Review of the tribe Chilocorini Mulsant from Iran (Coleoptera, Coccinellidae).

The Iranian checklist of the tribe Chilocorini Mulsant, 1846 (Coleoptera: Coccinellidae) is updated. In total, 13 species belonging to four genera (Brumoides Chapin, 1965, Chilocorus Leach, 1815, Exochomus Redtenbacher, 1843, and Parexochomus Barovsky, 1922) are listed from Iran. An identification key to all genera and species currently known from Iran is presented along with illustrations of adult specimens and male genitalia.

Introduction

The family , with nearly 6000 species and 360 genera, belongs currently to the superfamily (: ) (Robertson et al. 2015, Tomaszewska and Szawaryn 2016). It is divided into two subfamilies: and . The subfamily includes three tribes: , (including ) and (Escalona and Ślipiński 2012); the remaining taxa belong to the subfamily (Seago et al. 2011, Robertson et al. 2015, Szawaryn et al. 2015, Escalona et al. 2017).

The tribe Mulsant, 1846 contains approximately 250 species belonging to 27 genera (Łączyński and Tomaszewska 2012, Li et al. 2017), of which nine genera have hitherto been recorded from Palaearctic region including: Chapin, 1965, Leach, 1815, Sasaji, 2005, Redtenbacher, 1843, Barovsky, 1922, Weise, 1895, Kovár, 1995, Ahmad & Ghani, 1966 and Miyatake, 1970) (Kovář 2007).

Although most members of are coccidophagous (Giorgi et al. 2009, Escalona et al. 2017), aphidophagy is also present in some species (Ślipiński and Giorgi 2006); so, the members of this tribe have the potential to be effective biological control agents of coccids and aphids (Drea and Gordon 1990, Ponsonby and Copland 1997).

In the last classification of the former subfamily by Kovář (2007), the species of the genus Mulsant, 1850 were transferred to Redtenbacher and the subgenus of was considered as a valid genus, under the name of Barovsky, 1922. This classification was followed by Nedvěd and Kovář (2012). Moreover, according to Ślipiński (2007), the subfamily Mulsant was dissolved and all tribes were lumped into the subfamily . This classification was confirmed by subsequent morphological and molecular studies (Seago et al. 2011, Robertson et al. 2015). The number of genera and species of this tribe is continuously increasing (Ślipiński and Giorgi 2006, Łączyński and Tomaszewska 2009, Wang and Ren 2010, Łączyński and Tomaszewska 2012, Li et al. 2015, Li et al. 2017) and it is expected that this trend will be continuing.

Although a large number of species of this tribe have hitherto been reported from Iran (Duverger 1983, Kovář 2007, Moddarres-Awal 2012), there is no complete and comprehensive information on the Iranian . The checklist by Abdolahi Mesbah et al. (2016) differs from our view and does not include identification key, diagnosis, and synonymy. Our paper corrects the previous studies on the species of this tribe in Iran, in order to update the information about Iranian .

Materials and methods

This study was mainly based on review of the literature along with the samples collected by the first author. The samples were collected by hand, aspirator, or sweep net in the fields, orchards, and pastures of various provinces of Iran. The specimens were PageBreakexamined under Olympus stereomicroscope (SZ-ST). The specimens were first boiled in 10% KOH for a maximum of 20 min depending on the darkness of the body color/ sclerotization in order to dissect the genitalia. The dissected genitalia were then transferred into distilled water for a maximum of 10 min to rinse off the KOH. Finally, the slides were prepared using Canada balsam. The slides were examined under a microscope (Olympus CX21) and images were taken using a digital camera and edited in Photoshop software (Adobe Photoshop CS5.1). The specimens were identified to species using available keys and resources (Mader 1955, Fürsch 1961, Bielawski 1984, Kovář 1995, Raimundo and van Harten 2000, Raimundo et al. 2008).

Although the higher classification of Seago et al. (2011) was followed in this study, taxonomy at the species level is based on Kovář (2007). Morphological terminology follows that of Ślipiński (2007). All of the specimens collected and examined during this study are deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

Results and discussion

The Iranian coccinellid species list of the tribe is updated, which includes 13 species belonging to four genera (, , , and ).

Although there are some records of Thunberg, 1781 from Iran (Ansari pour and Shakarami 2011, Tavakol et al. 2014), re-examination of the voucher specimens of this species showed that these reports are misidentifications and these samples are actually (Goeze, 1777). is morphologically similar to but is distinguished from it by the male genitalia, and has not hitherto been reported from Palaearctic region (Kovář 2007). It is distributed in the northern states of USA (Gordon 1985) and south and west of Africa (Fürsch 1961).

Mahghari and Ostovan (2006) reported two ladybird species, L. and (Say, 1835), from the northern provinces of Iran (Gilan and Mazandaran province) as natural enemies of whiteflies. In coccinellid taxonomy, there is no known species under the name of , while has not been reported so far from Palaearctic region (Kovář 2007). According to our knowledge, the presence of these species in Iran is doubtful and not confirmed.

Barovsky (1922) reported Barovsky, 1922 from Iran (Shahrood, H. Christoph leg.). There are also specimens in Zoologichesky Institut (Akademii Nauk SSSR) in St. Petersburg, labeled as which had been collected from Iran (Shahrood, H. Christoph leg). Kovář (1995) however identified these specimens as Weise.

Data on the presence of in Iran are based on Kovář (2007). Since we do not have any information (particularly morphological) about this species, it is excluded from the identification key of Iranian species of .

Subfamily Latreille, 1807

Mulsant, 1846

Diagnosis.

Body size small to medium (2.0–8.0 mm), with downward directed head inserted into prothorax to some extent; dorsum usually without obvious pubescence. Head wider than long, flattened ventrally; clypeus variously expanded laterally and wholly concealing antennal insertions. Mandibles triangular, strong with an apical tooth and heavily developed molar teeth; maxillary palps relatively long, terminal palpomere parallel sided to weakly enlarged apically; labial palp clearly separated basally, inserted on ventral side of prementum. Antenna composed of 7–10 antennomeres, markedly short with a fusiform club composed of three terminal antennomeres. Prosternum fairly elongate in front of coxae; prosternal process narrow, parallel sided without carinae. Hind wings with large anal lobe. Elytra irregularly punctate, with epipleuron wide and complete to apex, frequently with foveae for receiving apices of femora. Abdomen with five or six ventrites; postcoxal lines at abdominal ventrite 1 variable, without associated pits and pores. Male genitalia with symmetrical tegmen, penis guide sometimes asymmetrical; parameres well developed, apically setose; penis a simple, single sclerite with sizeable basal capsule. Coxites triangular and faintly sclerotized, usually without styli; bursa copulatrix with infundibulum or fleshy lobe, with sperm duct composed of two parts of different diameter; spermatheca bean-shaped, sclerotised without well differentiated nodulus or ramus, with large accessory gland (after Ślipiński 2007).

Updated checklist of the Iranian species of

Chapin, 1965

Chapin, 1965: 237. Type species:

Body length 2.0–3.5 mm. Dorsum glabrous; yellowish or brown, elytra with dark markings. Eye distinctly emarginate. Antenna composed of 8 antennomeres; terminal antennomere small, partly embedded in penultimate one. Clypeus short; labrum exposed. Pronotal base bordered; prosternal process extremely narrow, without carinae; without hypomeral fovea. Fore tibia narrow, simple, middle and hind tibiae with two apical spurs; tarsal claws appendiculate or weakly thickened basally. Abdominal ventrite 6 visible in males; abdominal postcoxal lines separated medially, each arcuately recurving apically and reaching or nearly reaching midpoint of lateral line (after Ślipiński 2007).

Ecology.

Various species of have been associated with mealybugs (Ślipiński 2007), namely (: ), (: ), (: ), and (: ) (Gordon 1985, Gautam 1990, Hodek and Honěk 2009, Arif et al. 2012, Giorgi et al. 2014). Some species of this genus, such as (F.) feed on some whitefly species, such as (Maskell) (Inayatullah 1984, Hodek and Honěk 2009) in addition to feeding on some coccids, such as (better for development) and (better for oviposition) (Gautam 1990).

Fürsch, 1987

Figs 1 , 21 , 24–26

Figures 1–9.

Dorsal habitus of species. 1 Fürsch 2 Linnaeus 3 Crotch 4 Weise 5 Gebler 6 Fleischer 7, 8 Linnaeus 9 Weise.

Figures 21–31.

Morphological details and male genitalia of species. 21, 24–26 : 21 Abdominal postcoxal lines 24 Antenna 25 Tegmen 26 Penis apex 22 : Abdominal postcoxal lines 23, 28 : 23 Hind leg 28 Antenna 27 : Antenna 29–31 : 29 Tegmen, ventral view 31 Tegmen, lateral view 30 Penis apex.

Fürsch, 1987: 44.

General distribution.

Middle East (that includes Iran, Saudi Arabia, United Arab Emirates, Yemen) (Kovář 2007), Southern Africa (Łączyński and Tomaszewska 2012).

Distribution in Iran.

Iran (Kovář 2007) – no specific distribution known.

Remarks.

The species descriptions and photographs by Fürsch (1987) and Raimundo et al. (2008) were used with some modifications.

Leach, 1815

Leach, 1815: 116. Type species:

Body length 2.5–4.8 mm. Dorsal body glabrous; elytra black or brown with white or orange markings; eye clearly emarginate. Antennae short, composed of 8 antennomeres; with scape symmetrical; 8th antennomere either as long as or markedly longer than antennomere 7. Clypeus long; labrum partly exposed. Pronotal base unbordered; prosternal process narrow without carinae; hypomeral fovea absent. All tibiae flattened and angulate externally, without apical spurs; tarsal claws strongly appendiculate. Elytral margin not reflexed with indistinct bead; epipleural foveae weak. Abdominal ventrite 6 visible in males; abdominal postcoxal lines separated medially, each running parallel to hind margin of ventrite (after Ślipiński 2007).

Although various scale insects are primary hosts of (Escalona et al. 2017), some species at least accept aphids as prey (Gordon 1985, Drea and Gordon 1990, Ślipiński 2007, Hodek and Honěk 2009). Nonetheless, there are some reports about some species of this genus, such as (Say) which feed on some whitefly species, such as Ashby (Dowell and Cherry 1981, Hodek and Honěk 2009).

(Linnaeus, 1758)

Figs 2 , 16 , 15–20

Figures 14–20.

Morphological details and male genitalia of species. 14–20 : 14 Head 15 Abdominal postcoxal lines 16 Leg 17 Tegmen 18 Penis guide of tegmen 19 Penis apex 20 Penis.

Linnaeus, 1758: 367.

Müller, 1776: 68.

Börner, 1776: 250.

Thunberg, 1792: 105. [Homonym]

Florencourt Chassot, 1796: 214.

Fabricius, 1798: 79. [Homonym]

Costa, 1839: 104.

Sahlberg, 1903: 86.

Material examined.

8♂, 3♀: Iran, Lorestan province, V.2013, lgt. Amir Biranvand, det. Biranvand. 2♂, 1♀: Iran, Semnan province, V.2015, lgt. Mino Toozandejani, det. Biranvand.

Afrotropical region, Nearctic region, Palaearctic region (Mader 1955, Gordon 1985, Kovář 2007, Canepari 2011) and Oriental region (Poorani 2002).

Widely distributed (Duverger 1983, Moddarres-Awal 2012).

This species feeds on a wide range of species: (), (), (), (), (), (), (), (), (), (), (), (), (), (), (), (), () (Moddarres-Awal 2012) and other coccids, particularly armoured scales (Hodek 1973, Stansly 1984).

Redtenbacher, 1843

Redtenbacher, 1843:11. Type species:

Body length 2.8–5.5 mm. Dorsal body glabrous; elytra black, brown, or yellow, often with contrasting red or yellow markings; sometimes (in lighter coloured species) with black stripes along lateral margins of elytra. Antenna composed of 10 antennomeres, minute terminal antennomere embedded in penultimate one; pronotal basal margin completely bordered with submarginal line; prosternal process narrow, truncate apically, without carinae; elytral epipleura clearly narrowing, without foveae; abdominal postcoxal lines complete or nearly so, semicircular, reaching to inner end of lateral line; meso- and metatibiae each with two apical spurs; tarsal claws with or without basal tooth (after Li et al. 2015).

Most species of this genus are aphidophagous and coccidophagous (Gordon 1985, Kovář 1995, Magro et al. 2010). Nontheless, there are some reports about some species of the genus feeding on aleyrodids e.g., Mulsant which feeds on (Gennadius) (Yigit 1992, Leite et al. 2003, Hodek and Honěk 2009).

Barovsky, 1927

Barovsky, 1927: 200.

China, Iran, Kazakhstan (Kovář 2007).

Iran (Kovář 2007) – no specific distribution provided.

Crotch, 1874

Fig. 3

Crotch, 1874: 193.

Fairmaire, 1876: 94.

Fairmaire, 1876: 94.

Allard, 1870: 9.

Algeria, Iran, Morocco, Tunisia (Mader 1955, Duverger 1983, Kovář 2007).

Dasht Arzhanregion, Kerman, Nowshahr region (Duverger 1983).

We used the species descriptions and photographs of Mader (1955) with some modifications.

Weise, 1885

Figs 4 , 40–43

Figures 40–47.

Morphological details and male genitalia of species. 40–43 : 40 Tarsal claw 41, 42 Tegmen in ventral and lateral view 43 Penis apex 44–47 : 44–45 Tegmen in ventral and lateral view 46 Tip of penis 47 Penis.

Weise, 1885: 55.

5♂, 2♀: Iran, Yazd province, spring and summer 2013, lgt. Mehdi Zare Khormizi, det. Biranvand.

Afghanistan, Iran, Turkey (Kovář 2007).

Golestan, Semnan (Kovář 1995), Lorestan (Jafari and Kamali 2007), Fars (Moddarres-Awal 2012), Yazd (current study).

(Gebler, 1830)

Figs 5 , 37–39

Figures 32–39.

Morphological details and male genitalia of species 32–34 : 32, 33 Tegmen in ventral and lateral view 34 Penis apex 35–36 : 35 Tegmen, ventral view 36 Penis apex 37–39 : 37 Tarsal claw 38 Tegmen, ventral view 39 Penis apex.

Gebler, 1830: 225.

Motschulsky, 1840: 175.

Gebler, 1841: 376.

Weise, 1879: 135.

Fleischer, 1900: 118.

Afghanistan, Armenia, Azerbaijan, France, Iran, Iraq, Italy, Kazakhstan, Kyrgyzstan, Mongolia, Russia, Tajikistan, Turkmenistan, Turkey, Uzbekistan (Kovář 2007).

Khameshorkn region (Duverger 1983), Khorasan (Moodi and Mossadegh 1995, Yaghmaei and Kharrazi Pakdel 1995), Chaharmahal and Bakhtiari (Bagheri and Mossadegh 1996), East Azerbaijan, Gilan, Isfahan, Kerman, Qom, Tehran, Sistan and Baluchestan (Moddarres-Awal 2012).

This species feeds on the mealybugs and () (Moddarres-Awal 2012).

Fleischer, 1900

Figs 6 , 29–31

Fleischer, 1900: 118.

Roubal, 1926: 245.

Roubal, 1927: 135.

3♂, 8♀: Iran, Semnan province, VII.2015, lgt. Mino Toozandejani, det. Biranvand.

Caucasus, Iran, Lebanon, Syria (Duverger 1983), Armenia, Turkey (Kovář 2007).

Sagdar region (Duverger 1983), Kerman (Moddarres-Awal 2012), Semnan (current study).

(Linnaeus, 1758)

Figs 7–8 , 32–34

Linnaeus, 1758: 367.

Gmelin, 1790: 1662.

Fabricius, 1792: 288.

Donovan, 1798: 74.

Schrank, 1798: 444.

Brullé, 1832: 273

Motschulsky, 1837: 422.

Motschulsky, 1837: 423.

Costa, 1849: 62.

Schaufuss, 1862: 50

Kraatz, 1873:192

Weise, 1879: 133.

Weise, 1879: 134.

Schneider, 1881: 16

Fuente, 1910: 444

60♂, 75♀: Iran, Lorestan province, in all seasons, 2013, 2014, 2015, 2016, 2017, lgt. Amir Biranvand, det. Biranvand. 3♂, 3♀: Iran, Semnan province, V.2015, lgt. Mino Toozandejani, det. Biranvand.

Palaearctic Region, Oriental region, Australian region, Nearctic region (USA: California) (Canepari 2011, Li et al. 2015).

Widely distributed (Duverger 1983, Moddarres-Awal 2012).

This species feeds on various species of , namely: (), (), (), (), (), (), (), (), (), (), () (Moddarres-Awal 2012), and other aphids and (Uygun 1981, Ülgentürk and Toros 2001, Kaydan et al. 2006, Kaydan et al. 2012).

Weise, 1878

Figs 9 , 27 , 35–36

Weise, 1878: 93

10♂, 16♀: Iran, Lorestan province, in all seasons, 2013, 2015, 2016, lgt. Amir Biranvand, det. Biranvand.

Palestine (Mader 1955), Caucasus (Duverger 1983), Afghanistan, Azerbaijan, Egypt, Georgia, Iraq, Iran, Lebanon, Syria, Tajikistan (Kovář 2007).

Lorestan (Jafari and Kamali 2007), Chaharmahal and Bakhtiari, Fars, Isfahan, Kerman, Khorasan, Kohgiluyeh and Boyer-Ahmad, Qazvin (Moddarres-Awal 2012), Tehran (Ghanbari et al. 2012), Markazi (Ahmadi et al. 2012), Yazd (Zare Khormizi et al. 2016).

This species feeds usually on (: ) (Moddarres-Awal 2012).

Barovsky, 1922

Barovsky, 1922: 293. Type species:

:

Body length 3.0–3.5 mm. Dorsal body glabrous or pubescent, dark brown or black with lateral margins of pronotum or at least anterior angles yellow or red. Antenna composed of 10 antennomeres, minute terminal antennomere embedded in penultimate one; terminal maxillary palpomeres stout, nearly parallel-sided; pronotal basal margin entirely bordered with submarginal line; prosternal process narrow, rounded apically, without carinae; elytral epipleura clearly narrowing towards apex, without foveae; abdominal postcoxal lines complete and semicircular, reaching to middle of lateral line; meso- and metatibiae each with two apical spurs; tarsal claws with basal tooth (after Li et al. 2015).

The species of are aphidophagous or coccidophagous (Moddarres-Awal 2012).

(Zubkov, 1833)

Fig. 10

Figures 10–13.

Dorsal habitus of species. 10 Zubkov 11 Erichson 12 Goeze 13 Küster.

Zubkov, 1833: 339.

Mulsant, 1850: 1033.

Southern Russia, Caucasus (Mader 1955), Azerbaijan, Armenia, Bulgaria, Georgia, Iran, Kazakhstan, Tajikistan, Turkmenistan, Turkey, Uzbekistan (Kovář 2007).

Razavi Khorasan (Yaghmaei and Kharrazi Pakdel 1995), Lorestan (Jafari and Kamali 2007), Chaharmahal and Bakhtiari, Khorasan (Moddarres-Awal 2012), Kerman (Salehi et al. 2011), Hormozgan (Fallahzadeh et al. 2013).

(Erichson, 1843)

Figs 11 , 23 , 28 , 48–50

Figures 48–55.

Morphological details and male genitalia of species. 48–50 : 48 Tegmen, ventral view 49 Penis apex 50 Penis 51–55 : 51–53 Tegmen, ventral and lateral view 54 Penis 55 Penis apex.

Erichson, 1843: 267.

Fairmaire, 1864: 648.

10♂, 16♀: Iran, Lorestan province, VII.2014, lgt. Amir Biranvand, det. Biranvand.

Oriental region (Poorani 2002), Afrotropical region, Mediterranean region, Middle East (Kovář 2007).

Golestan (Montazeri and Mossadegh1995), Lorestan (Jafari and Kamali 2007), Gilan (Hajizadeh et al. 2003), Fars, Kerman, Khorasan, Khuzestan, Sistan, and Baluchestan (Moddarres-Awal 2012), Lorestan (current study).

This species feeds usually on the following hemipterans: (), (), () (Moddarres-Awal 2012).

(Goeze, 1777)

Figs 12 , 51–55

Goeze, 1777: 248.

Stephens, 1832: 375.

75♂, 90♀: Iran, Lorestan province, spring and summer 2013, 2014, 2015, 2016, 2017, lgt. Amir Biranvand, det. Biranvand. 3♂, 1♀: Iran, Semnan province, VI.2015, lgt. Mino Toozandejani, det. Biranvand.

Palaearctic region (Duverger 1983, Kovář 2007).

Widely distributed (Duverger 1983, Moddarres-Awal 2012).

This species feeds usually on the following species of : (), (), (), (), (), (), (), () (Moddarres-Awal 2012) and other aphids and (Uygun 1981, Atlıhan and Özgökçe 2002, Kaydan et al. 2012).

(Küster, 1848)

Figs 13 , 22 , 44–47

Küster, 1848: 94

Fairmaire, 1884: 59.

Sahlberg, 1903: 36.

Motschulsky, 1849: 155.

Mulsant, 1850b: 947. [Homonym]

Fairmaire, 1875: 540.

Mulsant, 1853: 194.

Sicard, 1929: 60

3♂, 5♀: Iran, Lorestan province, VII.2014, lgt. Amir Biranvand, det. Biranvand.

Oriental region, Palestine, Syria (Poorani 2002), Afghanistan, Algeria, Egypt, France, Greece, Iran, Israel, Italy, Libya, Morocco, Saudi Arabia, Spain, Tunisia (Kovář 2007).

Angohran region, Hormozgan, Tehran (Karaj), Khuzestan (Susangerd), Ramine region, Daran region, Sagdan region (Duverger 1983), Lorestan (Jafari and Kamali 2007), Fars, Kerman, Khorasan, Khuzestan, Sistan, and Baluchestan (Moddarres-Awal 2012).

This species feeds on (: ) and () (Moddarres-Awal 2012).

1	Fronto-clypeal plate emarginate anteriorly (Fig. 14). Postcoxal line on abdominal ventrite 1 merging with posterior margin of ventrite or running very close to it (Fig. 15). All tibiae with tooth at outer side; tibial spurs absent (Fig. 16). Elytron brown or reddish brown with 3 small orange discal spots in transverse row, usually partially fused (Fig. 2). Male genitalia with penis guide as long as parameres (Figs 17, 18), penis as in Figs 19, 20. (Body circular, strongly convex, 3.5–4.5 mm long)	Chilocorus bipustulatus Linnaeus
–	Fronto-clypeal plate not emarginate. Postcoxal line on abdominal ventrite 1 distant from posterior margin of ventrite (Figs 21, 22). Mid-and hind tibiae smoothly arcuate; with 2 apical spurs (Fig. 23)	2
2	Antenna composed of 8 antennomeres (Fig. 24). Body yellow with two small black spots on each elytron, one behind the other (Fig. 1). Male genitalia with parameres slightly longer than penis guide (Fig. 25); penis as in Fig. 26. (Body broadly oval, 2.0–2.5 mm long)	Brumoides adenensis Fürsch
–	Antenna composed of 10 antennomeres (Figs 27, 28)	3
3	Elytra black with red spots or red-brown with or without black spots. Body size 2.8–5.0 mm	Exochomus Redtenbacher 4
–	Elytra completely black. Body size 2.2–4.5 mm	Parexochomus Barovsky 10
4	Elytra black; each elytron with two small or medium sized, separated red spots	5
–	Elytra orange to red-brown, with or without black spots, or elytra black with large pale maculae of irregular shape	6
5	Each elytron with two similar and equally-sized rounded spots (Fig. 6). Male genitalia with penis guide approximately as long as parameres (Figs 29, 31); penis as in Fig. 30. Body oval, 3.5–4.5 mm long	E. quadriguttatus Fleischer
–	Each elytron with two differently sized and shaped spots (Figs 7, 8). Male genitalia with penis guide clearly shorter than parameres (Figs 32, 33); penis as in Fig. 34. Body subcircular, 3.5–4.0 mm long	E. quadripustulatus Linnaeus
6	Background of elytra black; elytral maculae large and of irregular shape, brown or orange	7
–	Background of elytra orange to red-brown; with or without contrasting markings	8
7	Humeral part with brown macula (Fig. 9); male genitalia with penis guide longer than parameres (Fig. 35); penis as in Fig. 36. Form oblong, body length 4.3–5.0 mm	E. undulatus Weise
–	Humeral part with orange macula surrounding a black round spot (Fig. 3). Body form oblong, 3.0–5.0 mm long)	E. ericae Crotch
8	Elytra brown without markings; (Body subcircular, 3.5–4.0 mm long)	E. quadripustulatus Linnaeus
–	Each elytron with 4 nearly equally sized, small, black spots similarly distributed	9
9	Pronotum reddish orange, with a medio basal ungulate black spot (Fig. 5). Tarsal claw simple (Fig. 37). Male genitalia with penis guide as long as parameres (Fig. 38); penis as in Fig. 39. Body nearly of spindle form, 2.8–4.5 mm long	E. octosignatus Gebler
–	Pronotum entirely black except for dark bordering oflateral and anterior margins (Fig. 4).Tarsal claw with small basal tooth (Fig. 40). Male genitalia with penis guide distinctly shorter than parameres (Figs 41, 42); penis as in Fig. 43. Body subcircular, 4.0–5.0 mm long	E. gebleri Weise
10	Body pubescent	11
–	Body glabrous	12
11	Body covered with dense, moderately long setae (Fig. 13). Male genitalia with penis guide shorter than parameres (Figs 44, 45); penis as in Figs 46, 47. Body short oval to nearly circular, 2.8–2.9 mm long	P. pubescens Küster
–	Body apparently glabrous, but actually with minute sparse setae particularly at pronotum (Fig. 10). Form oblong, 2.2–2.7 mm long	P. melanocephalus Zubkov
12	Pronotum yellow (Fig. 11). Male genitalia as in Figs 48, 49, 50. Body oval and highly convex, 3.8–4.2 mm long	P. nigripennis Erichson
–	Pronotum black with yellow lateral margins (Fig. 12). Male genitalia as in Figs 51–55. Body broadly oval, moderately convex, 3.1–4.5 mm long	P. nigromaculatus Goeze