Bembidion (Limnaeoperyphus) petromarinum Sundukov et Makarov, 2024

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 : 354-365

Bembidion (Limnaeoperyphus) petromarinum Sundukov et Makarov , sp. n.

( Figures 18–30 , 37–49 , 54–57 , 78–97 , 106–118 )

Holotype ♂ with labels: “Kunashir, Cape Prasolov, south coast, rocks, 44°21’15”N 146°00’19”E, 11.VIII.2017, leg. K. Makarov & Yu. Sundukov ” [ ZISP].

Paratypes. 8♂ 2♀, Southern Kuril Islands , Tanfiliev Island, Zorkaya Bay, seaside rocky cliffs, 7.VI.2017, leg. Yu. Sundukov [ FEB, ZMMU, NHM, MSPU] ; 1♂ 2♀, Southern Kuril Islands , Polonsky Island, Moryakov Bay, 2.IX.2019, leg. Yu. Sundukov & L. Sundukova [cBK, MSPU] ; 1♂ 1♀, Southern Kuril Islands , Shikotan Island, Tserkovnaya Bay, seaside rocks, 27.V.2012, leg. Yu. Sundukov [ MSPU] ; 6♂ 2♀, ibid, 18.VIII.2022, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 5♂ 2♀, Shikotan Island , in nests Larus sp. 14.VIII.56, leg. Violovich, [ ZISP] ; 18♂ 13♀, Southern Kuril Islands , Rogachev Island near Kunashir Island, seaside rocks, 28–29.VI.2017, leg. Yu. Sundukov [ FEB, ZMMU, ZISP, MSPU, cBK] ; 1♀, Southern Kuril Islands , Kunashir Island, mouth of the Ozernaya River, 43°53’07”N 145°27’44”E, 11.VIII.2009, leg. K. Makarov [ MSPU] ; 1♂ 2♀, Southern Kuril Islands , Kunashir Island, mouth of Tyatina River, 44°16’06’’N 146°09’30’’E, 6.IX.2009, leg. I. Melnik [ MSPU] ; 1♀, Southern Kuril Islands , Kunashir Island , sea coast between Filatova River and Rogacheva Cape, 44°11’11”N 146°01’57”E, 10.IX.2009, leg. I. Melnik [ MSPU] ; 1♂, Southern Kuril Islands , Kunashir Island , mouth of the Vodopadny Stream south of Ivanovsky Cape, 43°48’20’’N 145°24’59’’E, 11.VI.2011, leg. I. Melnik [ MSPU] ; 1♂ 2♀, Southern Kuril Islands , Kunashir Island, 0.3 km south-west from mouth of the Ozernaya River, 43°52’55”N 145°27’38”E, 21.VII.2011, leg. K. Makarov [ MSPU] ; 2♂, Southern Kuril Islands , Kunashir Island, Bely Utes Cape, seaside cliffs, 11.VIII.2013, leg. Yu. Sundukov [ MSPU] ; 1♂ 2♀, Southern Kuril Islands , Kunashir Island, Prasolov Cape, south coast, rocks, 44°21’15”N 146°00’19”E, 11.VIII.2017, leg. K. Makarov & Yu. Sundukov [ ZISP, NHM, MSPU] ; 3♂ 1♀, Southern Kuril Islands , Kunashir Island, shore of Mikhaylovskoye Lake, 44°18’01”N 145°58’07”E, 14.VIII.2017, leg. K. Makarov [ ZISP, MSPU] .

72 specimens were measured (including all types), and 26 preparations made.

Type locality. RUSSIA, Kuril Archipelago: Kunashir Island, Cape Prasolov.

Material not included in the typical series. RUSSIA: Kuril Archipelago : 2♂ 2♀, Yuri Island, western bay of Shirokaya Bay , sand and pebble beach, 26.VIII.2016, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♀, ibid, 1.IX.2016, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♂ 2♀, Polonsky Island, Chasovaya Bay , seaside rocky cliffs, 30.VIII.2017, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♀, Polonsky Island, Moryakov Bay , seaside rocky cliffs, 2.IX.2017, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♀, Polonsky Island, coast near Cape Yazykovy , seaside rocky cliffs, 5.IX.2017, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 5♂ 5♀, Polonsky Island, Udobnaya Bay , seaside rocks, 6.IX.2017, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♀, Polonsky Island, Cape Severny , seaside rocky cliffs, 9.IX.2017, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♂, Shikotan Island, Tserkovnaya Bay , seaside rocks, 25.V.2012, leg. Yu. Sundukov [ FEB] ; 2♂, Kunashir Island, rocks at Cape Prasolov , 11.VIII.2017, leg. Yu. Sundukov [ FEB] ; 1♀, Kunashir Island, mouth of the Tyatina River , river bank with large boulders, 21.VII.2014, leg. Yu. Sundukov & L. Sundukova [ FEB] ; 1♂, Kunashir Island, Cape Bely Utes , seaside cliffs, 11.VIII.2013, leg. Yu. Sundukov [ FEB]. Primorsky Territory : 1♂ 1♀, Khasansky district, Ostrovok Falshivy Peninsula , seaside rocks, 23.V.2022, leg. Yu. Sundukov [ FEB] .

Description of the holotype. Body slightly convex. Length 3.80 mm, width 1.55 mm.

Dorsal side black with a distinct bluish or bluish-greenish tint ( Fig. 41 ). Appendages of head: 2.5 basal antennal segments red-brown, the rest partially blackened; palps red-brown with slightly blackened apical segments; mandibles red-brown with blackened apices and ribs at base; labrum red-brown ( Figs 41 , 57 ). Ventral side black or brownish black ( Fig. 42 ). Epipleurae of elytra barely paler than body venter. Legs red-brown with pale brown tibiae and blackened trochanters, femora and tarsi; most of hind coxae black.

Dorsal side without punctures. Microsculpture of dorsal side of head (including clypeus and labrum) and along lateral margins of pronotum distinct, but not coarse, and consisting of more or less isodiametric meshes ( Figs 18, 19 ); on pronotal disc ( Fig. 20 ) and elytra (completely or 6–7 internal intervals) weakly transverse ( Figs 21, 22 ). Microsculpture on abdominal sternites is transverse ( Fig. 23 ), especially coarse on anal sternite ( Fig. 24 ).

Standard sizes (in mm): HW 0.8; HL 0.56; PA 0.75; PW 1.04; PB 0.73; PL 0.80; EW 1.54; EL 2.23; TL 3.80.

Head quite large, moderately convex, its width with eyes is 1.42 times its length. Eyes moderately convex ( Figs 41 , 57 ). Mandibles rather narrow, elongate, with pointed apices slightly curved inward. Labrum trapezoidal, with six setae along anterior margin. Clypeus trapezoidal, with two lateral setae removed from anterior edge by 1/3 length of clypeus ( Fig. 57 ). Tooth of mentum large, broadly triangular, rounded at apex, with two small setae at base. Submentum with two large setae each side, outer setae shorter. Gula in basal part with a short longitudinal depression. Temples short, slightly shorter than 1/4 longitudinal diameter of eye. Two supraorbital setae: one at the middle of eye and the other at its posterior margin. Frontal grooves parallel, widely groove-shaped, reaching the middle of posterior half of eye; grooves narrow and less deep on clypeus ( Fig. 57 ). Anterior tentorial pits quite deep. Space between frontal groove and lateral edge of forehead convex, weakly keeled; space between anterior and posterior supra-orbital pores with oblique wrinkles ( Fig. 57 ). Antennae long, reaching 2/5 length of elytra ( Fig. 41 ).

Pronotum moderately convex, distinctly cordate, weakly transverse (PW/PL = 1.30), slightly wider than head (PW/HW = 1.30), with the maximum width of 3/5 of base ( Figs 41 , 57 ). Anterior margin slightly and regularly concave, very finely edged at anterior angles. Anterior angles almost not protruding forward, their apices narrowly rounded. Base slightly concave in the middle, slightly sloping forward at basal foveae, without clear edging, slightly narrower than anterior margin (PB/PA = 0.97). Posterior angles not protruding to sides, rectangular, slightly skewed forward, their apices pointed. Sides moderately convex and regularly rounded, slightly concave in front of posterior angles. Lateral edges narrowly bordered all along. Two lateral setae each side: one in front of the maximum width and the other in posterior angles. Midline very narrow, reaching both anterior and posterior transverse impressions, these being weak. Basal foveae relatively small, delimited externally by a weak carina; their short central impression arcuate and concave from posterior angles to anterior edge. Area between foveae distinctly longitudinally rugose; weaker transverse wrinkles present along lateral margins in basal half of pronotum and longitudinal wrinkles present on disc along anterior depression.

Elytra oval ( Fig. 41 ), moderately convex, rather short (EL/EW = 1.43, EL/PL = 2.78, EW/PW = 1.50), 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 and uniformly narrow all along; an outer apical angle not expressed, gradually turning into lateral margin. Striae complete, moderately deep, with shallow punctures; 2 nd stria at apex widely depressed; 7 th stria well-developed all along; apical striole moderately deep, continuous, connected to 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 moderately convex. Subscutellar striole short, marked by 3 points on right elytron; located inside 1 st interval. Prescutellar pore located on top of fused 1 st and 2 nd striae. Third interval with two clearly visible discal pores lying at 3 rd stria: anterior approximately 1/3 of base, posterior one approximately 1/3 of apex. Lateral series (series umbilicate) consisting of seven setae: four in the humeral group, one in the middle, and two at apex.

Wing in the form of a narrow triangular plate, 9–10 times shorter than elytra. Bases of costal and subcostal veins and traces of longitudinal veins visible on wing.

Legs relatively short, slender; middle tibia 1.21 times as long as middle tarsus; hind tibia 1.16 times as long as hind tarsus.

Underside of body not punctured. Metasternal process bordered by a thin line smoothed anteriorly. Metepisternum shortened: 1.6 times longer along outer margin than width along anterior margin. Hind coxae with three setae. Metatrochanters each with one seta, in the middle. Abdominal sternites simple, with neither pubescence nor additional setae; apical sternite with two setae at apex.

Penis ( Figs 43, 45 ) thin and long, its ventral edge very slightly curved, lamella being short and widely rounded.

Left paramere ( Fig. 46 ) wide, with three setae at apex: middle one very long, 3–3.5 times as long as both lateral ones; right paramere ( Fig. 44 ) narrow, with three setae at apex (middle one very long, 3–3.5 times as long as lateral ones) and one short subapical seta on ventral side.

Endophallus armament ( Figs 25, 26 ) typical of Ocydromus . Dorsal plate (DP) short, without processes. Ribbon brush (RB) compact, subequal in length to dorsal plate. A flagellum (Fl) originating from left part of the central complex of sclerites, its length being subequal to diameter of penis tube. A sclerotized part of flagellar sheath (aFS) visible near apex of flagellum. Basal sclerite (BS) slightly shorter than ribbon brush, being strongly and asymmetrically curved in profile. Ventral sclerite (VS) small, with an oblong field of spines next to it.

Everted endophallus ( Figs 37–40 ) asymmetrical, strongly curved to the left. Both left and right laterobasal lobes small, round, regularly beset with spines ( Fig. 37 ). Unpaired dorso-apical lobe in front of the complex of central sclerites gentle, covered with similar spines ( Fig. 39 ). Two small, conical, apical lobes projecting in front of the complex of central sclerites.

Female paratypes. Antennae shorter, up to 1/3 elytra; apical sternite of abdomen with four setae at apex. Apical gonocoxite ( Fig. 48 ) crescent-shaped, slightly shorter than basal one. In young individuals, apex flattened, with a weak rib-like thickening. In addition to a pair of thin subapical setae, bearing a thick dorsal spine and three lateral ones. Spermatheca ( Fig. 49 ) large, fusiform, 2.7–3.5 times as long as wide. Distal part separated by a shallow groove with gentle edges.

Variability. Body length 3.80–4.68, width 1.56–1.93 mm ( Table 1 ). Sexual dimorphism less pronounced than in B. quadriimpressum , males are smaller than females, but significant differences (p<0.05) found only for total body length, was well as length and width of elytra. Colouration dark bronze, with a more or less strong greenish or bluish sheen ( Figs 78–97 ). 2 nd antennomere always light. Gula in basal part with a rather deep longitudinal depression, larger in males (1/3–1/4 gula) and smaller in females (sometimes reduced to a point).

Notch in front of posterior angles of pronotum often very weak; edges of pronotum sometimes narrowed to straight line towards posterior angles.

FIGURES 123–130. Typical habitats of Limnaeoperyphus species: 123, 125, 127, 129— B. quadriimpressum , 124, 126, 128, 130— B. petromarinum sp. nov., 123—Khasansky district, Gorshkov Bay, 124—Khasansky district, Ostrovok Falshivy Peninsula, 125—Kunashir Island, seashore norther Krugly Cape, 126—Kunashir Island, Prasolov Cape, 127—Yuri Island, Shirokaya Bay, 128—Yuri Island, Shiroky Cape, 129—Polonsky Island, Moryakov Bay, 130—Polonsky Island, Udobnaya Bay.

Legs relatively short, slender; mesotibia 1.22–1.29 (M 1.25) times longer than mesotarsus; hind tibia 1.24–1.29 (M 1.26) times longer than hind tarsus. Femora darkened to a greater or lesser extent, but tibiae not darker than the rest of femora.

Intervals moderately or slightly convex. Discal pores small, placed close to 3 rd stria, and never interrupting the striae. 2 nd stria of elytra deepened in front of apex and ending freely in half of the cases. Prescutellar striola quite long, with several deep points and, less often (in less than 20% cases), shortened.

No statistically significant differences in sizes and proportions are found between populations from the islands of Kunashir, Shikotan, Polonsky, Tanfiliev and Yuri.

No interpopulation differences are found in the structure of the spermatheca ( Figs 114–118 ), but in young individuals it is usually shorter and wider ( Fig. 117 ). The shape of the lamella of the penis is a little variable. The armament of the endophallus is quite constant; there are only minor variations in the length of the dorsal sclerite, but these are found in different parts of the distribution range.

Etymology. The name of the new species comes from the Latinized form of the Greek noun “ πέτρα ” (stone) and the Latin adjective “ marina ” (sea). It reflects the specific habitat of the new species—rocks, cliffs and stones on the seashore.

Distribution. Northwestern Pacific: known from the Kuril Archipelago (Kunashir, Shikotan, Polonsky, Yuri, Tanfiliev, and Rogacheva islands), southern Primorsky Territory, southern Khabarovsk Territory (Sikhote-Alin, Botchinsky Nature Reserve; personal communication from R.Yu. Dudko, Novosibirsk, Russia).

Habitat. Sheer cliffs of the supra-littoral zone (Figs 124, 126, 128, 130). As a rule, during the daytime, beetles are found in rock cracks or rocky-gravel-soil deposits in places where fresh water oozes out (small streams in cracks and small gorges, water streaks on rock slabs), staying close to or directly at the edge of a water stream. Sometimes they are found on large-block or large-pebble seashores, where they have possibly ended up by accident (falling, being washed away by the waves).

Differential diagnosis. The new species differs from Bembidion quadriimpressum in its relatively wide elytra ( Fig. 120 ; EW/PW ratio 1.47–1.61, vs 1.38–1.53 in B. quadriimpressum ) with a distinct bluish or bluish-greenish tint; darkened, sometimes almost black hind coxae, trochanters, partly dark femora, and the structure of the genitalia: the female has a large, elongated spermatheca, an apical gonocoxite with three lateral spines; in the male, the medial lobe is wider ( Fig. 121 ), the flagellum is longer ( Fig. 122 and Table 2 ), the ribbon brush and ventral sclerite are more strongly developed, the laterobasal lobes of the endophallus are slightly protruding. Bembidion petromarinum sp. nov. is on average larger than B. quadriimpressum ( Fig. 119 , Table 1 ).

Figures

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 41–49. Bembidion (Limnaeoperyphus) petromarinum sp. nov. (41, 43–49—Kunashir Island, Prasolov Cape, 42— Tanfiliev Island, Zorkaya Bay): 41–46—male, holotype, 47–49—female, paratype, 41, 47—habitus, dorsal view, 42—habitus, ventral view, 43—median lobe of aedeagus, right view, 44—right paramere, 45—median lobe of aedeagus, ventral view, 46— left paramere, 48—female genitalia, 49—spermatheca.

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 78–97. Variability of Bembidion (Limnaeoperyphus) petromarinum sp. nov.: 78–91—male, 92–97—female, 78, 80, 93, 94—Rogachev Island near Kunashir Island, 81, 82, 83, 84—Tanfiliev Island, Zorkaya Bay, 85, 95, 96—Polonsky Island, Moryakov Bay, 86, 87—Shikotan Island, Tserkovnaya Bay, 88, 89—Kunashir Island, Bely Utes Cape, 90, 97—Kunashir Island, Prasolov Cape, 91, 92—Khasansky district, Ostrovok Falshivy Peninsula.

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.