini) in the Neotropical region where head. The species in these genera are highly currently 26 species are known (Seevers, convex, strongly limuloid, and have little in- 1958; Herman, 2001a). Removing from con- tegumental ground sculpturing. Coproporus sideration the autapomorphic features that and Vatesus were included in the same clade 66 AMERICAN MUSEUM NOVITATES NO. 3430

TABLE 2 pigmented than the median portion, there ap- Synapomorphic Character States pears to be two lobes, not one. Fusion of the for the Megarthropsini two lobes is also found in some species of (Homoplasious states are preceded by an asterisk) Tachinus (Campbell, 1973a, 1988; Ullrich, 1975; Li, 1995a, 1995b; M. SchuÈlke, correspondence), all Sepedophilus and Eucono- soma (Campbell, 1976a, 1976b; M. SchuÈlke, correspondence) and Tachyporus (Campbell, 1979; M. SchuÈlke, correspondence). In Tach- inus the fused lobe may have one or two fanlike clusters of setae (Campbell, 1973; Ull- rich, 1975) and fusion evidently arose multiple times. Species of Tachyporus have one or two clusters of setae (Campbell, 1979); Sepedophilus and Euconosoma have one (Campbell, 1976a, 1976b). In some species of Tachyporus and Tachinus the fused lobe is slightly emarginate medially. The fanlike cluster of setae on sternum VIII is lacking in based on larval features (Ashe and Newton, the Mycetoporini and Deropini. 1993: 283). It seems probable that the two The antennal scape of most species of the genera will be combined in the same tribe Megarthropsini is more or less cylindrical eventually; if they are, then the name of the (®gs. 189, 194). In Nepaliodes (®g. 184) the tribe would be Vatesini. Tachyporus and scape is strongly tapered from the base to the Tachinus might be in different clades and per- apex and is a useful feature for identifying haps in different tribes (M. SchuÈlke, corre- the genus. In Megarthropsis djawaensis the spondence). Herein Tachinus is suggested to scape is moderately strongly tapered toward be the basal lineage of a clade with Deropini the apex, and in M. durga, M. parca, and M. and Megarthropsini (®g. 204). If other genera smetanai (®g. 171) it is slightly tapered in related to Tachinus in fact do comprise a some individuals. The sole specimen of M. clade separate from Tachyporus, then the empusa has a slightly tapered scape. The ta- group would be named Tachinusini and that pered scape was not used in the analysis be- including Tachyporus would be Tachyporini. cause of the intergrading variability. A few other structures of interest were ex- In Nepaliodes the lateral margin of the el- amined but not employed in the phylogenetic ytra is strongly explanate (®g. 183) and the analysis. lateral edge re¯exed forming a trough or gut- In Derops the anterior margin of sternite ter. This epipleural gutter is wide, the dorsal VII has a large median lobe on the internal surface is polished, includes a few scattered edge; this lobe is probably associated with punctures, and is characteristic of the genus. secretory glands. The lobe seems to be In Lacvietina (®g. 1) and Megarthropsis (®g. unique to the genus. 171) the epipleural gutter is narrower and Sternum VIII of the females of most Tach- less dramatic, but it is still present; only in yporini has a pair of median lobes with a Megarthropsis djawaensis is it wider and fanlike cluster of long setae on the posterior more prominent and similar to the condition margin (®gs. 7, 141) of each lobe. In Nepa- in Nepaliodes.InPeitawopsis (®g. 188) the liodes (®gs. 82, 89) and one species of Me- epipleural gutter is absent; instead, the lateral garthropsis, M. smetanai (®g. 68), that pair quarter to third of the elytral surface is con- of lobes is fused into one broad lobe. Ne- cave. An epipleural gutter is absent in other paliodes retains two clusters of fanlike setae, tachyporines. but for M. smetanai the clusters have coa- Typically setae taper apically from the lesced into one. Because the lateral portions base and are often grooved (®g. 182). Evi- of the median lobe of Nepaliodes each have dently unique to the Megarthropsini are spe- a fanlike cluster of setae and are more darkly cialized setae on the dorsal surface. The api- 2004 HERMAN: MEGARTHROPSINI 67 cally expanded setae are at the edge of deep land), Guillaume-Marie de Rougemont (Lon- punctures (®gs. 173, 175, 179, 180, 185±187, dinieÁres, France), Harald Schillhammer (Na- 190±193, 198±203) and in Nepaliodes they turhistorisches Museum Wien, Austria), Mi- are scalelike (®gs. 185±187). Some have chael SchuÈlke (Berlin, Germany), andAlesÏ grooves (®gs. 173, 190, 191, 193, 199), are Smetana (Agriculture and Agri-Food Cana- shoehornlike (®gs. 200), lea¯ike (®g. 199), da, Ottawa, Ontario, Canada). or daggerlike (®gs. 192). These setae are I thank Volker Assing (Hannover, Ger- barely visible with a stereoscopic or com- many), Marc Branham (American Museum pound microscope in Nepaliodes, Megar- of Natural History), and Michael SchuÈlke thropsis, and Peitawopsis, but can be seen (Berlin, Germany) who reviewed the manu- only with a scanning electron microscope in script and suggested improvements; Eleanor Lacvietina. Smetana (1983a: 142, 143) spec- Sterling and Cal Synder of the Center for ulated that the deep punctures and expanded Biodiversity and Conservation at the Amer- setae were associated with secretions from ican Museum of Natural History for organiz- exocrine glands. Figures 198 and 201 show ing the ®eldwork; Khuat Dang Long and pores at the base of the seta from which such Nguyen Tien Hiep of the Institute of Ecology secretions may emerge. Smetana (op. cit.) and Biological Resources, Hanoi, Vietnam, further surmised that the secretions were im- for logistical support and companionship; portant for accumulation of a protective coat- andAlesÏs Smetana for his friendship and gra- ing. Such a coating is well developed in Ne- cious hospitality during my numerous visits paliodes, Megarthropsis, and Peitawopsis, to Ottawa and for our many staphylinid dis- but is lacking in Lacvietina. Presumably the cussions. Special acknowledgment is due expanded setae help to wick the chemical Michael SchuÈlke who pointed out several over the integument in the three genera with species buried in Tachinus that should be ex- large setae. In Lacvietina these specialized amined for this project. Thanks to Steve setae may be too small to disperse secretions Thurston for helping to organize the pages of over the surface. The abdomen in all four illustrations and to Sarfraz Lodhi for carry- genera lacks the specialized setae and the ing out numerous miscellaneous tasks. Three deep, coarse punctures, but the surface is anonymous reviewers made useful sugges- coated in the species of Nepaliodes. In ad- tions for improvement; I thank them. dition to the expanded setae at the edge of The material from Vietnam is based on large punctures, Megarthropsis smetanai ®eldwork supported by the National Science (and perhaps all species in the genus) has Foundation under grant no. 9870232 and the minute, beautiful, convoluted (®gs. 176, 177, Center for Biodiversity and Conservation at 178) or tubular (®gs. 174, 181) setae in slight the American Museum of Natural History. depressions between the large punctures; their function is unclear. REFERENCES As yet the Tachyporinae has not been suc- Ashe, J.S., and A.F. Newton. 1993. Larvae of Tri- cessfully de®ned as a monophyletic group; chophya and phylogeny of the tachyporine however, there are interesting, perhaps infor- group of subfamilies (Coleoptera: Staphylini- mative, characters in adults, the distribution dae) with a review, new species and character- and use of which in a phylogenetic analysis ization of the Trichophyinae. Systematic Ento- might help support the composition and re- mology 18: 267±286. lationships of the tribes and perhaps even Assing, V., and P. Wunderle. 1995. A revision of help begin de®ning a subfamily. the species of the subfamily Habrocerinae (Co- leoptera: Staphylinidae) of the world. Revue Suisse de Zoologie 102(2): 307±359. ACKNOWLEDGMENTS Blackwelder, R.E. 1952. The generic names of the beetle family Staphylinidae, with an essay on I thank the following individuals for gen- genotypy. United States National Museum Bul- erously lending specimens from their private letin 200: i±iv, 1±483. collections or from the collections in their Cameron, M. 1919. New species of Staphylinidae care: Giulio Cuccodoro and Ivan LoÈbl (Mu- from Singapore. Part II. Transactions of the En- seÂum d'Histoire Naturelle, GeneÁve, Switzer- tomological Society of London 1918: 231±246. 68 AMERICAN MUSEUM NOVITATES NO. 3430

Cameron, M. 1921. New species of Staphylinidae species. Bulletin of the National Museum of from Singapore. Part IV (Conclusion). Trans- Natural Science 4: 47±70. actions of the Entomological Society of London Campbell, J.M. 1994a. A revision of the genus 1920: 347±413. Coprotachinus Cameron (Coleoptera: Staphy- Cameron, M. 1932. The fauna of British India in- linidae: Tachyporinae). Bulletin de l'Institut cluding Ceylon and Burma. Coleoptera. Sta- Royal des Sciences Naturelles de Belgique. En- phylinidae, vol. 3. London: Taylor and Francis, tomologie 64: 25±47. xiii ϩ 443 pp. Campbell, J.M. 1994b. Cileoporus, a new genus of Campbell, J.M. 1973a. A revision of the genus rove beetles from Central and South America Tachinus (Coleoptera: Staphylinidae) of North (Coleoptera, Staphylinidae, Tachyporinae). Stud- and Central America. Memoirs of the Ento- ies on Neotropical Fauna and Environment mological Society of Canada 90: 1±137. 29(3): 125±144. Campbell, J.M. 1973b. A revision of the genus Coiffait, H. 1977. Staphilinides reÂcolteÂs au NeÂpal Tachinomorphus (Coleoptera: Staphylinidae) of par le Professeur Franz (2e partie). Bulletin de North and Central America. The Canadian En- la SocieÂteÂ d'Histoire Naturelle de Toulouse tomologist 105: 1015±1034. 112(1976): 243±275. Campbell, J.M. 1975. A revision of the genera Coiffait, H. 1982. Contribution aÁ la connaissance Coproporus and Cilea (Coleoptera: Staphylin- des Staphylinides de l'Himalaya (NeÂpal, La- idae) of America north of Mexico. The Cana- dakh, Cachemire) (Insecta: Coleoptera: Staphy- dian Entomologist 107: 175±216. linidae). Senckenbergiana Biologica 62: 21±179. Campbell, J.M. 1976a. A revision of the genus Coiffait, H. 1984. Staphylinides (Col.) de la reÂ- Sepedophilus Gistel (Coleoptera: Staphylini- gion himalayenne et de l'Inde. II. Tachypori- dae) of America North of Mexico. Memoirs of nae, Oxytelinae et Aleocharinae. Entomologica Basiliensia 9: 116±157. the Entomological Society of Canada 99: 1±89. Hammond, P. 1975. The phylogeny of a remark- Campbell, J.M. 1976b. A review of the Tachy- able new genus and species of gymnusine staph- porinae genus Euconosoma Cameron (Coleop- ylinid (Coleoptera) from the Auckland Islands. tera: Staphylinidae) with a description of a new Journal of Entomology (B) 44(2): 153±173. species from Nepal. The Coleopterists Bulletin Hayashi, Y. 1991. Notes on Staphylinidae from 30(2): 139±145. Taiwan, VI. The Entomological Review of Ja- Campbell, J.M. 1979. A revision of the genus pan 46(1): 45±51. Tachyporus Gravenhorst (Coleoptera: Staphy- Herman, L.H. 2001a. Catalog of the Staphylinidae linidae) of North and Central America. Mem- (Insecta: Coleoptera). 1758 to the end of the oirs of the Entomological Society of Canada Second Millennium. II. Tachyporine group. 109: 1±95. Bulletin of the American Museum of Natural Campbell, J.M. 1988. New species and records of History 265: 651±1066. North American Tachinus Gravenhorst (Cole- Herman, L.H. 2001b. Nomenclatural changes in optera: Staphylinidae). The Canadian Entomol- the Staphylinidae (Insecta: Coleoptera). Bulle- ogist 120: 231±295. tin of the American Museum of Natural History Campbell, J.M. 1991. A revision of the genera 264: 1±83. Mycetoporus Mannerheim and Ischnosoma Ste- Herman, L., and A. Smetana. 2002. New species phens (Coleoptera: Staphylinidae: Tachypori- of Peitawopsis and a review of the genus (Co- nae) of North and Central America. Memoirs leoptera: Staphylinidae: Tachyporinae: Megar- of the Entomological Society of Canada 156: thropsini). Special Bulletin of the Japanese So- 1±169. ciety of Coleopterology 5: 197±204. Campbell, J.M. 1993a. A revision of the genera Jordan, M. 1993. Encyclopedia of Gods. Over Bryoporus Kraatz and Bryophacis Reitter and 2,500 deities of the world. New York: Facts on two new related genera from America North of File, ix ϩ 337 pp. Mexico (Coleoptera: Staphylinidae: Tachypori- Leach, M. 1992. Guide to the gods. Santa Bar- nae). Memoirs of the Entomological Society of bara, CA: ABC-CLIO, xi ϩ 995 pp. Canada 166: 1±85. Li, L.-Z. 1995a. A revision of the genus Tachinus Campbell, J.M. 1993b. A review of the species of Gravenhorst (Coleoptera, Staphylinidae) of Ja- Nitidotachinus new genus (Coleoptera: Staphy- pan, I. The Japanese Journal of Systematic En- linidae: Tachyporinae). The Canadian Entomol- tomology 1(1): 51±72. ogist 125: 521±548. Li, L.-Z. 1995b. A revision of the genus Tachinus Campbell, J.M. 1993c. A review of the genus Gravenhorst (Coleoptera, Staphylinidae) of Ja- Olophrinus Fauvel (Coleoptera: Staphylinidae: pan, II. The Japanese Journal of Systematic En- Tachyporinae) with descriptions of three new tomology 1(2): 201±216. 2004 HERMAN: MEGARTHROPSINI 69

Li, L.-Z., and N. Ohbayashi. 1996. The genus tera, Staphylinidae). Revista Brasileira de En- Tachinus (Coleoptera, Staphylinidae) from the tomologia 8: 181±202. Himalayas with descriptions of three new spe- Smetana, A. 1983a. The tribe Megarthropsini cies. Japanese Journal of Entomology 64(1): Cameron (Insecta: Coleoptera: Staphylinidae). 151±162. Senckenbergiana Biologica 64(1/3): 141±155. Li, L.-Z., M.-J. Zhao, and N. Ohbayashi. 2002. A Smetana, A. 1983b. The status of the staphylinid new species of the genus Tachinus (Coleoptera, genera Derops Sharp and Rimulincola Sander- Staphylinidae) from Mt. Emei, southwest Chi- son (Coleoptera). Entomologica Scandinavica na. Special Bulletin of the Japanese Society of 14(3): 269±279. Coleopterology 5: 205±208. Smetana, A. 1988. Revision of the tribes Quediini Li, L.-Z, M.-J. Zhao, and M. Sakai. 2000a. Two and Atanygnathini. Part II. The Himalayan re- new species of the genus Tachinus (Coleoptera, gion (Coleoptera: Staphylinidae). Quaestiones Staphylinidae) from Zhejiang Province, east Entomologicae 24(2): 163±464. China. The Japanese Journal of Systematic En- Smetana, A. 1992. Peitowopsis monticola, a new tomology 6(2): 299±302. genus and species of the tribe Megarthropsini Li, L.-Z, M.-J. Zhao, and M. Sakai. 2000b. A new from southern Taiwan (Coleoptera: Staphylini- genus of the subfamily Tachyporinae (Coleop- dae: Tachyporinae). Bulletin of the National tera, Staphylinidae) with description of a new Museum of Natural Science 3: 199±208. species from Japan. Japanese Journal of Sys- Smetana, A. 1995a. A second species of the en- tematic Entomology 6(1): 11±18. demic Taiwanese genus Peitawopsis Smetana, 1992 from southern Taiwan (Coleoptera: Sta- Newton, A.F., Jr., and M.K. Thayer. 1992. Current phylinidae: Tachyporinae). Bulletin of the Na- classi®cation and family-group names in Sta- tional Museum of Natural Science 5: 131±134. phyliniformia (Coleoptera). Fieldiana: Zoology Smetana, A. 1995b. Revision of the tribes Que- (n.s.) 67: 1±92. diini and Tanygnathinini. Part III. Taiwan. (Co- Pace, R. 1986. Phaenoctavius brasiliensis n. gen., leoptera: Staphylinidae). National Museum of n. sp. del Brasile e Tachinus (Tachinoderus) Natural Science. Special Publication Number 6: rougemonti n. sp. della Thailandia (Coleoptera 1±145. Staphylinidae). Bollettino del Museo Civico di Ullrich, W.G. 1975. Monographie der Gattung Storia Naturale di Verona 11: 45±51. Tachinus Gravenhorst (Coleoptera: Staphylini- Scheerpeltz, O. 1933. Staphylinidae VII. In S. dae), mit Bemerkungen zur Phylogenie und Schenkling (editor), Coleopterorum Catalogus. Verbreitung der Arten. Dissertation zur Erlan- 6(129): 989±1500. Berlin: Junk. gung des Doktorgrades der Mathematisch-Na- SchuÈlke, M. 2003. Eine neue Art der Gattung turwissenschaftlichen FakultaÈt der Christian- Symmixus Bernhauer, mit Bemerkungen zur Albrechts-UniversitaÈt zu Kiel, 365 pp., 61 pls. Tribus Symmixini Bernhauer (Coleoptera, Sta- Zheng, F.-k. 2002. The female of Derops ding- phylinidae, Tachyporinae). Linzer Biologische shanus Y. Watanabe, 1999 (Coleoptera, Staphy- BeitraÈge 35(1): 443±452. linidae, Tachyporinae). Special Bulletin of the Seevers, C.H. 1958. A revision of the Vatesini, a Japanese Society of Coleopterology 5: 193± tribe of Neotropical Myrmecophiles (Coleop- 195.

APPENDIX Paratachinus laticollis Cameron num VIII of the female. Cameron knew the species only by the females, and Ullrich saw only the Paratachinus laticollis Cameron is the type type series. Coiffait (1984: 113) included the spe- species of Paratachinus. When Ullrich (1975) cies in a key but provided no illustrations. Here- synonymized Paratachinus with Tachinus (Tach- tofore characters of the male have not been pub- inoderus), P. laticollis became a junior secondary lished. homonym of Tachinus laticollis Gravenhorst, Because Tachinus punctatissimus (Hayashi) 1802, and was replaced by Tachinus oblongo- was described in Paratachinus, but was found to punctatus Ullrich. be congeneric with a group of Southeast Asian After the original description nothing of sub- species, it was essential to examine the type spe- stance was written about Paratachinus laticollis cies of Paratachinus to determine the generic Cameron until Ullrich (1975: 313) redescribed name of the group. Paratachinus punctatissimus and transferred the species to Tachinus, designat- shares with species of Lacvietina the metasternal ed a lectotype, and illustrated the tergum and ster- pit, the coarse, dense cephalic, pronotal and ely- 70 AMERICAN MUSEUM NOVITATES NO. 3430

Figs. 205±211. Tachinus oblongopunctatus Ullrich. 205. Sternite VII, male. 206. Aedeagus, ventral. 207. Sternum VIII, female. 208. Tergum VIII, male. 209. Sternum VIII, male. 210. Tergum VIII, female. 211. Spermatheca. tral punctation, the re¯exed cephalic margin, the tissimus are not congeneric; the former remains in emarginate posterior margin of the elytron, and Tachinus (Tachinoderus) and the latter is trans- the asymmetrical aedeagus, and it differs from ferred to Lacvietina. Paratachinus laticollis by these same features. Following is a redescription with illustrations Paratachinus laticollis and Paratachinus puncta- of salient features of P. laticollis Cameron; in- 2004 HERMAN: MEGARTHROPSINI 71 cluded for the ®rst time are characters of the male. angle rounded, not produced. Mesosternum with I did not study type material of P. laticollis Cam- midlongitudinal basal carina; paramedial basal ca- eron; the description and discussion concerning rina absent. Metasternum without pit adjacent to this species are based on specimens identi®ed by apex of mesosternal process (see ®gs. 146, 197); Milton Campbell. The specimens are consistent circum-mesocoxal ridge moderately strongly de- with the descriptions published by Cameron veloped medially. (1932) and Ullrich (1975). As the name suggests, Abdominal segment II with one pair of para- the elongate pronotal punctation is a characteristic tergites; segment III with two pairs of parater- feature of the species. gites; segments IV±VII with one pair of paratergites. Sternum I present as narrow sclerite anterior Tachinus oblongopunctatus Ullrich to II. Sternite II narrow, fused to III, and with Figures 205±211 median carina. Sternite III with median carina basally. Tergite VII with palisade fringe on posterior Tachinus oblongopunctatus Ullrich, 1975: 313 margin. Tergites IX divided middorsally by ter- (replacement name). gum X. ÐCoiffait, 1984: 133.ÐL.-Z. Li and Ohbayashi, MALE: Sternite V unmodi®ed. Sternite VI with 1996: 160. feeble emargination of posterior margin; surface Paratachinus laticollis Cameron, 1932: 397. Type unmodi®ed. Sternite VII (®g. 205) with broad, locality: Darjeeling Dist.: Ghum, alt. 9000 feet. shallow median depression; depression with clus- Type not examined. Junior secondary hom- ter of about 16 peg setae; sternite with medially onym of Tachinus laticollis Gravenhorst, 1802. separated row of peg setae adjacent to posterior margin; apical row of peg setae bordered laterally DESCRIPTION: Color reddish brown. Head and by long, darkly pigmented seta; posterior margin pronotum usually darker than elytra and abdomen. with broad, deep median emargination and with Head and clypeus with feeble, scattered punc- long, stout, sinuate, blunt-tipped seta on lateral tation; surface polished. Head not explanate lat- ®fth of emargination. Sternum VIII (®g. 209) with erally and without re¯exed supraantennal or clyp- broad, deep median emargination; lateral margin eal margins; surface near antennal insertion shal- of emargination sinuate; emargination with nar- lowly impressed. Epistomal suture present, com- row, round base. Tergum VIII (®g. 208) with plete or interrupted medially, and gradually moderately deep, acute emargination between me- curved, not angulate medially; midcranial suture dian pair of apical lobes; emargination deeper absent. Dorsum without median impression or than wide. Aedeagus (®g. 206) more or less sym- midlongitudinal groove. Postocular lateral margin metrical in ventral view; parameres contiguous without ridge or carina extending posterolaterally along medial margin; parameres straight and of from eye; vertical postocular carina absent. Gular equal length. sutures widely separated and divergent from about FEMALE: Tergum VIII (®g. 210) with four apical middle. Submentum with scattered ®ne puncta- lobes; median pair of lobes shorter than lateral tion. Antenna moderately long, extending to about pair. Sternum VIII (®g. 207) with three pairs of basal quarter of elytra; scape more or less parallel- lobes. Spermatheca as in ®gure 211. sided from near base to apex, not tapered; base of DISTRIBUTION AND HABITAT: This species is insertion visible from above. known from India and Nepal and has been col- Pronotum strongly convex; anterior and poste- lected at 2000 to 3200 m elevation. rior angles broadly rounded; anterior angles pro- MATERIAL EXAMINED: Nine males and 9 fe- duced beyond median portion of anterior margin; males. Nepal: Khandbari District: Induwa Khola lateral margin evenly curved; dorsal surface cov- Valley, 2000 m, 14.IV.84, Smetana and LoÈbl (2 ered with moderately dense, elongate punctation; males, ASC), 16.IV.84 (1 male, 1 female, ASC), punctation ®ner medially and anteriorly and in- 2050 m, 16.IV.84 (1 male, 2 females, ASC); creasingly coarse laterally and posteriorly; punc- above Sheduwa, 3000 m, 31.III±1.IV.82, A. and tation most coarse near posterolateral margin; Z. Smetana (1 male, ASC); Bakan, W of Tashi- punctures distinct, not anastomosing. gaon, 3200 m, 5.IV.82, A. and Z. Smetana (2 Elytra convex, moderately convex medially and males, 1 female, ASC), 3100 m, 8.IV.82, A. and strongly convex laterally; dorsal surface evenly, Z. Smetana (1 female, ASC); Forest NE Kuwa- densely, and coarsely punctate; lateral margin nar- pani, 2400 m, 24.IV.84, Smetana and LoÈbl (1 rowly re¯exed from humeral angle to about mid- male, 1 female, ASC), 5.IV.84 (1 female, ASC), dle and with short, spinelike setae; posterior mar- 2500 m, 28.III.82, A. and Z. Smetana (1 male, 1 gin without emargination laterally; posterolateral female, ASC), 11.IV.82 (1 female, ASC). 72 AMERICAN MUSEUM NOVITATES NO. 3430

ADDENDUM

Two relevant articles arrived just before assumption that when examined the species the present study was due to be published. In will be found to have the pit. one, a new species is placed in Peitawopsis The illustrations published for L. takashii and another species is transferred to the same (®gs. 3±5 in Hayashi, 2003a) are similar to genus. The second article concerns Para- those for L. punctatissima (®gs. 161, 162, tachinus. 165 herein) and do not permit separation of the two species, but Hayashi (2003a: 118) Lacvietina takashii (Hayashi), discussed characters that differentiate them. new combination The presence of L. takashii on Iriomote Island slightly extends the range of Lacvie- Peitawopsis takashii Hayashi, 2003a: 114. Type locality: Japan, Okinawa, Iriomote Is., near tina from Vietnam and Taiwan. Kanbira Falls. Holotype, male, not examined. Lacvietina punctatissima (Hayashi) DISCUSSION: No specimens of this species were studied, nonetheless the habitus illus- DISCUSSION: Hayashi (2003a: 114) trans- tration clearly demonstates that it should not ferred the species to Peitawopsis, but for the be placed in Peitawopsis. reasons presented in an earlier section of the The posterior margin of the elytra of Pei- present work the species should be placed in tawopsis is broadly emarginate. The emar- Lacvietina. gination extends in a continuous arc from the lateral apical angle of one elytron to the other Paratachinus Blackwelder and the lateral third of the dorsal elytral sur- Hayashi (2003b: 133) treated Paratachin- face is concave (see ®g. 188 herein). These us as a genus, not a subgenus as was done characters are among those diagnostic for the in the present work, but he erroneously stated genus. The posterior elytral margin of L. tak- that Cameron was the author of the name. ashii is emarginate only near the lateroapical According to the International Code of Zoo- angle, the remainder of the posterior margin logical Nomenclature a genus-group name is a broad convex curve and the dorsal sur- proposed after 1930 without a type species face of the elytra is broadly convex to near designation is unavailable (Article 13.3) and the ``gutter'' of the lateral margin (see ®g. 1 the ®rst writer to satisfy the conditions of by Hayashi, 2003a). availability is the author of the name (Article The elytral features illustrated for L. tak- 50.1). Cameron did not cite designate a type ashii are found in species of Megarthropsis species so Paratachinus was unavailable; it and Lacvietina. The Japanese species cannot became available when Blackwelder desig- be included in Megarthropsis because it nated a type species so he is the author. lacks the laterobasal angle of the pronotum of species in that genus (see ®g. 171 herein). ADDENDUM REFERENCES The elytral and pronotal features presented Hayashi, Y. 2003a. Notes on Peitawopsis (Cole- by Hayashi for L. takashii are in accord with optera: Staphylinidae: Tachyporinae), with a those found in Lacvietina. Although the most description of a new species from Japan. The diagnostic character for Lacvietina, the inter- Entomological Review of Japan 58(2): 113± mesocoxal metasternal pit (see ®gs. 196, 197 119. Hayashi, Y. 2003b. Notes on Paratachinus (Co- herein), was neither illustrated nor mentioned leoptera: Staphylinidae) with a description of a in the description for L. takashii, the species new species. The Entomological Review of Ja- is transferred to Lacvietina with the implicit pan 58(2): 133±141.