Oxyporus Fabricius, 1775

Tokareva, Alexandra S., Konstantinov, Fedor V., Brunke, Adam J. & Solodovnikov, Alexey, 2021, DNA-barcode and endophallus morphology delimit congruent species in a systematic revision of the oxyporine rove beetles of Russia (Coleoptera: Staphylinidae: Oxyporinae), Contributions to Zoology 90 (2), pp. 344-407 : 377-385

Genus Oxyporus Fabricius, 1775

Oxyporus : Fabricius, 1775: 267. Type species by subsequent designation: Staphylinus rufus Linnaeus, 1758 (Latreille, 1810: 427).

Diagnosis. Antennae filiform or clavate, inserted at side of head near anterior margin of eyes; mandibles prominent, sickle-shaped; apical labial palpomere large, transverse, strongly securiform; procoxae large, conical, prominent, protrochantin broadly exposed; middle coxae widely separated; posterior coxae transverse; tarsi 5-5-5; abdomen with six visible sterna and two pairs of paratergites per segment. In Russia, the genus includes 10 species which cannot be confused with any other genus of rove beetles.

Note. Soon after Nakane and Sawada (1956) described the genus Pseudoxyporus for a number of Japanese species formerly in Oxyporus, Campbell (1969) , in his revision of the New World Oxyporinae , downgraded Pseudoxyporus to subgeneric rank. Since then, both concepts persist in literature, in parallel use. For practical reasons (see the last paragraph of the Discussion) here we use the genus Oxyporus in a broad sense, i.e., with Oxyporus s. str. and Pseudoxyporus as subgenera.

Key to the Oxyporus species of Russia

1. Elytra metallic blue with reddish-yellow humeri; body red, with some parts of head and abdomen black. Scutellum with faint punctation. Aedeagus: endophallus with dorsal distal lobe subdivided by transverse line ( figs 4B , 15C, D ) …..……………………………… O. cyanipennis

- Elytra of different coloration. Scutellum without punctation, smooth ( fig. 3A–G ). Aedeagus: dorsal distal lobe of endophallus not divided by transverse line…….………….…………………..………… 2

2. Pronotum red, or reddish-yellow, never black ( figs 3D , 4B–E )…………….…………. 3

- Pronotum black ( figs 3A–C, E–G , 4A ). ………………………………………………..……6

3. Head reddish or amber-yellow, of the same color as pronotum and elytra; abdomen black. Body on average larger (body length can reach 16 mm;the largest species in the Russian fauna). Aedeagus: median lobe with two-pointed apex, apical distal lobe of endophallus of same width along its entire length ( figs 4D , 14A, B )….………….........………... O. procerus

- Head black; elytra bicolored. Body on average smaller (body length at most 14 mm)………….……………………………… 4

4. Antennae relatively short, at most as long as head capsule; middle antennomeres transverse, wider than long; antennomeres with longitudinal asetose middle part forming bold line along antenna ( figs 3A–G , 4A, D ). Aedeagus: endophallus with one ventral, one pair of lateral and one pair of dorsal lobes; apical distal lobe of endophallus curved dorsad ( fig. 14C, D )………..……………………...… O. rufus

- Antennae relatively long, distinctly longer than head capsule; middle antennomeres longer than wide ( figs 4B, C, E ) ………….……………………...…………………. 5

5. Mesoventrite, abdomen and legs entirely - Mesoventrite black; abdomen orangeblack; elytra apically black; maxillary red with black apex only; legs yellow;

and labial palps black. Aedeagus: dorsal elytra with largely yellow humeri and lobe of endophallus narrow, vermiform, sutural angles; maxillary and labial palps covered with transverse rows of denti- yellow. Aedeagus: dorsal lobe of endo-

cles ( figs 4C , 16C, D )…….………….………… phallus without rows of denticles (figs ………….…………. O. (P.) melanocephalus . 4E, 16 Downloaded A, B)........…… from.. Brill …….com .. O. (P 12.) /12 dybowskii /2023 04:19:55PM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

6. Elytra always entirely black……............... 7 Subgenus Oxyporus Fabricius, 1775

- Elytra always bicolored….…….....……….. 8 Diagnosis. Labrum divided by membranous

7. Pronotum with transverse impression. median line. Frons smooth, seldom with weak Mesepisterna and mesepimera cha- impressions. Antennae distinctly shorter than greened, not punctured. Aedeagus: head, vestiture absent at central area of latapical distal lobe of endophallus eral surface of each segment, medial segments forms a sclerotized, narrow, weakly shorter than wide. Posterior pronotal angles curved hook ( figs 3B , 11C, D )…… broadly rounded or indistinct. Mentum trap- ………….……………….………………. O. niger ezoid, with membranous flaps on each side.

- Pronotum without transverse impres- Protarsomeres cylindrical with some scattered sion, smooth. Mesepisterna and mese- strong setae. Aedeagus: parameres acicular, dispimera not chagreened but punctured. tinctly shorter than half of median lobe; median Aedeagus: apical distal lobe of endo- lobe dorso-ventrally flattened, basal bulb and phallus membranous ( figs 3C , 11A , ventral wall membranous, median orifice open- B)…………………………… O. mannerheimii ing ventrally (Nakane & Sawada, 1956).

8. Pronotum smooth. Paratergites of the abdominal segments III and IV (first Oxyporus (Oxyporus) basicornis Cameron , and second visible) yellow in contrast to 1930 black tergites ( fig. 3E–G )….......………..… 9 O aequicollis Bernhauer, 1935 , syn. nov.

- Pronotum with transverse impression. O. parvus Lee et al., 2020 , syn. nov. ( figs 3E, F , Paratergites of the abdominal seg- 9A, B) ments III and IV (first and second visi- O. lewisii var. basicornis : Cameron 1930: ble) always of the same colour as their 184. tergites (yellow or black)...........………… 10 O. basicornis : Nakane and Sawada 1956: 67;

9. Aedeagus: apical sclerites of endophal- Shabalin 2012: 413. lus widened at apex, forming lamellate O. aequicollis : Bernhauer 1935: 6; Kirschenblatt triangle ( fig. 9C, D )……..…… O. germanus 1938: 529; Shavrin and Gildenkov 2009: 123

- Aedeagus: apical sclerites of endophal- O. parvus : Lee et al., 2020: 685. lus include a number of paired, arched spicules which form a distinctly scle- Type material examined. Oxyporus basicorrotised frame around the secondary nis: JAPAN: Lectotype, here designated: ♂, gonopore ( fig.9A, B ).………… O.basicornis ‘Type\ Japan; Chuzenji; AT 4000; 23.VII.’28;

10. Elytra at sutural area with distinct J.E.A. Lewis\ O. lewisi ; v. basicornis ; Type black triangle in basal half of elytral Cam.\ M. Cameron Bequest. B.M.1955–147.\ length. Aedeagus: median lobe with Oxyporus basicornis Cameron ; det. M.-H. Kim two-pointed apex, apical distal lobe of 2004\ Oxyporus ; basicornis; Cameron; det. M. endophallus with a spicule inside (figs Schülke 2019’ (NHM). 4A, 14A, B)………………………. O.triangulus Oxyporus aequicollis : RUSSIA: Possible

- Elytra at sutural area yellow or with holotype? ♀, ‘Baikal\ Sharp Coll.; 1905-313.\ black, narrow, non-triangular stripe. Oxyporus aequicollis Bernh ; M.Bernhauer Aedeagus: median lobe with rounded det.’ (NHM). apex; apical distal lobe of endophallus Other material examined. RUSSIA: Republic forms sclerotized, distinctly curved hook of Buryatia: 2 ♂, Pribaikalsky District, Selenga ( figs 3A , 13A–E ).............……. O. maxillosus river valley Downloaded , Diky Byk from cape Brill .com, 52.1405 12/12N /2023 107.1186 04:19:E 55, PM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

7.VIII.2019, I. Makhov leg. (ZISP); Primorsky only by CO1 barcode and endophallus struc- Territory: 3 ♂, Gornotaezhnoe, 43.6945N ture. In particular, the endophallus of O. 132.15203E, 2–21.IX.2018, in fungi, A. Tokareva basicornis has two additional spiculae and the leg. (ZISP); ♂, Suputinsky Nature Preserve, additional central sclerite missing in O. ger- [43.6327N 132.2902E], 3.IX.1969, Kryzhanovskij manus. From all other species of Oxyporus in leg. (ZISP); ♀, Posiet peninsula, 42.6622N Russia, O. basicornis can be distinguished by 130.7933E, 4.VIII.2019. A. Tokareva leg. (ZISP); the structure of endophallic sclerites and col- 4 ex., Pogranichny area, Pogranichny suburbs oration of head, thorax, and elytra. 44.4230N 131.3649E, 20.VIII.2019, A. Tokareva Distribution. Oxyporus basicornis is relileg. (ZISP). ably known from Japan (Honshu), South Diagnosis. Pronotum smooth or at most Korea (Lee & Ahn, 2020) and Russia, where with slight impression according to Lee et al. it is recorded here for the first time. Russian (2020; as O. parvus ), paratergites of first three records come from Primorsky Territory and or four visible abdominal segments yellow Buryatia; it is presumably widely distribwhile respective tergites and sternites black. uted in the Russian Far East and East Siberia Endophallus : distal lobe bent dorsad, con- ( fig. 10C ). sisting of dorsal, ventral and additional pair Bionomics. This species can be usually found of thin bow-like spicules and central sclerite in large fungal caps together with O. germanus of characteristic shape ( fig. 9A, B ), all covered and/or O. rufus , or alone in Suillus americanus with membrane; dorsal and ventral pairs of (Peck) Snell in the Russian Far East. thin spicules bent ventrad and then sharply Comments. Oxyporus basicornis was dorsad to form notable sclerotized frame sup- described from Japan as a variety of O. lewisi porting walls of secondary gonopore. ( Cameron, 1930) based on the dark median Redescription. Body length 5.7–10.1 mm line on the last six antennomeres and yellow (n = 10). Head subovate to round, smooth, paratergites on the first three (and not four as brown to black. Pronotum without transverse in O. lewisi ) abdominal segments. Nakane and elevation, smooth, brown to black. Elytra Sawada (1956) raised it to the status of a speblack with yellow pattern. Mesepisterna cies. In Russia, O. basicornis was recorded for and epimera with rough punctation, black. the first time in Primorsky Territory, diagnosed Abdominal tergites and sternites black, lat- only using color (Shabalin, 2012). Our study, eral abdominal sclerites of segments I-III or however, has revealed that O. basicornis over- IV yellow. Aedeagus: median lobe with round laps in coloration with O. germanus , another apex; endophallus with twin-coned spheri- species described from Japan, and distributed cal dorsal lobe covered with medium-sized in the Russian Far east as well (see below). Both denticles, pair of spherical lateral lobes, and species reliably differ from each other only by transverse ventral lobe covered with tiny gran- CO1 barcode and structure of the endophallus. ulation; apical distal lobe membranous, with Therefore, it was unclear which species was tiny granulation, bent dorsad, with four pairs recorded by Shabalin (2012) under the name O. of thin long bow-like sclerites under mem- basicornis , while our data here, based on a reexbrane and one thin ligament sclerite dorsally. amination of Shabalin’s material as well, are the Basal sclerotization disjunct, additional spicu- first reliable records of this species from Russia. lae of a peculiar form ( figs 3E, F , 9A, B ). Oxyporus aequicollis was described from Comparison. Oxyporus basicornis is identi- a single specimen sampled in ‘Irkutsk’ cal to O. germanus in coloration and body pro- (Bernhauer Downloaded , 1935) as from a species Brill .com close to 12/12/2023 O. man- 04:19:55PM portions. Both species can be distinguished via Open nerheimii Access. This is an. In open fact access, the article original distributed description under of the terms of the CC-BY 4.0 license.

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O. aequicollis is nearly identical to the original smaller body size and distinct longitudinal description of O. germanus , earlier described rows of punctation on elytra. The material from Japan. Kirschenblatt (1938) recorded available to us which matches the descrip- O. aequicollis from Irkutsk Prov., Republic of tions of O. germanus , O. aequicollis , and O. Buryatia, Zabaikalsky Terr., and Khabarovsk basicornis shows no hiatus among the speci- Terr. and noticed its close affinity to O. ger- mens in these and other characters as far as manus. He provided two characters to dif- external morphology is concerned. At the ferentiate O. aequicollis from O. germanus : same time Downloaded , based from on the Brill .com CO1 barcode 12/12/2023 and 04:19:55PM

via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license.

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structures of the endophallus, this material of the size, number and location of these can be unambiguously divided into two spe- patches even within a species, at least in O.

cies, one of which includes the type material rufus , O. germanus , O. basicornis , O. maxil-

of O. germanus , and another, O. basicornis . losus, O. procerus , O. mannerheimii , and O.

Because of previous taxonomic confusion, it niger . The fine scratch-like impression in is also unclear which species was recorded the anterior third of the pronotum of O. ger-

from Russia under the name O. aequicollis manus appears to be a varying trait as well.

in Kirschenblatt (1938). Unfortunately, we Unfortunately, Lee et al. (2020) did not comcould not examine the holotype of O. aequi- pare O. parvus with O. basicornis . Inclusion collis, which was not found either at the Field of their molecular data in our analysis, as Museum or at Vienna Museum of Natural well as matching the molecular clades with History (A.F. Newton and H. Schillhammer, the respective (not sequenced) type mate-

pers. comm., respectively). It is not even clear rial of O. basicornis via examination of the if the type is a male or female, and in the endophallus, clearly shows that O. parvus is latter case, may not be informative for this conspecific with O. basicornis and thus the taxonomic problem. Based on the sample former is placed in synonymy with the latter.

of examined material that includes a female from D. Sharp’s collection in NHM identified Oxyporus (Oxyporus) germanus Sharp, 1889

by M. Bernhauer as O. aequicollis and that ( figs 3G , 9C, D )

maybe in fact a holotype of this species, it is O. germanus : Sharp 1889: 409; Shavrin and clear that O. aequicollis must be a synonym of Gildenkov 2009: 123.

either O. germanus or O. basicornis . Since our data shows that reliably identified O. basicor- Type material examined: JAPAN: Lectotype,here nis has wider known distribution westwards designated. ♂, Type\ Japan.; G. Lewis.\ Sharp (found in Buryatia), while reliably identified Coll.; 190–313\ Oxyporus ; germanus; Type; O. germanus is so far restricted to the Far East, Sendai Jap.’ ( NHM); Paralectotypes: ♂, ‘Japan.;

we place O. aequicollis in synonymy with O. G. Lewis.\ Sharp Coll.; 1905-313\ Oxyporus ; gerbasicornis. manus; Sendai; Japan. Lewis’ ( NHM); ♀, Japan.; Oxyporus parvus was described from South G. Lewis.\ Sharp Coll.; 1905-313\ Oxyporus ; Korea (Lee et al., 2020). The authors showed germanus; Sendai; Japan. Lewis’ ( NHM).

the difference of O. parvus from O. germanus Other material examined. RUSSIA: based on CO1 and provided a scheme of the Primorsky Territory: ♂, Gornotaezhnoe, endophallus of their new species, also men- 43.6945N 132.15203E, 2–21.IX.2018, in fungi, A.

tioning and partly illustrating diagnostic Tokareva leg. ( ZISP); ♂, Kedrovaya Pad’ Nature characters of O. parvus , namely the presence Preserve, Kedrovaya river valley, [43.1029N

of the tomentose patches on tergites III and 131.4897E], 25.VIII.1998, O. Kireichuk leg.

IV in O. parvus (on tergite IV in O. germanus ), ( ZISP); 2 ex., South of Anisimovka village, brown abdominal tergites III – V in O. parvus 43.1504N 132.7893E, 27.VIII.2019, A. Tokareva ( III – VI in O. germanus ) and the presence of leg. ( ZISP); ♂, Vladivostok, Sputnik sta-

a shallow and broad depression on anterior tion, [43.2328N 132.0412E], 13.IX.2009, S.

third of pronotum in O. parvus (presuma- Shabalin leg. ( FEB RAS); ♂, Pogranichny area ,

bly absent in O. germanus studied by the Pogranichny suburbs 44.4230N 131.3649E, authors). Regarding the tomentose patches, 20.VIII.2019, A. Tokareva leg. ( ZISP); Sakhalin our sample of material shows variability Province: 3 Downloaded ♂, Sakhalin from Island Brill .com, Okha 12/12suburbs /2023 04:19,:55 PM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ near Medvezhye lake, [53.6139N 142.9568E], 28.VIII.1979, in fungi, Okulov leg. ( FEB RAS).

Diagnosis. Pronotum smooth, paratergites of first three or four visible abdominal segments yellow while respective tergites and sternites black. Endophallus: distal lobe of apical sclerites consisting of ventral pair of spiculae and dorsal pair of complex sclerites flattening and widening in the middle of distal lobe and forming characteristic structure supporting walls of distal lobe ( fig. 9C, D ).

Redescription. Body length 5.2–9.8 mm (n = 10). Head subovate to round, smooth, brown to black. Pronotum without transverse elevation, smooth. Elytra black with yellow pattern. Mesoepisterna and epimera with rough punctation, black. Abdominal tergites and sternites black, paratergites of segments III-V or III-VI yellow. Aedeagus: median lobe with round apex; endophallus with twin-coned spherical dorsal lobe covered with medium-sized denticles and transverse, relatively large ventral lobe covered with tiny granulation, pair of spherical lateral lobes, and apical distal lobe membranous, with tiny granulation, bent dorsad; one pair of thin long bow-like ventral spiculae and paired complex additional spiculae in center flattening and widening towards apex under membrane and small obscure ligament sclerite dorsally; basal sclerotization obscure ( fig. 3G , 9C, D ).

Comparison. Oxyporus germanus is identical to O. basicornis in coloration and body proportions. Both species can be distinguished only by CO1 barcode and endophallus morphology. In particular, the endophallus of O. germanus has paired additional central spiculae flattening and widening towards apex, which are absent in O. basicornis . From all other species of Oxyporus in Russia O. basicornis can be distinguished by the structure of endophallic sclerites and coloration of head, thorax, and elytra.

Distribution. Oxyporus germanus is reliably known from Honshu, Japan ( Sharp, 1889) and Russia (Primorsky Territory, new data) ( fig. 10C ).

Bionomics. This species can be usually found in large fungal caps together with O. basicornis and/or O. rufus , or alone in Suillus americanus (Peck) Snell in the Russian Far East.

Comments. Oxyporus germanus was described from the Japanese Honshu island ( Sharp 1889) and recently recorded from Russia ( Shavrin & Gildenkov 2009). Since, based on our data (see above), reliable discrimination of O. germanus from O. basicornis is impossible without DNA barcoding or study of the endophallus, Shavrin and Gildenkov’s (2009) record needs verification. Here, O. germanus is reliably recorded from Russia for the first time.

Oxyporus (Oxyporus) mannerheimii

Gyllenhal, 1827

( figs 3C , 11A, B )

O. mannerheimii : Gyllenhal 1827: 495; Kirschenblatt 1938: 531; Shabalin 2012: 413. (Also a number of published more or less precise records from European Russia in the faunistic literature).

Material examined. BELARUS: Vitebsk Province: ♂, Korolevo , [53.5988N 27.4470E], VII-VIII.1894, ex coll. Birulya ( ZISP) ; ♀, Korolevo , [53.5988N 27.4470E], VII-VIII.1894, ex coll. Birul’. ( ZISP) ; KAZAKHSTAN: Western Kazakhstan Province: ♀, inflow Tautekeli of river Kara-Koba , [48.9036N 85.6630E], 7.VIII.1989, V. Kastcheev leg. ( ZISP) ; ♂, Rakhmanovskie Kliuchi, Berel’ river , [49.5701N 86.4994E], 11.VIII.1988, V. Kastcheev leg. ( ZISP) ; Aqmola Province: ♂, near Stepnyak, Zhukey lake , [52.8931N 70.5845E], 20.VII.2002, V. Kastcheev leg. ( ZISP) ; MONGOLIA: 2 ♀, Tov Province, near Batsumber, Suzukte clove, s-w Kentay , [48.4122N 106.7590E], 20.VIII.1924, Kozlov leg. ( ZISP) ; RUSSIA: Leningrad Province: ♀, [59.59N 31.09E], 18. VI.1902, Borovsky leg. ( ZISP) ; Tver Province: ♂, Bologoye , [57.8868N 34.0709E]: 5.VII.1902, F. Zaytsev leg. ( ZISP) ; ♂, 1.VII.1902, F. Zaytsev leg. ( ZISP) ; Pskov Province: ♀, Shchepets , [58.7641N 28.2036E], 16. VI.1901, N. Ivanov leg. ( ZISP) ; Sebezhskiy District, near Osyno , [56.1495N 28.6672E], 25.VII.2009 , V.N. Prasolov leg. ( ZISP) ; Sverdlovsk Province: 4 ex. ,

Visimsky Nature Reserve [57.4296N 60.0287E], 16.VIII.2019, in fungi, N. Ukhova leg.( ZISP); 3 ♀, near Yekaterinburg, Uktus station , [56.7807N 60.6553E], 20. VI.1910, G. Jacobson leg. ( ZISP) ; ♀, Pivovarikha , [52.2803N 104.4665E], 1903, Soldatov leg. ( ZISP) ; Irkutsk Province: ♂, Listvyanka, near Baikal lake , [51.8589N 104.8725E]: 22.VII.1912, Kuligin leg. ( ZISP) , ♀, 12.VII.1912, Kuligin leg. ( ZISP) ; Kemerovo Province: ♂, near lake Berchikul , [55.6277N 88.3580E], 15–31.VII.1903, ( ZISP) ; Primorsky Territory: 2 ♂, Anisimovka, Litovka mountain , [43.10263N 132.78437E], 26.VIII.2019, in fungi, A. Tokareva leg. ( ZISP) ; 3 ♂, Gornotaezhnoe , 43.6945N 132.15203E, 2–21.IX.2018, in fungi, A. Tokareva leg. ( ZISP) ; 2 ♂, Lazo District, Tachingoiz Bay , [43.0214N 134.1298E], 20.VII.1975, V. Kastcheev leg. ( ZISP) .

Diagnosis. Entirely black, except yellowish tarsi, apical portion of labial palps, and lateral parts of antenna. Pronotum smooth, without transversal ridge or impression; mesepimera and mesepisterna smooth, shagreened, without punctation. Endophallus: apical distal lobe membranous; spiculae relatively short, additional pair of spiculae slightly flattened, located in apical portion of lobe ( fig. 11A, B ).

Redescription. Body length 8.5–10.7 mm (n = 10). Head subovate to round, smooth, brown to black. Pronotum without transverse elevation, smooth. Elytra black. Mesoepisterna and epimera shagreened, black. Abdomen black. Aedeagus: median lobe with round apex; endophallus with twin-coned spherical dorsal lobe covered with medium-sized denticles and transverse ventral lobe covered with tiny granulation, pair of spherical lateral lobes, and apical distal lobe membranous, with tiny granulation, bent dorsad; it includes one pair of relatively short dorsal and ventral spiculae and paired additional spiculae in center disposed in apical portion of lobe, and ligament sclerite closely connected with dorsal spiculae; basal sclerotization connected only to ligament sclerite ( figs 3C , 11A, B ).

Comparison. Among all other species of Oxyporus in Russia, O. mannerheimii is most similar in coloration to O. niger , from which it differs by the smooth pronotum and shagreened, not punctured surface of mesepisterna and mesepimera, and by the structure of the endophallus (compare fig. 11A, B and 11C, D ).

Distribution. Oxyporus mannerheimii is a Transpalaearctic species and its distribution is confirmed here by matching DNA barcodes of the European and Far Eastern specimens. In Russia, it occurs from Leningrad Region, through Urals and Irkutsk Region to Primorsky and Sakhalin Regions ( fig. 12A ).

Bionomics. In Leningrad Province, Russia, it was collected from fungal caps of Pleurotus spp.

Comments. This species appears to be phylogenetically remote from other Russian members of the subgenus Oxyporus (K2Pdistance 0.09–0.15).

Figures

FIGURE 3 Oxyporus of Russia. Habitus.A – male and female of O. (O.) maxillosus; B – male and female of O. (O.) niger; C – male and female of O. (O.) mannerheimii; D – male and female of O. (O.) rufus; E – male of O.(O.) aequicollis (new synonym of O.(O.) basicornis); F – female of O. (O.) basicornis; G – female of O. (O.) germanus. Scale bar:1 mm.

FIGURE 4 Oxyporus of Russia.Habitus.A – male and female of O. (O.) triangulus; B – male and female of O. (P.) cyanipennis; C – male and female of O. (P.) melanocephalus; D – male and female of O. (O.) procerus; E – male of O. (P.) dybowskii.Scale bar:1 mm.

FIGURE 9 Schemes and photos of everted endophallus of O. (O.) basicornis (A, B) and O. (O.) germanus (C, D). Acronyms: adsp – additional spiculae; ddl – dorsal distal lobe; dsp – dorsal spiculae; lgsc – ligament sclerite; ltl – lateral lobe; vl – ventral lobe; vsp – ventral spiculae. Scale bar: 0.1 mm.

FIGURE 10 Distributional maps of O. (P.) cyanipennis,O. (P.) melanocephalus, and O. (P.) dybowskii (A); O. (O.) procerus, O. (O.) triangulus, and O. (O.) niger (B); O. (P.) dybowskii O. (O.) basicornis and O. (O.) germanus (C).

FIGURE 11 Schemes and photos of everted endophallus of O. (O.) mannerheimii (A, B) and O. (O.) niger (C, D). Acronyms: adsp – additional spiculae; bsc – basal sclerotization; lgsc – ligament sclerite. Scale bar: 0.1 mm.

FIGURE 12 Distributional maps of O. (O.) rufus and O. (O.) mannerheimii (A); and O. (O.) maxillosus (B). Material examined abbreviated as ME.

FIGURE 13 Schemes and photos of everted endophallus of O. (O.) maxilosus (A, B) and its color forms described as separate species: O.(O.) aokii (C), O.(O.) basiventris (D), and O.(O.) japonicus (E). Acronyms:bsc – basal sclerotization; lgsc – ligament sclerite. Scale bar: 0.1mm.

FIGURE 14 Schemes and photos of everted endophallus of O. (O.) procerus (A, B) and O. (O.) rufus (C, D). Acronyms: adsp – additional spiculae; ddl – dorsal distal lobe; dsp – dorsal spiculae; lgsc – ligament sclerite; ltl – lateral lobe; vl – ventral lobe; vsp – ventral spiculae. Scale bar: 0.1 mm.

FIGURE 15 Schemes and photos of everted endophallus of O. (O.) triangulus (A, B) and O. (P.) cyanipennis (C, D).Acronyms: ddl – dorsal distal lobe; dsp – dorsal spiculae; lgsc – ligament sclerite; vsp – ventral spiculae. Scale bar: 0.1mm.

FIGURE 16 Schemes and photos of everted endophallus of O. (P.) dybowskii (A, B) and O. (P.) melanocephalus (C, D).Acronym:ddl – dorsal distal lobe.Scale bar:0.1 mm.