New species and additional records of Lomechusini from the Palaearctic region , primarily from China ( Coleoptera : Staphylinidae : Aleocharinae ) With 81 figures and 2 maps

Six species of Lomechusini from China and North India are (re-)described and illustrated: Drusilla yunnanensis Pace, 1993; D. watanabei Maruyama & Kishimoto, 2002; D. flagellata spec. nov. (China: Yunnan); D. palata spec. nov. (China: Yunnan); Amaurodera angularis spec. nov. (North India: West Bengal); Lomechusoides penicillatus spec. nov. (China: Gansu). Two synonymies are proposed: Drusilla canaliculata (Fabricius, 1787) = D. puncticollis Motschulsky, 1845, syn. nov.; Amaurodera smetanai Pace, 1992 = A. yaoana Pace, 1992, syn. nov. Additional records of 29 previously described species of Drusilla Leach, 1819, Pella Stephens, 1835, Amaurodera Fauvel, 1905, Tetrabothrus Bernhauer, 1915, and Lomechusoides Tottenham, 1939 are reported from the Palaearctic region, primarily from China, among them several new country records and the first record of Pella lugens (Gravenhorst, 1802) from the East Palaearctic. The previously unknown aedeagus of Pella reuteri Assing, 2008, the previously unknown female sexual characters of Amaurodera ilamica Assing, 2005, and the primary sexual characters of several Pella species recorded from China are illustrated. The distributions of the Drusilla species currently known from China are illustrated.

Material examined since the latest contribution to the Lomechusini of the Palaearctic region (Assing 2010) included several new species and records of zoogeographic interest.The genus Orphnebius and the Chinese species of Tetrabothrus Bernhauer, 1915 were treated separately (Assing 2015a, b).The present paper deals with species of Drusilla, Pella, Amaurodera, Tetrabothrus, and Lomechusoides, primarily from China.

Material and methods
The material treated in this study is deposited in the following collections: The morphological studies were conducted using a Stemi SV 11 microscope (Zeiss Ger many) and a Jenalab compound microscope (Carl Zeiss Jena).The images of the external characters were created using a photographing device constructed by Arved Lompe (Nienburg) and CombineZ software.A digital camera (Nikon Coolpix 995) was used for the remaining photographs.The maps were created using MapCreator 2.0 (primap) software.Body length was measured from the anterior margin of the labrum to the apex of the abdomen, the length of the forebody from the anterior margin of the labrum to the posterior margin of the elytra, head length from the anterior margin of the clypeus (without ante-clypeus) to the posterior constriction of the head, the length of the elytra along the suture from the apex of the scutellum to the posterior margin of the elytra, the length of the median lobe of the aedeagus from the apex of the ventral process to the base of the aedeagal capsule, and the length of the spermatheca is given as the maximal extension (measured from the apex of the distal portion of the capsule).
The "parameral" side of the median lobe of the aedeagus (i.e., the side where the sperm duct enters) is referred to as the ventral, the opposite side as the dorsal aspect.(2005a, b, 2006b, 2008a, 2009, 2010).However, similar character conditions have been observed also in specimens from some regions in Middle Asia and the West Palaearctic region (particularly Iran), suggesting that these differences should be interpreted as intraspecific, possibly clinal variation.Moreover, the presence of a distinct subspecies in Kamchatka appears most unlikely.Hence the synonymy proposed above.

Comment:
The original description is based on 16 type specimens from two localities in Hubei and Hunan provinces (Maruyama & Kishimoto 2002a).The authors did not find (and illustrate) a spermatheca, although the type series includes seven females.A full redescription is provided below.
Description: Relatively large species; body length 7.0-7.8mm; length of forebody 2.6-3.0 mm.Coloration: body black, with the posterior margins of the abdominal segments paler; legs with the tibiae and tarsi yellowish to dark-yellowish, the profemora blackish, the mesofemora blackish with the basal third yellowish, and the metafemora yellowish in basal half and infuscate in apical half; antennae reddish-yellow, with antennomere I weakly to distinctly infuscate; maxillary palpi dark-brown, with the terminal palpomere reddish-yellow.Head (Figs 1-2) with sexually dimorphic punctation and with fine and shallow microreticulation.Eyes distinctly shorter than distance from posterior margin of eye to posterior constriction of head.Antenna (Fig. 3) 2.6-2.9 mm long and slender; all antennomeres at least weakly oblong.Pronotum (Figs 1-2) 1.00-1.05times as long as broad and 1.17-1.18times as broad as head, broadest in anterior half; posterior angles weakly marked; near posterior margin with pronounced transverse impression; other   Abdomen (Fig. 4) distinctly broader than elytra; tergites III-VIII with sexual dimorphism; punctation sexually dimorphic; anterior impressions of tergites III-VI impunctate; tergites with very weak, nearly obsolete transverse microsculpture; posterior margin of tergite VII with palisade fringe.: lateral portions of dorsal surface of head with dense and rather coarse punctation, with the interstices narrower than diameter of punctures; frons, posterior portion, and the narrow midline of the head impunctate or nearly so (Fig. 1); pronotum in posterior half with pronounced longitudinal elevation on either side, between these elevations with broadly flattened impression; median portion of pronotum somewhat shiny and with moderately dense and moderately coarse punctation, with the interstices as broad as, or broader than, diameter of punctures; lateral portions of pronotum practically matt, with very dense and coarsely granulose punctation, with the interstices narrower than diameter of punctures (Fig. 1); elytra with dense and coarsely granulose punctation (Fig. 1); abdominal tergite III (Fig. 4) with a pronounced median tubercle of somewhat variable shape and with scattered fine punctures; tergites IV-VI each with a median pair of distinct granules near posterior margin, impunctate (or nearly so), except for some punctures at posterior margins; tergite VII with a pronounced median granule and numerous additional smaller granules in posterior half (Fig. 4); tergite VIII (Fig. 5) with densely granulose punctation and with distinct microreticulation, posterior margin broadly concavely excavate; sternite VIII (Fig. 6) weakly transverse and with nearly truncate posterior margin; median lobe of aedeagus large, 1.0-1.1 mm long, shaped as in Figs 7-8; paramere 0.9 mm long and shaped as in Fig. 9. : head (Fig. 2) with sparse and very fine punctation, interstices much broader than diameter of punctures; pronotum (Fig. 2) with weakly pronounced elevations in posterior half; punctation moderately fine, moderately dense, and non-granulose; elytra (Fig. 2) with weakly granulose puncation; abdominal tergites III-V with fine and sparse punctation; tergite VI impunctate except for a median pair of setiferous punctures bearing long dark setae and additional punctures at posterior margin; tergite VII with a median pair of setiferous punctures bearing long dark setae and with additional fine and sparse punctation in posterior half; posterior margin of tergite VIII somewhat truncate, only indistinctly concave in the middle (Fig. 10); sternite VIII (Fig. 11) distinctly shorter and more transverse than that of male, posterior margin truncate; spermatheca very small in relation to body size, 0.25 mm long, and shaped as in Figs 12-13.
Comparative notes: This highly distinctive species differs from other Drusilla species known from China by numerous characters, particularly its large body size, the long and slender antennae, the short elytra, the reduced hind wings, the male secondary sexual characters, and the male and female primary sexual characters.

Distribution and natural history:
The known distribution now includes a total of three localities in Hubei, Hunan, and Yunnan provinces.The examined specimens were sifted from litter, moss, and the roots of herbs in a terraced secondary pine forest at an altitude of 2620 m.Maruyama & Kishimoto (2002a)   : pronotum (Fig. 14) with extensive, but not very deep median impression with somewhat denser punctation; elytra in large males each with conspicuous oblong and oblique elevation (Fig. 14); tergite VIII (Fig. 18) strongly transverse and with distinctly serrate posterior margin; sternite VIII (Fig. 19) transverse, posterior margin truncate in the middle; median lobe of aedeagus (Figs 20-21) 1.0 mm long and of highly distinctive morphology, laterally somewhat flattened, with short ventral process, and with long flagellum in internal sac; paramere (Fig. 22) very small, approximately 0.55 mm long, little more than half as long as median lobe.: pronotum (Fig. 15) flattened, but not impressed in the middle; tergite VIII (Fig. 23) strongly transverse, posterior margin with a triangular projection on either side; sternite VIII (Fig. 24) distinctly shorter and more transverse than that of male, posterior margin convex; spermatheca long and of distinctive shape (Fig. 25).

Intraspecific variation:
Body size and the modifications of the male elytra are subject to remarkable intraspecific variation.The elevations on the male elytra are fully pronounced only in the holotype, weakly pronounced in one, barely noticeable in two, and absent in three males.
Comparative notes: This species is characterized particularly by the modifications of the male elytra, the shapes of the male and female tergites VIII, and above all by the conspicuous morphology of the aedeagus and the distinctive shape of the spermatheca.For additional characters separating it from the sympatric and syntopic D. palata see the comparative notes in the following section.

Distribution and natural history:
The known distribution is confined to two localities in West Yunnan.The specimens were collected on and near the bank of a stream and near a pond at altitudes of 1900 and 2070 m, in one locality together with D. palata.Two paratypes are slightly teneral.

Etymology:
The specific epithet is an adjective derived from the Latin noun pala (shovel) and alludes to the conspicuous shape of the posterior projection of the male sternite VIII.
Description: Body length 5.1-6.3mm; length of forebody 2.5-2.6 mm.Coloration: head, pronotum, and abdomen blackish; elytra dark-yellowish, with the scutellar region and an extensive lateral spot infuscate; legs uniformly pale-yellowish; antennae blackish-brown; maxillary palpi reddish to dark-brown, with the terminal palpomere yellow.Head (Fig. 26) approximately 1.2 times as broad as long; dorsal surface with median portion flattened or shallowly impressed; punctation fine to moderately coarse and of variable density, sparse to dense; interstices without microsculpture and glossy.Eyes large and bulging, much longer than distance from posterior margin of eye to posterior constriction of head.Antenna (Fig. 27) 2.1 mm long; antennomeres III-X gradually and very weakly increasing in width; preapical antennomeres weakly transverse.Pronotum (Fig. 26) 1.06-1.10times as broad as long and 1.06 times as broad as head, broadest at anterior angles; posterior angles obtusely marked; posteriorly with a very small and indistinct impression; midline with narrow, deep, and sharply delimited furrow extending from posterior impression cephalad, but not reaching anterior margin of pronotum; punctation dense and coarse, not granulose; interstices without microsculpture and glossy.Elytra (Fig. 26) approximately 0.9 times as long as pronotum; humeral angles marked; punctation dense, similar to that of pronotum.Hind wings fully developed.Metatarsomere I shorter than the combined length of II-IV.Abdomen (Fig. 28) narrower than elytra; tergites III-VII without sexual dimorphism; anterior impressions of tergites III-V and posterior margins of tergites III-VII with fine punctation, remainder of tergal surfaces with very sparse punctation; posterior margin of tergite VII with palisade fringe.: pronotum (Fig. 26) with extensive and pronounced median impression in posterior half; tergite VIII (Fig. 29) strongly transverse and with pronounced posterior concavity of nearly semi-circular shape; sternite VIII (Fig. 30) of highly distinctive shape, posteriorly with a pronounced truncate projection and with acutely projecting posterior angles; median lobe of aedeagus (Figs 31-32) 0.9 mm long and of weakly derived morphology, with pronounced crista apicalis, and with short flagellum and additional dark structures in internal sac; paramere (Fig. 33) approximately 0.75 mm long, only slightly shorter than median lobe.: unknown.
Comparative notes: From the externally (size, coloration, proportions, punctation) similar and syntopic D. flagellata, this species is readily distinguished by the uniformly yellowish legs, the deeper impression on the male pronotum, the slightly different coloration of the elytra, the absence of a sexual dimorphism of the elytra, the competely different shapes of the male tergite and sternite VIII, the completely different morphology of the median lobe of the aedeagus, and the distinctly larger and differently shaped paramere.Comment: In the original description, which is based on a male holotype from "Yunnan, Dali" and a female paratype from "Yunnan, Kunming", Pace (1993) describes the coloration as "...; uriti liberi primo e secondo giallo-rossicci; antenne brune con i due articoli basali di un rossiccio scuro; zampe gialle con ginocchia medie e posteriori di un rossiccio scuro".The other characters specified in the very short original description are too vague to be of taxonomic use.Despite the differences in the coloration (see the redescription below), the examined material is undoubtedly conspecific with the holotype, as can be inferred from the identical shape of the median lobe of the aedeagus (compare posterior margin of eye to posterior constriction of head.Antenna (Fig. 36) 2.0-2.1 mm long; antennomeres III-X gradually increasing in width; antennomeres VI-X transverse.
Pronotum  approximately 1.06 times as broad as long and 1.10-1.13times as broad as head, broadest at anterior angles; posterior angles obtusely marked; posteriorly with small transverse impression; midline with very fine line extending cephalad from posterior impression, but not reaching anterior margin of pronotum; punctation sexually dimorphic; interstices without microsculpture and glossy.Elytra (Figs 34-35) approximately 0.9 times as long as pronotum; humeral angles marked; punctation very dense, not very coarse, but much more distinct than that of pronotum.Hind wings fully developed.Metatarsomere I shorter than the combined length of II-IV.Abdomen (Fig. 37) approximately as broad as elytra; tergites III-VII without sexual dimorphism; anterior impressions of tergites III-V and anterior portion of tergite VI impunctate; tergite III with moderately sparse to very sparse fine punctation; tergites IV-VI with setiferous punctures at posterior margin, remainder of tergal surfaces with very sparse and minute punctures; tergite VII with sparse and fine punctation, posterior margin with palisade fringe.: postero-median portion of head with moderately sparse, somewhat coarser punctures, remainder of dorsal surface with fine and sparse punctation; pronotum (Fig. 34) with extensive, but not very deep median impression in posterior half; punctation of pronotum fine and moderately dense, postero-lateral edges of postero-median impression with coarser granulose sculpture; tergite VIII (Fig. 38) strongly transverse and with distinctly serrate posterior margin, postero-lateral angles with more pronounced tooth on either side; sternite VIII (Fig. 39) weakly transverse, posterior margin weakly convex in the middle; median lobe of aedeagus (Figs 40-41) relatively small, approximately 0.65 mm long; paramere 0.6 mm long and shaped as in Fig. 42.: head (Fig. 35) with extemely fine, barely noticeable sparse punctation; pronotum (Fig. 35) somewhat depressed in posterior half, but without distinct impression (aside from the small posterior transverse impression); punctation of pronotum rather dense and fine, but more distinct than that of head; tergite VIII (Fig. 43) strongly transverse, posterior margin weakly concave in the middle; sternite VIII (Fig. 44) distinctly shorter and more transverse than that of male, posterior margin weakly convex; spermatheca 0.33 mm long, with large, apically truncate, and strongly sclerotized distal portion (Fig. 45).
Comparative notes: Among the Drusilla species recorded from China, D. yunnanensis is characterized particularly by the dense punctation of the elytra, the fine punctation of the head and the pronotum, the impunctate anterior impressions of the abdominal tergites III-V, the male secondary sexual characters, the shape of the female tergite VIII, the morphology of the median lobe of the aedeagus, and the shape of the spermatheca.

Distribution and natural history:
The currently known distribution is confined to three localities in Yunnan.Maruyama & Kishimoto (2002a) collected the species from trails of Lasius (Dendrolasius) spathepus Wheeler, 1910 and L. (D.) capitatus Kuznetsov-Ugamsky, 1927.The examined specimens were sifted from litter and various debris at the margin of a secondary mixed forest at an altitude of 1810 m.

Comment:
The trans-Palaearctic distribution of P. limbata ranges from West Europe to East Siberia (Smetana 2004).

Comment:
The original description of this species is based on two females from northern Iraq (Assing 2008b).The male sexual characters were previously unknown.Based on the above male, they are as follows: posterior margin of tergite VIII weakly concave in the middle; posterior margin of sternite VIII broadly convex; median lobe of aedeagus 0.72 mm long and shaped as in Figs 46-47.
The male sexual characters are highly similar to those of P. ruficollis (Grimm, 1845), except for the slightly longer ventral process of the aedaegus.

Comment:
The distribution of this species is confined to the Caucasus region, from Turkey and Armenia across Georgia to South Russia and Azerbaijan (Smetana 2004).
Comment: This rather widespread species has been recorded from Japan, the Russian Far East and Korea (Maruyama 2006).
impressions and punctation sexually dimorphic; interstices without microsculpture and glossy.Elytra (Figs 1-2) short, approximately 0.65 times as long as pronotum; humeral angles moderately marked; punctation sexually dimorphic; interstices without microsculpture.Hind wings reduced to short stubs extending slightly beyond posterior margin of elytra.Metatarsomere I elongate, approximately as long as the combined length of II-IV.
The previously known trans-Palaearctic distribution of D. canaliculata extended from West Europe to the Russian Far East (Smetana 2004).The above specimens from Hokkaido represent the first records from Japan.