Revision of the subgenus Tinotus Sharp, stat. n., of the parasitoid rove-beetle genus Aleochara Gravenhorst (Coleoptera, Staphylinidae, Aleocharinae) from Japan, Taiwan, and the Russian Far East

Abstract The subgenus Tinotus Sharp, 1833, stat. n., of the genus Aleochara Gravenhorst, 1802 (Aleocharini: Aleocharina) from Japan, Taiwan, and the Russian Far East is revised. Tinotus is a new record from the latter two regions. Three species are recognized: Aleochara (Tinotus) morion Gravenhorst, 1802, comb. n. [Japan (new record), the Russian Far East (new record)], Aleochara (Tinotus) eoa nom. n. [replacement name for Tinotus japonicus Cameron, 1933; Japan, Taiwan (new record)], and Aleochara (Tinotus) takashii sp. n. (central Honshû, Japan). The systematic position of Tinotus is discussed. All species are (re-)described, keyed, and figured. A world checklist of Tinotus species, comprising 40 valid species, is provided in an appendix. Additional taxonomic changes are proposed, including a new synonymy, a revalidation, 13 new replacement names, and 27 new combinations.

On the other hand, the genus Tinotus Sharp, 1883 has recently been recognized as phylogenetically close to, or a possible member of, Aleochara (Maus et al. 2001;Hanley 2002;Osswald et al. 2013).Its taxonomic placement, including its tribal assignment, has been controversial for a long time, mainly due to its tarsal formula (4-5-5; see Hanley 2002).According to Maus et al. (2001), Tinotus is a taxon that should be considered within the genus Aleochara.The distributional range of Tinotus is quite wide, as it has been recorded from every zoogeographic region, except the Australian Region, with approximately 40 species (Hanley 2002;Klimaszewski et al. 2002).
Larvae of Aleochara and Tinotus act as ectoparasitoids on cyclorrhaphous Diptera, and the adults prey upon dipteran eggs and larvae (e.g., Klimaszewski 1984;Maus et al. 1998).Thus, they have been considered potential candidates for classic biological control of commercial crops against pest flies in Europe and North America (e.g., White and Legner 1966;Fournet et al. 2000).
Taxonomic knowledge of Tinotus in East Asia is still incomplete.In Japan, Taiwan, and the Russian Far East, just one species, Tinotus japonicus Cameron, 1933, has been originally described from Japan.In this study, we (re-)describe three Tinotus species distributed in these regions.We discuss the systematic position of Tinotus.We also provide a complete Tinotus species list, reflecting recent species additions, synonyms, and corrections.

Material and methods
We used the technical procedures and methods used by Maruyama (2006) and Yamamoto and Maruyama (2012).The terminology used for this study generally follows that of Hanley (2002), Klimaszewski et al. (2002), and Yamamoto andMaruyama (2012, 2013).For chaetotaxy of the mouthparts we followed Sawada (1972Sawada ( , 1987)).We followed Welch (1997) for genital terminology, especially that of females.In the descriptions, the number of macrosetae on tergite VIII and sternite VIII refers to one side of the body.Furthermore, minute setae were omitted or depicted only for one side of the body.
Abbreviations for measurements: BL, length of the body from clypeus to apex of the abdomen; EW, maximum width both elytra combined; HL, maximum length of the head; HW, maximum width of the head; PL, maximum length of the pronotum; PW, maximum width of the pronotum along midline.
Remarks.See other characters mentioned in detail by Hanley (2002) and Klimaszewski et al. (2002).
Systematic position.Sharp's (1833) original description of Tinotus placed this taxon in the group Myrmedoniina (= Lomechusini) due to its 4-5-5 tarsal formula.Since, Tinotus has also been placed in Hoplandriini (e.g., Seevers 1978) or Aleocharini (e.g., Lohse 1974), mainly based on the presence of a pseudosegment on the maxillary and labial palpi (see Hanley 2002 for a historical review).
In contrast to these ambiguities, recent studies have refuted all tribal placements other than Aleocharini.Hanley (2002) recognized Tinotus within Aleocharini, suggesting a close relationship with the genus Aleochara, based on the seven shared morphological characteristics of the genus, e.g., bifid to crescent-shaped apex of the ligula.According to the extensive molecular study of Aleochara by Maus et al. (2001), Tinotus was fully resolved within the "bilineata clade" of Aleochara.Similarly, Osswald et al. (2013), who used significantly fewer species (only four species of Aleochara, one of which is Tinotus) but analyzed them with much more molecular markers (4599 bp), also supported the assignment of Tinotus to Aleochara.
Diagnosis (see Klimaszewski et al. 2002).This species can be distinguished from most members of the subgenus Tinotus by the following combination of characters: body entirely black, rarely light brown (Fig. 1); median lobe of aedeagus of male with a basal protuberance in lateral view (Fig. 16: arrow); spermatheca with a simple and oblong spermathecal head, equally serrate inner walls inside spermathecal head, and with four coils at base (Fig. 19).Aleochara morion is extremely similar to A. (T.) rougemonti (Pace, 1993), comb.n. from China, including male-female genital structures, but the former species is discriminated from the latter by having longer sclerites inside a median lobe of the male aedeagus (Pace 1993: Fig. 160), and by having four coils of the female spermatheca (A.rougemonti with two coils; Pace 1993: fig.162).
Body (Fig. 1): fusiform, compact, and robust; dorsal surface moderately glossy and pubescent, covered with large micro-reticulation.Color (Fig. 1): usually uniformly black to blackish brown; antennomeres I-III dark brown, but segments IV to X darker with numerous minute whitish setae; mouthparts and legs yellowish brown to brown; pubescence yellowish brown to brown.
Head (Fig. 1): subquadrate, as long as width (HW/HL = 1.03, n = 30), widest at just behind eyes; setae on vertex rather dense, directed anteriomedially.Eyes: small, occupying approximately one third of head length, very slightly protruding laterally.Labial palpus (Fig. 5): setula a well-developed, situated near apical margin of labial palpomere I, while that of b and c strongly reduced.Maxilla (Fig. 6): lacinia with a distal comb consisting of dense-thin spines; galea rather short, as long as maxillary palpomere II; maxillary palpomere IV narrow and short, less than half length of that of III.Mentum (Fig. 8): anterior margin broadly emarginate.
Abdomen (Fig. 1): first three visible tergites rather shallowly impressed transversely at base; dorsal and ventral surface covered with setae densely.
Female.Tergite VIII (Fig. 13): basal suture fully developed; posterior margin very weakly serrate, insignificantly emarginate medially; dorsal surface covered with setae rather sparsely, with five macrosetae.Sternite VIII (Fig. 15): basal suture fully developed; posterior margin rounded; ventral surface covered with setae densely, with approximately seven macrosetae.Spermatheca (Fig. 19): L-shaped; spermathecal head and neck fused together, forming a narrowly elongate capsule; attachment of spermathecal duct inconspicuous; basal part of spermathecal stem moderate in size, slightly longer than spermathecal neck, with four coils attached at base; each part of spermatheca entirely and very weakly sclerotized; inner wall of spermathecal head and neck, along border with head, finely and densely striate.
Distribution.This species has a wide range in distribution covering the entire Holarctic region, mainly Europe and North Africa (Schülke and Smetana 2015).The records in the Nearctic region are regarded as those species introduced from Europe (Klimaszewski et al. 2002).In Russia, the easternmost record reported is from West Siberia (Schülke and Smetana 2015).We provide new distributional records of A. morion as follows: new country record, Japan (Hokkaidô, Honshû); new regional record, the Russian Far East.Bionomics.SY collected eight specimens from one Hokkaido brown bear (Ursus arctos) dung found on the roadside of a mixed needleleaf and broadleaf forest in Hokkaidô, Japan (Fig. 36).Aleochara morion has been found among various habitats, such as decaying organic matter, including fungi, compost, animal excrement, and carrion (Horion 1967).In addition, this species is found in moss, bark debris, straw, hay, and on sandy soil (Klimaszewski et al. 2002).
Diagnosis.This species is distinguished from the other congeneric species of the subgenus by a following combination of character states: body reddish brown to dark brown (Fig. 2); median lobe of aedeagus of male with a coiled flagellum, and with two pairs of characteristic sclerites (Figs 24,25); spermatheca with a curved spermathecal head, unequally serrated inner walls inside spermathecal head, and with multiple coils at base (Fig. 26).Aleochara eoa is the most similar externally to A. (T.) rougemontiana (Pace, 1999a), comb.n., from mainland China, differing from it additionally by having much less coiled spermatheca in the female (Pace 1999a: Fig. 183).
Body (Fig. 2): fusiform, compact, and robust; dorsal surface somewhat strongly glossy and pubescent, covered with small and inconspicuous micro-reticulation.
Color (Fig. 2): usually uniformly dark reddish brown to dark brown; antennomeres I-IV much lighter, but segments V to XI darker with numerous minute whitish setae; mouthparts and legs light-yellowish brown to reddish brown; pubescence yellowish brown to brown.
Abdomen (Fig. 2): first three visible tergites rather shallowly impressed transversely at base; dorsal and ventral surface covered with setae densely.
Male.Tergite VIII (Fig. 20): basal suture fully developed; posterior margin very weakly serrate, insignificantly emarginate medially; dorsal surface covered with setae rather sparsely, with six macrosetae.Sternite VIII (Fig. 21): basal suture fully developed; posterior margin rounded to only weakly produced; ventral surface covered with short setae sparsely, with approximately nine macrosetae.Median lobe of aedeagus (Figs 24 & 25): ovular in lateral and limuloid in parameral view; apical lobe rather slender, gently curved paramerally, weakly narrowing apically in parameral but with weakly dilated apex in lateral view; without a protuberance at base of apical lobe; a pair of sclerites S-shaped, longer than half length of apical lobe; flagellum strongly developed, much longer than median lobe, coiled 1.5 times at base.
Female.Tergite VIII (Fig. 21): basal suture fully developed; posterior margin very weakly serrate or almost truncate; dorsal surface covered with setae rather sparsely, with six macrosetae.Sternite VIII (Fig. 23): basal suture fully developed; posterior margin rounded; ventral surface covered with setae rather sparsely, with approximately nine macrosetae.Spermatheca (Fig. 26): deformed M-shaped; spermathecal head curved at middle; attachment of spermathecal duct inconspicuous; basal part of spermathecal stem moderate in size, clearly longer than spermathecal neck, with approximately ten coils attached complicatedly at base; each part of spermatheca entirely and very moderately sclerotized; inner wall of spermathecal head and neck, along border with head, finely and densely striate irregularly.
Etymology.The replacement name is derived from "Eos" of the Greek mythology which is a Titaness and the goddess of the dawn because "Nippon" (= Japan, type locality) means a country of the dawn.
Distribution.Japan, Taiwan (new record).Bionomics.One individual was caught with a flight interception trap (FIT).Host records.No host record is available.
Remarks on type materials.Five syntypes were found.Among them, a male specimen (Fig. 35) labeled "Tinotus / japonicus / TYPE Cam [HW]" is designated as the lectotype herein.
Abdomen (Fig. 3): first three visible tergites deeply impressed transversely at base; dorsal and ventral surface covered with setae rather sparsely.
Male.Tergite VIII (Fig. 27): basal suture not fully developed, suture partially disappeared laterally (Fig. 27: arrow 1); posterior margin very weakly serrate, insignificantly emarginate medially or truncate; dorsal surface covered with setae rather densely, with six macrosetae.Sternite VIII (Fig. 29): basal suture completely lost (Fig. 29: arrow 3); posterior margin rounded; ventral surface covered with short setae dense- ly, with approximately eight macrosetae.Median lobe of aedeagus : narrowly elongate in lateral, and limuloid narrowly in parameral view; apical lobe slender and long, as long as basal capsule, moderately narrowing apically in lateral view, with deeply notched apex medially in parameral view (Fig. 33); without a protuberance at base of apical lobe; a pair of sclerites curved just above middle, long, approximately 2/3 length of apical lobe; flagellum developed, shorter than median lobe, without any coils at base.
Etymology.The species name is dedicated to its collector, Mr. Takashi Watanabe (Kanagawa, Japan).
Bionomics.Most specimens were caught with flight interception traps (FIT).Host records.No host record is available.
Remarks.This new species is distinct among the species of Tinotus.In particular, reduced or non-developed basal sutures on tergite and sternite VIII of both sexes are notable character states .Furthermore, the apical lobe of the median lobe of male aedeagus, [i.e., deeply notched medially at apex in parameral view (Fig. 33)], is also a remarkable character state even among the subfamily.Since other morphological characters correspond fully to that of the subgenus Tinotus, we assign this species to Tinotus without hesitation.
Key to species of the subgenus Tinotus (genus Aleochara) from Japan, Taiwan, and the Russian Far East Body black to blackish brown including pronotum and elytra (Fig. 1); median lobe of aedeagus of male with a basal protuberance in lateral view (Fig. 16); spermatheca coiled four times basally, with oblong and simple spermathecal head (Fig. 19) ..

Discussion
We recognize three Tinotus species from Japan.This species count is clearly lower than those for mainland China (6 spp.: Pace 2013a; Schülke and Smetana 2015) and the United States (6 spp.: Klimaszewski et al. 2002;Gusarov 2003).In contrast, it surpasses the counts of adjacent countries, e.g., South and North Korea (0 sp.: Schülke and Smetana 2015) and Russia, including the Far East (1 sp.; Schülke and Smetana 2015; present study).Surprisingly, all of the continental European countries has no, or at most, only one Tinotus species (A.morion; Schülke and Smetana 2015).Within central Honshû, Japan, we found all of the three species.Therefore, Japan, especially central part of the country, is considered to possess a relatively high species diversity of Tinotus.
Only one species has been confirmed in Taiwan and the Russian Far East, respectively, which may reflect potentially low species diversity in these regions or merely insufficient accumulation of materials.Since taxonomic studies and records of Tinotus in East Asia, including these two regions, are still lacking, further discussions are avoided here.The finding of A. eoa in Taiwan implies a wide distributional range of this species, and it may be discovered on mainland China.
recorded A. morion on the mainland of China.