Fig. 3
Pterostichus marginatus: Matsumura (1911): 111 (original description), type locality: “Tonnaitcha, Ushoro” [written as “Oshiyoro” on the labels of syntypes, but both corresponding to the same locality, Okhotskoe, southern Sakhalin]; Bousquet (2003): 515; Bousquet (2017): 747.
Pterostichus (Lyperopherus) subrugosus: Straneo (1955) (original description): 88, type locality “M. Daisetsu, Hokuchindake” [Mt. Hokuchin-dake, the Daisetsuzan Mountains, Hokkaidô]; Jedlička (1962): 266.
Pterostichus (Lyperopherus) wellschmiedi: Kirschenhofer (1985): 224 (original description), type locality: “Süd-Sachalin, Umgeb. Nowoalexandrowsk, Berg Tschkowa” [Mt. Chekhov, southern Sakhalin]. Syn. nov.
Pterostichus (Lenapterus) subrugosus: Morita (1995): 4; Bousquet (2003): 497.
Pterostichus (Steroperis) wellschmiedi: Kryzhanovskij et al. (1995): 105.
Pterostichus (Lenapterus) wellschmiedi: Berlov & Berlov (1997d): 39; Bousquet (2003): 497 (synonym of subrugosus); Bousquet (2017): 718.
Materials examined.Pterostichus marginatus marginatus, 9♂3♀, Sakhalin (P. marginatus Matsumura, 1911syntypes1♂, “/ 11/VIII 1906 Oshiyoro” [partly in Japanese] and 1♀, “ 11/VIII 1906 Oshiyoro” [partly in Japa- nese], “P. margina?? n sp.” in HUM; 3♂1♀, Makarov distr., Porechje envir., 10–15 km from sea shore, larch forest, 28.vii.1991, E. Berlov leg. in BO coll., 1♂1♀, Yuzhno-Sakhalinsk distr., Novoaleksandrovsk envir., Mt. Chekhov, 19.vi.1990, E. Berlov leg. in BO coll.; 3♂, Yuzhno-Sakhalinsk distr., Novoaleksandrovsk envir., Mt. Chekhov, 400–600 m, 10.vi.1988, leg. E. Berlov leg. in BO coll.); P. marginatus subrugosus, 2♂2♀, Kumonodaira, the Daisetsuzan Mountains, Hokkaidô, Japan, 5.viii.2005, N. Yasuda leg. in KS coll.
Other consubgeneric species examined. The following species were investigated through literature: P. cancellatus (Motschulsky) by Sundukov (2005); P. vermiculosus (Ménétriés) by Sasakawa & Kubota (2005).
Notes.Pterostichus marginatus was described from southern Sakhalin but has been forgotten since its description and treated as incertae sedis in the Palaearctic Catalogue (Bousquet 2003, 2017). Our comparison of the syntypes of P. marginatus with a photo of a female paratype of P. wellschmiedi (Kirschenhofer 1985, fig. 9) and with specimens treated as P. wellschmiedi revealed that P. wellschmiedi is a synonym of P. marginatus. Moreover, comparison of this Sakhalin species with P. subrugosus from Hokkaidô revealed the relationship between the two taxa. Although the previously reported differences in external characters between the two (e.g., flatter elytra with less frequently interrupted intervals, and the absence of metallic luster of dorsal surface in P. subrugosus; Sundukov 2005) were confirmed, no conspicuous differences were found in the structure of the endophallus. Thus, P. subrugosus is downgraded from a distinct species to subspecies P. marginatus subrugosusstat. nov. This species was recorded from the summit of Mt. Rishiri-dake based on two female specimens (Morita 1995, as P. subrugosus). However, the subspecies identity of the Rishiri population remains undetermined because specimens from the locality were not examined in this study.