Bembidion (Limnaeoperyphus) quadriimpressum ( Motschulsky, 1860 )

Makarov, Kirill V. & Sundukov, Yurii N., 2024, A revision of the subgenus Limnaeoperyphus Nakane, 1963 (Coleoptera: Carabidae, Bembidion), with the description of a new species, Zootaxa 5493 (4), pp. 345-370 : 347-354

Bembidion (Limnaeoperyphus) quadriimpressum ( Motschulsky, 1860)

( Figures 1–17 , 31–36 , 50–53 , 58–77 , 98–105 )

Lymnaeum quadriimpressum Motschulsky, 1860: 90–91 , pl. vi, fig. 8. Type locality: “les bords de l’Océan pacifique dans la baie d’Avatcha au Kamtschatka et sur les iles Kourilles” (original citation), Avacha Bay on the east coast of Kamchatka and the Kuril Islands , Russian Far East.

Lectotype, ♂ with labels: illegible text [red rectangle; handwritten], “ Lymnaeum 4 impressum Motsch. Kamtschatka ” [white rectangle; handwritten], “ Holotypus Bembidion (Limnaeum) quadriimpressum Motsch. 73 Lafer det.” [red rectangle; handwritten], “ LECTOTYPUS Lymnaeum quadriimpressum Motschulsky, 1860 design Yu.Sundukov et K.Makarov, 2024.” [red rectangle; printed] [ ZMMU].

Paralectotype, ♀ with labels “Ins. Kuril.” [folded white rectangle; printed], “Scher (illegible text) 2.19.” [white rectangle; handwritten], “ Phila 4 impressum Menet. Urupa.” [white rectangle; handwritten], “Holotypus B. (Limnaeum) quadriimpressum Mot. ” [red rectangle; handwritten], “PARALECTOTYPUS Lymnaeum quadriimpressum Motschulsky, 1860 design Yu.Sundukov et K.Makarov, 2024.” [red rectangle; printed] [ ZISP]. The specimen is severely damaged: 4–11 left antennomeres, the left fore and right hind legs are missing, the right elytra is split in the middle .

Others material. RUSSIA: Kuril Archipelago: Zelyonyi Island [Khabomai Arch.], western coast 1.5 km N of Glushnevsky Cape, single-sand beach under wracks, 6.VIII.1994, leg. K. Eskov, 1 ex [ MSPU]; Tanfiliev Island, Zorkaya Bay, sandy beach between the sea and a freshwater lake, 7.VI.2017, leg. Yu. Sundukov, 14 ex [ FEB, MSPU]; Yuri Island, western bay of Shirokaya Bay, sand and pebble beach, 26.VIII.2016, leg. Yu. Sundukov & L. Sundukova, 9 ex [ FEB]; ibid, 1.IX.2016, Yu. Sundukov & L. Sundukova, 2 ex [ FEB]; Yuri Island, Katernaya Bay, sand and pebble beach, 28.VIII.2016, Yu. Sundukov & L. Sundukova, 8 ex [ FEB]; Yuri Island, eastern bay of Shirokaya Bay, sand and pebble beach, 30.VIII.2016, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; Yuri Island, ocean coast, sand and pebble beach, 31.VIII.2016, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; Polonsky Island, Chasovaya Bay, sandy beach, 30.VIII.2017, leg. Yu. Sundukov & L. Sundukova, 27 ex [ FEB, MSPU]; Polonsky Island, Moryakov Bay, sandy beach, 2.IX.2017, leg. Yu. Sundukov & L. Sundukova, 41 ex [ FEB]; Polonsky Island, coast near Cape Yazykovy, sandy beach, 5.IX.2017, leg. Yu. Sundukov & L. Sundukova, 19 ex [ FEB]; Polonsky Island, Udobnaya Bay, sand-pebble beach, 6.IX.2017, leg. Yu. Sundukov & L. Sundukova, 48 ex [ FEB]; Polonsky Island, Cape Severnyi, sandy beach, 9.IX.2017, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; Shikotan Island, in nests Larus sp. 14.VIII.56, leg. Violovich, 11 ex [ ZISP]; Shikotan Island, mouth of Vesolyy Stream, 43°49’36’’N 146°54’02’’E, 26.VI.2011, leg. I. Melnik, 16 ex [ MSPU]; Shikotan Island, Malaya Tserkovnaya Bay, peat shore in the high tide zone, 16.VIII.2022, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; Shikotan Island, Tserkovnaya Bay, sand-pebble beach at the mouth of the stream, 25.V.2012, leg. Yu. Sundukov, 3 ex [ FEB]; ibid, 3.VI.2012, leg. Yu. Sundukov, 52 ex [ FEB, MSPU]; ibid, 7.VI.2012, leg. Yu. Sundukov, 11 ex [ FEB]; ibid, 26.VIII.2012, leg. Yu. Sundukov & L. Sundukova, 2 ex [ FEB]; ibid, 12.VIII.2016, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; ibid, 18.VIII.2022, leg. Yu. Sundukov & L. Sundukova, 46 ex [ FEB]; Shikotan Island, mouth of the Anama River, muddy shore in the high tide zone, 13.VIII.2022, leg. Yu. Sundukov & L. Sundukova, 14 ex [ FEB]; Shikotan Island, Malokurilskaya Bay, sand-pebble beach, 20.VIII.2016, leg. Yu. Sundukov, 1 ex [ FEB]; Kunashir Island, Veslo Cape, 43°39’12”N 145°32’22”E, 21.VII.2008, leg. I. Melnik, 3 ex [ MSPU]; Kunashir Island, Veslovsky Peninsula, muddy shore in the high tide zone, 27.VII.2017, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; Kunashir Island, Veslovsky Peninsula 1.5 km north of Veslo Cape, 43°39’57”N 145°32’12”E, 20– 22.VII.2008, leg. K. Makarov, 1 ex [ MSPU]; Kunashir Island, Belozerskie lakes, sandy beach, 19.VIII.2017, leg. Yu. Sundukov, 1 ex [ FEB]; ibid, 23.VIII.2017, leg. Yu. Sundukov, 8 ex [ FEB]; Kunashir Island, mouth of the Ricorda River, muddy shore in the high tide zone, 19.VIII.2017, leg. Yu. Sundukov, 1 ex [ FEB]; Kunashir Island, env. Golovnino, 43°44’05”N 145°31’15”E, 14.VII.2008, leg. I. Melnik, 1 ex [ MSPU]; Kunashir Island, Golovnino, on the seashore under seaweed, 8.IX.1976, leg. V. Kuznetsov, 1 ex [ FEB]; Kunashir Island, mouth of the Golovnina River, sandy beach, under seaweed, 20.VIII.2017, leg. Yu. Sundukov, 1 ex [ FEB]; ibid., leg. K. Makarov, 11 ex [ MSPU]; Kunashir Island, interfluve of the Golovnin and Khlebnikov rivers, fine gravel beach, under seaweed, 7. VI.2015, leg. Yu. Sundukov, 12 ex [ FEB]; ibid, 30.IX.2015, leg. Yu. Sundukov, 5 ex [ FEB]; Kunashir Island, mouth of the Khlebnikov River, under the algae, 16.VI.2015, leg. Yu. Sundukov, 12 ex [ FEB]; Kunashir Island, mouth of the Khlebnikov River, 4 km east of Paltusovo, 43°43’59’’N 145°29’33’’E, 11.VIII.2011, leg. K. Makarov, 5 ex [ MSPU]; Kunashir Island, mouth of Sennaya River ca. 2.5 km eastern Paltusovo, 43°43’44”N 145°28’34”E, 11.VIII.2011, leg. K. Makarov, 11 ex [ MSPU]; Kunashir Island, env. Paltusovo, 43°43’31”N 145°26’48”E, 15.VI.2011, leg. A. Matalin, 4 ex [ MSPU]; Kunashir Island, mouth of the Bystry Creek, sandy beach, 10.VII.2013, leg. L. Sundukova, 2 ex [ FEB]; Kunashir Island, mouth of the Bystry Stream, 43°47’18’’N 145°25’19’’E, 18.VIII.2011, leg. K. Makarov, 3 ex [ MSPU]; Kunashir Island, mouth of the Temnaya River, sandy beach, 6.VI.2015, leg. Yu. Sundukov, 5 ex [ FEB]; Kunashir Island, ca. 2 km north of Bolotny Stream, 43°49’13’’N 145°24’32’’E, 18.VIII.2011, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, 3 km south of Ivanovsky Cape, mouth of the stream, 43°48’53’’N 145°24’44’’E, 15.VI.2011, leg. I. Melnik, 2 ex [ MSPU]; Kunashir Island, mouth of the Vodopadny Stream south of Ivanovsky Cape, 43°48’20’’N 145°24’59’’E, 11.VI.2011, leg. A. Matalin, 13 ex [ MSPU]; ibid, 18.VIII.2011, leg. K. Makarov, 9 ex [ MSPU]; Kunashir Island, cliff betweeh Ivanovsky Cape and Vodopadny Stream, 43°48’55”N 145°24’41”E, 15.VI.2011, leg. A. Matalin, 32 ex [ MSPU]; Kunashir Island, small stream 2 km south of Grozovoye, sandy beach, 2.VI.2015, leg. Yu. Sundukov, 8 ex [ FEB]; Kunashir Island, Cape Ivanovsky, sandy beach, 25.V.2013, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; ibid, 16.VI.2013, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; ibid, 14.VII.2013, leg. Yu. Sundukov & L. Sundukova, 3 ex [ FEB]; ibid, 2.IX.2013, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; ibid, 4.IX.2013, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; Kunashir Island, mouth of the Krivonozhka River, sandy beach, 28.V.2013, leg. Yu. Sundukov & L. Sundukova, 5 ex [ FEB]; Kunashir Island, sea coast north of the Krivonozhka River, sandy beach, 11.VII.2013, leg. Yu. Sundukov & L. Sundukova, 2 ex [ FEB]; Kunashir Island, stream north of the Krivonozhka River, pebble beach, 19.IX.2013, leg. Yu. Sundukov & L. Sundukova, 10 ex [ FEB]; Kunashir Island, mouth of stream, 1.3 km north of Krivonozhka River, 43°50’55’’N 145°26’08’’E, 16.VIII.2011, leg. K. Makarov, 5 ex [ MSPU]; Kunashir Island, mouth of stream, southern of Blizhny Island, 43°51’05’’N 145°26’37’’E, 26.VII.2011, leg. K. Makarov, 4 ex [ MSPU]; Kunashir Island, mouth of stream, 1 km north of Blizhny Island, 43°51’22’’N 145°26’51’’E, 16.VIII.2011, leg. K. Makarov, 8 ex [ MSPU]; Kunashir Island, mouth of the Ozernaya River, 43°53’07”N 145°27’44”E, 7.VII.2008, leg. K. Makarov, 1 ex [ MSPU]; Kunashir Island, 1.5 km south-west mouth of the Ozernaya River, sea coast, 43°52’27”N 145°27’16”E, 26.VII.2011, leg. K. Makarov, 3 ex [ MSPU]; Kunashir Island, 6 km SW of Alekhin Cape, 43°53’58’’N 145°28’31’’E, 20.VIII.2009, leg. I. Melnik, 1 ex [ MSPU]; Kunashir Island, bay south-west of the Odinoky Creek, 43°54’36”N 145°30’25”E, 1.VI.2009, leg. I. Melnik, 2 ex [ MSPU]; ibid, 1.VI.2011, leg. A. Matalin, 3 ex [ MSPU]; Kunashir Island, mouth of Odinoky Stream 1.5 km south of Alekhino, 43°54’47”N 145°30’46”E, 3.VIII.2011, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, mouth of the Odinokiy Creek, sandy bank of the stream, 10.IX.2014, leg. Yu. Sundukov, 2 ex [ FEB]; Kunashir Island, 1 km south-west of Alekhino, coastal rockery, 43°54’58’’N 145°31’00’’E, 4.VIII.2011, leg. K. Makarov, 5 ex [ MSPU]; Kunashir Island, near Alekhino, fumarole, 6.VIII.1970, leg. Loktin, 1 ex [ ZISP]; Kunashir Island, near Alekhino, sea shore under algae, 3.VIII.1970, leg. Loktin, 32 ex [ ZISP]; Kunashir Island, near Alekhino, sandy beach, 14.IX.2014, leg. Yu. Sundukov, 1 ex [ FEB]; ibid, 5.VII.2015, leg. L. Sundukova, 3 ex [ FEB]; ibid, 7.VII.2015, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; ibid, 23.VII.2015, leg.Yu.Sundukov &L.Sundukova, 1 ex [ FEB]; Kunashir Island, mouth of Alekhina River, 43°55’15”N 145°31’59”E, 2.VIII.2011, leg. K. Makarov, 5 ex [ MSPU]; Kunashir Island, flood-plain of the Alekhina River near mouth, 43°55’14’’N 145°32’09’’E, 2.VIII.2011, leg. K. Makarov, 1 ex [ MSPU]; Kunashir Island, Cape Puzanov, sandy beach, 16.VIII.2014, leg. Yu. Sundukov & L. Sundukova, 6 ex [ FEB]; ibid, 26.VIII.2014, leg. Yu. Sundukov & L. Sundukova, 8 ex [ FEB]; Kunashir Island, north coast of Puzanova Cape, 43°52’23’’N 145°36’23’’E, 21.VIII.2011, leg. K. Makarov, 5 ex [ MSPU]; Kunashir Island, northern coast of Chetverikova Cape, 43°53’01”N 145°37’30”E, 10.VII.2008, leg. K. Makarov, 14 ex [ MSPU]; Kunashir Island, mouth of the Andreevka River, sandy beach, 9.VIII.2014, leg. Yu. Sundukov & L. Sundukova, 5 ex [ FEB]; Kunashir Island, mouth of the Andreevka River, sandy river bank, 22.VIII.2014, leg. Yu. Sundukov, 1 ex [ FEB]; Kunashir Island, mouth of Andreevka River, 43°53’13.9”N 145°37’32.6”E, 7.VII.2008, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, mouth of the Belkina River, 43°53’26’’N 145°37’39’’E, 20.V.2011, leg. I. Melnik, 5 ex [ MSPU]; Kunashir Island, eastern shore of Glukhoe Lake, 43°54’11”N 145°38’12”E, 9.VII.2008, leg. I. Melnik, 7 ex [ MSPU]; Kunashir Island, mouth of Tyurina River, 43°54’33’’N 145°41’06’’E, 22.VIII.2011, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, south coast of Vodopadny Cape, 43°54’39’’N 145°39’31’’E, 22.VIII.2011, leg. K. Makarov, 6 ex [ MSPU]; Kunashir Island, Vodopadny Cape, 43°54’27”N 145°40’40”E, 22.VIII.2011, leg. K. Makarov, 5 ex [ MSPU]; Kunashir Island, Krugly Cape, ca. 1.5 km NE of Tretyakovo, 44°00’16”N 145°39’21”E, 26.VIII.2017, leg. K. Makarov, 3 ex [ MSPU]; Kunashir Island, coast of Okhotsk Sea, south of Znamenka Cape, 43°56’10”N 145°33’13”E, 4.VIII.2009, leg. K. Makarov, 12 ex [ MSPU]; ibid, 22.VIII.2009, leg. K. Makarov, 4 ex [ MSPU]; Kunashir Island, shore of Peshchanoe Lake, 43°56’36”N 145°35’22”E, 5.VI.2011, leg. A. Matalin, 4 ex [ MSPU]; Kunashir Island, western shore of Lake Peshchanoe, sandy shore of the lake, 4.IX.2014, leg. Yu. Sundukov, 10 ex [ FEB]; ibid, 7.IX.2014, leg. Yu. Sundukov, 24 ex [ FEB]; ibid, 6.VIII.2011 leg. K. Makarov, 2 ex [ MSPU]; ibid, 8.VIII.2008 leg. K. Makarov, 2 ex [ MSPU]; ibid., 30.VII.2011, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, south-west shore of Peshchanoe Lake, 43°56’33”N 145°33’20”E, 22.VIII.2009,A. Zaitsev, 26 ex [ MSPU]; Kunashir Island, north-west shore of Peshchanoe Lake, 43°56’41”N 145°35’39”E, 6.VIII.2011, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, Danilovo Natural Boundary, sea shore, 43°57’15”N 145°35’29”E, 7.VIII.2011, leg. K. Makarov, 9 ex [ MSPU]; Kunashir Island, cordon Alekhinsky, 43°57’17”N 145°35’34E, 17–18.IX.2009, leg. I. Melnik, 5 ex [ MSPU]; Kunashir Island, mouth of the Asin Stream, 43°58’59’’N 145°37’33’’E, 8.VIII.2009, leg. K. Makarov, 2 ex [ MSPU]; Kunashir Island, Tretjakovo, 10.VIII.1973, leg. Kerzhner, 3 ex [ ZISP]; Kunashir Island, mouth of Valentiny Stream, 43°59’34”N 145°38’47”E, 10.VIII.2011, leg. K. Makarov, 21 ex [ MSPU]; Kunashir Island, mouth of Zmeiny Stream, 44°00’40”N 145°40’27”E, 27.VII.2008, leg. I. Melnik, 3 ex [ MSPU]; Kunashir Island, mouth of the Stolbovskoy Stream, 44°00’39’’N 145°40’29’’E, 9.VI.2011, leg. A. Matalin, 2 ex [ MSPU]; Kunashir Island, Pervukhina Bay, 44°03’18”N 145°44’36”E, 15.VIII.2011, leg. K. Makarov, 4 ex [ MSPU]; ibid, 23.VII.2013, leg. K. Makarov, 4 ex [ MSPU]; Kunashir Island, env. Yuzhno-Kurilsk, ocean shore, 15–19.VI.1990, leg. K. Makarov, 6 ex [ MSPU]; Kunashir Island, env. Yuzhno-Kurilsk, 44°01’49”N 145°51’13”E, 17–18.IX.2009, leg. I. Melnik, 5 ex [ MSPU]; Kunashir Island, Yuzhnokurilsky Cape, 44°01’14’’N 145°51’52’’E, 22.VII.2013, leg. K. Makarov, 4 ex [ MSPU]; Kunashir Island, env. Yuzhno-Kurilsk, Golovnina Bay, 44°02’29”N 145°51’50”E, 28.VIII.2009, leg. I. Melnik, 3 ex [ MSPU]; Kunashir Island, southern coast of Serebryanoe Lake, 44°03’05”N 145°49’12”E, 11.IX.2009, leg. A. Zaitsev, 1 ex [ MSPU]; Kunashir Island, shore of Mikhaylovskoye Lake, 44°18’01”N 145°58’07”E, 14.VIII.2017, leg. K. Makarov, 23 ex [ MSPU]; Kunashir Island, mouth of the Filatova River, sand-pebble beach, 26.VI.2013, leg. Yu. Sundukov, 10 ex [ FEB]; Kunashir Island, Filatova River, 16.VIII.1981, leg. O. Kabakov, 1 ex [ ZISP]; Kunashir Island, sea coast between Filatova River and Rogacheva Cape, 44°11’11”N 146°01’57”E, 10.IX.2009, leg. I. Melnik, 5 ex [ MSPU]; Kunashir Island, mouth of the Medny Creek, seaside sandy beach, 16.VIII.2017, leg. Yu. Sundukov, 26 ex [ FEB]; ibid, 27.VI.2008, leg. K. Makarov, 3 ex [ MSPU]; Kunashir Island, sea coast near Valentina Lake , 44°16’34”N 145°56’27”E, 16.VIII.2017, leg.K.Makarov,24 ex [ MSPU]; Kunashir Island, mouth of the Saratovskaya River, sandy river bank, 2.VII.2014, leg. Yu. Sundukov, 17 ex [ FEB]; ibid, 12.VII.2014, leg. Yu. Sundukov & L. Sundukova, 17 ex [ FEB]; ibid, sandy beach, 4.VII.2014, leg. Yu. Sundukov & L. Sundukova, 7 ex [ FEB]; Kunashir Island, lower reaches of the Saratovskaya River, swampy river bank in the high tide zone, 14.VII.2014, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; Kunashir Island, mouth of the stream ~ 250 m north of the mouth of the Saratovskaya River, sand and pebble beach, 16.VI.2016, leg. Yu. Sundukov, 7 ex [ FEB]; Kunashir Island, env. cordon Tyatinsky, 44°16’20”N 146°10’20’’E, 3.VII.2011, leg. I. Melnik, 1 ex [ MSPU]; Kunashir Island, mouth of the Tyatina River, pebble river bank, 21.VII.2014, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; Kunashir Island, Lovtsova Peninsula, Spokoiny Bay, sandy beach, 21.VIII.2015, leg. Yu. Sundukov, 1 ex [ FEB]; Kunashir Island, tip of Cape Lovtsov near Pico Island, sandy beach, 12.VIII.2015, leg. Yu. Sundukov & L. Sundukova, 17 ex [ FEB]; Iturup Island, Burevestnik—Iodny, stream mouths, 26.VIII.1975, leg. S. Storozhenko, 18 ex [ FEB]; Iturup Island, Medvezhyi Peninsula, Slavnaya Bay near the Slavnaya River mouth, 20.VIII.1994, leg. K. Eskov, 2 ex [ MSPU]; Urup Island, Natalia Bay, Irina Bay , 46°05’43”N, 150°09’49”E, rocks in the tidal zone, 22.VIII.2021, leg. D.R. Zhigir, 5 ex [ ZMMU]; Urup Island, Natalia Bay, Irina Bay , 46°05’45”N, 150°10’04”E, rocks in the tidal zone, 23.VIII.2021, leg. D.R. Zhigir, 1 ex [ ZMMU]; Paramushir Island, Krasheninnikov Bay, 4.VIII.1997, leg. A. Lelej & S. Storozhenko, 1 ex [ FEB]; Paramushir Island, Shelikhovo, 13.VIII.1997, leg. A. Lelej & S. Storozhenko, 1 ex [ FEB]; Paramushir Island, near Severo-Kurilsk, 10.VIII.1997, leg. Yu. Marusik, 1 ex [ FEB]. Sakhalin Island: Kuznetsova Cape, 22.VIII.1988, leg. S. Korolev, 2 ex [ MSPU]; Aniva Bay, Taranai, sea shore, 7.VII.1947, leg. E. Guryanova, 6 ex [ ZISP]; Aniva district, ca 1.8 km SE from Novikovo, Evstafiya Cape, 8.VI.1990, leg. K. Makarov, 2 ex [ MSPU]; Ptichye Lake, 16.VIII.1974, leg. M. Kuporosov, 5 ex [ ZISP]; Solovyovka, muddy seashore in the high tide zone, 7.VIII.2018, leg.Yu.Sundukov&L. Sundukova, 5ex [ FEB];Okhotskoe,mouth of the Krasnoarmeysky channel, muddy shore in the high tide zone, 1.VIII.2022, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; Yasnomorsky north of Nevelsk, mouth of the Yasnomorka River, 21.VII.2019, leg. M. Sergeev, 1 ex [ FEB]. Primorsky Territory: Vityaz Bay, Andreevka, 11.IV.1974, leg. A.B. Egorov, 1 ex [ FEB]; Khasansky district, Vityaz Bay, intertidal zone, 14.VII.1959, leg. R.A. Vasiliev, 4 ex [ ZISP]; Khasansky district, sea shore, 9.VII.1959, leg. R.A. Vasiliev, 1 ex [ ZISP]; Khasansky district, Gorshkov Bay, sandy beach at the mouth of the stream, 28.V.2022, leg. Yu. Sundukov, K. Makarov & A. Matalin, 25 ex [ FEB, MSPU]; Ryazanovka, sandy sea beach, 7.VII.1985, leg. G. Lafer, 8 ex [ FEB]; Ryazanovka, sea beach, 4.X.1998, leg. B. Kataev, 1 ex [ ZISP]; Khasansky district, coast between the mouths of the Zaklyuchennaya and Ryazanovka rivers, sandy beach, 21.V.2023, leg. Yu. Sundukov, 4 ex [ FEB]; Khasansky district, Kedrovy Cape, shore of the Amur Bay, 43°04’04’’N 131°36’17’’E, 27.V.2015, leg. A. Matalin, 5 ex [ MSPU]; Khasansky district, mouth of the Kedrovaya River, sandy beach, 43°04’47’’N 131°36’47’’E, 25.V.2015, leg. K. Makarov & A. Matalin, 16 ex [ MSPU]; Khasansky district, Melkovodnaya Bay, beach, 19.VII.1973, leg.G.Lafer, 9 ex [ ZISP];Khasansky district, mouth of the Kedrovaya River, sandy beach, 22.VIII.2018, leg.Yu. Sundukov, 15 ex [ FEB, MSPU]; Vladivostok, Shamara Bay, VII.1976, leg. G. Lafer, 8 ex [ FEB]; Vladivostok env., 28.VIII.1969, leg. O. Kryzhanovskij, 4 ex [ ZISP]; near Bolshoi Kamen, Andreeva Bay, 27.V.2023, leg. M. Sergeev, 6 ex [ FEB]; Lazovsky district, Sudzukhe, 25.VIII.1961, leg. Tryapitsyn, 1 ex [ ZISP]; Lazovsky district, Lake Chekhunenko at the mouth of the Kievka River, pebble beach, 1.VII.2022, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; Lazovsky Reserve, Petrov Bay, sandy beach, 16–19.VI.2008, leg. V. Shokhrin, 2 ex [ FEB]; Lazovsky Reserve, env. cordon Petrova, 1th Olenevod, 20.IX.1998, leg. B. Kataev, 2 ex [ ZISP]; Lazovsky district, Sokolovka Bay, seaweed discharges at the mouth of the stream, 14.VII.2002, leg. Yu. Sundukov, 7 ex [ FEB]; ibid, 28.VII.2005, leg. Yu. Sundukov, 7 ex [ FEB]; ibid, leg. K. Makarov, 1 ex [ MSPU]; Lazovsky Reserve, mouth of the Proselochnaya River, sandy river bank, 14.VII.2005, leg. Yu. Sundukov & V. Shokhrin, 1 ex [ FEB]; ibid, 12.VII.2005, leg. K. Makarov, 2 ex [ MSPU]; Olginsky district, mouth of the Avvakumovka River, sandy beach at the mouth of the river, 30.VII.2004, leg. Yu. Sundukov & L. Sundukova, 4 ex [ FEB]; ibid, 1.IX.2019, leg. Yu. Sundukov & L. Sundukova, 1 ex [ FEB]; Kamchatka: river Kireun [=Kirevna], valley of Kamchatka River, 9.VIII.1929, leg. Novograblenov, 1 ex [ ZISP].

* 92 specimens were measured, and 29 preparations made.

Redescription of the lectotype. The body is slightly convex. Length 3.9 mm, width 2.3 mm.

Head and pronotum dark bronze, elytra dark brown with faint bronze tint. Appendages of head: basal antennal segment red-brown, the rest partially blackened (one basal segment of the left antenna and three basal segments of right antenna preserved); palps and mandibles more or less uniformly brown (left maxillary palp absent); labrum dark brown. Underparts dark brown with barely noticeable greenish tint. Epipleurae of elytra and pronotum brownish yellow. Legs light brown, coxae and trochanters slightly darker, brown.

Dorsal side without punctures. Microsculpture of dorsal side of head (including clypeus and labrum), pronotum and most of elytra isodiametric, very weakly transverse around basal foveae of pronotum and on the 1 st to the 6 th inner intervals of elytra.

Standard sizes (in mm): lectotype —HW 0.90; HL 0.60; PA 0.87; PW 1.11; PB 0.78; PL 0.86; EW 1.57; EL 2.30; TL 3.9; paralectotype —HW 0.95; HL 0.60; PA 0.89; PW 1.17; PB 0.83; PL 0.91; EW 1.73; EL 2.53; TL 4.2.

Head large, moderately convex, its width with eyes 1.56 times its length. Eyes normal, moderately convex. Mandibles rather narrow, elongate, with pointed apices slightly curved inward. Labrum trapezoidal, with six setae along anterior margin. Clypeus trapezoidal, with two lateral setae, distant from anterior edge by 1/3 length of clypeus. Tooth of mentum large, broadly triangular, rounded at apex, with two small setae at base. Submentum with two large setae each side. Gula in basal third with a narrow longitudinal groove. Temples very short, ~ 1/5 eye diameter. Two supra-orbital setae, one each at the middle of eye and at its posterior margin. Frontal grooves not bifurcated, parallel, widely groove-shaped, expressed from the middle of posterior half of eye to clypeus; from posterior edge of clypeus to its setae, grooves narrow and less deep; immediately behind clypeal suture, a pore-like formation on each groove. Space between frontal groove and lateral edge of forehead convex, weakly keeled; space between anterior and posterior supra-orbital pores with three oblique wrinkles. Antennae: one basal segment of left antenna and three basal segments of right antenna preserved.

Pronotum moderately convex, distinctly cordate, weakly transverse (PW/PL = 1.28), slightly wider than head (PW/HW = 1.21), with the maximum width of 2/3 from base. Anterior margin very weakly and regularly concave, very finely edged at anterior angles. Front angles almost not protruding forward, their apices narrowly rounded. Base slightly concave in the middle, at basal foveae clearly sloping forward, without clear edging, slightly narrower than anterior margin (PB/PA = 0.92). Posterior corners almost not protruding, rectangular, slightly skewed forward, pointed at apex. Sides moderately convex and regularly rounded, slightly concave in front of posterior angles. Lateral edges narrowly bordered entire. Two lateral setae each side: one in front of the maximum width and the other in posterior angle. Midline very narrow, distinct, reaching both anterior and posterior transverse impressions, these being weak. Basal foveae relatively small, externally delimited by an indistinct carina; their short central impression arched from posterior angles to anterior edge. Area between foveae distinctly and longitudinally rugose.

Elytra oval, moderately convex, short (EL/EW = 1.34, EL/PL = 2.73, EW/PW = 1.56), with the maximum width approximately at middle. Shoulders broadly rounded, not protruding. Basal border very short, gradually turning into lateral margin, reaching the apex of 5 th stria. Lateral edge flattened, uniformly narrow all along; an outer apical angle not expressed, gradually turning into lateral margin. Striae complete, moderately deep (deeper in anterior half), finely punctured; 2 nd stria at apex clearly connected to 1 st stria; 7 th stria developed all along; apical stria moderately deep, continuous, connected to the apex of 5 th stria, and bearing two setigerous pores: one at apex, the second opposite the junction of 4 th and 5 th striae. Intervals in anterior half moderate, in posterior half slightly convex. A subscutellar striola absent. Prescutellar pore is located on top of fused 1 st and 2 nd striae. Third interval with two clearly visible discal pores lying inside 3 rd stria: anterior approximately 1/3 of base, posterior approximately 1/3 of apex. Lateral series (series umbilicate) consisting of eight setae: four in humeral group, two in the middle, and two at apex. Wings reduced to a small and narrow plate.

Legs relatively short, slender; middle tibia 1.10 times as long as middle tarsus; hind tibia 1.04 times as long as hind tarsus.

Underside of body not punctured. Metasternal process weakly and indistinctly bordered. Metepisternae short. Hind coxae with three setae. Hind trochanters each with one seta, in the middle. Abdominal sternites simple, with neither pubescence nor additional setae; apical sternite with two setae at apex.

Penis ( Fig. 4 ) thin, 5.6 times longer than thick, ventral margin almost straight, slightly curved in front of apex, lamella short and rounded.

Left paramere ( Fig. 5 ) wide, with three setae at apex: middle one 2.5 times as long as lateral ones; right paramere ( Fig. 6 ) narrow, with three setae at apex, a subapical seta absent.

Endophallus armament ( Fig. 7 ) well-developed. Dorsal plate (DP) small and without processes. Ribbon brush (RB) rounded and slightly shorter than dorsal plate. Flagellum (Fl) short, approximately twice as long as ribbon brush. The sclerotized part of the flagellar sheath (aFS) is approximately half as long as the flagellum. Basal sclerite (BS) longer than ribbon brush, rather weakly and symmetrically curved in profile. Ventral sclerite (VS) short, without spines.

Additional morphological information (based on non-type specimens).

Wing reduced down to a narrow triangular plate with visible remains of costal and subcostal veins at base and faint traces of 1–2 longitudinal veins distally, these probably corresponding to radial and medial veins.

Everted endophallus ( Figs 33–36 ) asymmetrical, with large, clearly defined, laterobasal tubercles, unpaired dorso-apical tuberosity small, all tubercles covered with spines. Two small apical tubercles in front of central sclerite complex.

In female, apical gonocoxite crescent-shaped, shorter than basal one ( Fig. 9 ). In juveniles, apex slightly expanded laterally; in beetles that have laid eggs, apex strongly worn out and devoid of its characteristic shape. Subapical setae thin, dorsal spine short, lateral spines large, usually two, rarely three (three spines usually found on only one, left or right, gonocoxite). Spermatheca almost symmetrical, short, 1.1–1.5 times as long as wide. Basal part of spermatheca conical, separated from distal part by a deep groove ( Fig. 10 ).

Variability. Body length 3.40–4.45, width 1.38–1.75 mm ( Table 1 ). Sexual dimorphism well-expressed, females larger than males in all measurements (p<0.01). Colouration from light to dark bronze, sometimes with greenish or bluish sheen ( Figs 58–77 ). Legs always lightly coloured, only knees slightly darkened. Antennae always darkened colouration from 3 rd segment; in most specimens, 2 nd segment also partially darkened. Anterior depression of pronotum well-expressed, rarely (in approximately 10% beetles) smoothed. Subscutellar striola always developed, rather long (not shorter than width of scutellum), usually with several punctures, rarely without punctures. Discal pores of elytra large; in approximately every fifth beetle they interrupt adjacent intervals. Sutural stria at apex of elytra usually connected with 2 nd stria and noticeably deepened there; less often ending up freely.

Populations from different islands of the Kuril Archipelago fail to differ in colouration, size or proportions (p<0.05).

Structure of spermatheca constant in different parts of the range. Shape of aedeagus and armament of endophallus little variable.

Diagnosis. It differs from B. petromarinum sp. nov. in the light colouration of the legs, the lightened integument, the relatively narrow elytra, the short spermatheca, the strongly extended laterobasal lobe of the endophallus (see below), and the short flagellum ( Table 2 , Figs 119–122 ).

Note. Bembidion quadriimpressum was first mentioned by V.I. Motschulsky (1850) under the name Phila quadriimpressa in his catalogue of ground beetles, indicating the authorship of E. Ménétriés. At the same time, he pointed out ( Motschulsky 1850: 14) that four specimens were known: one from Kamchatka and three from the Urup Island, Kuril Islands. Ten years later, Motschulsky published a description of this species, without indicating the number of specimens, but mentioning material from Kamchatka and the Kuril Islands: “... habite les bords de l’Océan pacifique dans la baie d’Avatcha au Kamtschatka et sur les îles Kourilles” ( Motschulsky 1860: 91). Thus, it can well be assumed that Motschulsky had several syntypes at his disposal. Subsequently, some of the specimens were lost or moved, and only one specimen is mentioned from Motschulsky’s collection in the Moscow Museum, i.e., a male labeled “ Holotypus. Kamtschatka ” ( Keleinikova 1976). The second specimen, a significantly damaged female, is kept in the collection of ZISP RAS, St. Petersburg. As lectotype, we designate the male from the ZMMU collection in accordance with the recommendations of Art. 74 ICZN. The type area, according to clause 76.2 of the ICZN, is the Kamchatka Peninsula.

Distribution. Northwestern Pacific: the coasts of the Sea of Okhotsk and the Sea of Japan from Kamchatka to the Korean Peninsula, as well as the Kuril Islands (all), Sakhalin, Moneron Island, Hokkaido and Honshu.

Habitat. Coastal areas: sandy and pebble beaches, muddy shores of the tidal zone, river mouths (Figs 123, 125, 127, 129). Beetles are found under discarded algae, boards, logs and other objects in the supra-littoral zone; more abundant near fresh waters discharging into the sea.

Figures

FIGURES 1–10. Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860): 1–7—male, lectotype, 8–10—female (Tanfiliev Island, Zorkaya Bay): 1—labels of the lectotype; 2, 8—habitus, dorsal view, 3—habitus, ventral view, 4—median lobe of aedeagus, 5—left paramere, 6—right paramere, 7—sclerite of endophallus, 9—female genitalia, 10—spermatheca. Abbreviations see in text.

FIGURES 11–24. Microsculpture: 11–17—Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860), 18–24— Bembidion (Limnaeoperyphus) petromarinum sp. nov., 11, 18—frontal furrow, 12, 19—forehead, 13, 20—disk of pronotum, 14, 21—3rd elytral interval near discal pore, 15, 22—4th elytral interval, 16, 23—first visible sternite, 17, 24—last visible sternite.

FIGURES 25–32. Median lobe of aedeagus:25–30—Bembidion (Limnaeoperyphus) petromarinum sp. nov. (25–26—holotype, Kunashir Island, Prasolov Cape, 27–28—Tanfiliev Island, Zorkaya Bay, 29–30—Khasansky district, Ostrovok Falshivy Peninsula), 31–32—Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860), from Kunashir Island, Serebryanoe Lake, 25, 27, 29, 31—right view, 26, 28, 30, 32—ventral view. Abbreviations see in text.

FIGURES 33–40. Everted endophallus: 33–36—Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860), Kunashir Island, Sennaya River, 37–40—Bembidion (Limnaeoperyphus) petromarinum sp. nov., Shikotan Island, Tserkovnaya Bay, 33, 37—ventral view, 34, 38—right view, 35, 39—left view, 36, 40—dorsal view.

FIGURES 50–57.Variation of the head and pronotum: 50–53—Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860), 54–57—Bembidion (Limnaeoperyphus) petromarinum sp. nov., 50–51, 53–57—male, 52—female, 50—Khasansky district, Gorshkov Bay, 51—Khasansky district, Kedrovaya River, 52, 53—Polonsky Island, Moryakov Bay, 54—Rogachev Island near Kunashir Island, 55—Tanfiliev Island, Zorkaya Bay, 56—Khasansky district, Ostrovok Falshivy Peninsula, 57— Kunashir Island, Prasolov Cape.

FIGURES 58–77. Variability of Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860): 58–67—male, 68– 77—female, 58, 69—Khasansky district, Gorshkov Bay, 59, 67, 73, 77—Polonsky Island, Moryakov Bay, 60, 61, 68, 71— Khasansky district, Kedrovaya River, 62, 74—Kunashir Island, Sennaya River, 63—Kunashir Island, Serebryanoe Lake, 64, 66—Shikotan Island, Tserkovnaya Bay, 65, 72, 76—Kunashir Island, Krugly Cape, 70—Tanfiliev Island, Zorkaya Bay, 75— Kunashir Island, Medny Creek.

FIGURES 98–118. Variation of the genitalia in Bembidion (Limnaeoperyphus) spp.: 98–105—Bembidion (Limnaeoperyphus) quadriimpressum (Motschulsky, 1860), 106–118—Bembidion (Limnaeoperyphus) petromarinum sp. nov., 98–113—median lobe of aedeagus; 114–118—spermatheca, 98—Kamchatka, lectotype, 99—Kunashir Island, Serebryanoe Lake, 100—Kunashir Island, Krugly Cape, 101, 102, 108, 109, 117—Polonsky Island, Moryakov Bay, 103, 110, 111—Tanfiliev Island, Zorkaya Bay, 104, 105—Khasansky district, Kedrovaya River, 106, 114, 115—Kunashir Island, Prasolov Cape, 107—Kunashir Island, Bely Utes Cape, 112—Shikotan Island, Tserkovnaya Bay, 113, 118—Khasansky district, Ostrovok Falshivy Peninsula, 116— Rogachev Island near Kunashir Island.

FIGURES 119–122. Morphometric differences between species of the Limnaeoperyphus: 119—length of body from clypeus to apex of elytra, 120—elytra length/elytra width ratio, 121—aedeagus length/width ratio, 122—aedeagus length/flagellum length ratio.