Heterocerus heydeni Kuwert, 1890

Litovkin, Stanislav V., Sazhnev, Alexey S. & Jr, Fedor Čiampor, 2019, Validation of Heterocerus heydeni Kuwert, 1890 based on morphology and DNA barcoding, with notes on the problems of classification of the Heteroceridae (Coleoptera), Zootaxa 4614 (1), pp. 160-172 : 163-169

Heterocerus heydeni Kuwert, 1890 stat. resurr.

( Figs. 2–4 , 6–16 , 18–20 , 24 )

Heterocerus hauseri Kuwert, 1893 ? syn.n.

Material examined. 1583 specimens. KAZAKHSTAN. Akmola Oblast: Esil environs, near Ishim River, 51.95°N 66.32°E, at light, 25 May 2005, leg. A. Shapovalov, 1 specimen ( CL); Aktobe Oblast: 25 km SW Emba, Peski Kumzhargan , 48.64408°N 57.91611°E, 25 Apr. 2012, leg. A. Ivanov, 4 specimens ( IBIW) ; 3.3 km NE Khlebodarovka , 50.5340°N 56.9579°E, 207 m, hills, steppe, at light, 21–22 Apr. 2013, leg. S. Litovkin, 2 specimens ( CL) ; 24 km WSW Yrgyz , 48.58361°N 60.90620°E, 167 m, at light, 28–29 May 2017, leg. S. Litovkin, 6 specimens ( CL) ; Almaty Oblast: Sarkand Distr., 15 km N Lepsy, SE shore of Balkhash Lake , 46.359741°N 78.878415°E, sands, at light, 14 May 2014, leg. S. Kolov, 2 specimens ( CL) ; 26 km SE Koktal, Ili Riv. , 43.96364°N 79.60528°E, 506 m, 01 July 2015, leg. S. Korb, 1 specimen ( IBIW) ; E Kazakhstan , 43.24294°N 78.88006°E, at light, 25 May 2016, leg. A. Ivanov, 1 specimen ( IBIW) ; SE Kazakhstan, 8 km N Kanshengel, Peski Taukum , 44.41222°N 75.55675°E, 402 m, 02 June 2016, leg. A. Ivanov, 1 specimen ( IBIW) ; Atyrau Oblast: Atyrau ( Guryev ), 02 Apr. 1911, leg. P. Shmidt, 1 specimen ( ZIN) ; Kostanay Oblast: Aksuat Lake , 250 km S of Kostanay, [51.5°N 64.5°E], 19 July 1947, leg. For- mozov, 1 specimen ( ZIN) ; Kyzylorda Oblast: Baygekum near Dzhulek , 04 June 1907, leg. D. Glazunov, 4 speci- mens ( ZIN) ; Dzhulek [44.29°N 66.43°E], Orenburg–Tashkent railway, Syrdarja , 01 July 1910, leg. I. Kozhantshi- kov, 1 male ( ZIN) ( Fig. 1 ) ; same locality data, 14 July 1910, leg. I. Kozhantshikov, 4 specimens ( ZIN) ; near of Kyzylorda ( Perovsk ), 19 May 1916, leg. N. Pulikovskaya, 1 specimen ( ZIN) ; 42 km SE Aralsk , 46.45306°N, 61.91000°E, 26 Apr. 2011, leg. A. Shapovalov, 1 specimen ( CL) ; 15 km SE Kyzylorda , 44.71183°N 65.75561°E, 127 m, at light, 30 Apr. 2012, leg. A. Ivanov, 2 specimens ( IBIW) ; 2.7 km NE Maylibash , 45.8409°N 62.6555°E, 79 m, desert, at light, 24–25 Apr. 2013, leg. S. Litovkin, 1 specimen ( CL) ; Karatau Mts., 18.5 km ENE Birlik, Akuyik River valley , 43.937°N 67.680°E, 380–400 m, mountain river valley, at light, 29 Apr.–02 May 2013, leg. S. Litovkin, 76 specimens ( CL, IBIW) ; 21 km NW Aralsk , 46.9638°N 61.4853°E, 79 m, sand-soil desert, at light, 07–08 May 2013, leg. S. Litovkin, 6 specimens ( CL) ; same locality data, at light, 01–03 May 2015, leg. S. Litovkin, 8 specimens ( CL) ; 67 km N Chiili , 44.77786°N 66.96811°E, 21 Apr. 2014, leg. A. Ivanov, 5 specimens ( IBIW) ; 52 km SSE Aralsk , [46.36°N 61.95°E], at light, 20–21 Apr. 2015, leg. A. Prosvirov, 7 specimens ( IBIW) ; 30 km SE Kazaly , [45.54°N 62.35°E], at light, 21–22 Apr. 2015, leg. A. Prosvirov, 9 specimens ( IBIW) ; 42 km NE Kyzylorda, Tartogay , [44.4°N 66.2°E], 09–11 May 2015, leg. A. Prosvirov, 3 specimens ( IBIW) ; 4.3 km SE Tartogay, 44.41040°N 66.27504°E, 145 m, tugay woodland, near Syrdarja River , at light, 13–14 June 2015, leg. S. Litovkin, 12 specimens ( CL) ; same locality data, at light, 23–24 July 2017, leg. S. Litovkin, 4 specimens ( CL) ; 42 km SSE Aralsk , 46.44609°N 61.89872°E, 56 m, sand-soil desert, at light, 25–26 July 2017, leg. S. Litovkin, 1 specimen ( CBRB) ; Mangystau Oblast: Ustyurt Nature Reserve, cordon Kenderli , 42.95742°N 54.69278°E, 19 May 2011, leg. A. Ivanov, 1 specimen ( IBIW) ; Mangystau Reg. , 9 km E Shetpe, 44.15078°N 52.26886°E, 04–06 June 2017, leg. D. Po- tanin and A. Potanina, 2 specimens ( IBIW) ; Karakiyansky Reg. , 55 km NE Senek, 43.07639°N 53.92858°E, 07–09 June 2017, leg. D. Potanin and A. Potanina, 1 specimen ( IBIW) ; Karakiyansky Reg. , 5 km NW Senek, 43.39089°N 53.33494°E, 09–11 June 2017, leg. D. Potanin and A. Potanina, 2 specimens ( IBIW) ; Tupkaragan Reg., Taushik , 44.40786°N 51.07508°E, 12–13 June 2017, leg. D. Potanin and A. Potanina, 1 specimen ( IBIW) ; Turkestan Oblast (former Southern Kazakhstan Oblast): 60 km ESE Turkestan, 14 km SSW Mynbulak , [43.103°N 68.936°E], 23–24 Apr. 2015, leg. A. Prosvirov, 2 specimens ( IBIW) ; Karatau Mts. , 11 km NW Aksumbe, 44.51456°N 67.42278°E, 376 m, 16 May 2015, leg. A. Ivanov, 2 specimens ( IBIW) ; 7.3 km NE Baltakol, 43.15872°N 67.84753°E, ~ 180 m, sandy shore of Syrdarja River , 08 June 2016, leg. S. Litovkin, 1 specimen ( CL) ; 7.4 km NE Baltakol, 43.15871°N 67.85095°E, ~ 180 m, near Syrdarja River , at light, 08–10 June 2016, leg. S. Litovkin, 50 specimens ( CL, IBIW, CBRB) ; 7.3 km NE Baltakol, 43.15673°N 67.84992°E, ~ 180 m, tugay woodland, near Syrdarja River , at light, 09–10 June 2016, leg. S. Litovkin, 5 specimens ( CL) ; 9 km NE Baltakol, 43.17198°N 67.86345°E, ~ 180 m, sandy shore of Syrdarja River , 10 June 2016, leg. S. Litovkin, 1 specimen ( CL) ; 3 km W Baltakol, N border of Kyzylkum Desert , 43.11790°N 67.71356°E, 181 m, at light, 13–14 June 2016, leg. S. Litovkin, 9 specimens ( CL) ; 6 km NE Baltakol , 43.16040°N 67.83648°E, tugay, at light, 22–23 July 2017, leg. S. Litovkin, 55 specimens ( CL) ; Zhambyl Oblast: Mujunkum [= Moiynkum], identification labels “ Heterocerus parallelus Reitter det.”, “ Heterocerus parallelus pale form”, July 1907, leg. E. Fischer, 2 specimens ( ZIN) ; Taraz ( Aulie-Ata ), identification label “ Heterocerus hauseri ”, leg. Willberg, 4 specimens. ( ZIN) ; Moiynkum Distr., Aksuyek , [44.620°N 74.511°E], 07–11 Jule 1993, leg. A. Tilly, 1 specimen ( CL) ; 42 km NE Karatau, Akkol , [43.443°N 70.787°E], 02–03 May 2015, leg. A. Prosvirov, 4 specimens ( IBIW) ; vicinity of Kyzylkol Lake , 43.77602°N 69.51012°E, 335 m, saline soil, at light, 04–07 May 2015, leg. S. Litovkin, 882 specimens ( CL, IBIW) ; same locality data, at light, 22–23 May 2015, leg. S. Litovkin, 48 specimens ( CL) ; same locality data, at light, 04–06 June 2015, leg. S. Litovkin, 24 specimens ( CL) ; same locality, but 43.77°N 69.50°E, at light, 31 May–01 June 2017, leg. S. Litovkin, 2 specimens ( CL) ; along N bank of Kyzylkol Lake , 43.75995°N 69.50293°E, 330 m, sand, gravel, algae, 05–06 May, leg. S. Litovkin, 4 specimens ( CL) ; same locality data, 04 June 2015, leg. S. Litovkin, 10 specimens ( CL) ; Peski Moiynkum (= Moiynkum Desert ), 44.29262°N 70.16508°E, 315 m, sandy desert, at light, 09–11 May 2015, leg. S. Litovkin, 9 specimens ( CL) ; same locality data, but 44.29290°N 70.17608°E, 310 m, at light, 31 May–02 June 2015, leg. S. Litovkin, 35 specimens ( CL) ; 7 km NE Kumkent , 43.81614°N 69.69615°E, 300 m, saline soil, at light, 13–14 May 2015, leg. S. Litovkin, 160 specimens ( CL, IBIW) ; W bank of Akkol Lake , 43.40997°N 70.62446°E, 405 m, at light, 17–18 May 2015, leg. S. Litovkin, 37 specimens ( CL) ; vicinity of Ashchykol Lake (hypersaline lake), 43.53851°N 70.63009°E, 378 m, at light, 18–19 May 2015, leg. S. Litovkin, 4 specimens ( CL) ; same locality data, at light, 02–04 June 2015, leg. S. Litovkin, 7 specimens ( CL) ; Karatau Mts. , 13 km ENE Bayzhansay, 43.21416°N 70.06335°E, 930 m, mountain stream valley, at light, 28–30 May 2015, leg. S. Litovkin, 1 specimen ( CL) . RUSSIA. Astrakhan Oblast: Astrakhan Nature Reserve, to UV light, 26 June / 17 July 1956, leg. Breev, 1 specimen ( ZIN) ; Krasnoyarsk Distr., 2 km E Dosang , at UV light, 09–12 June 2005, leg. A. Tilly, 1 specimen ( CL) ; Krasnoyarsk Distr., Dosang environs, coast of Akhtuba River , 06 May 2009, leg. Ya. Kovalenko, 1 specimen ( IBIW) ; Krasnoyarsk Distr., 14 km NE Dosang, dune Tuvayak , 01 May 2011, leg. A. Prosvirov, 2 specimens ( IBIW) ; Liman Distr., Yango Asker , at light, 30 June 2010, leg. A. Prosvirov, 1 specimen ( IBIW) ; Liman Distr., Stantsiya Basinskaya , at light, 03 July 2010, leg. A. Prosvirov, 1 specimen ( IBIW) ; Crimea: Donuzlav Bay, Krasnoyarskoe , [45.50°N 33.26°E] 24 July 2016, leg. V. Shaporinsky, 4 specimens ( IBIW) ; Dagestan: Kizlar Distr. , vicinity of Bryansk, 44.3292°N 46.9797°E, 10 June 2009, 1 specimen ( IBIW) ; Makhachkala , 20 June 2009, leg. E. Il’ina, 1 specimen ( IBIW) ; Tarumovsky Distr., Staryy Biryuzak, Kuma Riv. , 04 June 2015, leg. E. Il’ina, 3 specimens ( IBIW) ; Kalmykia: 25 km SE from Elista , to light at night, 24 July 1986, leg. Makarova, 4 specimens ( MSPU) ; Orenburg Oblast: Western part of Orenburg Oblast, 3 km N Kurlin, 10–11 June 2004, leg. A.Tilly, 1 specimen ( CL) ; Guberlya environs, 51°16’47’’N 58°09’37’’E, at light, 17 May 2005, leg. A. Shapovalov, 1 specimen ; Gaysky Distr., Guberlinsky sovkhoz, Khmelyovka , [51.245°N 57.868°E], at light, 07–08 May 2012, leg. A. Ukrainsky, 5 specimens ( IBIW) ; Akbulak Distr., 18.5 km SE Akbulak, 50.93025°N 55.89042°E, near Karabutak River , at light, 15–16 June 2015, leg. S. Litovkin, 4 specimens ( CL) ; Saratov Oblast: Krasnokutsky Distr. , vicinity of D’yakovka, [50.731°N 46.769°E], June 2004, leg. A. Ukrainsky, 3 specimens ( IBIW) ; Volgograd Oblast: Palassovsky Distr., Elton Settlm. environs, at UV light, 21–28 May 2006, leg. A.V. Matalin, 4 specimens ( IBIW) ; same locality data, at UV light, 27–28 June 2006, leg. A.V. Matalin, 2 specimens ( IBIW) . TURKMENISTAN. Ahal Velayat: South Turkmenia, Tedzhen environs. 17–24 Apr. 1985, leg. A. Tilly and V. Nikulin, 1 specimen ( CL) ; LebapVelayat: Farap , 01 June 1905, leg. G.G. Sumakov, 3 specimens ( ZIN) ; Birata ( Darganata ), 03 July 1910, leg. E. Fisher, 1 specimen ( ZIN) ; Repetek , 10 Mar. 1982, leg. Krivokhatsky, 1 specimen ( ZIN) ; Mary Velayat: Ýolöten , 07 June 1926, leg. V. Kizeritsky, 1 specimen ( ZIN) ; Merv , 17 May 1976, leg. G. Medvedev, 1specimen ( ZIN) ; UZBEKISTAN. Khorezm Oblast: Khiva , 03 July 1910, leg. E. Fisher, 1 specimen ( ZIN) ; Qashqadaryo Oblast: Qamashi on NO by Guzar , 12 June 1932, leg. Pogg, 1 specimen ( ZIN) ; Qamashi , 22 July 1932, leg. V. Gussakov, 1 specimen ( ZIN) ; Mirishkor Reg., Urta-Bulak deposit environs, at light, 14 Apr. 2014, leg. Ya. Kovalenko, 1 specimen ( CK) .

Additional material without detailed label data from ZIN. A several dozens specimens. KAZAKHSTAN. Aktobe Oblast: Shalkar (Chankara), [47.8°N 59.6°E]; Temir (Kok-Dzhizha), [49.1°N 57.1°E]; Atyrau Oblast: In- der Lake, [48.5°N 51.8°E]; Kyzylorda Oblast: Kazalinsk, [45.8°N 62.1°E]; Baygekum, [44.3°N 66.5°E]; Western Kazakhstan Oblast: Khan Ordasy (Khanskaya Stavka), [48.8°N 47.4°E]; Uralsk, [51.2°N 51.4°E]. RUSSIA. Volgograd Oblast: Sarepta, [48.5°N 44.6°E].

Redescription. Body length from the apex of labrum to the apex of elytra 3.6–5.2 mm, males and females do not differ.

Ground colour yellow; a combination of black-brown, almost black blotches on the pronotum, elytra and abdomen form a characteristic pattern. Head and mandibles black-brown. Antennae vary from completely yellow to brownish on the club. Pronotum black-brown with a more or less wide yellow edging on the sides and narrow yellow stripe along the front edge; there with often a reddish or yellow basomedial spot, its size females is usually larger than in males, additionally it can stretch into a poorly distinguishable middle line in females ( Fig. 11 ). Scutellum black-brown. Yellow pattern of each elytron normally ( Fig. 4 ) forms edging, S-shaped basal mark, M-shaped medial mark, apical spot and suture stripe. Basal marks close in along the suture forming a solid yellow field; with a dark space between them like in H. flexuosus for a few specimens. Dark pattern of elytra ranges from black-brown to brownish almost blending with the yellow background, often reduced in basal third (e.g. Figs. 6, 11 ); only in some specimens the dark pattern of elytra dominates over the light pattern ( Figs. 9, 13 ). Prosternum black-brown with yellow episternae and yellowish front margin or mostly yellow. Mesepisternum and metepisternum black-brown. Abdomen black-brown with yellow pattern; only the rear corners of ventrites and part of hypopygidium are yellow in males ( Fig. 15 ), while females possess a more or less wide yellow edge ( Fig. 16 ). Legs, except dark metacoxae, yellow; front legs with brownish knees and margins of tibiae.

Body moderately elongated, compressed dorsoventrally.

Head possess structure typical for the family; relatively wider in large and medium males compared with females; covered with dense, whitish hairs including labrum. Clypeus transversal, with front edge widely emarginate; gently sloping towards the labrum in females and smaller males, forming a rung over it in larger males; with a pair of tubercles on the sides of the medial line in larger males (hardly visible under the hairs) ( Fig. 14 ). Structure of mandibles of males and females slightly different; mandibles of larger males are more elongate, their dorsal arista possesses a denticle pointed upwards or inwards. Antennae 11-segmented with 7-segmented club.

Pronotum not bordered basally; slightly narrower or comparable in width with base of elytra in females and smaller males, in medium and large males slightly wider than elytral base ( Figs. 6–7 ). Hairs on disc dense, short, pressed, long and standing on the sides; whitish and yellowish. Puncturation dense, fine and matted.

Elytra moderately wide, k ≥0.60; covered with different lengths of hairs variously raised; long hair length is comparable with the length of first spines of mesotibiae; hairs slightly longer along sides and near the elytral apex; yellowish hairs alternate with more rare dark ones. Puncturation double, dense, matted; punctures slightly depressed; the diameters of large and small punctures vary in different degrees from different specimens.

Abdomen ( Figs. 15–16 ) covered with elevated yellowish hairs of different length; punctures with rasp-like edges; microsculpture represented by transversal creases. Postmetacoxal lines absent. Stridulatory ridge (plectrum) in males with few distinct striae in the basal third, smooth or with small creases in females.

Legs of structure typical for family, covered with yellowish or whitish hairs.

Aedeagus ( Figs. 18–20 ) vaginate, characteristic wedge-shaped with truncate top, length 0.77–0.86 mm, tegmen surface with nested penis oriented ventrally. Lateral lobes of phallobase cuneiformly narrowed, continued into narrow lateral walls in the apical part of the tegmen ( Fig. 18 ). Apical edge of phallobase {-shaped (bow-shaped), often with very small medial notch ( Figs. 19–20 ). Sclerotization of dorsal surface of tegmen usually uniform ( Fig. 20 ). Penis elongate, almost parallel-sided, narrowing in the basal third. Penis apex is asymmetric, comes forward of the endophallus loop in a shape of short, rounded lobe. Endophallus rather long, carrying a dense brush of large teeth along its length.

Judging by the general structure and orientation of the aedeagus, the morphology of the male clypeus and pro- notum, and the elytral pattern, H. heydeni belongs to the flexuosus group (sensu Charpentier 1965).

Comparison. Characteristic for H. heydeni : the apex of the tegmen is unsplit, characteristic bow-shaped form; legs are completely yellow, with the exception of small darkening on the front knees and tibiae; elytral pattern characteristic of flexuosus group of species, with yellow pattern predominant or equal in area to dark pattern.

H. heydeni and H. flexuosus are very similar externally ( Figs. 4–5 ); they possess similar pattern and pubescence of elytra; they lack distinct sexual dimorphism; their aedeagi are similar in shape and structure. H. flexuosus is different in that it has dark body coloration; black-brown, almost black pattern predominate on the pronotum and elytra; bright pattern of elytra is either yellow or ochre, often reduced to a different degree; only some specimens (from 851) light elytron pattern prevails over dark (e.g. Fig. 17 ); scutellum is completely surrounded by a dark field; legs from black-brown to brownish, but not yellow; the elytral puncturing is double, large punctures usually coarser than small; aedeagus ( Figs. 21–23 ) slightly larger; lateral walls in tegmen apical part are wide ( Fig. 21 ); apical edge of phallobase rounded, with a medial short split (seen as a dark, elongate umbilicus at low magnification) ( Figs. 22–23 ); sclerotization of dorsal side of tegmen uneven ( Fig. 23 ); penis fusiform; penis apex protrudes significantly forward of the endophallus loop, looks like a tongue-like lobe; endophallus relatively short; teeth of endophallus are significantly smaller in size. Dark specimens of H. heydeni are similar in appearance with H. flexuosus ; their legs remain yellow.

H. heydeni and H. parallelus are similar in coloration of body and legs. H. parallelus is larger; has elongate elytra with k <0.60; elytra with short and dense pubescence; bright pattern of elytra often greenish-yellow; sexual dimorphism is distinct; hypermandibular males are common; phallobase apex is two-lobed.

Biology. Poorly studied. A few specimens were collected in sandy-gravel substrate on the shore of saline lake Kyzylkol ( Kazakhstan) together with mass H. parallelus . Two specimens were collected on sandy shore of Syrdarja River ( Kazakhstan) together with Augyles cf. flavidus (Rossi, 1794) and H. fenestratus (Thunberg, 1784) . Prevalent part of material is attracted to artificial light. Attraction of beetles to light was observed in saline habitats, sandy and sand-saline deserts, riparian forests (tugay forest), in stream valleys at elevations up to 930 m a.s.l., in steppes and once in a steppe forest plantation. Beetles were observed at distances of up to 12 km from permanent large water bodies. The following species were attracted to light together with H. heydeni : H. parallelus , H. flexuosus , H. fenestratus , H. mus Charpentier, 1965 , H. obsoletus Curtis, 1828 , H. fusculus Kiesenwetter, 1843 , Augyles . cf. flavidus, A. turanicus (Reitter, 1887) , A. dilutissimus (Reitter, 1887) . It is one of the most common species in southern Kazakhstan.

Distribution. Dzhizak and Alai ( Zaitzev 1908). Former Zakaspiyskaya, Samarkand and Fergana oblasts (Ja- cobson 1913), presently—territories of Uzbekistan, Turkmenistan and Kyrgyzstan (catalog data need clarification), Tadzhikistan ( Hebauer 1991);? SW Iran ( Skalický 1998) (material study required). South of the European part of Russia east to the Ural Mountains, Crimean Peninsula, Kazakhstan excluding the north-east, Turkmenistan, Uzbekistan ( Fig. 24 ).

Figures

FIGURE 1. Maximum likelihood tree of Heteroceridae samples. Colour of branches: blue—Heterocerus, orange—Tropicus, green—Augyles, black—undetermined Heteroceridae. New samples in red, number next to nodes—bootstrap support (only if> 0.50). Samples with retained generic and species names as in BOLD data systems database.

FIGURE 24. Distribution of Heterocerus heydeni. Circles—studied material: red circles—records only H. heydeni; orange circles—records H. heydeni and H. flexuosus; square—type locality (Dzhizak, Uzbekistan).

FIGURES 2–5. Heterocerus spp. 2—male H. heydeni from collection ZIN (photo by A. Kovalev); 3—labels of this specimen: handwriting by F.A. Zaitsev (ink), handwriting by R. Charpentier (pencil); 4—H. heydeni general view, male (South Kazakhstan); 5—H. flexuosus general view, male (South Kazakhstan). Scale bar = 1 mm.

FIGURES 6–17. Variability and sexual dimorphism of Heterocerus spp. 6–9—H. heydeni, males (South Kazakhstan); 10– 13—H. heydeni, females (South Kazakhstan); 14—H. heydeni, tubercles on clypeus of male, hairs removed; 15—H. heydeni, male abdomen; 16—H. heydeni, female abdomen; 17—H. flexuosus, female with extremely light elytra (Crimea). Scale bars = 1 mm (Figs. 6–13, 15–17) and 0.2 mm (Fig. 14).

FIGURES 18–23. Male genitalia of Heterocerus spp. 18—aedeagus H. heydeni, ventrally; 19—tegmen apex H. heydeni, dorsally; 20—tegmen sclerotization H. heydeni, dorsal; 21—aedeagus H. flexuosus, ventrally; 22—tegmen apex H. flexuosus, dorsally; 23—tegmen sclerotization H. flexuosus, dorsal. On Figs. 18 & 21 most of the sclerites and membranes connecting the lateral lobes of tegmen and penis base are omitted. Scale bar = 0.1 mm.