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Eight species attributed to Tmeticus are reviewed; five are redescribed and illustrated: Tmeticus affinis (Blackwall, 1885), Tmeticus bipunctis (Bösenberg & Strand, 1906), Tmeticus nigriceps Kulczyński, 1916, Tmeticus ornatus (Emerton, 1914) and Tmeticus tolli Kulczyński, 1908. The new genus, Paratmeticus gen. n. is erected for Tmeticus bipunctis, and a new combination is established: Paratmeticus bipunctis (Bösenberg & Strand, 1906), comb. n. Three species names: Gongylidium vile Kulczyński, 1885, syn. n., Tmeticus difficilis Kulczyński, 1926, syn. n. and Tmeticus dubius Kulczyński, 1926, syn. n., are synonymized with Tmeticus tolli Kulczyński, 1908. Although Gongylidium vile has date priority over Tmeticus tolli it is synonymized because of the lack of usage. Three species from Japan attributed to Tmeticus: Tmeticus neserigonoides Saito & Ono, 2001, Tmeticus nigerrimus Saito & Ono, 2001 and Tmeticus vulcanicus Saito & Ono, 2001 are not related to Tmeticus affinis, the type species of the genus, and their affinities remain unclear. The male of Tmeticus nigriceps is described for the first time.
spiders, Erigoninae, Palaearctic, Nearctic, new synonym, new combination
Tmeticus is a small Erigoninae genus with nine species restricted to the Holarctic Region (
While studying the Siberian and Far Eastern Linyphiidae, we encountered certain difficulties in identifying Tmeticus species. Only two of the four species occurring in northern Asia, Tmeticus affinis (Blackwall, 1885) and Tmeticus tolli Kulczyński, 1908, were properly illustrated. Thus, the main purposes of this study are to provide diagnostic illustrations for each Asian species and to describe a new genus.
Material and methodsPictures of the general appearance and copulatory organs
were made using an Olympus SZX16 stereomicroscope, with an Olympus
E-520 camera, and prepared using CombineZM software. Photographs were
taken in dishes of different sizes with paraffin in the bottom.
Different sized holes were made in the bottom to retain the specimens in
the desired position. Scanning electron micrographs were made using a
SEM JEOL JSM-5200 scanning microscope. SEM and digital photographs were
made in the Zoological Museum, University of Turku. The terminology of
the copulatory organs follows
IBPN Institute for Biological Problems of the North, Russian Academy of Sciences, Magadan (curator Yu.M. Marusik).
PSU Department of Zoology, Perm State University (curator S.L. Esyunin).
ZMMU Zoological Museum of the Moscow State University (curator K.G. Mikhailov).
ZMUT Zoological Museum, University of Turku (curator S. Koponen).
Tmeticus leptocaulis Menge, 1868 (= Tmeticus affinis (Blackwall, 1855)).
Males of this genus are easily recognized by possessing a mastidion (large tooth on frontal part of chelicera) and by their elongate palp with patella longer than cymbium, ventral terminal tooth on patella, and thin bulbus (as wide as terminal part of tibia). Females are recognized by their flat epigyne without a cavity. Males may be confused only with the trans-Palaearctic Hylyphantes graminicola (Sundevall, 1830) because it also has a mastidion and a patellar tooth. However, the males of Hylyphantes have shorter palp, undivided embolic division and screw-like embolus.
The females of Tmeticus may be confused with those of several genera, such as Oedothorax Bertkau, 1883 or with Donacochara speciosa (Thorell, 1875). However, Oedothorax females have a different colour pattern, and Donacochara speciosa is notably larger.
Small to medium-sized (2.5–4.1), light to dark-coloured erigonines. Male carapace unmodified and without sulci, it may be uniformly coloured or with a darker cephalic region. Abdomen unmodified, dark, of uniform colour. Male chelicera modified by possessing a mastidion (Ma, promarginal tooth). Maxilla with apical-retrolateral spine. Tibial spines 2-2-1-1. TmIV present. TmI 0.65–0.8. Male palp elongate. Femur, patella and tibia longer than wide. Patella with conical, ventral terminal tooth (Tt). Tibia with two apophyses (Ta). Paracymbium large, with or without (Tmeticus affinis) distinct apical pocket. Tegulum with distinct sac (Ts) and large (Tmeticus tolli) or small protegulum (Pt). Radix with straight apical process (Ap), tailpiece (Tp) without extension, embolus (Em) short and straight, or long and forming a semicircle; embolic membrane (Me) large. Epigyne without distinct fovea or openings. Median (=dorsal, sensu Hormiga 2002) plate plain or with ridges.
According to Platnick’s (2010) catalogue eight species are listed in this genus: Tmeticus affinis (Blackwall, 1855) (Holarctic), Tmeticus bipunctis (Bösenberg & Strand, 1906) (Far East Asia), Tmeticus neserigonoides Saito & Ono, 2001 (Japan), Tmeticus nigerrimus Saito & Ono, 2001 (Japan), Tmeticus nigriceps (Kulczyński, 1916) (Northern Siberia), Tmeticus ornatus (Emerton, 1914) (USA & Canada), Tmeticus tolli Kulczyński, 1908 (Siberia) and Tmeticus vulcanicus Saito & Ono, 2001 (Japan). In fact, there are three more species names within this genus: Tmeticus difficilis Kulczyński, 1926, Tmeticus dubius Kulczyński, 1926 and Gongylidium vile Kulczyński, 1885. Of these, the first two are listed under Centromerus, and the last one under Oedothorax. These three names were included in Tmeticus by
On the basis of the present study, we conclude that Tmeticus encompasses four species: Tmeticus affinis, Tmeticus nigriceps, Tmeticus ornatus and Tmeticus tolli. A new genus has been erected for Tmeticus bipunctis. Tmeticus neserigonoides might be correctly placed in this genus, but as we failed to re-examine its specimens, we treat it as incertae sedis (see below). Two other Japanese species belong elsewhere, but their correct assignments require further study.
Tmeticus is unusual in the Erigoninae because all its species can be recognized by their carapace colour pattern. Three sibling species: Tmeticus nigriceps, Tmeticus ornatus and Tmeticus tolli cannot be recognized by their embolic division, but the females of these species have distinctly different epigynes.
Tmeticus affinis differs from the three other species by the shape of the paracymbium, the straight embolus and the high protegulum with papillae. It also possesses a different type of the tibial apophysis, not originating at the terminal edge of the tibia as in other Erigoninae and other Tmeticus, but slightly aside of the edge.
In general appearance, male palp structure and cheliceral dentition, the members of this genus are similar to Hylyphantes graminicola, but the latter has a different type of embolic division and epigyne. When
The embolic division of Tmeticus is similar to those in Phaulothrix hardyi (Blackwall, 1850) (cf.
Judging from the drawings (Figs 35.110, 35.111 in
The males of Tmeticus nigriceps and Tmeticus ornatus cannot be distinguished.
| 1. | Carapace uniformly coloured (Figs 10–11, 25–26) | 2 |
| – | Cephalic part darker than red/orange thoracic part (Figs 21–24) | 3 |
| 2. | Carapace red, orange/yellow; occurs in Siberia and the Far East | Tmeticus tolli |
| – | Carapace reddish brown, with slightly lighter posterior part (Figs 10–11), tibia with two small claw-like apophyses (Figs 1–2, 5, 7), median plate of epigyne square-shaped (Figs 4, 8–9, 19); distributed throughout the Holarctic | Tmeticus affinis |
| 3. | Cephalic part dark brown (Figs 23–24), epigyne with extended median plate (Figs 18, 43–44); occurs in the tundra zone of Siberia | Tmeticus nigriceps |
| – | Cephalic part brown (Figs 21–22), epigyne without extension (Figs 17, 47, 48); occurs in southern Canada and the northern United States | Tmeticus ornatus |
Figs 1–11, 19, 49, 61
FINLAND: 2♀ (ZMUT), Turku Ruissalo, 14.11.1966 (M. Saaristo); 2♀ (ZMUT), Turku Ruissalo, sea shore litter, 27.10.1966 (M. Saaristo); 1♂ (ZMUT) Turku Hirvensalo Illoinen, 30.5.1966 (P.T. Lehtinen); 1♀ (ZMUT), Turku Kärsämöki Pomponrahka, 30.05.1967 (M. Saaristo); 1♀ (ZMUT) Pori Yyteri, 16.10.1966 (M. Saaristo); 1♀ (ZMUT), Pudasjärvi Hirvaskoski, 12.08.1959 (P.T. Lehtinen); 1♂ (ZMUT), Kuusamo Torankijärvi, 7.7.1966 (M. Saaristo); 1♀ (ZMUT), Kajaani, Koutaniemi, 16.07.1972 (P.T. Lehtinen); 1♂ (ZMUT) Inari Repojoki, 9.7.1961 (O.V. Lindqvist); 1♀ (ZMUT) Utsjoki Kevo, birch forest on lake shore, 20.06.-20.07.1970 (E.T. Linnaluoto). RUSSIA: Krasnoyarsk Province, 1♂ 2♀ (ZMMU), Mirnoye, Yenisei River left bank, 23.06.1978 (K.Yu.Eskov); 2♀ (ZMMU), Mirnoye, Yenisei River left bank, 27.07.1979 (K.Yu. Eskov). Yakutia, 2♂ 2♀ (ZMUT), El’gyay, big ”alas” pond, 24.07.1977 (S. Koponen); 1♂ (ZMMU), western Yakutia, Kempendyay River 80 km up stream from the mouth, riverside meadow, 1–15.08.1988 (K.Yu. Eskov). Kamchatka Peninsula, 1♂ 1♀ (IBPN), Talovskoye Lake, Kuyul River, 16.08.1990 (M.B. Skopets). Chukotka: 1♂ (ZMMU), Markovo, July 1986 (G. Chernova). CANADA, Alberta: 1♂ (only the photo provided by D.J. Buckle has been studied), Caribou Mountain Wildlands, Wentzel Lake, 50°02N; 114°28W, sweeping horsetail meadow, 16.07.2003 (T. Johnson).
Copulatory organs of Tmeticus affinis. 1 male palp, retrolateral view 2 palpal tibia, dorso-lateral view 3 whole male palp, retrolateral view 4 epigyne, ventral view. (scale bar 0.1 mm).
Copulatory organs of Tmeticus affinis. 5 male palp, retrolateral view 2 male palp, prolateral view 7 male palp, dorsal view 8 epigyne, caudal view 9 epigyne, ventral view. Abbreviations: Pt - protegulum; Ta - tibial apophysis;Ts - tegular sac.
Habitus and epigyne of Tmeticus affinis (10–11, 19), Paratmeticus bipunctis (12–14), Tmeticus tolli (15–16, 20), Tmeticus ornatus (17) and Tmeticus nigriceps (18). 10, 12, 14–16 male habitus, dorsal view 11, 13 female general appearance, dorsal view 14, 17–20 epigyne, ventral view 15–16 difference in the size between males from the same sample. (scale bar 0. 2 mm, if not otherwise indicated).
This species is easily recognized by its brownish carapace with a darker cephalic region. Males are easily recognized by their palp, which has a characteristic tibial apophysis and embolic division with the anterior radical process equal in length to the embolus proper (embolus longer than anterior radial process in other species). Females are easily recognized by the shape of the epigyne.
For detailed description see
This species is known all over Eurasia, from
western Europe to Kamchatka. In the Nearctic Region, it has been
reported from Alberta (
Figs 15–16, 20, 25– 34, 50, 56
RUSSIA: Krasnoyarsk Province: 1♂ 2♀ (ZMMU), Mirnoye, Yenisei River left bank, 23.06.1978 (K.Yu. Eskov). Evenkiya: 40♂♀ (ZMMU), Taimura River, Neptene River mouth, riparian spruce forest with alder, Summer 1982 (K.Yu. Eskov); 2♀ (ZMMU), Chambe River mouth, meteorological station “Kerbo”, floodplain willow stand, litter, 21.08.1982 (K.Yu. Eskov). Khabarovsk Province: 2♀ (IBPN), Okhotski Dist., Gyrbykan R. (Ul’ya River basin), 20.08–15.09.1986 (I.D. Sukatcheva); 1♂ 3♀ (IBPN), Khetana River (tributary of Amka River, Ulya River basin), Agust 1985 (V.V. Zherikhin). Maritime Prov.: 3♂ 2♀ (IBPN), [05], Khanka Lake CW shore, Sosnovy Isl & peninsula nearby, 44°52N; 132°07E, 17.07.1998 (Yu.M. Marusik). 1♀ (IBPN), [03], Khanka Lake, CE shore, 44°39N; 132°34E, 15–16.07.1998 (Yu.M. Marusik). Magadan Area: 3♂ 2♀ (IBPN), Motykley Bay, 59°30N; 148°50E, Summer 1994 (E. Izergina); 1♂ 1♀ (IBPN), 137th km of Kolyma Hwy, 60°25N; 151°30E, Ola River, valley forest, 28.09.1994 (Yu.M. Marusik); 1♂ 2♀ (IBPN), ca 50 km N of Magadan, Khasyn River, environs of Splavnaya Vil., 28.05.1988 (Yu.M. Marusik); 25♂♀ (IBPN), 30km N of Magadan, Snow Valley Vil., Dukcha River valley, 7.10.1984 (Yu.M. Marusik). Sakhalin Island: 1♂ 6♀ (IBPN), Okha Dist., Ten’ga River, May 1987 (A.M. Basarukin); 4♀ (IBPN), Tomari Dist., Ainskoye Lake, Ptichya river, 24.05.-10.06.1984 (A.M. Basarukin); 1♀ CE part, Leonidovka River, 8 km SE of Leonidovo Vil., 49°16.506N; 142°58.390E, 9.08.2001 (Yu.M. Marusik). Kamchatka Peninsula: 2♀ (ZMMU), 40 km from Ust’-Kamchatsk, 09.1973 (A.S. Glikman); 1♂ 4♀ (IBPN), 10–12 km N of Paratunka Vil., Yelizovo Forestry, 53.050°N; 158.225°E, 15–28.07.2004 (A.S. Ryabukhin). MONGOLIA: Arkhangai Aimak: 2♂ 2♀ (IBPN) [12], Ondrer-Ulaan, Tsakhir, Chulut gorge 48°07N; 100°22E, 2100 m, 10–13.06.1997 (Yu.M. Marusik). Central (=Tov) Aimak: 1♀ (IBPN), Terelzh Mt., south exposed slope (about 80 km NE of Ulan-Bator, 1988 (S. Heimer).
Habitus of Tmeticus ornatus (21–22), Tmeticus nigriceps (23–24) and Tmeticus tolli (25–26). 21, 23, 25 male, dorsal view 22, 24, 26 female, dorsal view.
Copulatory organs of Tmeticus tolli. 27 male palp, retrolateral view 28 palpal tibia, dorso-lateral view 29 bulbus, prolateral view 30 – whole male palp, retrolateral view 31 epigyne, ventral view.
Copulatory organs of Tmeticus tolli. 32 male palp, retrolateral view 33 epigyne, ventral view 34 epigyne, caudal view. Abbreviations: Ap – anterior radial process; Me - embolic membrane;Ts - tegular sac.
Tmeticus tolli is easily distinguished from the similar Tmeticus ornatus and Tmeticus nigriceps by having a uniformly coloured carapace in both sexes.
Both sexes were described in detail by
Tmeticus difficilis Kulczyński, 1926 was described on the basis of the female holotype from Lake “Kurarotschnoje” (=Kurazhechnoye, ca. 56°10N; 161°45E, collected 9.06.1909) and Tmeticus dubius
Kulczyński, 1926 was described on the basis of two females from Lake
“Klutschevskoje” (= maybe Klyuchi Vil., c. 60 km from Kurarochnoye
Lake). In his descriptions
Although
This species is distributed east of Yenisei (
Figs 18, 23–24, 35–44, 52, 55
RUSSIA: Arkhangel’sk Area: 1♂ (IBPN), Barents Sea, Dolgiy Ilsand, 69°12'N, Summer 2004 (O.L. Makarova). Polar Ural: 1♂ (ZMUT), Oktyabrskij, Ob River shore, Salix viminalis vegetation, 12.-13.7.1994 (S. Koponen); 1♀ (PSU-95), North Ural expedition by Fridolin, sample 36, Sob’ River right bank, 4.07.1925 (V. Fridolin). Yamal Peninsula: 1♂ 5♀ (ZMMU), Yorkugayakha River, environs of “Canary” trading station, riparian willow stand, 08.07.2002 (D. Osipov); 4♂ 1♀ (ZMMU), south Yamal, Shchuchye Vill, Shchuchya River (A.L. Tikhomirova); 1♂ (PSU-96), South Yamal, Khadyta-Yakha River, meadow valley, pitfall traps, 8.08.1982 (S.L. Esyunin); 1♂ 1♀ (PSU-97), same locality, river bank, drift, 26.07.1981 (S.L. Esyunin). Taimyr Peninsula: 1♂ 1♀ (ZMMU), Taimyr Reserve, Novaya River, Ary-Mas Site, 25.07.1992 (A.B. Ryvkin); 10♂ 1♀ (ZMMU), SW Taimyr, Nyapan’ Ridge, 70°09N; 87°47E, Carex-moss bog, pitfall trapping, 1–10.08.1999 (D. Osipov); 3♂ (ZMMU), NW shore of Pyasino Lake, 70°04'54N; 87°32'12E, Carex bog with sphagnum tussocks, 10–20.07.1999 (D. Osipov); 3♂ 2♀ NW shore of Pyasino Lake, 70°04'54N; 87°32'17E, Carex bog with sphagnum tussocks, Summer 1999 (D. Osipov); 2♀ (ZMMU), NW shore of Pyasino Lake, Lazannakh Lake, sandy beach, 70°05'47N; 87°26'28E, 1–10.07.1999 (D. Osipov). Yakutia: 2♀ (IBPN), Yana River down flow, environs of Kular Village, 70°35N; 134°34E, grass-herb- Arctagrostis meadow on the former open mine, Summer, 2000 (N.K.Potapova). Chukotka: 2♂ (IBPN), western Chukotka, Chaun River mouth part, 68.810°N; 170.432°E, Summer, 1986 (A.S. Ryabukhin); 1♂ (IBPN), western Chukotka, Chaun Bay, Pucheveyem River mouth, 25.07.1985 (A.S. Ryabukhin); 1♂ (IBPN) Lamutskoye Vil., 65°32'39N; 168°51'08E, along creek, 17.08.1968 (Novikova); 1♂ (IBPN), western Chukotka, Markovo Town, July 1986 (G. Chernova); 1♂ (IBPN), Vulvyveyem River upper flow, Gytlenumkuum Stand, 67°10N; 178°E, 8.06.1988 (Yu.M. Marusik).
Copulatory organs and male carapace of Tmeticus nigriceps. 35 male palp, retrolateral view 36 palpal tibia, dorso-lateral view 37 whole male palp, retrolateral view 38 male carapace, dorsal view 39 epigyne, ventral view. (scale bar 0.1 mm).
Copulatory organs of Tmeticus nigriceps. 40 male palp, retrolateral view 41 male palp, prolateral view 42 male palp, dorso-prolateral view 43 epigyne, ventral view 44 epigyne, caudal view. Abbreviations: Ap – anterior radial process; Em - embolus proper; Me - embolic membrane;Ts - tegular sac.
Copulatory organs of Tmeticus ornatus. 45 male palp, retrolateral view 46 male palpal patella and tibia, retrolateral view 47 epigyne, caudal view 48 epigyne, ventral view. Abbreviations: Tt – patellar tooth.
Retrolateral view of the male palp of Tmeticus affinis (49), Tmeticus tolli (50), Tmeticus ornatus (51), Tmeticus nigriceps (52) and Paratmeticus bipunctis (53). (scale bar 0.1 mm).
Tmeticus nigriceps is easily distinguished from the other Palaearctic species by the dark cephalic region contrasting with the reddish thoracic area. Only the Nearctic Tmeticus ornatus has a similar colour pattern. The male palp is almost undistinguishable from the Siberian Tmeticus tolli and the Nearctic Tmeticus ornatus. In addition to the carapace pattern, females can be distinguished by the shape of their epigyne.
♀ 2.9–3.3, ♂ 2.3–2.7. TmI 0.69–0.72. Carapace orange with dark, blackish cephalic region (Figs 23–24, 38) and chelicera. Legs orange. Abdomen black. Palp as in Figs 35–37, 40–42, 52, epigyne as in Figs 18, 39, 43–44.
This species is known from Dolgiy Island and the Polar Urals to Chukotka (
Embolic division of Tmeticus ornatus (54), Tmeticus nigriceps (55), Tmeticus tolli (56), Oreoneta sp. (57), Lophomma punctatum (58), Paratmeticus bipunctis (59–60) and Tmeticus affinis (61). Abbreviations: Ap – anterior radial process; Em - embolus proper; Ma – mastidion; Me - embolic membrane;Pt - protegulum; Sa - distal suprategular apophysis;Tp - tailpiece;Ts - tegular sac.
Copulatory organs of Paratmeticus bipunctis. 62 male palp, retrolateral view 63 male palpal tibia, dorso-retrolateral view 64 whole male palp 65 epigyne, ventral view.
Copulatory organs and male chelicera of Paratmeticus bipunctis. 66 male palp, retrolateral view 67 male palp, ventral view 68 male chelicera inner view 69 male palp, from above 70 male palp, prolateral view 71 male palp, dorsal view 72–73 epigyne, ventral and caudal view. Arrows show the cheliceral teeth. Abbreviations: Ap – anterior radial process; Em - embolus proper; Ma – mastidion; Me – embolic membrane;Sa - distal suprategular apophysis;Tp - tailpiece;Ts - tegular sac.
Figs 17, 21–22, 45–48, 51, 54
CANADA: 4♂ 4♀ (ZMMU), Saskatchewan, Lady Lake, sedge tops – flooded marsh, 13–15.04.1971 (D.J. Buckle); 3♂ 3♀ (ZMUT), same locality, marsh, late April, 1978 (J.V. Buckle).
Differs from Tmeticus affinis, which also occurs in the Nearctic Region, by the carapace colour (black cephalic region and red-orange thoracic area in Tmeticus ornatus, carapace uniformly brown in Tmeticus affinis). The males are easily separated by their tibial apophyses (one apophysis with a claw-like processes in Tmeticus affinis and two separate apophyses in Tmeticus ornatus); the females have distinctly different epigynes.
♂ 2.5–3.3, ♀ 2.8–35. TmI 0.73–0.78. Carapace orange with darker cephalic region. Abdomen dark. Palp as in Figs 45–46, 51, 54. Epigyne as in Figs 17, 47–48.
This species has a trans-Nearctic distribution, recorded from British Columbia to Quebec and south to New York (
The three species from Japan assigned to Tmeticus remain unstudied and belong elsewhere (see ‘Comments’ below).
Judging from the available figures, this species might belong in Tmeticus, The male has a long palp with a patellar tooth. However, the chelicera appears to lack a mastidion and the tibial apophyses are absent. Figures of the male palp are unclear, TmI index (0.59) is lower than in Tmeticus species (>0.63).
This species is clearly not related to Tmeticus affinis or other members of the genus due to the short palpal patella lacking a tooth in the male, embolic division of a different shape, the relatively long tibial apophysis, lack of a mastidion, epigyne with a septum and some other additional characters. The correct generic placement remains unclear.
This species is clearly not related to Tmeticus affinis or other members of the genus due to the short palpal patella lacking a tooth in the male, embolic division of a different shape (anterior radical process absent), and some other characters. The correct generic placement remains unclear.
urn:lsid:zoobank.org:act:3F57381F-374C-4D90-9312-24A8419BF422
Oedothorax bipunctis Bösenberg and Strand, 1906.
Prefix “Para”- indicates the resemblance of this genus to Tmeticus The gender is masculine.
The new genus is easily distinguished from the similar Tmeticus by lacking distinct tibial apophyses, and in having the papillate tegular sac larger than the protegulum, a slightly twisted embolic division, a sharply pointed embolic membrane and a large distal suprategular apophysis, longer than the embolic division. In contrast to Tmeticus, themedian plate of the epigyne in the new genus is widest in the anterior region, rather than in the posterior region.
Medium-sized erigonine spiders. Uniformly coloured, male carapace without modifications, male chelicera with mastidion, inner row with 4 inner teeth and 5 outer teeth (all smaller than inner teeth). TmI 0.63–0.65. Male palp elongate, with patella as long as tibia, tibia lacks apophyses, distal suprategular apophysis longer than embolic division; embolic division slightly twisted with two arms: anterior radical process and embolus proper; embolus parallel to process with lamellate basal process; epigyne without cavity, median plate widest anteriorly.
The type species only.
Figs 12–14, 53, 59–60, 62– 73
RUSSIA: Sakhalin Island: 1♂ 4♀ (IBPN), Okha Dist., 5–7 km N of Kolendo Vil., 22–23.08.1991 (A.M. Basarukin); 1♂ 2♀ (IBPN), Pil’tun Bay, 06.-0.7.1991 (A.M.Basarukin); 1♂ 1♀ (IBPN), Okha Dist., Ten’ga River, May 1987 (A.M. Basarukin); 10♂ 26♀ (IBPN), Korsakov Dist., Tunaiga Lake south shore, 26.09.1991 (A.M.Basarukin). Kuril Isles, 4♂ 3♀ (IBPN), Paramushir Isl. NE shore, environs of Severo-Kuril’sk, 50°40N; 156°06E, 10.08–15.09.1996 (Yu.M. Marusik); 2♂ 1♀ (ZMMU), Iturup Island, Dobroye Nachalo Bay, Lesozavodskoye, mixed forest, 14.08.1994 (K.Yu. Eskov). Kamchatka Peninsula, 6♂ 2♀ (IBPN), 10–12 km N of Paratunka Vil., Yelizovo Forestry, 53.050°N; 158.225°E, 15–28.07.2004 (A.S. Ryabukhin).
Well described by
Kamchatka (south part), 8 islands in Kuril Archipelago (Shikotan, Kunashir, Iturup, Urup, Simushir, Ketoi, Shiashkotan, Paramushir, but seems to occur on all large islands); Sakhalin and Japan (Hokkaido, Honshu and Kyushu).
The material used in this study became available thanks to K.Yu.Eskov, K.G. Mikhailov, D. Osipov (all from Moscow, Russia), D.I. Berman, A.S. Ryabukhin (both from Magadan, Russia) and D.J. Buckle (Saskatoon, Canada). S.L. Esyunin helped us with the information about Tmeticus in the Urals. The English of the final draft was edited by D. Penney (Manchester, UK). The study was supported in part by financial help provided by RFFI grant 09–04–01365.