Megarthropsis Cameron, 1918

HERMAN, LEE H., 2004, Revision of the Asian Tribe Megarthropsini (Coleoptera: Staphylinidae: Tachyporinae), American Museum Novitates 3430, pp. 1-72 : 7-23

Megarthropsis Cameron

Figures 2–73 , 171–182

Megarthropsis Cameron, 1919: 231 . Type species: Megarthropsis decorata Cameron , fixed by monotypy.

— Cameron, 1921: 355, 406 (characters; checklist).— Scheerpeltz, 1933: 1478 (catalog).— Blackwelder, 1952: 236 (type species).— Smetana, 1983a: 144 (characters; illustrations).— Smetana, 1992: 204 (characters; illustrations; habitus).— Herman, 2001a: 678 (catalog).

DIAGNOSIS: Megarthropsis can be separat­

ed from Nepaliodes by the angulate postocular carina (fig. 172), the rounded anterior angle of the pronotum (figs. 66, 172), and, except for M. djawaensis , the narrowly explanate lateral margin of the elytra (fig. 171). The first antennal segment of most species of Megarthropsis is parallel­sided to slightly tapered (fig. 171), but in one species, M. djawaensis , the scape is moderately strongly tapered apically and similar to that of Nepaliodes . Megarthropsis is distinguished from Peitawopsis and Lacvietina by the distinct postocular carina (fig. 172) and the angulate basal angle of the pronotum (fig. 66); the latter two genera lack both features.

DESCRIPTION: Length 3.8–4.2 mm; width 1.1–1.4 mm.

Color reddish brown with yellowish brown pronotal lateral margins; head usually dark reddish brown to nearly black.

Head (fig. 171) with dorsum densely and coarsely punctate. Clypeal surface largely impunctate, but with a few punctures present. Head (fig. 171) with lateral margin strongly reflexed from antenna to anterior margin of clypeus; anterior margin of clypeus with feeble ridge. Epistomal suture complete and angulate at middle; midcranial suture rudimentary, minute, and poorly developed (best seen in lightly pigmented or cleared specimens). Dorsum of head without midlongitudinal groove; surface with or without broad, shallow, median depression. Postocular lateral margin of head with carina extending posteromedially from eye; carina angulate near eye (fig. 172); postocular vertical carina present and extending ventrally from lateral carina. Gular sutures widely separated. Submentum impunctate. Antennae long, slender, and reaching beyond posterior margin of elytra; scape more or less parallel­sided from near base to apex (see M. decorata , M. deverra , M. frazerensis ) or slightly (see M. durga , M. empusa , M. parca , M. smetanai ) to moderately (see M. djawaensis ) tapered apically; dorsal surface of scape with short, moderately dense pubescence and ventral surface sparsely pubescent.

Pronotum (figs. 66, 172) with anterior angle broadly rounded and extending beyond median portion of anterior margin; lateral margin unevenly curved to basal angle; bas­ al angle strongly angulate; median threequarters of notum coarsely and moderately densely punctate and strongly convex; lateral eighth explanate and with coarse, moderately dense punctation.

Elytra convex (fig. 171), slightly convex medially and strongly convex laterally; surface evenly, densely, and coarsely punctate; lateral sixteenth to fifth explanate and with margin reflexed; lateral margin with row of short spinelike setae extending from humeral angle to near basal angle; setae of epipleural ridge increasingly short posteriorly; posterior margin (figs. 25, 69) slightly, moderately, or strongly emarginate laterally; posterolateral angle slightly, moderately, or strongly produced (figs. 25, 69). Mesosternum with short, thick, midlongitudinal carina extending posteriorly from basal margin; paramedial carina rudimentary, present only basally. Metasternum with shallow depression adjacent to apex of mesosternal process; circum­mesocoxal ridge enlarged medially.

Procoxa without carina on medial surface.

MALE: Sternite VII (figs. 4, 11, 21) with wide emargination of posterior margin; surface with deep to shallow median depression and with or without median peg setae. Tergum VIII (figs. 26, 36, 39, 67) with four apical lobes.

Aedeagus (figs. 5, 10, 49, 59) with parameres straight or slightly curved; parameres of nearly equal length and width; median lobe with deep groove adjacent to midline of ventral surface of base (fig. 48).

FEMALE: Sternum VIII with five (fig. 68) or six lobes (figs. 7, 19); median lobe or pair of lobes with one fanlike cluster of setae. Tergum VIII with four (figs. 9, 16) or five (fig. 28) apical lobes; lateral lobe with small secondary lobe on lateral edge.

Spermatheca (figs. 8, 57) with apex of capsule enlarged and globose, then abruptly constricted and tapered proximally to spermathecal duct; juncture of capsule with spermathecal duct not enlarged.

DISCUSSION: Cameron (1919: 231) described the genus in detail; Smetana (1983a: 144; 1992: 204–206) supplemented it with new characters and numerous illustrations.

KEY TO SPECIES OF MEGARTHROPSIS

1. Sternum VIII (fig. 6) with posterior margin emarginate (males)................. 2

— Sternum VIII with posterior margin entire, not emarginate (females)............... 9

2(1). Sternite VII without peg setae (figs. 21, 40) ............................... 3

— Sternite VII with peg setae (figs. 4, 61).. ............................... 4

3(2). Antennal scape tapered anteriorly from base; cephalic postocular vertical carina obliquely oriented........................... M. djawaensis , new species

— Antennal scape of more or less equal diameter throughout; cephalic postocular vertical carina dorsoventrally oriented.............. M. empusa , new species

4(3). Sternites V (fig. 63) and VI (fig. 65) with dense mat of pubescence in median depression.... M. smetanai , new species

— Sternites V and VI without dense median mat of pubescence............... 5

5(4). Sternite V (fig. 45) with small peg setae medially..... M. frazerensis , new species

— Sternite V without peg setae......... 6

6(5). Sternite VII (fig. 11) with broad, deep emargination; parameres (fig. 10) obliquely truncate apically (in ventral or dorsal view)....... M. deverra , new species

— Sternite VII (figs. 4, 33, 53) with broad, moderately deep emargination; parameres acute or subacute apically, not obliquely truncate (in ventral or dorsal view).. 7

7(6). Sternite VII (fig. 53) without peg setae on disc.......... M. parca , new species

— Sternite VII (figs. 4, 33) with peg setae on disc........................... 8

8(7). Sternite VII (fig. 33) with sinuate posterior margin........ M. durga , new species

— Sternite VII (fig. 4) with evenly curved posterior margin... M. decorata Cameron

9(1). Sternum VIII (fig. 68) with one median lobe........... M. smetanai , new species

— Sternum VIII (fig. 7) with two median lobes .............................. 10

10(9). Tergum VIII (fig. 28) with five lobes on apical margin............................. M. djawaensis , new species

— Tergum VIII (figs. 9, 34, 71) with four lobes on apical margin.......... 11

11(10). Tergum VIII (fig. 55) tumescent at base of median lobes............................. M. parca , new species

— Tergum VIII without median tumescence, surface (in lateral view) straight from base to apex................. 12

12(11). Tergum VIII (fig. 16) with pair of short, broad, widely separated median lobes.......... M. deverra , new species

— Tergum VIII (figs. 9, 34) with pair of long narrowly separated median lobes... M. decorata Cameron , M. durga , new species

DESCRIPTION OF SPECIES Megarthropsis decorata Cameron Figures 2–9

Megarthropsis decorata Cameron, 1919: 232 . Type locality: Singapore: Mandai. Lectotype, male, designated by Smetana (1983a: 146) in The Natural History Museum (London); not examined.

— Cameron, 1921: 379, 406 (characters; Singapore).— Scheerpeltz, 1933: 1478 (catalog).— Smetana, 1983a: 146 (lectotype designation; characters; figures 1–7 [only, the other illustrations refer to M. parca ]; bionomics; Singapore).

DIAGNOSIS: Megarthropsis decorata has no one feature that immediately separates it from all of its congeners; various characters must be used to distinguish it. The males of M. decorata can be separated from M. smetanai by the absence of a dense mat of setae on sternites V and VI, from M. frazerensis by the absence of peg setae on sternite V, from M. deverra by the shallower emargination of sternite VII (fig. 4) and the more slender parameres (fig. 5), from M. durga by the evenly curved emargination of sternite VII (fig. 4), and from M. djawaensis , M. empusa , M. parca , and M. durga by the large cluster of discal peg setae on sternite VII (fig. 4).

Females of M. decorata have two median lobes on sternum VIII (fig. 7) and a pair of straight, basally broad median lobes on tergum VIII (fig. 9). Tergum VIII is straight from the base to the apex; the median region is not tumid.

DESCRIPTION: Length 3.2–3.8 mm; width 1.1–1.3 mm. Color reddish brown and yellowish brown. Head dark reddish brown, nearly black. Pronotum with convex median region reddish brown, darker anteriorly than posteriorly; lateral, explanate region yellowish brown. Elytra dark reddish brown. Abdomen reddish brown. Antennae and legs pale reddish brown.

Head with deep, moderately large punctation dorsally; postocular vertical carina dorsoventrally oriented. Antennal scape more or less parallel­sided.

Pronotum with deep, moderately large punctation.

Elytra with deep moderately large punctation; lateroapical emargination moderately deep; epipleural gutter moderately wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V with shallow median impression; surface without peg setae or dense mat of setae medially; posterior margin with broad, shallow, indistinct emargination.

Sternite VI (fig. 3) with shallow median impression; impression with dense cluster of setae near posterior margin; posterior margin with broad, shallow emargination.

Sternite VII (fig. 4) with broad, shallow median depression; posterior margin with broad, deep, evenly curved, median emargination; emargination without median notch; apical region of disc with two medially separated clusters of peg setae that also extend to and laterally along posterior margin; peg setae along posterior margin arranged in even or slightly sinuate row.

Sternum VIII (fig. 6) with deep, broad, median emargination; emargination strongly constricted near middle and with basal por­ tion of emargination strongly narrowed; margins of emargination strongly sinuate.

Aedeagus (figs. 2, 5) with parameres straight, slender, and tapered to subacute apex.

FEMALE: Tergum VIII (fig. 9) with median pair of lobes straight, tapered apically, and broad basally; surface without tumescence at base of middle lobe. Sternum VIII (fig. 7) with three pairs of apical lobes.

Spermatheca as in figure 8.

DISCUSSION: The illustrations cited in 1992 as M. decorata ( Smetana, 1992: figs. 30–42) are of M. smetanai and in 1983 the illustrations of the female abdominal segments ( Smetana, 1983a: figs. 8–10) are of M. parca .

Illustrations of the male of M. decorata from Singapore were published by Smetana (1983a: 1–7). The figures of M. decorata in the present article were taken from a specimen from Borneo.

DISTRIBUTION AND HABITAT: The species was described from Singapore. The type series was collected from Mandai; later two more specimens were collected from the Nee Soon swamp forest, which is in the Mandai region.

A male collected on Mt. Kinabalu, Borneo, is indistinguishable from the specimens from Singapore. Two females were collected with the male from Borneo. This disjunction of the distribution of M. decorata is surprising given the diversity of species in the Kinabalu region. However, because most species discussed herein are known from only one collecting site or are from different elevations on the same mountain, little can be said about the distribution of the species.

Specimens of the type series were collect­ ed in wet rotting leaves on the edge of a jungle stream ( Cameron, 1919: 233) and from flood debris ( Smetana, 1983a: 146). Smetana (op. cit.) concluded that the species lives in leaf litter and other debris in very wet habitats. Although no elevational data were included on the locality labels, the Mandai region is at low elevation (the highest point in Singapore is about 175 m). The specimens from Mt. Kinabalu were collected from forest floor litter near a stream in the vicinity of the headquarters of the National Park at an elevation of about 1500 m.

MATERIAL EXAMINED: Five specimens: 3 males, 2 females. Singapore: Nee Soon Swamp forest , 24­V­68, R.W. Taylor (1 male, MHNG; 1 male, NHMW) . Malaysia: Sabah: near Kinabalu N.P. headquarters, forest floor litter near Liwagu Stream, 29­VII­82, G.­M. de Rougemont (1 male, 2 females, GRC) .

Three males and one female were dissect­ ed for genitalic and abdominal features.

Megarthropsis deverra , new species

Figures 10–19

HOLOTYPE: Male. [ Malaysia:] ‘‘SABAH: Poring Hot Springs, Langanan Falls, 900–950 m 12.V.1987 Burckhardt—Löbl’ ’. Deposited in the Muséum d’Histoire Naturelle , Genève, Switzerland ( MHNG).

PARATYPES: One male, 1 female. Same data as holotype (1 female, MHNG). Malaysia: Sabah: Crocker Range, Mawar Waterfall env. (9c), 17.6.1996, vegetation debris and forest litter around fallen trees (1 male, NHMW) .

DIAGNOSIS: The males of M. deverra can

be distinguished from males of all other species by the broad deep emargination of sternite VII (fig. 11) and the short, broad, obliquely truncate parameres of the aedeagus (fig. 10).

The females can be distinguished from those of other species by the short, widely separated median pair of lobes of tergum VIII (fig. 16).

DESCRIPTION: Length 3.7–4.2 mm; width 1.2 mm. Color dark reddish brown, reddish brown, and yellowish brown. Head dark reddish brown, nearly black. Pronotum with explanate lateral region yellowish brown; convex median region dark reddish brown, nearly black anteriorly and paler reddish brown posteriorly. Elytra dark reddish brown, nearly black. Abdomen reddish brown. Antennae reddish brown. Legs pale reddish brown.

Head with deep, moderately large punctation dorsally; postocular vertical carina dorsoventrally oriented. Antennal scape more or less parallel­sided.

Pronotum with deep, moderately large punctation.

Elytra with deep moderately large punctation; lateroapical emargination moderately deep; epipleural gutter moderately wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V with feeble, shallow median impression; surface without peg setae or dense mat of setae; posterior margin feebly emarginate.

Sternite VI with broad, shallow depression; surface with cluster of setae on posterior margin near lateral margin of impression, but without dense discal mat of setae; posterior margin with broad moderately deep emargination.

Sternite VII (fig. 11) with broad, moderately deep, median depression; posterior margin with broad, deep emargination; emargination truncate or slightly rounded medially and without median notch; apical region of disc with two medially separated clusters of numerous peg setae; peg setae along posterior margin arranged in evenly curved row.

Sternum VIII (fig. 15) with deep, broad, median emargination; emargination strongly constricted near middle and with basal portion strongly narrowed; margins of emargination strongly sinuate.

Aedeagus (figs. 10, 12) with parameres slightly bent to right (in ventral view); par­ ameres short, broad medially, and obliquely truncate apically.

FEMALE: Tergum VIII with median pair of lobes broad, short, and widely separated (fig. 16). Sternum VIII with three pairs of apical lobes (fig. 19).

Spermatheca as in figures 17, 18.

ETYMOLOGY: The name of this species is from the Latin for the Roman guardian of newborn children (Deverra) and is symbolized by a broom for sweeping away evil influences ( Jordan, 1993); used in apposition.

DISTRIBUTION AND HABITAT: This species is known from two localities, one in the Mt. Kinabalu National Park at about 900 m, the other in the nearby Crocker Range.

MATERIAL EXAMINED: Three specimens: holotype male and male and female paratypes.

All specimens were dissected for genitalic and abdominal characters.

Megarthropsis djawaensis , new species

Figures 20–28

HOLOTYPE: Male. ‘‘ INDONESIA: W Java ‘ Ranca Upas’ ca. 1300m 10 km S Ciwidey lg. Schuh 9.8.1994 ’’. Deposited in the Naturhistorisches Museum Wien , Wien , Austria ( NHMW).

PARATYPES: Two males, 3 females. Same data as holotype but collected from ‘‘forest litter’’ (1 male, 2 females, NHMW). Indonesia: W. Java, Situ Lembang , ca 1500 m, 15 km N. Bandung, lg. Schuh, 7.8.1994 (1 male, 1 female, NHMW) .

DIAGNOSIS: This species is separated from all others of the genus by the apically tapered antennal scape, the oblique orientation of the postocular vertical carina, and the strongly produced posterolateral angle of the elytra (fig. 25). The punctation of the pronotum and dorsal surface of the head and elytra is coars­ er and deeper and the punctation of the pronotal disc is more sparse than for any of the other species of the genus. Only this species and M. empusa lack peg setae on sternite VII (fig. 21); these two species can be separated by the features listed in the preceding two sentences.

DESCRIPTION: Length 3.3–3.8 mm; width 1.3–1.4 mm. Color reddish brown and yellowish brown. Head dark reddish brown to nearly black. Pronotum with convex median region dark reddish brown anteriorly and reddish brown posteriorly; lateral, explanate region reddish brown. Elytra dark reddish brown. Abdomen reddish brown with apical segments darker reddish brown. Antennae and legs reddish brown.

Head with large, deep punctures on dorsal surface. Postocular vertical carina obliquely, dorsoposteriorly to ventroanteriorly, orient­ ed. Antenna with first segment moderately tapered from base to apex.

Pronotum with large, deep punctures.

Elytra with large, deep punctation; lateroapical emargination deep (fig. 25); epipleural gutter wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V without median impression, peg setae, or dense mat of setae medially; posterior margin without emargination.

Sternite VI with feeble median impression; surface without special cluster of setae; posterior margin without emargination.

Sternite VII (fig. 21) with broad, shallow, median impression; impression without setae medially, but with short setae along lateral margins; posterior margin with moderately deep, broadly V­shaped emargination; posterior margin evenly curved, not sinuate, to median emargination; surface without peg setae.

Sternum VIII (fig. 20) with deep, broad, median emargination; emargination more strongly narrowed near basal quarter; base of emargination narrowly rounded; margins of emargination weakly sinuate.

Aedeagus (figs. 22, 23) with parameres straight, slender, tapered, and narrowly rounded apically.

FEMALE: Tergum VIII (fig. 28) with paramedial lobes separated by smaller median lobe. Sternum VIII (fig. 27) with three pairs of apical lobes.

Spermatheca as in figure 24.

ETYMOLOGY: The name for this species is taken from a variant of the Indonesian name (Djawa) for Java.

DISTRIBUTION AND HABITAT: The species is known only from western Java where specimens were collected from forest litter at 1300 m and 1500 m in August.

MATERIAL EXAMINED: Six specimens: ho­ lotype male, 2 paratype males, and 3 paratype females.

Two males and one female were dissected for characters of the abdomen and genitalia.

Megarthropsis durga , new species

Figures 29–36

HOLOTYPE: Male. [ Malaysia]: ‘‘SABAH Mt. Kinabalu 1580 m, 27.IV.1987 Burckhardt—Löbl’ ’. Deposited in the Muséum d’Histoire Naturelle , Genève, Switzerland ( MHNG).

PARATYPES: Two males. Same data as holotype (1 male, MHNG). [ Malaysia]: Sabah: Poring Hot Springs , Langanan Falls , 900–950 m, 12­V­ 1987, Burckhardt and Löbl (1 male, MHNG) .

DIAGNOSIS: Males of M. durga can be separated from other species by the broad, moderately deep, sinuate posterior margin of sternite VII (fig. 33) and the sinuate row of peg setae near the margin. The disc of sternite VII has only a few peg setae medially.

Females are separated from other species, except M. decorata and M. smetanai , by the pair of median lobes of tergum VIII (fig. 34) that are straight and broad at the base; the surface adjacent to the base of the median lobes is not tumescent. This pair of median lobes is broader basally in M. durga than in M. decorata (fig. 9). Sternum VIII (fig. 35) of M. durga has a pair of median lobes in contrast to the single median lobe of M. smetanai . Denser cephalic, pronotal, and elytral punctation separate M. durga from M. djawaensis .

DESCRIPTION: Length 3.4–3.8 mm; width 1.4 mm. Color reddish brown and yellowish brown. Head reddish brown to dark reddish brown to nearly black. Pronotum with convex median region reddish brown to dark reddish brown; lateral explanate region yellow brown. Elytra and abdomen reddish brown. Antennae and legs pale reddish brown.

Head with deep, moderately large punctation dorsally; postocular vertical carina dorsoventrally oriented. Antennal scape more or less parallel­sided to slightly tapered apically.

Pronotum with deep, moderately large punctation.

Elytra with deep moderately large punctation; lateroapical emargination moderately deep; epipleural gutter moderately wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V with feeble median impression; surface without peg setae or dense mat of setae; posterior margin without emargination.

Sternite VI with feeble median impression; impression with slightly denser row of setae along posterior margin, but without dense discal mat of setae; posterior margin with broad, shallow emargination.

Sternite VII (fig. 33) with broad, shallow median impression; posterior margin with broad, moderately deep, sinuate emargination; emargination without median notch; surface with small, median cluster of peg setae on disc and with row of peg setae along posterior margin; peg setae of posterior margin arranged in sinuate row and row narrowly divided medially.

Sternum VIII (fig. 31) with deep, broad, median emargination; emargination slightly more strongly constricted from about apical quarter and gradually convergent to base; margins of emargination moderately sinuate.

Aedeagus (figs. 29, 30) with parameres slightly curved to right (in ventral view), long, slender, and tapered to acute apices.

FEMALE: Tergum VIII (fig. 34) with median pair of lobes straight, tapered apically, and broad basally; surface without tumescence near base of median lobes. Sternum VIII (fig. 35) with three pairs of apical lobes.

Spermatheca as in figure 32.

ETYMOLOGY: Named for the many armed Hindu goddess Durga, who is depicted as a woman riding on a lion or tiger and in whose hands are weapons. In later Hindu tradition, particularly in northern India, Durga is linked with the fertility of crops and is celebrated at harvest time in the Durga Puja ( Jordan, 1993); used in apposition.

DISCUSSION: The females identified as M. durga are not designated as paratypes because they cannot be distinguished reliably from M. decorata .

DISTRIBUTION AND HABITAT: This species is known only from Mt. Kinabalu in Borneo where it was collected at 900–1580 m.

MATERIAL EXAMINED: Six specimens: holotype male, 2 paratype males, 3 females. The females are from the type locality and were collected with the holotype.

Three males and one female were dissect­ ed for genitalic and abdominal characters.

Megarthropsis empusa , new species

Figures 37–41

HOLOTYPE: Male. [ Malaysia]: ‘‘ SABAH: Crocker Ra. 1600m, km 51 rte Kota Kinabalu­Tambun­

an, 18.V.87 Burckhardt—Löbl’’. Deposited in

the Muséum d’Histoire Naturelle, Genève,

Switzerland ( MHNG).

DIAGNOSIS: The absence of peg setae on sternite VII (fig. 40) will separate the males of this species from the others of the genus except M. djawaensis . This species is distinguished from M. djawaensis by the denser cephalic and pronotal punctation, the nearly cylindrical first antennomere, the dorsoventral orientation of the vertical postocular carina that extends ventrally from the dorsolateral postocular carina, the less strongly produced posterolateral elytral angle, and the narrower epipleural gutter. The emargination of sternite VII of the male of M. empusa (fig. 40) is deeper than in M. djawaensis (fig. 21).

The female is unknown.

DESCRIPTION: Length 3.4 mm; width 1.2 mm. Color reddish brown and yellowish brown. Head dark reddish brown. Pronotum with convex median region dark reddish brown anteriorly and reddish brown to yellowish brown posteriorly; explanate lateral region yellowish brown. Elytra dark reddish brown. Abdomen reddish brown with darker apical segments. Antennae and legs reddish brown.

Head with moderately large punctation on dorsal surface; postocular ventral carina dorsoventrally oriented. Antennal scape more or less parallel­sided to slightly tapered apically.

Pronotum with moderately large punctation.

Elytra with moderately large punctation; posterolateral emargination moderately deep; epipleural gutter narrow.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V slightly flattened medially; surface without peg setae or dense mat of setae medially; posterior margin with moderately wide feeble median emargination.

Sternite VI with wide, feeble median impression; impression with slightly denser pubescence near posterior margin; posterior margin with wide, shallow median emargination.

Sternite VII (fig. 40) with broad, shallow, median impression; impression without setae medially near posterior margin, but with short setae along lateral margin; posterior margin with broad, sinuate emargination; emargination deeper and more or less broadly V­shaped medially; surface without peg setae.

Sternum VIII (fig. 41) with broad, deep, median emargination; emargination strongly constricted from about middle and with basal portion of emargination strongly narrowed; margin of emargination strongly sinuate.

Aedeagus (figs. 37, 38) with paramere straight, moderately broad, and tapered to subacute apex.

FEMALE: Unknown.

ETYMOLOGY: The name is from the Greek (Empousa) for hobgoblin or a mischievious goblin, and is used for the species found hidden in a long series of M. smetanai .

DISTRIBUTION AND HABITAT: This species is known only from the Crocker Range in northern Borneo where it was collected at 1600 m with M. smetanai .

MATERIAL EXAMINED: One specimen: holotype male.

Megarthropsis frazerensis , new species

Figures 42–48

HOLOTYPE: Male. [ Malaysia]: ‘‘ PAHANG Frazer’s Hill 11.III.90 Rougemont’’// ‘‘ Megarthropsis decorata Cam. det Rougemont’’ [labels handwritten]. In collection of G.­M. de Rougemont ( GRC) to be deposited in Natural History Museum, London.

PARATYPE: One male. Malaysia: Prov. Johor, Gg. Ledang (14), Hutan Lipur , 200 m, 4.2.1992, leg. Schillhammer (1 male, NHMW) .

DIAGNOSIS: The male of Megarthropsis frazerensis can be separated from the males of all other species of the genus by the unique presence of a cluster of peg setae on the middle of the disc of sternite V (fig. 45). The emargination of the posterior margin of sternite VII is notched medially and the peg setae are divided into two clusters (fig. 44).

The female is unknown.

DESCRIPTION: Length 3.4 mm; width 1.2 mm. Color reddish brown and yellowish brown. Head dark reddish brown, nearly black. Pronotum with explanate lateral region yellowish brown; convex median region reddish brown, darker anteriorly and paler posteriorly. Elytra and abdomen reddish brown. Antennae and legs pale reddish brown.

Head with deep, moderately large punctation dorsally; postocular vertical carina dorsoventrally oriented. Antennal scape more or less parallel­sided.

Pronotum with deep, moderately large punctation.

Elytra with deep moderately large punctation; lateroapical emargination shallow; epipleural gutter moderately wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V (fig. 45) with broad, shallow emargination of posterior margin; disc with scattered cluster of small peg setae in feeble impression.

Sternite VI (fig. 47) with broad, shallow, median impression; posterolateral edge of impression with small dense cluster of setae; posterior margin with moderately deep, rounded median emargination.

Sternite VII (fig. 44) with broad, shallow median depression; posterior margin with broad, deep median emargination; emargination with small median notch; median region of disc with peg setae arranged in two dense, medially separated clusters; peg setae of anterior region scattered; peg setae along posterior margin aligned in more or less even, medially separated rows.

Sternum VIII (fig. 46) with deep, broad, median emargination; emargination wide apically and strongly constricted near middle and with basal portion of emargination strongly narrowed; margin of emargination strongly sinuate.

Aedeagus (figs. 42, 43) with parameres feebly bent to left (in ventral view) and tapered to subacute apex.

FEMALE: Unknown.

ETYMOLOGY: The name for this species is taken from the type locality, Frazer’s Hill in West Malaysia .

DISTRIBUTION AND HABITAT: This species is known by only two Malaysian specimens and both lack further collecting data.

MATERIAL EXAMINED: Two specimens: holotype and paratype males.

Both specimens were dissected for genitalic and abdominal characters.

Megarthropsis parca , new species

Figures 49–58

HOLOTYPE: Male. [ Malaysia]: ‘‘SABAH Mt. Kinabalu 1900 m, 26.IV.1987 Burckhardt—Löbl’ ’. Deposited in the Muséum d’Histoire Naturelle , Genève, Switzerland ( MHNG).

PARATYPES: Three males, 5 females. Same data as holotype (2 males, 3 females, MHNG). [ Malaysia]: Sabah: Mt. Kinabalu, 6200 ft, 22– 26.VI.68, R.W. Taylor (1 male, MHNG). [ Malaysia]: Borneo: Sabah: Mt. Kinabalu N.P., Summit Trail, 1890 m, [26.IV.86], A. Smetana (1 female, ASC). [ Malaysia]: Sabah Mt. Kinabalu 1580 m, 27.IV.1987 Burckhardt—Löbl (1 female, MHNG) .

— Smetana, 1983a: 146 (figs. 8–10; misidentified as M. decorata ).

DIAGNOSIS: The evenly curved posterior margin of sternite VII (fig. 53), the single row of peg setae near the posterior margin, and absence of discal peg setae will separate the males of M. parca from other species. The ventral surface of each paramere is oblique to the midsagittal plane of the ae­ deagus; this configuration is unique among the known species.

Females of M. parca can be separated from other species by the tumid apical third of tergum VIII (fig. 55). Each lobe of the median pair of lobes of tergum VIII curves laterad from the midlongitudinal line and the lobes are slender (fig. 54).

DESCRIPTION: Length 3.7–4.1 mm; width about 1.4 mm. Color reddish brown; head, convex median region of pronotum, elytra, and abdomen with darker infusions; head darker than remainder of body; pronotum with explanate lateral portion paler reddish brown; antennae and legs pale reddish brown.

Head with deep, moderately large punctation dorsally; postocular vertical carina dorsoventrally oriented. Antennal scape more or less parallel­sided to slightly tapered apically.

Pronotum with deep, moderately large punctation.

Elytra with deep moderately large punctation; lateroapical emargination moderately deep; epipleural gutter moderately wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V with feeble median impression; surface without peg setae or dense setal mat; posterior margin with broad feeble emargination.

Sternite VI (fig. 52) with broad, shallow median depression; impression with moderately dense cluster of setae near posterior margin, but without dense setal mat; posterior margin with broad, shallow emargination.

Sternite VII (fig. 53) with broad, shallow median depression; posterior margin with broad, deep, evenly curved, median emargination; emargination without median notch; disc without peg setae; peg setae arranged in curved row adjacent to posterior margin; row continuous or slightly separat­ ed medially.

Sternum VIII (fig. 51) with deep, broad, median emargination; emargination moderately constricted near middle and gradually convergent to narrow base; margins of emargination moderately strongly sinuate.

Aedeagus (figs. 49, 50). Parameres more or less straight (in ventral view), moderately long, moderately broad, and rounded apically; ventral surface oriented obliquely to midsagittal plane; apex with small point on ventral surface.

FEMALE: Tergum VIII (fig. 54) with median pair of lobes slightly curved laterally; median margin curved; lobes slender and moderately tapered apically; surface near base of median lobes strongly tumescent (fig. 55). Sternum VIII (fig. 56) with three pairs of lobes.

Spermatheca as in figures 57 and 58.

ETYMOLOGY: The name of this species is from the Latin (Parca) for the goddess of destiny and birth ( Leach, 1992); used in apposition.

DISCUSSION: Megarthropsis parca was reported as M. decorata from East Malaysia, Sabah, Mt. Kinabalu ( Smetana, 1983a: 146 and figs. 8–10).

DISTRIBUTION AND HABITAT: This species is known only from Mt. Kinabalu in Borneo where it was collected at 1580 to 1900 m.

MATERIAL EXAMINED: Nine specimens: holotype male and three male and five female paratypes.

Three males and three females were dissected for genitalic and abdominal features.

Megarthropsis smetanai , new species

Figures 59–73 , 171–182

HOLOTYPE: Male. [ Malaysia]: ‘‘SABAH Crocker Ra. 1600 m, km 51 rte Kota Kinabalu­Tambun­ an, 18.V.87 Burckhardt—Löbl’ ’. Deposited in the Muséum d’Histoire Naturelle , Genève, Switzerland ( MHNG).

PARATYPES: Thirty­two males, 16 females. [ Malaysia]: Same data as holotype (15 males, 5 females, MHNG) . Sabah: Mt. Kinabalu , 1430 m, 22­V­1987, Burckhardt and Löbl (3 males, MHNG) ; 1500 m, 30­IV­1987 (1 male, MHNG) ; 1500 m, 21­V­1987 (1 male, 2 females, MHNG) ; 1550 m, 28­IV­1987 (2 males, 2 females, MHNG) ; 1550 m, 29­IV­1987 (5 males, 3 females, MHNG) ; 1550–1650 m, 24­ IV­1987 (1 male, 2 females, MHNG; 1 male, MSC) ; 1750 m, 27­IV­1987 (1 female, MHNG) . Borneo: Sabah: Mt. Kinabalu National Park HQ, Liwagu Riv. Tr., 1520 m, 12­VIII­ 88, A. Smetana [B106] (1 male, AMNH) ; 11­ VIII­88 [B100] (1 female, AMNH) . Borneo: Sabah Mt. , Kinabalu National Park HQ, Silau­ Silau Tr., 1550 m, 4­IX­88, A. Smetana [B177] (1 male, AMNH) ; 2­IX­88 [B171] (1 male, AMNH) .

— Smetana, 1992: 204–206 (figs. 30–42; misidentified as M. decorata ).

DIAGNOSIS: Males of M. smetanai can be distinguished from their congeners by the dense mat of pubescence in the large median depression of sternites V and VI (figs. 63, 65). Sternite VII (fig. 61) has a deep, rounded median depression, the peg setae are arranged in a dense cluster along the posterior margin, and the posterior margin is sinuate with a small, rounded median notch (fig. 62); this configuration is unique to M. smetanai .

The females of M. smetanai are distinguished from those of all other species by the presence of only five lobes on sternum VIII, the median lobe of which is broad (fig. 68).

DESCRIPTION: Length 3.3–4.6 mm; width 1.3–1.4 mm. Color reddish brown and yellowish brown. Head dark reddish brown to nearly black. Pronotum convex, median region dark reddish brown to reddish brown, posterior portion usually paler; explanate lateral region pale reddish brown to yellowish brown. Elytra and abdomen reddish brown. Antennae and legs pale reddish brown.

Head with deep, moderately large punctation dorsally; postocular vertical carina dorsoventrally oriented. Antennal scape more or less parallel­sided to slightly tapered apically (fig. 171).

Pronotum with deep, moderately large punctation.

Elytra with deep moderately large punctation; lateroapical emargination moderately deep; epipleural gutter moderately wide.

Wings fully developed. Tergite VII with well­developed palisade fringe on posterior margin.

MALE: Sternite V (fig. 63) with large, moderately deep, median depression; depression with dense mat of setae and depression nearly as long as segment; surface without peg setae; posterior margin with broad, shallow median emargination.

Sternite VI (fig. 65) with large, deep, median depression; depression nearly as long segment and surface covered with dense mat of setae; posterior margin with broad, moderately deep, median emargination.

Sternite VII (fig. 61) with large, deep, rounded, median depression; depression polished medially and without setae in middle, but with short setae laterally and basally; surface with dense cluster of peg setae along posterior margin; cluster of peg setae slightly separated medially; posterior margin with broad sinuate emargination; emargination with small, rounded median notch (fig. 62).

Sternum VIII (fig. 64) with broad, deep median emargination; emargination moderately strongly constricted at about apical third then gradually convergent to narrow base; margins of emargination moderately sinuate.

Aedeagus. Parameres (figs. 59, 60) bent moderately to left (in ventral view) and tapered to subacute apex.

FEMALE: Tergum VIII (fig. 71) with median pair of lobes straight and broad basally. Sternum VIII (fig. 68) with five lobes, one lateral and one paralateral pair of lobes and one broad median lobe.

Spermatheca as in figure 70.

ETYMOLOGY: This species is named to hon­ or my friend and colleagueAleš Smetana for his many sterling contributions to our understanding of the Staphylinidae and who, over the years, I have come to admire for the quality of his work and depth of his knowledge. In large part his studies and collections of the Megarthropsini made the present work possible.

DISCUSSION: Figures 30–42 published by Smetana (1992) and identified as M. decorata are illustrations of M. smetanai .

Sternum VIII (fig. 68) of the females has one broad median lobe, but other members of the tribe, except the species of Nepaliodes , have a median pair of lobes. The broad lobe in M. smetanai is clearly the result of a fusion of this pair of median lobes.

DISTRIBUTION AND HABITAT: This species is known only from Mt. Kinabalu and the nearby Crocker Range where it was collected at elevations of 1430–1750 m. According to field notes for specimens collected by A. Smetana this species has been collected from fleshy mushrooms and soft polyporus­type mushrooms growing on a fallen rotting tree, and from fresh and rotting fruits of the ‘‘kerosene tree’’.

MATERIAL EXAMINED: Forty­nine specimens: holotype male, 32 male and 16 female paratypes.

Six males and one female were dissected for genitalic and abdominal characters; one male was disarticulated.

Figures

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.

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).

Figs. 16–19. Megarthropsis deverra. 16. Tergum VIII, female. 17, 18. Spermatheca. 19. Sternum VIII, female.

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

Figs. 26–28. Megarthropsis djawaensis. 26. Tergum VIII, male. 27. Sternum VIII, female. 28. Tergum VIII, female.

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

Figs. 34–36. Megarthropsis durga. 34. Tergum VIII, female. 35. Sternum VIII, female. 36. Tergum VIII, apex, male.

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).

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.

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

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

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).

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.

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.

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.