The species of the Eastern Palaearctic region east of the Himalayas ( Coleóptera : Staphylinidae )

Types and additional material of Othius from the Eastern Palaearctic region east of the Himalayas as well as from southern China and Taiwan are revised. 31 valid species and 2 subspecies are recognized; the identity of a further 3 species, whose types were not made available, remains dubious. The following new synonymies are established: O. medius Sharp = O. medius kusuii Ito, syn. n., = O. mediusyakushimanus Ito, syn. n.; O. rosti Bernhauer = O. ussuriensis Coiffait, syn. n.; O. latus Sharp = O. stoetzneri Bernhauer, syn. n., = O. chongqingensis Zheno, syn. n., = O. latus ozakii ITO, syn. n.; O. punctatus BERNHAUER = O. arisanus (Shibata), syn. n. Lectotypes are designated for O. medius Sharp, O. rosti Bernhauer, O. stoetzneri Bernhauer, and O. ruflpennis Sharp. 19 new species and one new subspecies are described: O. bordonii sp. n. (China), O. latus gansuensis ssp. n. (China: Gansu), O. atavus sp. n. (China: Yunnan), O. lubricus sp. n. (China: Yunnan), O. discrepans sp. n. (China: Sichuan), O. excisus sp. n. (China: Sichuan), O. sculptipennis sp. n. (China: Sichuan), O. tuberipennis sp. n. (China: Yunnan), O. Jibulifer sp. n. (China: Sichuan), O. wrasei sp. n. (China: Sichuan), O. smetanai sp. n. (Taiwan), O. aequabilis sp. n. (Taiwan), O. hamatus sp. n. (Taiwan), O. acutus sp. n. (Taiwan), O. reticulatus sp. n. (Taiwan), O. praecisus sp. n. (Taiwan), O. collapsus sp. n. (Taiwan), O. contumax sp. n. (Taiwan), and O. coniceps sp. n. (Taiwan). Comparative diagnoses are given for the species and subspecies treated and complemented by comments on phylogenetic relationships; revised biogeographical and the available bionomical data are compiled. The primary and secondary sexual characters are figured; for several species, the distribution and intraspecific morphological variation are illustrated. A diagnostic key is presented for the Othius species of the Eastern Palaearctic region east o f the Himalayas. There is some evidence that both the Western Palaearctic and the Eastern Palaearctic species of Othius are monophyla; the former is distributed west and the latter east of the Caspian Sea, the only exception being the Western Himalayan O. kashmirica subgroup, which is closely related to the Western Palaearctic O. laeviusculus group. In the region east of the Himalayas, species diversity is highest in Yunnan and Sichuan (mainland China), and in Taiwan. The phylogenetics of the Taiwanese species are discussed, and conclusions regarding the colonization of the island are presented.


Introduction
After the Othius species of the Western Palaearctic, the Central Asian and of the Himalayan region have been revised in six previous contributions (A s s in g , 1997a, b; 1998a, b; A s s in g & So l o d o vn ik o v , 1998; A s s in g & W u n d e r l e , 1995), the object of the present paper is to treat the species of the remainder of the Palaearctic region, i. e. the Russian Far East, the Japanese archipelago, northern China, and adjacent regions. Strictly speaking, the mountainous areas of southern China and Taiwan belong to the Oriental region. Nevertheless, they are here treated as Palaearctic, because at higher altitudes, a considerable proportion of the flora and fauna is of Palaearctic affiliations; this is also true for the genus Othius, as will be shown below. Apart from the Palaearctic and adjacent Oriental regions, species of Othius have only been reported from New Zealand (Kl im a s z e w s k i et al., 1996); these species, however, have not yet been revised. Such an amphipolar distribution would suggest that the genus is phylogenetically old, particularly since, in Coleoptera, amphipolar vicariance patterns are usually observed at the level of tribus or subtribus (C r o w s o n , 1980). B e r n h a u e r & Sc h u b e r t (1914) list four species from the Eastern Palaearctic region east of the Himalayas, all of them from Japan. Since then, 16 further species and three subspecies have been described: nine species from China (Ber n h a u e r , 1916,1922,1931; Ca m e r o n , 1939b; Z h e n g , 1995), five species from Taiwan (It o , 1989; Sh ib a t a , 1973), one species and three subspecies from Japan (ITO, 1993), and one species from the Russian Far East (C o if f a it , 1974). One further species, O. javanus, was described from Java by Ca m e r o n (1937), but shortly afterwards referred to Metolinus Ca m e r o n of the Xantholinini (C a m e r o n , 1939a).

Measurements and abbreviations
The m easurem ents in the descriptions and the key are indicated in mm and abbreviated as follows: HW : maximal head width HL: head length from front margin o f clypeus to neck PW: maximal width of pronotum PL: length of pronotum along median line EL: length of elytra from apex of scutellum to elytral hind margin TiL: length of metatibiae (external aspect, from knee to insertion o f first metatarsomere) TaL: length of metatarsi (claws not included) TL: total length from apex o f mandibles to hind margin o f tergum VUI.
For the calculation o f the relative eye size, the postgenae were measured from the hind margin o f the eye to the neck (lateral view).

The O thius species of the Eastern Palaearctic region east of the Himalayas
As has already been shown for the previously revised species from other Palaearctic regions (AssiNG, 1997a), intraspecific variation in the genus is remarkable. This particularly applies to the widespread species, in which gene flow is hampered by a distribution which is both extensive and fragmented by various barriers (ocean, stretches of land without suitable habitats). (Note, however, that the sea barriers between and within the Eastern Palaearctic archipelagos and those separating them from China, South Korea and the Ussuri region of the East Asian mainland (e. g. Tatarskiy Proliv, Korea Strait, Yellow Sea) are either very narrow or mostly less than 100m deep, so as to allow colonization even of short-distance dispersers in various directions at least during the glacial periods of the Quaternary period.) All the synonymies recognized below refer to such species; they are in most cases a result of an underestimation of the intraspecific variation of external characters and, at the same time, of the fact that sexual characters, especially the structures in the internal sac of the aedeagus, were not or not sufficiently considered.
Evidently, a taxonomic revision is strongly dependent on the study and availability of type mate rial. There has been no difficulty in arranging loans of types deposited in Europe, Japan, and North America. Unfortunately, the same cannot be said for the types of the six species described by Z h e n g (1995), which are, according to the original descriptions, deposited in the Sichuan Teachers College, China. Repeated requests to the author for a loan of the type material have remained unanswered. It was A. B o r d o n i who provided me with paratypes of two of the species previously sent to him by Z h e n g ; a third species could be identified based on the illustration of the outline of the aedeagus in the original description. The remaining three species, however, had to be neglected and their identity must remain dubious, also because the descriptions mainly rely on variable and/or subjective external characters and because the rough sketches of the aedeagi are inadequate.

O. medius kusuii
Comments: According to the original description of O. medius kusuii, which is based on 2<f<f and 3 $ 9 from Okushiri island, the subspecies is characterized by a "slightly coarsened" ventral process of the aedeagus, the wider and posteriorly dilated head, the coarser punctation of the elytra, the more regular and more distinct microsculpture of head and pronotum, and finally the larger size. The distinguishing characters indicated for O. medius yakushimanus, whose description is based on lcf and 1 ¥ from Yakushima island, are a "shallower and less clearly defined" ventral depres sion of the aedeagus, relatively shorter parameres, the denser and coarser elytral punctation and slightly shorter penultimate antennomeres (ITO, 1993). A comparison of the holotypes of both taxa with numerous specimens collected in other Japanese localities, South Korea and China, however, showed that, apart from the slightly larger size of O. medius kusuii, the distinguishing characters indicated above are within the range of intraspecific variation of O. medius SHARP. I have seen specimens with posteriorly dilated and relatively even wider heads (see measurements) also from the Japanese mainland and from China. Elytral punc tation, the microsculpture of the forebody, and the shape of the antennomeres are highly variable in O. medius. The shape of the aedeagus (including the ventral carinae, surface sculpture, and the relative length of the parameres), too, is subject to some intraspecific variation, a phenomenon also encountered in other species of Othius (ASSING, 1997a). The slightly larger size of the head and the greater pronotal width of the holotype of O. medius kusuii (no overlap in HL, HW, and PW; see measurements) is here confirmed; in addition, the aedeagus was found to be somewhat larger than in average O. medius. On the other hand, size shift is a well-known phenomenon in island populations (G r a n t , 1998), and measurements showed that the size of the median lobe is subject to some allometric variation. The length of the median lobe was 1. Distribution and bionomics: O. medius is one of the most widespread species of the Eastern Palaearctic region. It is apparently common in Japan from Hokkaido in the north to Kyüshü in the south and is also known from Tsushima island in the Korea Strait and from various smaller islands (It o , 1993; W a t a n a b e , 1995, 1996; material examined). The species is here for the first time recorded from South Korea (Cheju-do island) and from northern and eastern China (Gansu, Zhejiang). Altitudes were only rarely indicated both on the labels of the examined specimens and in the literature; those available range between 4 5 m and 400m.  2 -11.8, 9.9. Colour variable; usual coloration: head, pronotum and most of abdomen dark brown to blackish; elytra, hind margins of abdominal segments VII and VIII, antennae, legs and mouthparts except for the apices of the mandibles lighter, yellowish to reddish brown; elytra often darkened near scutellum. Head of variable shape, but usually slender, 1.09 -1.22x (mean: 1.14) longer than wide and narrower than pronotum (HW/PW: 0.79 -0.92; mean: 0.86); eyes of similar relative size as in O. médius, ca. 1/3 -1/4 the length of temples; postgenae mostly slightly widened, sometimes ± parallel or even slightly narrowed in dorsal view; microsculpture of dorsal surface highly variable, usually predominantly composed of transverse meshes of variable length, often mixed with isodiametric meshes or transverse striae; sometimes predominantly composed of isodiametric meshes, rarely of transverse striae (particularly in posterior region of head); punctation similar to O. médius; frontal furrows variable, distinct -occasionally reaching front margin -to indistinct; antennae of similar shape as in O. médius, but antennomere V usually not oblong. Pronotum of variable shape, 1.16 -1.32x (mean: 1.23) longer than wide; maximal width in anterior half, but usually at some distance behind anterior angles; in median line posteriorly with scratch-like impression; dorsal surface with microsculpture similar to head, also very variable, in posterior half often more transverse than in anterior half; discal punctures as in O. médius. Elytra relatively short, but length subject to some variation (see measurements and Fig. 6); EL/PL: 0.53 -0.69, mean: 0.61; punctation generally less dense than in O. médius, but highly variable, often rugose near suture in anterior half and near scutellum; ± irregular short striae may be present near anterior margin and near scutellum; surface without distinct microsculpture, ± shining. Scutellum with microsculpture composed of short transverse and/or isodiametric meshes, punctu res small or indistinct; hind wing development apparently dimorphic, usually brachypterous; proportions of legs similar to O. medius; TiL/TaL: 1.09 -1.40, mean: 1.28. Abdomen with microsculpture similar to O. medius, but isodiametric microsculpture near anterior margins of terga usually weaker; surface often weakly iridescent; punctation on average less dense and finer than in O. medius-, palisade fringe at hind margin of tergum VII present, o': protarsomeres I -IV strongly dilated; sternum V and VI each antero-medially with small (smaller than in O. medius) ± oval flattened area, with tomentose long pubescence; sternum VII with shallow median impression increasing in depth and width posteriorly and with weakly concave hind margin; hind margin of sternum VIII centrally flattened to weakly impressed, its shape variable, concave to ± truncate; lateral tergal sclerites of segment IX apically acute, but shorter than in O. m edius, not reaching hind margin of tergum X, apically without spine-like process (Fig. 5e); shape of hind margin of sternum IX somewhat variable, but deeply concave, hind angles with spine-like processes (Fig. 5d); ventral process of aedeagus with pair of folds, which are weaker than in O. medius, and with basal carina; internal sac with 5 long sclerotized structures (basal median structure absent) and with apical membranous structure of characteristic shape; flagellum short, with 5 -7 coils, wide and distally darkened (Figs 5 a -c). 9: protarsomeres I -IV dilated; extent of dilatation somewhat variable, but always less than in o', sexual dimorphism pronounced; lateral tergal sclerites of segment IX apically without spine-like processes; tergum X with narrow basis, posteriorly with ca. 3 -10 rather long, apically curved modified setae; inner margin of styli with with fringe of numerous long, rather stout setae (Fig. 51).
Comments: B e r n h a u e r (1907) based his original description on three specimens from "Nemoro und Sapporo auf Jesso" without specifying a holotype. Therefore, and because of the possibility that the undiscovered third syntype may refer to a different species, a lectotype designation was deemed necessary. As the syntype B e r n h a u e r had labelled "Typus" has a somewhat deformed pronotum, the second syntype in his collection was selected as lectotype.
In the original description of O. ussuri ensis, Co if f a it (1974) compared the species only with O. chtysurus R e it t e r , but did not refer to any of the species described from Japan. A request for a loan of the holotype in the Khnzorian collection in Yerevan has remained unanswered. The paratype from the Coiffait collection, however, is without doubt conspecific with O. rosti B e r n h a u e r , so that O. ussuriensis is here treated as a junior synonym of that species.
Comparative notes: From all its congeners, O. rosti is distinguished by the characteristic internal structures of the aedeagus. From the often similar O. medius, it is in addition separated by on average smaller size, by the different secondary sexual characters on the <f sterna V and VI, and especially by the absence of spine-like processes on the lateral tergal sclerites IX. In O. latus, the body and especially the head are larger and broader, the colour is on the whole darker, and the d* sterna V and VI are unmodified.
Distribution and bionomics: O. rosti is a widespread species, but less so than 0 . m edius and with a somewhat more northern distribution, which ranges from Primorskiy Kray and adjacent regions, Sakhalin, and the southwestern Kuril Islands in the north to Honshu in the south (W a t a n a b e , 1994, 1995, 1996; material examined). It is apparently absent from the southern islands of the Japanese archipelago. Specimens from the south of the range of distribution tend to be larger than those from the north (Fig. 6). The labels of the material examined indicate lower to intermediate altitudes (300 -1280m). The species has been collected from April through November; several specimens taken in May (3 ex.), July (4 ex.) and August (1 ex.) were teneral. The ovaries of three dissected $ 9 collected in July and August each contained a mature egg.  Comments: Without specifying a holotype, Sh a r p (1874) based his original description of O. latus on two 9 9 from Japan ("Copper Temple, Nagasaki") and a < ? from Northern China, which he believed to belong "no doubt to the same species". Consequently, all three specimens were originally syntypes. In stating that he examined the holotype [sic!], probably because of the type label attached to the specimen, and giving a photograph of the labels, Ito (1993) designated a lectotype. B e r n h a u e r (1931) based his original description of O. stoetzneri on at least two syntypes kept in his own collection and in the "Sammlung des Dresdener Staatsmuseums". Since the possibility cannot be completely ruled out that further unexamined syntypes exist, which are not conspeciflc with the examined syntypes, the undamaged 9 syntype in the collection of the SMTD is here designated as lectotype. The description of O. stoetzneri (BERNHAUER, 1931), in which the author compares O. stoetzneri with O. latus emphasizing the different shape of the head and the different punctation, is apparently a result of B e r n h a u e r 's erroneous interpretation of O. latus. All the specimens in the Bernhauer collection identified by him as "0 . latus" were in fact O. médius.
It o (1993) indicates eye size, th e w id th o f th e an ten n ae, th e m icrosculpture o f th e in terstices on th e elytra, th e less shiny a n d d a rk e r ap p earan ce, a n d th e m ore slen d er a n d sm aller body as An examination of the types of the taxa treated here and of additional material from further localities revealed considerable intraspecific variation in O. latus, both within and between populations (Figs 8,9), but yielded constant differences neither in external morphology nor in the primary and secondary sexual characters, including the internal structures of the aedeagus. Specimens from China and southern Japan (types of O. stoetzneri, O. chongqingensis, O. latus and additional material) tend to be larger and have longer elytra than those from the northeastern part of the range of distribution (Sakhalin, Hokkaido, northern Honshu) (Figs 8,9). Wing length poly-and dimorphism is not uncommon in the genus (A ssin g 1997a, 1998b), and the length of the elytra is apparently extremely variable in O. latus (see measurements and Fig,9). Considerable intraspecific variation was also observed in elytral punctation, head width, pronotal shape, microsculpture and colour. In addition, there is some -probably allometric -variation in the size of the median lobe, which is usually larger in the Chinese material than in specimens from the northeast of the area of distribution. The shape (length, width, outline) of the internal structures may somewhat differ especially between (and apparently also within) populations, but the general pattern is very similar (Figs 7 c -f). Constant differences in the secondary sexual characters were not observed. Pronounced intraspecific variation, especially between populations, is what should be expected with a species like O. latus. Both the characteristics of the area of distribution, which is vast and at the same time fragmented by barriers (water and stretches of land without suitable habitats), and the probably low dispersal power of the species are bound to impede gene flow. Judging from the length of the elytra most, if not all the specimens examined from the northwest of the range of distribution (Sakhalin, northern Honshu, Hokkaido) were incapable of flight. In the absence of convincing morphological and biogeographical evidence in favour of the hypothesis that the different population samples should represent distinct species or subspecies, they are here regarded as subpopulations of one polytypical species. This view is also supported by the fact that O. Distribution and bionomics: The area of distribution of O. latus latus ranges from Sakhalin in the northeast to eastern central China in the southwest and includes the Japanese archipelago (Hok kaido, Honshü, Kyüshü). Apparently it occurs at lower altitudes; the altitudes specified on the labels of the examined specimens range from 300 to 900m. Adult specimens were collected in May and from July through November. One 9 taken in July (Sakhalin) was teneral; the ovaries of a 9 collected in August (Hokkaido) contained a mature egg. Distribution and bionomics: The subspecies is known only from several localities in Gansu, China, where it was collected in July and August. Apparently, it inhabits higher altitude habitats than 0. latus latus; the altitudes specified on the labels range from 2200 to 3600m.  lateral tergal sclerite IX and tergum X in lateral view (g); outline of o' sternum VUI (h); long setae partly or completely omitted in f -h. Scales: 0.5 mm.
Colour of head, pronotum and abdomen dark brown to blackish; mandibles, femora and tibiae reddish to dark brown; antennae, except for the sometimes darkened basal antennomeres, ferrugineous; elytra light reddish; tarsi and posterior margin of abdominal segment VIII yellowish.
Head slender, distinctly oblong (see ratios); eyes large and prominent, ca. half the length of postgenae in dorsal view, in lateral view 0.60 -0.67x (in the specimen from China only 0.5x) the length of postgenae; temples behind eyes ± parallel or narrowed in dorsal view; transverse mi crosculpture on dorsal surface fine; punctation rather coarse and dense not only on sides, but also near occiput; absent in central dorsal area; frons with two pairs of large punctures in ± quadrate arrangement, the anterior punctures deeper, larger, situated in oval grooves and accompanied by further -partly confluent -punctures; frontal furrows rather shallow, not reaching front margin; antennae long and slender, penultimate antennomeres not transverse. Pronotum comparatively slender (see ratio); maximal width a short distance anterior to middle or nearer to anterior angles; in median line posteriorly with a scratch-like impression; dorsal surface with microsculpture similar to head; position of discal punctures similar to O. medius. Elytra very long (see ratio); punctation well-defined, not very coarse, but very dense, interstices ap proximately equal to diameter of punctures; rugose punctation only near scutellum; surface without distinct microsculpture, shining; scutellum with distinct microsculpture composed of isodiametric and transverse meshes, and with several punctures; macropterous; legs long and slender. Abdomen with the area near the anterior margins of the terga mat due to distinct and extensive isodiametric microsculpture; the remaining tergal surfaces at most with very indistinct micro sculpture and weakly iridescent; punctation dense and relatively fine, finer and sparser on terga VII -VIII than on anterior terga; palisade fringe at hind margin of tergum VII present. <f: protarsomeres I -IV moderately dilated; sternum V and VI each centrally with small (much smaller than in O. medius) oval area with long, tomentose yellowish pubescence and rasp-like sculpture; sternum VII posteriorly weakly flattened and weakly concave to truncate hind margin; hind margin of sternum VIII truncate or weakly concave (Fig. llh ); lateral tergal sclerites of segment IX long, reaching, or slightly protruding over hind margin of tergum X, with one terminal seta, and apically without spine-like process ( Comments: The specimen from the Emai Shan differed from the two types in the slightly smaller eyes, the relatively shorter elytra, and in the lighter colour of the basal antennomeres. In addition, the dr sternum IX was somewhat less oblong, the aedeagus was slightly larger, the ventral process somewhat broader, and the relative length of the sclerotized structures in the internal sac of the aedeagus differed slightly (Figs 11 a -e). These differences, however, may well be within the range of intraspecific variation, and in general, the arrangement and shapes of the internal structures are quite similar. Slight differences in the shape of the ventral process were also observed in the two type specimens, and the size and shape of the aedeagus is quite variable in O. punctatus, too (see section 4.8). Therefore, the specimen from China is here regarded as conspecific with the type material of O. ruflpennis.
Distribution and bionomics: O. ruflpennis has become known only from the type locality, "Orakami Marsh", which is in Nagasaki City (Japan: Kyushu) and from the Emai Shan in Sichu an, China (Sh a r p , 1874; material examined; see also Z h e n g , 1995). Remarkably, further records are unknown, although the external morphology of the species (length of wings, eye size etc.) suggests that it is capable of flight. Head distinctly oblong and narrower than pronotum (cf. ratios), posteriorly subparallel; eyes large, ca. 0.5x the length of postgenae; macropunctation rather coarse and extensive, present on whole posterior half of dorsal surface; posterior pair of frontal punctures absent; frons anteriorly with shallow impression with 10 mostly large punctures; frontal furrows shallow; microsculpture fine and rather shallow, distinctly transverse; antennae long, antennomeres IV weakly oblong, V subquadrate, and VI -X weakly transverse. Pronotum with maximal width in anterior half, but nearer to middle than to anterior angles, posteriorly weakly tapering; lateral margins in posterior half indistinctly concave in dorsal view; microsculpture and shine similar to head; arrangement and position of dorsal punctures similar to O. rufipennis, posterior puncture very close to hind angles; posterior median scratch-like im pression distinct.
Elytra long (see measurements and ratio); punctures relatively small, but well defined, much sparser than in O. rufipennis, interstices mostly distinctly wider than punctures; microsculpture similar to O. opacipennis, composed of distinct micropunctation; micropunctures slightly less dense, surface therefore with more shine than in O. opacipennis-, scutellum mat due to very distinct microreticulation, several punctures present, though barely noticeable because of the microsculpture; hind wings present. Abdomen with punctation very fine and distinctly less dense than in O. rufipennis-, microsculpture near anterior and lateral tergal margins very distinct and isodiametric, on central and posterior surface of terga shallow and transverse; palisade fringe at hind margin of tergum VII present, o': protarsomeres I -IV moderately dilated; sternum V and VI centrally with subcircular area with minute punctation and very sparse pubescence; sternum VII truncate, not flattened posteriorly; sternum VIII very oblong, with dark pubescence and posteriorly distinctly convex (shape similar to ¥ sternum VIII in other species); sternum IX very long and slender, with mostly dark pu bescence, posteriorly weakly concave and with numerous long dark setae (Fig. 12d); lateral tergal sclerites of segment IX similar to O. rufipennis, very long and apically acute, with one terminal seta, and just reaching hind margin of tergum X (Fig. 12e); aedeagus of remarkable morphology (shape similar to that in Atrecus and Parothius): ventral process extremely long and slender, basal carina short and indistinct; parameres very long, apically weakly dilated; internal sac with 5 very weakly sclerotized structures: an almost unsclerotized median apical, and two pairs of ± fused lateral structures; flagellum rather wide and with ca. 10 -15 coils (Figs 12 a -c). $: unknown.
Derivatio nominis: The name (lat.: ancestor) alludes to the morphology of the aedeagus, which somewhat resembles that in Atrecus and Parothius and may therefore represent either an ancestral or an atavistic character.
Comparative notes and phylogenetics: From all its Palaearctic congeners O. atavus is di stinguished by the remarkable morphology of the aedeagus. From those Eastern Palaearctic species with a palisade fringe at the hind margin of tergum VII, it is in addition separated by the following character combination: head without posterior pair of frontal punctures, elytra dark reddish and with distinct and dense micropunctation. The external and the male secondary sexual characters (slender head, microsculpture and puncta tion of head and pronotum, and the shape and chaetotaxy of the lateral tergal sclerites IX) suggest a closer relationship with O. rufipennis. The flagellum is also rather short and wide, but otherwise the morphology of the aedeagus is completely different.
Distribution: The species is only known from the type locality in Yunnan, China. Its external morphology suggests that it is more widely distributed. Colour of body ± uniformly dark to blackish brown, with the antennae only slightly lighter; hind margin of abdominal segment VIII, mouthparts, and legs yellowish to light brown, with the internal faces of the tibiae and parts of the femora in mature specimens bright reddish or infúscate to various degrees. Head with shape similar to O. rufipennis, distinctly narrower than pronotum; temples behind eyes usually ± parallel, occasionally weakly widened or narrowed in dorsal view; microsculpture shallow, predominantly composed of transverse striae; macropunctation variable, on average denser in material from the Chinese mainland than in specimens from Taiwan; eyes slightly smaller than in O. rufipennis, but size highly variable, 0.40 -0.58x the length of postgenae in lateral view; posterior pair of frontal punctures present, sometimes accompanied by additional punctures; frontal furrows in most specimens weakly impressed, sometimes ± absent; antennae slender, penultimate antennomeres ± subquadrate, not distinctly transverse. Pronotum with shape similar to O. rufipennis, maximal width in anterior half, often nearer to middle than to anterior angles; lateral margins in posterior half usually straight to weakly convex; microsculpture weak and composed of transverse striae; position of discal punctures similar to O. rufipennis, but posterior puncture more distant from hind angles (Fig. 131). Elytra relatively long, but absolute and relative length subject to considerable intraspecific varia tion (see measurements and Fig. 15); punctation very variable, usually well-defined and interstices on average of similar width as punctures, near scutellum often with confluent punctation; inter stices sometimes with traces of shallow microsculpture; macropterous; scutellum with distinct microsculpture usually predominantly composed of isodiametric and/or transverse meshes, and with several punctures; legs, especially tarsi, shorter than in O. rufipennis.  (j); tergum X o f ¥ from Taiwan (k); schematic representation of pronotal punctation pattern (I); long setae partly or completely omitted in i, j. Scales: 0.5 mm; 1 without scale.
o' : protarsomeres I -IV moderately dilated; sternum V and VI with rather large, usually ellipsoid, sometimes subcircular central callosities of somewhat variable size and shape; callosities usually shallowly impressed, with dark tomentose pubescence, which is directed postero-mediad and which is sometimes denser on sternum V than on sternum VI, rasp-like microsculpture indistinct; sternum VII posteriorly concave and distinctly flattened or shallowly impressed; sternum VIII with dense and rather long black pubescence, posteriorly weakly concave, centrally with shallow, ± triangular impression, without row of marginal setae; sternum IX with dense and mostly blackish pubescence, posteriorly broadly and moderately to deeply concave (Fig. 13h); lateral tergal sclerites of segment IX with numerous long and one or two terminal setae, apically acute, shorter than in O. rufipennis, nearly reaching hind margin of tergum X in lateral view (Fig. 13i); aedeagus with apically dilated parameres; ventral process without basal carina, but with pair of pronounced and long lateral folds and with pair of less distinct apical folds of variable length; internal sac with 6 sclerotized structures: a long median basal, a somewhat triangular apical, a pair of longer straight lateral, and a pair of shorter straight lateral pieces; flagellum thin and with ca. 30 coils (Figs 13 a -g). ¥: protarsomeres I -TV weakly dilated, sexual dimorphism usually pronounced; tergum X with very variable number (0 -7) of modified setae (Figs 13 j -k); inner margin of styli with fringe of numerous setae, which are short and thin near apices of styli and very long and stout near the bases.
Com parative n o te s: For distinction from O. rufipennis and O. maculativentris Z h e n g see the diagnosis above and the diagnosis of the latter species, respectively. From the externally also similar O. atcrvus, O. punctatus is distinguished especially by the presence of the posterior pair of frontal punctures on the head, by the absence of distinct elytral microsculpture, and by the primary and secondary sexual characters. from Alishan and that from Houhuanshan. The specimens from Gansu, Sichuan and Shaanxi also differed from the material of other localities in that the colour of the legs was bright reddish and that the ventral process was slightly wider. Considerable intraspecific variation in external cha racters, especially in size, is what can be expected in a widely distributed species, particularly in island populations, and is a well-known phenomenon in several widespread Othius species both from the Eastern Palaearctic and from the Western Palaearctic region, e. g. O. medius, O. latus, O. rostí (see above), and O. lapidicola M ä r k e l & K ie s e n w e t t e r (A s s in g , 1997a). The shorter elytra in the material from Taiwan (Fig. 15) are hardly surprising; in many island Staphylinidae, particularly in soil-dwelling and more or less K-selected species, the wings are reduced as a result of selection operating against active dispersal by flight. I have been unable to find any further constant differences between the material from the Chinese mainland and that from Taiwan, neither in external morphology nor in the primary and secondary sexual characters. Therefore, 0. arisanus Sh ib a t a is here regarded as junior synonym of O. punctatus B e r n h a u e r .
Distribution and bionomics: O. punctatus is widespread in China (Sichuan, Gansu, Shaanxi, Shandong, Hunan, Zhejiang) and Taiwan. In Taiwan, O. punctatus is apparently the most com mon and widely distributed Othius species (Map 1, p. 85). In addition to the localities indicated above it was also collected on the Houhuanshan, Nantou (ITO, 1989). The species seems to inhabit a wide range of altitudes; the material examined was collected from as low as 450m (China) up to 2600m (Taiwan). The type specimens of O. arisanus "were collected among the roots of grasses near the margin of a pond" (S h ib a t a , 1973); some of the material from Shaanxi, too, was collec ted near water. Adult specimens have been observed from April through May and from M y through October (ITO, 1989; Sh ib a t a , 1973; and material listed above). Teneral specimens were collected in April and July; the ovaries of a dissected ¥ collected in July contained a mature egg. Pronotum on the whole similar to O. punctatus, but lateral margins in posterior half in most specimens weakly concave; microsculpture shallow, usually composed of transverse striae, sometimes with interspersed isodiametric or transverse meshes; pronotal punctation pattern similar to O. punctatus, but posterior puncture very close to hind angles (Fig. 16h). Elytra long, but absolute and relative length subject to some intraspecific variation; punctation similar to O. punctatus', interstices usually ± opaquely shining, sometimes with traces of shallow microsculpture; length of hind wings variable, in the material examined ca. 1.5 -2.5x the length of elytra; scutellum with ± distinct microsculpture usually predominantly composed of transverse meshes and striae, and with several punctures. Abdomen with punctation and microsculpture similar to O. punctatus-, palisade fringe at hind margin of tergum VII present. o": dilatation of protarsomeres I -IV variable, usually pronounced, occasionally moderate; sternum V and VI with small subcircular, weakly elevated central callosities of somewhat variable size and shape, with not very dense dark pubescence and shallow microsculpture; callosity of sternum V even smaller than that of sternum VI; sternum VII posteriorly weakly concave, only weakly flattened; sternum VIII with dense and rather long black pubescence, its hind margin weakly concave to truncate, centrally ± flattened, and at most with few short marginal setae (Fig. 16d); sternum IX with dense and mostly blackish pubescence, posteriorly broadly and moderately to deeply concave; lateral tergal sclerites of segment IX similar to O. punctatus, but broader in lateral view (Fig. 16e, f); aedeagus completely different from that in O. punctatus: ventral process broad and short, apically weakly pointed, without distinct lateral folds, but with median apical carina; basal carina indistinct; dorsal membranous region very small and situated much more distally than in O. punctatus and related species; parameres apically weakly dilated and basally with ± irregular outline; internal sac with 6 sclerotized stmctures: a long median basal, an approximately equally long median apical, a pair of thin long lateral, and a pair of stouter bifid lateral pieces; flagellum thin and with ca. 15 coils (Figs 16 a -c). $: dilatation of protarsomeres I -IV variable, weak to moderate, in some 9 ¥ as strongly dilated as in d V with relatively weakly dilated protarsomeres; sternum VIII posteriorly of variable shape, weakly concave to weakly convex; tergum X more distinctly pointed than in O. punctatus, poster iorly with several long, rather thin and apically curved modified setae; inner margin of styli with fringe of numerous setae, which are short and thin near apices of styli and very long and stouter near the bases (Fig. 16g).
Com parative notes: From the externally similar O. punctatus, O. maculativentris is readily distinguished by the larger and wider head, the darker legs, the different pronotal punctation pattern (posterior pronotal puncture very close to hind angles), the posteriorly usually concave pronotal lateral margins, and especially by the completely different shape and internal structures of the aedeagus.
Comments: Repeated requests for a loan of the types of the species have remained unanswered. Nevertheless, there is little doubt that the material which the above diagnosis is based on in fact refers to O. maculativentris. The most conclusive evidence is provided by the drawing of the aedeagus in Z h e n g (1995); the internal structures are not illustrated, but the shape of the aedeagus is very characte ristic. In addition, the (few) external characters indicated in the original description are in good agreement with the material studied. Finally, O. maculativentris was described from Sichuan (Wolong), not far from where the specimens indicated above were collected, and both the external morphology (eye size, length of hind wings, presence of palisade fringe) and the altitude at which the species was taken (see below) suggest that it is probably more widely distributed.
Distribution and bionomics: The species is presently known only from Wolong and Gongga Shan in Sichuan, western China, where it was collected at altitudes between 1920 -3300m (Z h e n g , 1995; material examined). Many of the specimens taken at the end of May and in July (see above) were teneral. In external appearance similar to and apparently the sister species of O. apicalis C a m e r o n ; for a detailed diagnosis of that species see A s s in g (1998a). Colour of body black, legs dark brown to blackish, tarsi, palpi and hind margins of abdominal segments VII and VIII light brown to reddish brown. Head smaller, more slender and with denser macropunctation than in O. apicalis; eyes larger, in lateral view 0.58 -0.66x the length of postgenae; outline of head weakly widened posteriorly; microsculpture composed of transverse striae and transverse meshes, but very shallow, surface therefore shining; posterior pair of frontal punctures present and relative large; frontal furrows relatively shallow; antennae slender, penultimate antennomeres weakly transverse. Pronotum of sim ilar shape and with similar punctation pattern as in O. apicalis (cf. Fig. lg in A s s in g , 1998a); microsculpture reduced to very indistinct traces visible only at higher magnifica tions, surface therefore very shiny. Elytra relatively long (see measurements); microsculpture indistinct; punctation much denser than in O. apicalis, diameter of interstices clearly smaller than that of punctures; scutellum with distinct microsculpture composed of transverse meshes and with several punctures. Abdomen with clearly denser punctation than in O. apicalis; microsculpture both near anterior margins of terga and on posterior tergal area very weak and shallow; palisade fringe at hind margin of tergum VII present. o": protarsomeres I -IV strongly dilated; sternum V and VI without modifications; sternum VII weakly concave posteriorly; sternum VIII with not very dense black pubescence, its hind margin ± truncate, centrally with small and shallow impression of ± triangular shape, without marginal setae; sternum IX with relatively sparse, long and mostly blackish pubescence, posteriorly deeply concave (Fig. e); lateral tergal sclerites of segment IX slender almost reaching hind margin of ter gum X (Fig. 17f), with one terminal seta. General morphology of aedeagus similar to that of O. apicalis-, ventral process without distinct carinae or folds, apically acutely pointed and with sinuate lateral margins in ventral view; parameres apically weakly dilated, relatively shorter than in O. apicalis; internal sac with 6 sclerotized structures: a long and distally wide median basal, an almost equally long median apical, a pair of thin long lateral, and a pair of stouter, characteristically shaped lateral pieces; flagellum thin and with ca. 25 coils (Figs 17 a -d). 9: protarsomeres I -IV distinctly dilated, only slightly less so than in d", sexual dimorphism therefore not very pronounced; hind margin of sternum VIII weakly convex and with numerous yellowish apical setae; tergum X posteriorly broadly rounded, setae unmodified; inner margin of styli with fringe of numerous long and stout setae (Fig. 17g).
Derivatio nominis: The name (lat.: smooth, slippery) refers to the remarkably shiny surface of the head and especially of the pronotum.
Com parative notes: From the similar O. apicalis, O. lubricus is readily distinguished by the denser punctation of head, elytra and abdomen, by the presence of a pair of frontal punctures, by the distinctly larger eyes, and by the < ? primary and secondary sexual characters. O. taiwanus It o , a species with an equally shining pronotum, is much larger, its head is distinctly more massive, the eyes are relatively smaller, the pronotum is more strongly widened anteriorly, the cf primary sexual characters (size and shape of aedeagus, internal structures) are completely different, and the $ tergum X is equipped with modified setae posteriorly.
Comments: As can be inferred from the drawings of the aedeagal outlines -the internal structures are either not or inadequately illustrated -and from the external characters referred to, none of the species described by Z h e n g (1995), all of them from Sichuan, is conspecific with O. lubricus.
Distribution and bionomics: The species is known only from Yunnan, southwestern China. Both its external morphology (eye size, wing development) and the relatively low altitude (1800 -2300m) of the localities, where the material was collected, suggest a wider distribution. Two specimens taken in May were teneral. Colour of body ± uniformly reddish brown to dark brown, the head slightly darker.
Head rather large and distinctly oblong, behind eyes ± subparallel; eyes relatively small, 0.28 -0.33x the length of postgenae in lateral view; macropunctation relatively dense and extensive, covering the whole posterior half of dorsal surface (nontype specimen) or nearly so; posterior pair of frontal punctures present; frontal furrows shallowly sulcate. Microsculpture on most of dorsal surface predominantly composed of isodiametric and short transverse meshes, near occiput and laterally also of transverse striae and meshes; antennae long and slender, antennomeres HI ca. 3x longer than wide, IV -VII distinctly oblong, and X ± subquadrate. Pronotum of veiy characteristic shape, very slender, as wide as head (see ratios) and dorsally flattened, so that the full length o f both lateral margins is visible when viewed from above (in other species of Othius the lateral margins are usually not visible at and near the anterior angles in dorsal view); outline of pronotum only weakly narrowed posteriorly; lateral margins in posterior half weakly sinuate; posterior angles more distinct than is usually the case in Othius, and hind margin only weakly convex; discal puncture near lateral margin, but distinctly distant from hind angles (Fig. 18b); microsculpture fine, predominantly composed of transverse striae and long transverse meshes; posterior median scratch-like impression present. Elytra relatively long (see measurements); punctation well-defined but not veiy coarse and moderately dense, interstices on average wider than punctures; microsculpture indistinct; surface with weak opaque shine; scutellum with rather shallow transverse microsculpture and with several punctures; hind wings fully developed. Abdomen with distinct, but fine punctation; microsculpture extremely shallow, barely visible even at higher magnifications, transverse microsculpture only slightly more distinct near anterior margins of terga; palisade fringe at hind margin of tergum VII present, a": not available for examination. ?: protarsomeres I -IV moderately dilated; hind margin of sternum VIII centrally obtusely pointed and with numerous yellowish apical setae; tergum X posteriorly broadly rounded, setae near hind margin long and unmodified; inner margin of styli with fringe of not particularly dense setae, which are rather short and thin posteriorly, and long and stout anteriorly (Fig. 18a).
Com parative notes: O. badius is distinguished from other Chinese congeners based on external characters alone: the ± uniform brown colour, the long antennae, the relatively small eyes in combination with the presence of a palisade fringe and fully developed wings, the extensive punctation of the head, and especially the characteristic shape and punctation pattern of the pronotum. Head and abdomen dark brown; pronotum, elytra, and antennae reddish brown; legs testaceous. Head distinctly oblong and narrower than the pronotum (cf. ratios); outline of head subparallel; eyes small, in lateral view 0.25x the length of postgenae; macropunctation present on dorsal surface of most of the posterior half, distinctly sparser than in O. badius\ posterior pair of frontal punctures present; both anterior frontal punctures in HT accompanied by 2 additional punctures; frontal furrows very shallow; microsculpture of whole dorsal surface distinct and isodiametric; antennae shorter and less slender than in O. badius, antennomeres IV ± subquadrate, and V -X weakly transverse. Pronotum widest in anterior half, maximal width nearer to middle than to anterior angles, poster iorly moderately tapering; microsculpture similar to head, also isodiametric, but shallower; pronotal punctation pattern very characteristic: with five discal punctures on either side; posterior pronotal puncture clearly distant from hind angles and from lateral margins (Fig. 19f); posterior median scratch-like impression distinct and long. Elytra very short (see measurements and ratio); microsculpture weak, but surface only with little, slightly opaque shine; punctation rather shallow and not very dense, interstices wider than punctures; anterior part of elytra and scutellum with a wide and rather deep transverse impression, which in normal conditions is apparently covered by pronotum, so that only the apex of the scutellum is visible; scutellum therefore in cross-section not flat, but distinctly impressed; scutel lum with shallow transverse microsculpture and without punctation, surface with distinctly more shine than that of elytra; hind wings reduced. Abdomen with moderately dense and fine punctation; microsculpture both near anterior margins of terga and on posterior tergal area transverse and extremely shallow, surface shining; palisade fringe absent. o': protarsomeres I -IV distinctly dilated; sternum V and VI without distinct callosities, but centrally with small subcircular areas with dense punctation and pubescence; sternum VII weakly concave posteriorly; sternum VIII with rather sparse and short dark pubescence, its hind margin convex and without marginal setae; sternum IX with relatively sparse, mostly yellowish pubescence, posteriorly deeply concave (Fig. 19d); lateral tergal sclerites of segment IX apically acute, just reaching hind margin of tergum X (Fig. 19e), with one terminal seta; ventral process with short basal carina, but without lateral folds, apically pointed and with curved lateral margins in ventral view; parameres apically weakly dilated; internal sac with 6 sclerotized structures: a very long median basal, a much shorter median apical, a pair of thin long lateral, and a pair of stouter, weakly curved and basally dilated lateral structures; flagellum thin and with ca. 30 coils (Figs 19 a -c). Derivatio nominis: The name (lat.: deviating, differing) refers to the remarkable pronotal puncta tion pattern.
Com parative notes and comments: From all its known congeners, O. discrepans is separated by its pronotal punctation pattern. The species is also characterized particularly by the distinctly isodiametric microsculpture of head and pronotum, by the narrow and oblong head, and by the internal structures of the aedeagus. In addition, among the other species without a palisade fringe from the Chinese mainland, it is distinguished by its smaller size and lighter colour. Distribution and bionomics: The species is known only from the type locality in Sichuan, where, judging from its external morphology (eye size, reduction of wings and palisade fringe), it is probably more or less endemic. No further data, apart from those indicated on the labels of the holotype, are available. Head and pronotum blackish; elytra and abdomen, except for the slightly lighter tergal and sternal hind margins, blackish brown; tibiae and femora ± dark brown; tarsi and antennae reddish brown. Head distinctly oblong and narrower than the pronotum (cf. ratios); lateral margins of head subparallel; eyes of moderate size, in lateral view 0.25x the length ofpostgenae; macropunctation in lateral areas of posterior half rather sparse, in central dorsal region absent; posterior pair of frontal punctures present, but small; anterior frontal punctures small and accompanied by 1 -2 additional punctures; frontal furrows shallowly sulcate; microsculpture as in O. discrepans on whole dorsal surface distinct and isodiametric; antennae slender, antennomeres IV -X subequal in length and oblong. Pronotum rather massive (cf. measurements and ratios), wider than head and elytra, widest in anterior half, maximal width nearer to middle than to anterior angles, posteriorly weakly tapering; microsculpture similar to head, also isodiametric; pronotal punctation pattern characteristic: posterior puncture at considerable distance from hind angle and from lateral margin, intermediate puncture accompanied by an additional puncture (Fig. 20f); posterior median scratch-like impres sion distinct and long. Elytra short (see measurements and ratio); microsculpture absent, surface with distinct, slightly opaque shine; punctation ± well-defined and rather dense, interstices approximately as wide as, in some areas smaller than punctures; scutellum only weakly impressed, with shallow transverse microsculpture and with some small punctures; hind wings reduced.
Abdomen with rather dense and fine punctation; microsculpture near anterior tergal margins transverse and very shallow, on posterior tergal area ± absent, surface therefore with distinct shine; palisade fringe on hind margin of tergum VII absent. o': protarsomeres I -IV strongly dilated; sternum V with minute subcircular, indistinctly elevated central area with condensed punctation; sternum VI with small subcircular area with ± tomentose dark and long pubescence, this area slightly larger than that of sternum V; sternum VII weakly concave and only indistinctly flattened posteriorly; sternum VIII very oblong, with rather dense and long dark pubescence, its hind margin distinctly concave and with short marginal setae; sternum IX with not very dense dark pubescence, posteriorly deeply excised (Fig. 20d); lateral tergal sclerites of segment IX very long and apically acute, projecting beyond hind margin of tergum X (Fig. 20e); ventral process long and slender, with short sclerotized basal carina, but without lateral folds; parameres apically weakly dilated; internal sac with 5 sclerotized structures: a short median apical, a pair of thin long lateral, and a pair of stouter and shorter lateral structu res; flagellum moderately thin and with ca. 15 coils (Figs 20 a -c). $: unknown.
Derivatio nominis: The name (lat.) refers to the deeply excised posterior margin of the cf sternum IX.
Com parative notes and phylogenetics: 0. excisus is apparently closely related to 0. discrepans, with which it shares the following synapomorphies: the distinctly isodiametric microsculpture of the head and pronotum, the position of the posterior pronotal puncture, the slender head, the reduction of wings and palisade fringe, and the reduction of the cr secondary sexual characters on tergum V and VI. In addition, the external morphology of the aedeagus is similar. It is, however, readily distinguished from that species by the distinctly larger size and darker colour, the larger eyes, the different pronotal punctation pattern, and by the cf primary and secondary sexual charac ters. For distinction from the following brachypterous congeners from the Chinese mainland see below those species.
Comments: The only completely dark species with the head and the elytra narrower than the pronotum in Zh e n g (1995) is the Taiwanese O. parvipennis, so that O. excisus is very unlikely to be conspecific with any of the species described in that study.
Distribution and bionomics: The species is known only from the type locality in southern Sichuan. As can be inferred from its external morphology (eye size, reduction of wings and palisade fringe), it is probably more or less endemic. No further data, apart from those indicated on the labels of the holotype, are available.
4.14. Othius sculptipennis sp. n.  Head oblong and slightly narrower than pronotum (cf. ratios), of characteristic oval shape, hind angles obsolete, postgenae in dorsal view evenly curving towards neck; eyes relatively small, in lateral view 0.28x the length of postgenae; macropunctation extensive, present on all of posterior b half of dorsal surface, denser and less coarse than in O. excisus; posterior pair of frontal punctures absent; anterior frontal punctures accompanied by 2 -3 additional punctures; frontal furrows dis tinctly sulcate; whole dorsal surface mat due to fine, but remarkably distinct and isodiametric microsculpture; antennae long, antennomeres IV weakly oblong, V subquadrate, and VI -X weak ly transverse. Pronotum with maximal width near anterior angles, posteriorly distinctly tapering; microsculpture similar to head, also isodiametric, much more distinct than in O. excisus, surface therefore mat; posterior puncture relatively close to lateral margin, but at some distance from hind angle; intermediate puncture with 1 -2 small additional punctures; anterior puncture in HT absent on left side (Fig. 21b); posterior median scratch-like impression distinct. Elytra short (see measurements and ratio); microsculpture very characteristic, mostly composed of fine and dense, ± distinct rugae of variable length and direction, in which the large, though shallow punctures are barely visible; surface mat; scutellum with microsculpture composed of transverse and isodiametric meshes, and with some small punctures; hind wings reduced. Abdomen with fine punctation; microsculpture very fine, but distinct, composed of very transverse meshes and transverse striae; near anterior margins of terga slightly more distinct and composed of shorter transverse and isodiametric meshes; surface with reduced shine; palisade fringe at hind margin of tergum VII absent, d": unknown. 9: posterior margin of sternum VIII concave and with fringe of dense, long and rather stout yellow and with interspersed long black setae; lateral tergal sclerites IX apically very acute; tergum X posteriorly with membranous appendage and with some relatively short modified setae (Fig. 21a).

Figs 21 a -b
Derivatio nominis: The name (lat.) refers to the characteristic microsculpture of the elytra.
Com parative notes and phylogenetics: O. sculptipennis is readily distinguished from all its known congeners based on external characters alone, especially by the remarkably distinct micro reticulation of head and pronotum, the very characteristic microsculpture of the elytra, and the shape of the head. In the Palaearctic region, the only further species of similar size and without a palisade fringe on the hind margin of tergum VII is O. strigulosus W o l l a s t o n from Madeira. O. sculptipennis is apparently most closely related to O. excisus and O. discrepans, with which it shares the isodiametric microsculpture of the head and pronotum and the reduction of wings and palisade fringe, and which also occur in Sichuan. In addition to the characters indicated above, it differs from them in the absence of a posterior pair of frontal punctures on the head and in the different shape and punctation o f the pronotum.
Distribution and bionomics: The species is known only from the type locality in Sichuan. Both its external morphology and the altitude at which it was collected (3500m) suggest that it may be more or less endemic. The ovaries of the holotype contained a mature egg. CAMERON, 1939 Fig. 22a Othius opacipennis CAMERON, 1939b: 162f.  Head slightly longer than wide; eyes large, 0.45 -0.50x the length of postgenae in lateral view; temples behind eyes weakly widened in dorsal view; dorsal surface somewhat shiny, but with distinct transversely striate micros culpture; without noticeable micropunctation; punctation in posterolateral area and near oc ciput rather coarse and distinctly denser than in O. medius; irons with posterior pair of punctures absent, the anterior punctures ac companied by 2 -5 additional punctures; fron tal furrows shallow. Pronotum dilated anteriorly, with maximal width ca. halfway between anterior angles and middle; posterior scratch-like impression di stinct; dorsal surface with fine and dense microsculpture composed of long transverse meshes; pronotal punctation pattern similar to O. medius, but first puncture separated from anterior margin by at least its diameter. Elytra rather long EL/PL: 0.67 -0.75; punctu res ± regularly spaced, not veiy deep, and mo derately dense, the interstices on average wi der than the diameter of the punctures; surface mat due to very dense micropunctation, which at lower magnifications may be mistaken for microreticulation; legs relatively long and slender. Abdomen more shining than forebody; punctation rather dense and fine; microsculpture near anterior margins of terga composed of shallow ± transverse and isodiametric meshes, on remain der of tergal surfaces transverse and very weak; palisade fringe at hind margin of tergum VII present, c?: unknown. 9: tergum X very slender, at and near hind margin with numerous (20 -30) apically curved, long modified setae; styli slender, their inner margin with fringe of dense long setae (Fig. 22a).

.1 5 . O th ius o p a cip en n is
Com parative notes: Owing to the mat appearance of the elytra, O. opacipennis externally somewhat resembles the almost equally large 0. ruficornis CAMERON, but is readily distinguished from that species by the darker colour, the distinctly larger eyes, the longer elytra and different microsculpture of the elytral surface (in O. ruficornis predominantly isodiametric), and by the presence of a palisade fringe at the hind margin of tergum VII. For distinction from the similar O. fibulifer and O. wrasei see below those species.
Comments: According to a footnote in the original description, the holotype is deposited in the collection of J. Clermont (C a m e r o n , 1 939b). It was looked for, but not found, in the collections of the M N H N P (B e r t i , in litt. Head rather massive, wider than long and only slightly narrower than pronotum (cf. ratios), posteriorly weakly widened; eyes relatively small, in lateral view 0.27x the length of postgenae; macropunctation rather coarse, but sparse, absent in central dorsal area; posterior pair of frontal punctures present; anterior frontal punctures accompanied by 2 -5 additional punctures; frontal furrows shallow; whole dorsal surface with subdued shine due to fine, but distinct microsculpture composed of isodiametric and short transverse meshes; antennae long, antennomeres IV -V weakly oblong, V -X ± subquadrate.

b
Pronotum with maximal width in anterior half, but nearer to middle than to anterior angles, posteriorly distinctly tapering; lateral margins in posterior half distinctly concave in dorsal view; microsculpture and shine similar to head; discal punctures somewhat asymmetric in HT (Fig. 23b); posterior median scratch-like impression distinct. Elytra short (see measurements and ratio); dorsal surface uneven, with large, but shallow protube rances; punctation large, shallow and sparse, interstices distinctly wider than punctures; mi crosculpture characteristic, mostly composed of wide ± isodiametric or scale-shaped meshes, which are several times as wide as those of pronotum; scutellum with rather weak ± transverse m i crosculpture; hind wings reduced. Abdomen with fine punctation; microsculpture very shallow, on tergum VII barely noticeable, near anterior tergal margins less clearly transverse and slightly more distinct than on posterior tergal surface; surface ± shiny; palisade fringe absent, o'; unknown. 9: posterior margin of sternum VIII convex and with fringe of dense, long and rather stout yellow setae; lateral tergal sclerites IX apically extremely acute; tergum X very long and apically almost pointed; all the posterior setae were rubbed off, but judging from the size of the insertions, the hind margin was equipped with numerous modified setae; styli very long and posteriorly distinctly tapering (Fig. 23a).
Derivatio nominis: The name (lat.) refers to the characteristic uneven surface of the elytra.
Comparative notes: From its Palaearctic congeners O. tuberipennis is distinguished by the elytral surface (flat protuberances, microsculpture) alone. In addition, it is characterized by its size, the reduced hind wings and palisade fringe on the hind margin of tergum VII, the massive and broad head, the presence of frontal punctures, the shape and punctation of the pronotum, and by the 9 secondary sexual characters.
Distribution: The species is known only from the type locality in Yunnan. As can be inferred from its external morphology it may have a restricted area of distribution. Head, pronotum and abdomen black; elytra sometimes slightly lighter; tibiae and femora dark brown to blackish, usually with the external faces somewhat paler; tarsi light brown; antennae dark brown with the basal antennomeres sometimes black. Head weakly to distinctly oblong, narrower than the pronotum (cf. ratios); lateral margins of head posteriorly subparallel or weakly dilated in dorsal view; eyes in lateral view ca. 0.38 -0.40x the length of postgenae; macropunctation rather dense, in central dorsal region absent or very sparse; posterior pair of frontal punctures usually present, but in most specimens relatively small, only in one of the paratypes absent, sometimes with additional punctures; anterior frontal punctures accompanied by 1 -4 additional punctures; frontal furrows distinctly sulcate; microsculpture rather shallow, usually predominantly isodiametric, sometimes composed of a mixture of isodiametric and transverse meshes, surface somewhat shining; antennae with antennomeres IV and V ± subquadrate and VI -X weakly to moderately transverse. Pronotum distinctly oblong (cf. ratios), maximal width approximately halfway between anterior angles and middle or nearer to anterior angles, posteriorly weakly tapering; microsculpture predominantly composed of transverse striae and meshes, sometimes partly mixed with inter spersed isodiametric meshes; pronotal punctation similar to O. opacipennis, posterior puncture slightly more distant from hind angles; posterior median scratch-like impression present. Elytra moderately short (see measu rements and ratio); microsculpture distinct, composed of± irregular and partly confluent micropunctation and microrugae, but surface not comple tely mat; punctation rather dense, distinctly more so than in O. opaci pennis, and relatively fine, intersti ces on average as wide as, partly wi der than punctures; scutellum ± flat, with shallow microsculpture compo sed of isodiametric and short trans verse meshes and with some punctu res; hind wings of reduced length, slightly longer than elytra. Abdomen with rather dense and fine punctation; microsculpture veiy shal low and mostly transverse, near ante rior tergal margins slightly more di stinct and composed of transverse and isodiametric meshes; palisade fringe at hind margin of tergum VU present, but weakly developed, o": protarsomeres I -IV strongly di lated; sternum V and VI without mo difications; sternum VII weakly con-cave and only indistinctly flattened posteriorly; sternum VIII distinctly oblong, with rather dense and short dark pubescence, with two transverse rows of 4 long setae, its hind margin ± truncate or weakly concave; sternum IX with mostly short dark pubescence, posteriorly distinctly concave and with broadly rounded hind angles (Fig. 24d); lateral tergal sclerites of segment IX slender, apically moderately acute, just reaching hind margin of tergum X (Fig. 24e); ventral process of aedeagus long and slender, apically pointed, without distinct basal carina, but with weak lateral folds; parameres apically distinctly dilated; internal sac with 6 sclerotized structures: a characteristically shaped median basal, a rather wide and basally weakly sclerotized median apical, a pair of stout long, and a pair of more slender and lateral structures; flagellum thin and with ca. 25 -30 coils (Figs 24 a -c). 9 : protarsomeres I -IV moderately dilated, sexual dimorphism distinct; sternum VIII with 2 transverse rows of 4 long setae, posteriorly convex and with fringe of yellow setae; tergum X posteriorly pointed and w ith weakly modified, not very stout and apically uncurved setae; inner margin of styli with fringe of numerous long and stout yellowish setae (Fig. 24f). Derivatio nominis: The name (lat.) refers to the shape of the basal median structure in the internal sac, which somewhat resembles a clip.
Comparative notes: From the similar O. opacipennis, O. flbulifer is distinguished by the predo minantly isodiametric microsculpture of the head, the distinctly denser punctation and the much more irregular microsculpture of the elytra, which in O. opacipennis is composed of well-defined dense micropunctation. From O. tuberipennis it differs in the less massive head, the larger eyes, the different microsculpture especially of the pronotum and the elytra, the longer elytra, the lack of elytral protuberances and the presence of a palisade fringe at the hind margin of tergum VII. For distinction from O. wrasei sp. n. and remarks on phylogenetics see comparative notes below that species (section 4.18).
Comments: The only species among those described by ZHENG (1995) that seems to be similar in external appearance is O. quadratus, but the shape of the ventral process especially in lateral view is different (cf. Fig. 2b in Zheng, 1995).
D istribution and bionomics: The species is known only from western Sichuan, where it was collected at altitudes of 2700 -ca. 3900m in May and July. Two of the three specimens collected in May were slightly teneral. o': protarsomeres I -IV moderately dilated; sternum V and VI without modifications; sternum VII weakly concave and only indistinctly flattened posteriorly; sternum VIII as in O. flbulifer; sternum IX with mostly short dark pubescen ce, of similar shape as in O. flbulifer, but posteriorly more broadly and deeply concave, and hind angles less broadly rounded (Fig. 25d); lateral tergal sclerites of segment IX as in O. flb u lifer (Fig. 25e); aedeagus of similar morphology as in O. flbulifer, but smaller, ventral process less slender and less acutely pointed apically, and parameres only mo derately dilated; internal sac with 6 sclerotized structures: a very weakly sclerotized, barely visible fibuliform median basal, a rather wide and basally weakly sclerotized median apical, a pair of basally dilated and somewhat twisted late ral, and a pair of straight lateral structures; flagellum with ca. 10 coils (Figs 25 a - terminalia similar to O. fibulifer, tergum X also of similar shape, but posteriorly apparently with fewer weakly modified -not very stout, apically not bent -setae (such a seta was discovered only in one 9, most of the setae of the ? paratypes were rubbed off) (Fig. 25f).
D erivatio nominis: The species is dedicated to David Wrase, Berlin, who collected most of the types, including the holotype.
Com parative notes and phylogenetics: Externally, O. wrasei is distinguished from other Eastern Palaearctic congeners, except O. coniceps from Taiwan, by the posteriorly strongly dilated head. Among the known species, it is most closely related to its presumed sister species O. fibulifer, with which it not only shares the similar general appearance (proportions, colour, punctation, mi crosculpture) and a similar external morphology of the aedeagus, but also the following hypo thesized synapomorphies: the characteristic microsculpture of the elytra (see diagnosis of O. fib u lifer), the shape of the hind margin of the o' sternum IX, the fibuliform median basal structure in the internal sac of the aedeagus, and the shape and chaetotaxy of the 9 tergum X. Both species, however, differ in various characters, especially in size, head shape, the microsculpture and punctation of the head, the position of the posterior dorsal puncture of the pronotum, the length of the hind wings, the development of the palisade fringe at the hind margin of tergum VII, and the primary and secondary sexual characters (see diagnosis above).
Distribution and bionomics: The species is known only from the Gongga Shan in western Sichuan, where it was collected at an altitude of 2 100m and 2750m in May and July. Three of the four type specimens taken in May were at least slightly teneral. Colour blackish brown to black, with the tarsi, the bases of first three antennomeres, the hind margins especially of the posterior abdominal segments, and sometimes the elytra lighter. Head 1.13 -1.19x longer than wide, approximately as wide as pronotum (HW/PW: 0.99 -1.03); eyes large, 0.40 -0.48x the length of postgenae in lateral view; temples behind eyes ± parallel or slightly widened; dorsal surface shining, with extremely shallow and tine microsculpture, which is visible only at higher magnifications, micropunctation absent; macropunctation in posterolateral area and near occiput moderately dense and rather coarse; irons with posterior pair of punctures present (3 specimens examined), absent (1 specimen) or reduced to one puncture (2 specimens), the anterior punctures accompanied by 1 -4 additional punctures; frontal furrows distinctly sulcate; antenna long, with antennomere III ca. 3x longer than wide and ca. 1.5x longer than II, the following antennomeres decreasing in length, but even X noticeably oblong. Pronotum distinctly dilated anteriorly, with maximal width ca. halfway between middle and anterior angles, lateral margins ± sinuate, 1.24 -1.27x longer than wide; posterior scratch-like impression minute or absent; dorsal surface very shiny, at most with traces of transverse mi crosculpture; pronotal punctation as in Fig. 26f. Elytra relatively long, EL/PL: 0.72-0.76; punctures rather dense and well-defined, interstices on average subequal in width to punctures, shining and without distinct microsculpture; scutellum with distinct transverse microsculpture and several rather large and well-defined punctures; hind wings apparently fully developed; legs slender, TiL/TaL: 1.24 -1.37. Abdomen with relatively fine, ± evenly spaced punctation; microsculpture fine and rather shallow, near anterior margins of terga somewhat more distinct and with transverse meshes; palisade fringe at hind margin of tergum VII present.
a": protarsomeres I -IV mode rately dilated, less so than in O. medius and related species; sternum V and VI centrally with subcircular areas with tomentose greyish to yellowish pubescence and rasp-like mi crosculpture; sternum VII po steriorly weakly concave and slightly flattened; hind margin of sternum VIII weakly conca ve, centrally flattened, and with long black setae; sternum IX posteriorly deeply concave, hind margin with predomi nantly dark setae (Fig. c); late ral tergal sclerites of segment IX long and acute, almost rea ching hind margin of tergum X in lateral view (Fig. 26d); hind margin of tergum X with long black setae; ventral process of aedeagus sometimes wider than shown in Fig. 26b, with weak, but long median carina; parameres apically hardly dilated; internal sac with 6 characteristic sclerotized structures: a doubly bifid median basal piece, a hook-shaped median apical piece with long proximal processes, a pair of stout and apically curved oblong lateral pieces, and a pair of straight oblong lateral pieces; flagellum relatively wide and with ca. 15 coils (Figs 26 a -b). 9: protarsomeres I -IV slightly less dilated than in o', sexual dimorphism rather indistinct; sternum VIII ± truncate to weakly convex posteriorly, hind margin with a mixture of black and brownish setae; tergum X near hind margin with few stout, apically weakly curved, modified setae (Fig. 26e); inner face of styli with fringe of numerous long and apically bent setae. Colour of body ± uniformly reddish brown. Head 1.18x longer than wide, slightly (1.03 -1.06x) wider than pronotum; eyes reduced, with <20 ommatidia, 0.15x the length of postgenae in lateral view; temples behind eyes remarkably straight, head posteriorly weakly widened in dorsal view; dorsal surface with subdued shine due to distinct isodiametric microsculpture, without noticeable micropunctation; macropunctation in posterolate ral area and near occiput moderately dense and coarse; irons with posterior pair of punctures present, the anterior punctures accompanied by 2 -3 additional punctures; frontal furrows di stinctly sulcate; antenna long, with antennomeres II 2x longer than wide, III 2.5x longer than wide, IV distinctly and V weakly oblong, and X weakly transverse. Pronotum dilated anteriorly, with maximal width near anterior angles and remarkably straight lateral margins, 1.34 -1.36x longer than wide; posterior scratch-like impression present; dorsal surface with microsculpture composed of long transverse meshes and transverse striae; pronotal punctation as in Fig. 27f. Elytra short, EL/PL: 0.57; punctures moderately dense, rather large, but shallow; interstices with shallow traces of microsculpture and micropunctation; scutellum with distinct transverse mi crosculpture, without macropunctation; hind wings reduced; metatibia in the basal third thin, in the apical two thirds distinctly and somewhat abruptly widened; TiL/TaL: 1.25 -1.29. Abdomen with distinct, ± evenly spaced punctation; microsculpture near anterior margins of terga composed of distinct isodiametric meshes, on remainder of tergal surfaces transverse and shallow; palisade fringe absent. d": protarsomeres I -IV moderately dilated, distinctly less so than in O. medius and related species and also somewhat less so than in O. punctatus\ sternum V and VI centrally with subcircular callosities without conspicuous pubescence; sternum VII posteriorly weakly concave and shallowly impressed; hind margin of sternum VIII weakly concave, centrally somewhat impressed, and with row of yellowish setae; sternum IX posteriorly broadly and moderately concave, hind margin with long yellowish setae (Fig. 27d); lateral tergal sclerites of segment IX apically rounded, not reaching hind margin of tergum X in lateral view (Fig. 27e); aedeagus with rather short, apically weakly dilated parameres; ventral process with distinct median carina and a pair of long parallel folds; internal sac with 6 sclerotized structures: a long median basal, a shorter, slightly hook shaped median apical, a pair of basally dilated and oblong lateral, and a pair of oblong, weakly sclerotized lateral pieces; internal sac also with two pairs of clusters of dark long spines; flagellum thin and with ca. 25 coils (Figs 27 a -c). 9: unknown.
Com parative notes and phylogenetics: Among the Taiwanese Othius species, O. shibatai is characterized by the reduced eyes, the light colour, the slender pronotum (which is not wider than the head), the veiy short elytra, the absence of conspicuous pubescence on the o' sterna V and VI, and the presence of clusters of dark long spines in the internal sac, apomorphic characters which this species only shares with O. yushanus, its adelphotaxon, and which characterize both species as a monophyletic group. For distinction from O. yushanus see the following diagnosis.
Distribution and bionomics: The species has become known only from the type locality in central Taiwan (Map 1, p. 85), where it was collected under stones at an altitude of 2200 -2300m (Ito, pers. comm.; additional material examined) in May and M y. The light body colour, the reduced eyes, the slender pronotum and the reduced length of the elytra are presumably adaptations to a subterranean habitat. Externally very similar to O. shibatai, but on the whole smaller (see measurements); colour of body ± uniformly reddish brown. Head 1.12 -1.16x longer than wide, about as wide as pronotum (HW/PW: 0.97 -1.02); eyes small, but larger than in O. shibatai, with ca. 25 -30 ommatidia, 0.20 -0.24x the length of postgenae in lateral view; shape of head and punctation similar to O. shibatai, but posterior pair of frontal punctures smaller; microsculpture less distinctly isodiametric, partly transverse; antennae clearly shorter than in O. shibatai, with antennomeres II ca. 1.5x longer than wide, III 2x longer than wide, IV subquadrate, V transverse and X strongly transverse. Pronotum of similar shape (PL/PW: 1.27 -1.36), microsculpture and punctation pattern as in O. shibatai, but with weakly concave lateral margins. Elytra short, EL/PL: 0.56 -0.61; punctation slightly denser than in 0. shibatai, interstices with shallow, ± irregular microsculpture and micropunctation particularly near scutellum and near suture; scutellum with distinct transverse and/or isodiametric microsculpture, and with few small punctures; hind wings reduced; metatibia shaped as in O. shibatai, but legs on the whole shorter, TiL/TaL: 1.30 -1.46. Abdomen with punctation and microsculpture similar to O. shibatai, but microsculpture near anterior margins of terga predominantly composed of ± transverse meshes; palisade fringe at hind margin of tergum VII absent. o": dilatation of protarsomeres as in O. shibatai', callosities of sternum V and VI very weak, reduced to subcircular areas with dense microsculpture and therefore less shine than the surroun ding sternal area; sternum VII and VIII as in O. shibatai', sternum IX similar to 0. shibatai (Fig.  28d); lateral tergal sclerites of segment IX apically more acute than in O. shibatai (Fig. 28e); aedeagus on the whole as in O. shibatai, but smaller, apex of ventral process in lateral view broader and in ventral view of different shape, and parameres relatively longer; internal sac with differently shaped clusters of long dark spines, apically more acute and basally more dilated median apical piece, and with basally more dilated dark lateral pieces; flagellum as in O. shibatai (Figs 28 a -c). 9: protarsomeres I -IV weakly dilated, sexual dimorphism pronounced; sternum V and VI centrally instead of callosities with subcircular areas with denser, not distinctly transverse mi crosculpture and therefore less shine; hind margin of sternum VIII convex and with marginal row of yellow setae; lateral tergal sclerites IX each with one terminal seta; tergum X with unmodified, long yellow setae posteriorly; inner face of styli with fringe of long, dense and apically weakly curved yellowish setae (Fig. 28f).  Colour of body brown, with the elytra, the hind margins of abdominal segments VII and VIII, parts of segments IX and X, and th e legs yellowish brown. Head weakly oblong, slightly narrower than pronotum (cf. ratios); eyes ca. 0.33 -0.38x the length of postgenae in lateral view; temples behind eyes straight, head posteriorly subparallel or very weakly widened in dorsal view; dorsal surface with subdued shine due to distinct, but fine mi crosculpture predominantly composed of transverse meshes of variable length, which are on average shorter in the anterior th an in the posterior region of head; sparse micropunctation visible only in posterior and lateral area; niacropunctation in posterolateral area and near occiput modera tely dense and coarse; irons w ith posterior pair of punctures present, but much smaller than the anterior punctures, which are accompanied by 1 -3 additional punctures; additional punctures may also be present near posterior pair of frontal punctures; frontal furrows distinctly sulcate; antennae moderately long, moderately incrassate apically; antennomeres II ca. 1.5x longer than wide, HI 2 -2.3x longer than wide, IV and V subquadrate, and VII -X transverse. Pronotum with maximal width in anterior half, but nearer to middle than to anterior angles, on the whole shorter and with more convex surface than in O. taiwanus; posterior scratch-like impression present; dorsal surface with microsculpture composed of long transverse meshes and transverse striae; posterior discal puncture clearly distant from hind angles, much more distant from lateral margins than in O. taiwanus (cf. Fig. 30f). Elytra short (cf. ratio); punctures dense, rather large and well-defined; interstices shining, without distinct microsculpture; scutellum shining, microsculpture very shallow and composed of large meshes, with macropunctation; hind wings reduced. Abdomen with distinct, ± evenly spaced punctation, which is finer on posterior than on anterior terga; microsculpture very weak, transverse, near anterior margins of terga only slightly more distinct and not reticulate; surface with weakly iridescent shine; palisade fringe at hind margin of tergum VII absent.
o': protarsomeres I -IV strongly dilated; central subcircular callosities on sternum V and VI large, larger than in O. yushanus and approximately of similar relative size as in O. medius, especially anteriorly more distinctly delimited from surrounding sternal area than in those two species, and with less dense, blackish pubescence; sternum VII posteriorly weakly concave and centrally weakly flattened; hind margin of sternum VIII ± truncate, centrally somewhat flattened, without row of marginal setae; sternum IX with mostly blackish pubescence, posteriorly broadly and deeply concave (Fig. 29d); lateral tergal sclerites of segment IX apically acute, just reaching hind margin of tergum X in lateral view (Fig. 29e); aedeagus with apically weakly dilated parameres; ventral process with distinct basal carina and pair of long apical folds; internal sac with 5 sclerotized structures: a long median apical, a pair of long curved lateral, and a pair of long straight, basally somewhat dilated lateral pieces; near apex of median apical structure with membranous structure of characteristic shape; basally with a pair of small membranous structures equipped with ± sclerotized short spines; flagellum thin and with ca. 20 coils (Figs 29 a -c). $: sternum VIII posteriorly convex and with fringe of dense and rather long yellow setae; tergum X with very narrow basis, near hind margin with several long, apically weakly curved modified setae; inner margin of styli with fringe of dense and relatively long setae (Fig. 29f). Colour darker, dark brown to blackish, with the legs and part of the basal two antennomeres light brown or reddish; body on the whole smaller (see measurements). Head with less distinct, predominantly transverse microsculpture; posterior pair of frontal punctu res larger. Pronotum with discal punctures as in O. parvipennis (Fig. 3 Of). Abdomen with slightly denser punctation, on the whole less shiny, but more intensely iridescent, o': protarsomeres I -IV somewhat less dilated than in O. parvipennis; sternum V and VI centrally with subcircular callosities with tomentose brownish pubescence, the callosities distinctly smaller (less than half the area) than in O. parvipennis; sternum VII posteriorly weakly concave, not distinctly flattened; hind margin of sternum VIII weakly convex, laterally with fewer dark setae; sternum IX similar to O. parvipennis (Fig. 30d); lateral tergal sclerites IX as in Fig. 30e; aedeagus smaller; ventral process more slender, pair of apical folds shorter and less distinct; internal sac with similar arrangement of sclerotized and membranous structures, but lateral pieces thinner and less dilated basally (probably due to the fact that the only d" available was teneral, the pair of dark basal membranous structures were not distinctly visible); flagellum moderately thin (thinner than in O. parvipennis) and with only ca. 12 coils (Figs 30 a -c). 9: unknown.
Derivatio nominis: This species is dedicated to Ales Smetana, to whom I am indebted especially for his remarkable material from Taiwan, which contained type material of five Taiwanese species described here for the first time.
Com parative notes and phylogenetics: For separation from the preceding Taiwanese   Externally highly similar to O. parvipennis, but distinguished from this species as follows: Colour on the whole slightly darker, ± uniformly dark brown, legs and antennal base reddish brown; body smaller than in O. parvipennis (see measurements). Head with microsculpture predominantly composed of short transverse and isodiametric meshes; posterior pair of frontal punctures relatively large; antennae distinctly shorter than in O. parvipen nis, antennomeres V -X (PT) or VI -X (HT) clearly transverse. Pronotum with microsculpture composed of transverse meshes and transverse striae; position of posterior discal puncture as in O. parvipennis and O. smetanar, position of anterior punctures apparently variable (Fig. 3 lg). Elytra of similar shape as in the related species, but punctation finer; scutellum with more distinct microsculpture composed of transverse meshes of variable length and with several punctures. Abdomen with similar microsculpture and shine as in O. parvipennis, but punctation somewhat finer. o": protarsomeres I -IV slightly less dilated than in O. parvipennis-, sternum V and VI centrally with large circular callosities with tomentose, dense brownish pubescence, the callosities of similar size and almost equally delimited as in O. parvipennis-, sternum VII posteriorly weakly concave and weakly flattened; hind margin of sternum VIII ± weakly convex, without marginal pubescen ce, pubescence on the whole similar to O. smetanar, sternum IX of similar shape and pubescence as in O. parvipennis, but much less strongly concave posteriorly (Fig. 3 Id); lateral tergal sclerites of segment IX apically as in O. parvipennis, but slightly longer, reaching beyond tergum X (Fig.  3 le); aedeagus with ventral process slender, apically less acute than in O. parvipennis-, apical folds indistinct, basal carina weak, not sclerotized; parameres apically weakly dilated; internal sac with 6 sclerotized structures: a median basal and an approximately equally long median apical, and two pairs of almost equally long and ± straight lateral pieces; near apex of median apical structure with membranous structure, which is less dark and distinct than in O. parvipennis-, pair of dark basal membranous structures apparently absent; flagellum thin and with ca. 15 coils (Figs 31 a -c). $: protarsomeres I -IV dilated, but distinctly less so than in cf; sternum VIII convex, posteriorly with marginal row of yellow setae; lateral tergal sclerites IX apically rounded; tergum X poster iorly with several rather short and apically weakly curved modified setae; inner margins of styli with fringe of long, rather stout setae, which are anteriorly shorter and less curved than in O. taiwanus (Fig. 3 (Fig. 32d); lateral tergal sclerites IX relatively short, not reaching hind margin of tergum X (Fig. 32e); aedeagus o f similar morphology as in O. smetanai, but ventral process broader in ventral view, and in lateral view apically narrower; apical folds absent, basal carina distinctly sclerotized; parameres apically weakly dilated; internal sac with sclerotized structures as in O. smetanai, but apical median piece more strongly curved and lateral pieces stouter; near apex of median apical structure with membranous structure, its shape slightly different from that in 0. smetanai; flagellum thin and with ca. 15 coils (Figs 32 a -c). $: protarsomeres I -IV dilated, but distinctly less so than in a"; sternum VIII convex, posteriorly with marginal row of rather dense yellow setae; lateral tergal sclerites IX apically acute, with two terminal setae, but without apical process; tergum X posteriorly with several rather short and apically weakly curved modified setae (Fig. 32f); styli with fringe of dense, stout, not or only weakly curved setae, posteriorly decreasing in length. hamatus, but apical median structure apically shorter and basally less distinctly constricted; near apex of median apical structure with membranous structu re; basal membranous structure with two small areas with distinctly sclerotized spines; flagellum long and thin, and with ca. 25 coils (Figs 33 a -c). $: unknown.
Derivatio nominis: The name (lat.: acute) refers to the pointed apex of the median lobe.
Com parative notes and phylogenetics: The species is distinguished from O. hamatus, which has a similar morphology of the aedeagus, by larger size, a broader head and pronotum, larger eyes, more massive and longer antennae, lighter colour, the larger tomentose areas on the d" sterna V and VI, an apically more strongly pointed median lobe and the different shape of the apical median structure in the internal sac. From O. parvipennis it is separated by smaller size, shorter antennae, different microsculpture of the head, the absence of a pair of median folds on the ventral process and the apically different shape of the aedeagus, and by the different shapes of the internal structures, especially the apical median structure. In O. smetanai, the antennae are distinctly longer, the colour is darker, the microsculpture of head and pronotum is transverse, and the o' primary and secondary sexual characters (shape and internal structures of the aedeagus, modifica tions of sterna V and VI) are different. In O. aequabilis the basal median structure in the internal sac is absent. For distinction from O. reticulatus see below that species. The similar morphology of the aedeagus (shape of ventral process, reduction of pair of median folds, shape of internal structures) suggests that O. acutus may be most closely allied to O. hamatus. A sister species relationship is also supported by the vicinity of the localities, from which the two species are known. The presence of distinct micropunctation on the head, the punctation and sculpture of the elytra (if they are constant characters), and the broad shape of the apical median structure in the internal sac of the aedeagus appear to be autapomorphic.  (Fig. 34d); lateral tergal sclerites IX relatively short, not reaching hind margin of tergum X (Fig. 34e); aedeagus of similar morphology as in O. parvipennis, also with distinct pair of long median folds on ventral process, but in ventral view more slender; basal carina distinctly sclerotized; parameres apically weakly dilated; internal sac with sclerotized structures similar to O. parvipennis, but apical median piece shorter and in ventral view wider, and long lateral structures more slender; near apex of median apical structure with membranous structure; basal membranous structure with two small areas with distinctly sclerotized spines; flagellum similar to O. parvipennis, with ca. 25 coils (Figs 34 a -c). 9: terminalia similar to O. parvipennis, but tergum X posteriorly distinctly membranous and with shorter modified setae (Fig. 34f).
Derivatio nominis: The name (lat.) refers to the distinct microreticulation of the head and pronotum, an external character distinguishing O. reticulatus from its most closely related con geners.
Com parative notes and phylogenetics: From all the closely related species allied to O. parvipen nis, O. reticulatus is distinguished by the distinct microreticulation of head and pronotum. From O. hamatus and O. aequabilis it is also separated by larger body size, by the shape of the ventral process of the aedeagus, the presence of a pair of distinct median folds on the ventral process, and by the internal structures of the aedeagus. From O. smetanai, it is additionally distinguished by the lighter colour, by the shorter antennae, by the the larger and more clearly delimited tomentose areas on the a" sterna V and VI, by the more distinct pair of median folds of the ventral process of the median lobe, and by the shape of the internal structures of the aedeagus. From the similar O. parvipennis it differs also in the shorter antennae, the more slender ventral process of the aedeagus and the shape of the internal structures, especially the much shorter apical median structure. In O. acutus, the ventral process of the aedeagus is much broader and more strongly pointed apically, and it lacks a pair of distinct median folds. O. reticulatus shares several conspicuous characters with O. parvipennis, i. e. the relatively broad head and pronotum, the similar modifications of the d" sterna V and VI, the presence of a pair of distinct median folds on the ventral process of the aedeagus as well as a long flagellum in the internal sac. Since these characters, however, are likely to represent the plesiomorphic states, it is uncertain whether both species are in fact adelphotaxa.  . 35c); lateral tergal sclerites of segment IX long, slightly reaching beyond tergum X, and apically with short, stout and weakly curved process (Fig. 35d); tergum X posteriorly with broad membranous appendage; aedeagus with ventral process of characteristic shape, in ventral view apically not acute, but broadly rounded; apical folds indistinct, basal carina weak, not sclerotized; surface of ventral process with a mixture of longitudinal and scale-like microsculpture and punctures; parameres apically distinctly dilated and with broad bases; internal sac with 6 sclerotized structures: a small median basal (absent in O. parvipennis and O. smetanai), a very large median apical, a pair of short and basally dilated lateral, and a pair of longer thin lateral pieces; flagellum slightly wider than in the related species and with ca. 15 -20 coils, inner coils widened and slightly darker (Figs 35 a -b). $: protarsomeres I -IV dilated, but somewhat less so than in d", sexual dimorphism moderate; sternum VIII convex, posteriorly with marginal row of yellow setae (similar to O. aequabilis)', lateral tergal sclerites IX with short and acute apical process (much shorter than in O. medius)\ tergum X posteriorly with several rather short, apically curved modified setae; inner margins of styli with fringe of dense, anteriorly long and stout, and posteriorly thin and rather short setae (Fig. 35e).
Derivatio nominis: The name (lat.: cut off, broken off, shortened) refers to the characteristic shape of the ventral process, which lacks the acute apex of the related species.
Com parative notes and phylogenetics: see below the following species. more distinctly delimited from surrounding sternal area, ± impressed, strongly microsculptured, that of sternum V either similar, or smaller, more weakly delimited, with relatively sparse pubescence and with weaker microsculpture; sternum VII posteriorly weakly concave, flattened and with ± dense dark pubescence directed postero-mediad; hind margin of sternum VIII ± truncate; sternum IX of similar shape and pubescence as in O. praecisus, also anteriorly weakly bifid, but posteriorly less deeply concave (Fig. 36c); lateral tergal sclerites of segment IX long, slightly reaching beyond tergum X, and apically with long, spine-like process (Fig. 36d); tergum X posteriorly w ith broad membranous appendage (cf. Fig. 37e); aedeagus much larger than in O. praecisus; ventral process of somewhat variable width, apically pointed, without apical folds, basal carina indistinct, microsculpture as in O. praecisus-, parameres apically di stinctly dilated; internal sac with arrangement of sclerotized structures similar to O. praecisus: a smaller median basal, a very large and long median apical, a pair of ± transverse, short, basally dilated and apically acute lateral, and a pair of longer thin lateral structures; flagellum with ca. 30 coils, inner 2 coils widened and darkened (Figs 36 a -b). 9: protarsomeres distinctly dilated, but less so than in d", sexual dimorphism moderate; sternum VIII posteriorly convex and with numerous yellowish marginal setae; lateral tergal sclerites IX apically with acute process (similar to O. medius)-, hind margin of tergum X broadly convex and with several relatively short and apically weakly bent modified setae; inner margin of styli with fringe of numerous long setae, which are very stout anteriorly and thin posteriorly (Fig. 36e).  (Fig. 37c); lateral tergal sclerites of segment IX long, slightly reaching beyond tergum X, on the whole similar to O. collapsus, but apically somewhat truncate and with much shorter process (Fig. 37d); tergum X posteriorly with broad membranous appendage (Fig. 37e); aedeagus of similar mor phology as in O. collapsus, but long apical median structure more abruptly narrowed apically and less acute in ventral view, basal median structure slightly shorter, and short lateral structures of different shape, more distinctly sclerotized and in less transverse position; ventral process of somewhat variable width; flagellum with ca. 30 coils, inner 2 coils widened and darkened (Figs 37 a -b). $: unknown. collapsus, but anteriorly more strongly bifid and hind angles less acute (Fig. 38c); lateral tergal sclerites of segment IX, slightly reaching beyond tergum X, broad and apically without acute process (Fig. 38d); tergum X posteriorly with broad membranous appendage; aedeagus with ventral process of similar shape as in O. collapsus, without apical folds, but with weak lateral elevations; basal carina indistinct; microsculpture as in O. praecisus-, parameres apically weakly dilated; internal sac with 5 sclerotized structures: a very large and long median apical, a pair of short, basally dilated and apically acute lateral, and a pair of longer thin lateral pieces; basal median structure absent; flagellum with ca. 30 coils, inner 2 -3 coils widened and darker (Figs 38 a-b). 9: unknown.
Derivatio nominis: The name (lat.) refers to the remarkably coniform shape of the head. but as yet it has not been recorded from outside Taiwan. The remaining 12 species are not only certain to be island endemics, as can be inferred from their external morphology (i. e. the reduc tion of elytra, hind wings, eye size, and palisade fringe), they apparently even have a very re stricted distribution within the island. All of them are known only from a single mountain or mountain range; only O. reticulatus was collected in two localities, but they are only some 2 0 km apart (Map 1). As can be concluded from the altitudes indicated on the labels, these local endemics inhabit the cool and the cold temperate coniferous forest zones, and the subalpine coniferous forest zone sensu Sm e t a n a (1 9 9 5 ). Of the total of 36 specimens of brachypterous Othius from Taiwan examined, 34 belonged to 12 species, 9 of them described here for the first time, and several species are represented only by single specimens. The remaining two females could not be attributed to any of the species, since no corresponding males were available, and may well represent two undescribed taxa. Therefore, it seems safe to assume that the inventory of Taiwanese Othius species is far from complete and that, with respect to Othius, Taiwan is among the most diverse areas of the Palaearctic region, probably more so than the Atlantic islands with 13 known endemics (A s s in g , 1998b), the Cauca sus with 12 endemic taxa (A s s in g , 1997a; ASSING & So l o d o v n ik o v , 1998), and exceeded only by the Himalayas with 24 known endemics (A s s in g , 1998a). The Othius fauna of the mountain ranges of Sichuan and Yunnan appears to be very diverse, too; these areas have only been poorly studied and are likely to be inhabited by a far greater number of species than are presently known.
The Taiwanese Othius fauna can be subdivided into five species groups. Two of them are represen ted by a single species each, O. punctatus and O. taiwanus. The former is widespread and also present in mainland China, so that -by geological standards -a recent colonization can be assumed. Judging from external characters (body shape and proportions, punctation, microsculptu re, male secondary sexual characters etc.) and the shape and internal structures of the aedeagus, the species seems to be most closely related to O. rufipennis. Like O. punctatus, O. taiwanus does not show the morphological adaptions -i. e. reductions of characters associated with flight activity -usually found in Othius species with a restricted distribution, especially island endemics (see A s s in g , 1998b), which also suggests a comparatively recent colonization. Among the known Eastern Palaearctic congeners, there is no obvious adelpho taxon. On the one hand, there is some similarity with O. fibulifer from Sichuan in the morphology of the aedeagus and the shape of the internal structures, especially the median structures, but on the other hand the possibility that these character states represent homoplasies cannot be ruled out, particularly since both species differ in many external characters and O. fib u lifer is apparently more closely related to O. wrasei. The three remaining species groups are monophyla whose respective ancestors seem to have colonized Taiwan in earlier geological periods. The monophylum O. shibatai + O. yushanus is constituted by several distinct synapomorphies, the reduced eyes, the light colour, the slender pronotum, the very short elytra, the absence of conspicuous pubescence on the 6" sterna V and VI, and the presence of clusters of dark long spines in the internal sac. There is no obvious adelphota xon among the Eastern Palaearctic congeners. The only apomorphy suggesting a closer relations hip to the species allied to O. parvipennis (SHIBATA) is the position of the posterior dorsal punctu re of the pronotum, a character state, however, which may well be homoplastic and which also occurs in other Eastern Palaearctic species groups. Together with O. smetanai sp. n., O. ham atus sp. n., O. reticulatus, O. acutus and O. aequabilis sp. n., O. parvipennis forms a monophyletic group supported by the following hypothesized synapomorphies: the pronotal punctation pattern with the posterior punctures clearly distant from the posterior angle, the presence o f an apical membranous structure of characteristic shape in the internal sac, the ± reduced number of coils of the flagellum, and the very narrow basis of the 9 tergum X. Within this group, a closer relationship of O. parvipennis, O. reticulatus, O. sm etanai, O. acutus and O. hamatus can be inferred from the apomorphic absence of the median basal structure and the presence of a basal pair of small membranous structures covered with ± sclerotized short spines in the internal sac. The phylogenetic affinities among the species of this group, however, are difficult to assess, partly because of the low degree of character divergence in these apparently extremely closely related species, a situation also encountered in the Othius intermedius subgroup of the Canary Islands (A s s in g , 1998b). In addition, most of the hypothesized apomorphies are either autapomorphies of single species or in conflict with other synapomorphies. Finally, the species treated here are very likely to represent only a fraction of the actual number of species present in the mountain ranges of Taiwan. Close affinities between O. acutus and O. hamatus are suggested by the strongly pointed ventral process of the aedeagus, the similar shape of the internal structures and especially by the apomorphic reduction of the pair of median folds of the aedeagus. The sister taxon of the group of species allied to O. parvipennis seems to be O. rosti B e r n h a u e r from the northeast of the Eastern Palaearctic region (Japan, Russian Far East). Hypothesized synapomorphies supporting the sister group relationship are the presence of an apical membranous structure in the internal sac, the absence of a basal median structure, the reduced number of flagellar coils, and the remarkably narrow basis of the ¥ tergum X.
Of the numerous characters shared by O. praecisus, O. collapsus, O. contumax and O. coniceps, at least the broad membranous appendage of tergum X, the anteriorly weakly bifid sternum IX, the remarkably large median apical structure, the characteristic shape of the short lateral structures of the internal sac, and the somewhat darkened and widened inner coils of the flagellum can be considered synapomorphies supporting the monophyly of the four taxa. Like the species related to O. parvipennis, they apparently have a very limited area of distribution and are confined to higher altitude habitats. O. collapsus, O. contumax, and O. coniceps share a long flagellum and an apically pointed ventral process; in O. praecisus, O. collapsus, and O. contumax, on the other hand, the tergal sclerites IX have a distinct apical process, and the internal sac contains a basal median structure. These character states, however, are not very conclusive, as they are very likely to be plesiomorphic. O. collapsus and O. contumax are without doubt adelphotaxa, not only because of their similar outward appearance and their probably synapomorphic large size, but also because of the very similar morphology of the internal structures of the aedeagus, especially of the basal median structure. In addition, a close relationship is supported by the vicinity of the localities where they were collected. It is somewhat more difficult to decide whether the monophylum O. collapsus + O. contumax is more closely allied to O. praecisus or to O. coniceps. Although biogeographic evidence would favour the latter, the former hypothesis (0. coniceps + (O. pra eci sus + (O. collapsus + O. contumax))) appears to be more likely, as it is supported by two probable synapomorphies, i. e. the anteriorly more weakly bifid < ? sternum IX, and the apically distinctly dilated parameres. Both O. praecisus and O. coniceps are characterized by several obvious autapomorphies: the former by the rounded apex of the ventral process of the aedeagus and the shorter and wider flagellum, the latter by the posteriorly distinctly dilated head, the reduced apical processes of the lateral tergal sclerites IX, and particularly by the absence of a basal median structure in the internal sac. O. collapsus and allied species are apparently the sister group of O. m edius + O. bordonii. The monophyly of these two species groups is supported by the presence of an apical process on the lateral tergal sclerites IX, a character which is absent in all other known species of Othius and therefore very likely to represent a synapomorphy. Based on the phylogenetic observations and conclusions, at least four, probably five successful colonization events can be assumed for Taiwanese Othius. At least one of them, the ancestor of O. parvipennis and allied species, is likely to have originated from the northeast. The probably most recent addition to the Othius fauna of Taiwan, O. punctatus, almost certainly came from mainland China, since this species does not seem to occur in Japan and the islands linking Japan and Taiwan. Although there is some doubt as to the origin of the remaining three species groups, either because no obvious sister group could be identified {O. shibatai + O. yushanus, O. taiwanus) or because the hypothesized adelphotaxon is widespread (O. collapsus and allied species), a coloniza tion from mainland China would seem more likely, not only due to the relatively low distance, but also because of the greater number of species present and the probably larger populations.
6. Key to the species o f Othius o f the Eastern Palaearctic region east o f the Himalayas The species from southern China and Taiwan are here considered Palaearctic elements and included below (see section 1). Note that three of the species described by ZHENG (1995) -O. quadratus, O. goui, and O. rudis -had to be neglected and are not accounted for in the key, because the types were not available and the original descriptions are inadequate for a safe identification of the species (see remarks in the introduction to section 4). For explanations of abbreviations and measurements see section 3.

5.
Posterior pronotal punctures nearer to posterior angles (Fig. 351); pronotum relatively broader (PL/PW: <1.25); lateral tergal sclerites IX with apical process, which may be reduced, in which case the sclerites are apically ± truncate. o' : ventral process of aedeagus without distinct pair of median folds and without distinct basal carina, internal sac in most species with six sclerotized structures of characteristic shape; apical median structure strongly enlarged; On average larger species; antennae slightly longer, antennomere IV weakly oblong; legs longer (TiL: >1.6; TaL: >1.2). cf: lateral tergal sclerites IX with long and very acute apical process (Fig. 36d); internal structures of aedeagus of different shape: apical median structure apically gradually tape ring, basal median structure slightly longer, and short lateral structures less distinctly sclerotized and in transverse position (Figs 36 a - cf: lateral tergal sclerites IX with short apical process (Fig. 37d); internal structures of aedeagus: apical median structure apically ± abruptly narrowed, basal median structure slightly shorter, and short lateral structures more sclerotized and in less transverse position (Figs 37 a - Colour darker, dark brown to blackish, with the legs and part of the basal two antennomeres light brown or reddish; antennae very long; microsculpture of head and pronotum very shallow and predominantly transverse. o': protarsomeres I -IV moderately dilated; sternum V and VI centrally with small areas with condensed pubescence; aedeagus with slender ventral process, pair of median folds short and not very pronounced; flagellum thin and with only ca. 12 coils (Figs 30 a -

11.
Microsculpture of head and pronotum pronounced and distinctly isodiametric. <?: ventral process of aedeagus slender and with distinct median folds (Fig 34 a - (Shibata) 13. Eyes smaller than in other species allied to O. parvipennis; elytra relatively shorter (EL/ PL: <0.56). sternum V and VI centrally with small suboval callosities; lateral tergal sclerites IX relatively short, not reaching hind margin of tergum X; sternum IX posteriorly strongly concave (Fig. 3 2d); ventral process of aedeagus in ventral view with curved lateral margins, pointed apex distinctly sclerotized basal carina; internal sac with 5 sclerotized structures and near apex of median apical structure with distinct membranous structure (Figs 32 a - o': posterior margin of sternum VII weakly concave, with shallow median impression and less dense pubescence; sternum VIII with convex hind margin; lateral tergal sclerites of segment IX longer, more slender and apically more acute (Fig. lOf); sternum IX more slender and with different pubescence (Fig. lOe); aedeagus smaller and more slender, ventral process longer, and basally more constricted; sclerotized internal structures of characteristic shape (Figs 10 a -d). 9: tergum X as in Fig Highly variable species; colour of legs reddish brown to brown; body on average larger; head on average relatively longer, eyes usually relatively smaller, d": sternum VII with broad but shallow median impression increasing in depth and width posteriorly and with weakly concave hind margin, the impression with dense and long, almost tomentose pubescence; hind margin of sternum VIII moderately concave and laterally with long dark setae (Fig. 7r); lateral tergal sclerites of segment IX short an broad in lateral view, apically obtusely rounded (Figs 7 n -p); sternum IX broader and with pubescence as in Fig. 7h; aedeagus larger and wider, ventral process shorter; sclero tized internal structures of different shape (Figs 7 a -f). 9: terminaba as in Fig. 7t