Checklist
Print
Checklist
Review of the tribe Chilocorini Mulsant from Iran (Coleoptera, Coccinellidae)
expand article infoAmir Biranvand, Wioletta Tomaszewska§, Wenjing Li|, Vincent Nicolas, Jahanshir Shakarami#, Lida Fekrat¤, Shahram Hesami«
‡ Islamic Azad University, Khorramabad, Iran
§ Polish Academy of Sciences, Warsaw, Poland
| Guangdong Academy of Agricultural Sciences, Guangzhou, China
¶ Unaffiliated, Saint-Junien, France
# Lorestan University, Lorestan, Iran
¤ Ferdowsi University of Mashhad, Mashhad, Iran
« Islamic Azad University, Shiraz, Iran
Open Access

Abstract

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.

Keywords

checklist, Chilocorini, Coccinelloidea, Iran, review

Introduction

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

The tribe Chilocorini 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: Brumoides Chapin, 1965, Chilocorus Leach, 1815, Chujochilus Sasaji, 2005, Exochomus Redtenbacher, 1843, Parexochomus Barovsky, 1922, Phaenochilus Weise, 1895, Priscibrumus Kovár, 1995, Simmondsius Ahmad & Ghani, 1966 and Xanthocorus Miyatake, 1970) (Kovář 2007).

Although most members of Chilocorini 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 Chilocorinae by Kovář (2007), the species of the genus Brumus Mulsant, 1850 were transferred to Exochomus Redtenbacher and the subgenus Parexochomus of Exochomus was considered as a valid genus, under the name of Parexochomus Barovsky, 1922. This classification was followed by Nedvěd and Kovář (2012). Moreover, according to Ślipiński (2007), the subfamily Chilocorinae Mulsant was dissolved and all tribes were lumped into the subfamily Coccinellinae. 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 Chilocorini. 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 Chilocorini.

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 examined 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 Chilocorini is updated, which includes 13 species belonging to four genera (Brumoides, Chilocorus, Exochomus, and Parexochomus).

Although there are some records of Exochomus flavipes 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 Parexochomus nigromaculatus (Goeze, 1777). Parexochomus flavipes is morphologically similar to P. nigromaculatus but is distinguished from it by the male genitalia, and P. flavipes 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, Brumus undecempunctata L. and Chilocorus stigma (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 Brumus undecempunctata, while Chilocorus stigma 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 Exochomus kiritshenkoi Barovsky, 1922 from Iran (Shahrood, H. Christoph leg.). There are also specimens in Zoologichesky Institut (Akademii Nauk SSSR) in St. Petersburg, labeled as E. kiritshenkoi which had been collected from Iran (Shahrood, H. Christoph leg). Kovář (1995) however identified these specimens as E. gebleri Weise.

Data on the presence of E. bifasciatus 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 Chilocorini.

Subfamily Coccinellinae Latreille, 1807

Chilocorini 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).

Key to the Iranian species and genera of Chilocorini

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

Updated checklist of the Iranian species of Chilocorini

Brumoides Chapin, 1965

Brumoides Chapin, 1965: 237. Type species: Coccinella suturalis Fabricius, 1798, by original designation.

Diagnosis

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 Brumoides have been associated with mealybugs (Ślipiński 2007), namely Coccidohystrix insolita (Hemiptera: Pseudococcidae), Dactylopius confusus (Hemiptera: Dactylopiidae), Ferrisia virgata (Hemiptera: Pseudococcidae), and Phenacoccus solenopsis (Hemiptera: Pseudoccocidae) (Gordon 1985, Gautam 1990, Hodek and Honěk 2009, Arif et al. 2012, Giorgi et al. 2014). Some species of this genus, such as Brumoides suturalis (F.) feed on some whitefly species, such as Aleurolobus barodensis (Maskell) (Inayatullah 1984, Hodek and Honěk 2009) in addition to feeding on some coccids, such as F. virgata (better for development) and Planococcus pacificus (better for oviposition) (Gautam 1990).

Brumoides adenensis Fürsch, 1987

Figs 1, 21, 24–26

Brumoides adenensis 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.

Chilocorus Leach, 1815

Chilocorus Leach, 1815: 116. Type species: Coccinella cacti Linnaeus, 1767, by monotypy.

Diagnosis

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).

Ecology

Although various scale insects are primary hosts of Chilocorus (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 Chilocorus stigma (Say) which feed on some whitefly species, such as Aleurocanthus woglumi Ashby (Dowell and Cherry 1981, Hodek and Honěk 2009).

Figures 1–9. 

Dorsal habitus of Chilocorini species. 1 Brumoides adenensis Fürsch 2 Chilocorus bipustulatus Linnaeus 3 E. ericae Crotch 4 E. gebleri Weise 5 E. octosignatus Gebler 6 E. quadriguttatus Fleischer 7, 8 E. quadripustulatus Linnaeus 9 E. undulatus Weise.

Chilocorus bipustulatus (Linnaeus, 1758)

Figs 2, 16, 15–20

Coccinella bipustulata Linnaeus, 1758: 367.

Coccinella fasciata Müller, 1776: 68.

Coccinella transversoguttata Börner, 1776: 250.

Coccinella frontalis Thunberg, 1792: 105. [Homonym]

Coccinella testudo Florencourt Chassot, 1796: 214.

Coccinella strigata Fabricius, 1798: 79. [Homonym]

Chilocorus olivetorum Costa, 1839: 104.

Chilocorus minor 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.

General distribution

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

Distribution in Iran

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

Ecology

This species feeds on a wide range of Hemiptera species: Agonoscena pistaciae (Psyllidae), Aonidiella orientalis (Diaspididae), Bemisia tabaci (Aleyrodidae), Chrysomphalus dictyospermi (Diaspididae), Eulecanium prunastri (Coccidae), Euphyllura olivina (Psyllidae), Salicola kermanensis (Diaspididae), Lepidosaphes malicola (Diaspididae), Leucaspis pusilla (Diaspididae), Maconellicoccus hirsutus (Pseudococcidae), Ommatissus binotatus lybicus (Tropiduchidae), Parlatoria blanchardi (Diaspididae), Parlatoria oleae (Diaspididae), Phloeomyzus passerinii (Aphididae), Planococcus citri (Pseudococcidae), Pseudaulacaspis pentagona (Diaspididae), Psylla pyricola (Psyllidae) (Moddarres-Awal 2012) and other coccids, particularly armoured scales (Hodek 1973, Stansly 1984).

Figures 10–13. 

Dorsal habitus of Chilocorini species. 10 Parexochomus melanocephalus Zubkov 11 P. nigripennis Erichson 12 P. nigromaculatus Goeze 13 P. pubescens Küster.

Exochomus Redtenbacher, 1843

Exochomus Redtenbacher, 1843:11. Type species: Coccinella quadripustulata Linnaeus, 1758, by subsequent designation of Thomson, 1859.

Diagnosis

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).

Ecology

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., Exochomus bimaculosus Mulsant which feeds on Bemisia tabaci (Gennadius) (Yigit 1992, Leite et al. 2003, Hodek and Honěk 2009).

Figures 14–20. 

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

Exochomus bifasciatus Barovsky, 1927

Exochomus bifasciatus Barovsky, 1927: 200.

General distribution

China, Iran, Kazakhstan (Kovář 2007).

Distribution in Iran

Iran (Kovář 2007) – no specific distribution provided.

Exochomus ericae Crotch, 1874

Fig. 3

Exochomus ericae Crotch, 1874: 193.

Chilocorus nigropictus Fairmaire, 1876: 94.

Chilocorus picturatus Fairmaire, 1876: 94.

Exochomus anchorifer Allard, 1870: 9.

General distribution

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

Distribution in Iran

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

Remarks

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

Figures 21–31. 

Morphological details and male genitalia of Chilocorini species. 21, 24–26 Brumoides adenensis: 21 Abdominal postcoxal lines 24 Antenna 25 Tegmen 26 Penis apex 22 Parexochomus pubescens: Abdominal postcoxal lines 23, 28 P. nigripennis: 23 Hind leg 28 Antenna 27 Exochomus undulatus: Antenna 29–31 E. quadriguttatus: 29 Tegmen, ventral view 31 Tegmen, lateral view 30 Penis apex.

Exochomus gebleri Weise, 1885

Figs 4, 40–43

Exochomus gebleri Weise, 1885: 55.

Material examined

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

General distribution

Afghanistan, Iran, Turkey (Kovář 2007).

Distribution in Iran

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

Exochomus octosignatus (Gebler, 1830)

Figs 5, 37–39

Coccinella octosignata Gebler, 1830: 225.

Coccinella deserta Motschulsky, 1840: 175.

Coccinella desertorum Gebler, 1841: 376.

Brumus lasioides Weise, 1879: 135.

Brumus conjunctus Fleischer, 1900: 118.

General distribution

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

Distribution in Iran

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).

Ecology

This species feeds on the mealybugs Phenacoccus aceris and Planococcus citri (Pseudococcidae) (Moddarres-Awal 2012).

Figures 32–39. 

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

Exochomus quadriguttatus Fleischer, 1900

Figs 6, 29–31

Exochomus quadriguttatus Fleischer, 1900: 118.

Exochomus cordiformis Roubal, 1926: 245.

Exochomus illaesicollis Roubal, 1927: 135.

Material examined

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

General distribution

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

Distribution in Iran

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

Exochomus quadripustulatus (Linnaeus, 1758)

Figs 7–8, 32–34

Coccinella quadripustulata Linnaeus, 1758: 367.

Coccinella lunulata Gmelin, 1790: 1662.

Coccinella quadriverrucata Fabricius, 1792: 288.

Coccinella cassidoides Donovan, 1798: 74.

Coccinella varia Schrank, 1798: 444.

Coccinella distincta Brullé, 1832: 273

Coccinella iberica Motschulsky, 1837: 422.

Coccinella floralis Motschulsky, 1837: 423.

Exochomus haematideus Costa, 1849: 62.

Exochomus unicolor Schaufuss, 1862: 50

Exochomus sexpustulatus Kraatz, 1873:192

Exochomus bilunulatus Weise, 1879: 133.

Exochomus koltzei Weise, 1879: 134.

Exochomus reitteri Schneider, 1881: 16

Exochomus vittatus Fuente, 1910: 444

Material examined

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.

General distribution

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

Distribution in Iran

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

Ecology

This species feeds on various species of Hemiptera, namely: Aonidiella orientalis (Diaspididae), Aphis fabae (Aphididae), Callaphis juglandis (Aphididae), Chromaphis juglandicola (Aphididae), Eriosoma lanigerum (Aphididae), Eulecanium prunastri (Coccidae), Euphyllura olivina (Psyllidae), Maconellicoccus hirsutus (Pseudococcidae), Parlatoria oleae (Diaspididae), Psylla pyricola (Psyllidae), Saissetia oleae (Coccidae) (Moddarres-Awal 2012), and other aphids and Coccidae (Uygun 1981, Ülgentürk and Toros 2001, Kaydan et al. 2006, Kaydan et al. 2012).

Figures 40–47. 

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

Exochomus undulatus Weise, 1878

Figs 9, 27, 35–36

Exochomus undulatus Weise, 1878: 93

Material examined

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

General distribution

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

Distribution in Iran

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).

Ecology

This species feeds usually on Euphyllura olivina (Hemiptera: Psyllidae) (Moddarres-Awal 2012).

Parexochomus Barovsky, 1922

Exochomus (Parexochomus) Barovsky, 1922: 293. Type species: Exochomus pubescens Küster, 1848, by subsequent designation of Chapin 1965.

Parexochomus: Kovář 2007: 595.

Diagnosis

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).

Ecology

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

Parexochomus melanocephalus (Zubkov, 1833)

Fig. 10

Coccinella melanocephala Zubkov, 1833: 339.

Exochomus russicollis Mulsant, 1850: 1033.

General distribution

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

Distribution in Iran

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).

Parexochomus nigripennis (Erichson, 1843)

Figs 11, 23, 28, 48–50

Chilocorus nigripennis Erichson, 1843: 267.

Exochomus xanthoderus Fairmaire, 1864: 648.

Material examined

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

General distribution

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

Distribution in Iran

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).

Ecology

This species feeds usually on the following hemipterans: Acanthococcus abaii (Eriococcidae), Agonoscena pistaciae (Psyllidae), Bemisia tabaci (Aleyrodidae) (Moddarres-Awal 2012).

Figures 48–55. 

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

Parexochomus nigromaculatus (Goeze, 1777)

Figs 12, 51–55

Coccinella nigromaculata Goeze, 1777: 248. Coccinella testudinare Geoffroy in Fourcroy, 1785: 151. Coccinella aurita Scriba, 1791: 101. Coccinella humerale Townson, 1800: 167.

Chilocorus rufipes Stephens, 1832: 375. Exochomus collaris Küster, 1849: 100. Exochomus pyrenaeus Kraatz, 1873: 194.

Material examined

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.

General distribution

Palaearctic region (Duverger 1983, Kovář 2007).

Distribution in Iran

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

Ecology

This species feeds usually on the following species of Hemiptera: Agonoscena pistaciae (Psyllidae), Aonidiella orientalis (Diaspididae), Bemisia tabaci (Aleyrodidae), Diuraphis noxia (Aphididae), Eulecanium prunastri (Coccidae), Euphyllura olivina (Psyllidae), Maconellicoccus hirsutus (Pseudococcidae), Therioaphis maculata (Aphididae) (Moddarres-Awal 2012) and other aphids and Coccidae (Uygun 1981, Atlıhan and Özgökçe 2002, Kaydan et al. 2012).

Parexochomus pubescens (Küster, 1848)

Figs 13, 22, 44–47

Exochomus pubescens Küster, 1848: 94

Exochomus apicatus Fairmaire, 1884: 59.

Exochomus circumcinctus Sahlberg, 1903: 36.

Platynaspis flavilabris Motschulsky, 1849: 155.

Platynaspis flavilabris Mulsant, 1850b: 947. [Homonym]

Exochomus gestroi Fairmaire, 1875: 540.

Exochomus lugubrivestis Mulsant, 1853: 194.

Exochomus saharae Sicard, 1929: 60

Material examined

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

General distribution

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

Distribution in Iran

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).

Ecology

This species feeds on Bemisia tabaci (Hemiptera: Aleyrodidae) and Tetranychus turkestani (Acari) (Moddarres-Awal 2012).

Acknowledgements

Adam Ślipiński (Australian National Insect Collection, CSIRO, Canberra, Australia) and Oldřich Nedvěd (Faculty of Science, University of South Bohemia, České Budějovice, Czech Republic) read an early version of this manuscript providing valuable suggestions. Two anonymous reviewers are acknowledged for their valuable comments.

References

  • Abdolahi Mesbah R, Nozari J, Allahyari H, Zare Khormizi M (2016) Checklist and distribution of lady beetles (Coleoptera: Coccinellidae) in Iran. Iranian Journal of Animal Biosystematics 12: 1–35.
  • Ahmadi A, Jafari R, Vafai R (2012) The faunistic survey of ladybird (Cole., Coccinellidae) in orchards and crops Markazi and shrub. Proceedings of the 20th Iranian Plant Protection Congress (Shiraz), 139 pp.
  • Ansari pour A, Shakarami J (2011) Study of ladybirds (Col: Coccinellidae) in Khorramabad district and the first report of Hyperaspis quadrimaculata (Redtenbacher, 1844) for Iranian fauna. Life Sciences 8(3): 488–495.
  • Arif MI, Rafiq M, Wazir S, Mehmood N, Ghaffar A (2012) Studies on cotton mealybug, Phenacoccus solenopsis (Pseudococcidae: Homoptera), and its natural enemies in Punjab, Pakistan. International Journal of Agricultureand Biology 14: 557–562.
  • Atlıhan R, Özgökçe MS (2002) Development, fecundity and prey consumption of Exochomus nigromaculatus feeding on Hyalopterus pruni. Phytoparasitica 30: 443–450. https://doi.org/10.1007/BF02979748
  • Barovsky V (1922) Revisio specierum palaearticarum Coccinellidarum generis Exochomus Redtb. Annuaire du Musée Zoologique de l’Académie des Sciences de Russie 23: 289–303.
  • Barovsky V (1927) Revisio specierum palaearcticum Coccinellidarum generis Brumus Muls. (Coleoptera). Annuaire du Musée Zoologique de l’Académie de Sciences de l’URSS 28: 194–200.
  • Bagheri MR, Mossadegh MS (1996) Fauna of Coccinellid beetles (Col.: Coccinellidae) in Chaharmahal and Bakhtiari province, One part of subfamilies: Coccinellinae and Chilocorinae. Journal of Agricultural Sciences Islamic Azad University 7: 29–35.
  • Bielawski R (1984) Coccinellidae (Coleoptera) of Mongolia. Annales Zoologici 38: 281–460.
  • Boerner ICH (1776) Beschreibung eines neuen Insects, des Coccinella transversopunctata. Der Ekonomische Nachricht der Gesellschaft in Schlesien 4: 250.
  • Brullé GA (1832) IVe Classe. Insectes. In: Brullé GA, Guérin-Menéville FM (Eds) Expédition scientifique de Morée. Section des sciences physiques. Tome 3.I.re partie. Zoologie. Deuxième section. Des animaux ariculés. FL Lefraut, Paris et Strasbourg, 400 pp. [27–52 pls]
  • Canepari C (2011) Contribution to the knowledge of the Coccinellidae of Sardinia (Coleoptera). Conservazione Habitat Invertebrati 5: 501–516.
  • Chapin EA (1965) The genera of the Chilocorini (Coleoptera, Coccinellidae). Bulletin of the Museum of Comparative Zoology 133: 227–271.
  • Costa A (1849) Coleotteri Trimeri. Famigliadegli Coccinellidae. In: Fauna dell regno di Napoli ossia enumerazone di tutti gli animali, che abitano le diverse regioni di questo regno e le acque che le bagnano contenente la descrizionede’nuovi o poco esattamente conosciuti noc figure ricavate da originali viventi e dipinte al nature di Oronzio-Gabrielle Costa. Colleotteri. Parte I.a. Gaetano Sautto, Napoli, 364 pp. [families separately paginated, 1–112, 7 pls]
  • Costa OG (1839) Monographia degli insecti dell’ulivo e delle olive. In: Corrispondenza Zoologica destinata a diffondere nel rogno delle due Sicilie tutto ciò che si va discuoprendo entro e fuori Europa risguardante la Zoologica in generaleredatta. Vol. 1. Azzolino, Napoli, 104 pp.
  • Donovan E (1798) The natural history of British Insects; explaining them in their several states, with the periods of their transformations, their food, economy and c. together with the history of such minute insects as require investigations by the microscope. Vol. VII. Rivington, London, 96 pp. [pls 217–251]
  • Drea JJ, Gordon RD (1990) Predators. Coccinellidae. In: Rosen D (Ed.) The Armored Scale Insects, their Biology, Natural Enemies and Control. Elsevier, Amsterdam, 19–40.
  • Duverger C (1983) Contribution à la connaissance des Coccinellidae d’Iran. Nouvelle Revue d’Entomologie, Paris, Nov. Ser 13(1): 73–93.
  • Erichson WF (1843) Beitrag zur Insecten-Fauna von Angola, in Besonderer Beziehung zur geographischen Verbreitung der Insecten. Archiv für Naturgeschichte 9: 199–267.
  • Escalona HE, Ślipiński A (2012) Generic revision and phylogeny of Microweiseinae (Coleoptera: Coccinellidae). Systematic Entomology 37: 125–171. https://doi.org/10.1111/j.1365- 3113.2011.00601.x
  • Escalona HE, Zwick A, Li HS, Li J, Wang X, Pang H, Hartley D, Jermiin LS, Nedvěd O, Misof B, Niehuis O, Ślipiński A, Tomaszewska W (2017) Molecular phylogeny reveals extreme food plasticity in evolution of true ladybird beetles (Coleoptera: Coccinellidae: Coccinellini). BMC Evolutionary Biology 17(151): 1–11. https://doi.org/10.1186/s12862-017-1002-3
  • Fabricius JC (1792) Entomologia systematica emendate et aucta. Secundum classes, ordines, genera, species, adjectis synonymis, locis, observationibus, descriptionibus, I, 2. Hafniae, 586 pp. https://doi.org/10.5962/bhl.title.125869
  • Fabricius JC (1798) Supplementum Entomologia systematicae. Hafniae, 572 pp.
  • Fairmaire L (1864) Coléoptères d’Algérie rapportés par M. Ogier de Baulny. Annales de la Société Entomologique de France (4)3[1863]: 637–648.
  • Fairmaire L (1875) Coléoptèresde la Tunisie récoltés par Mr. Abdul Kerim. Annali del Museo Civico di Storia Naturale in Genova 7: 475–540.
  • Fairmaire L (1876) Diagnoses de coléoptères du nord de l’Afrique. Petites Nouvelles Entomologiques 2[1876–1879]: 93–94.
  • Fairmaire L (1884) Descriptions de coléoptères recueillis par le Baron Bonnaire en Algérie. Bulletin ou Comptes–Rendus des Séances de la Société Entomologique de Belgique 28: lix-lxx.
  • Fallahzadeh M, Abdimaleki R, Saghaei N (2013) Contribution to the knowledge of the ladybird beetles (Coleoptera, Coccinellidae), predators of mealybugs (Hemiptera, Pseudococcidae) in Hormozgan province, Southern Iran. Linzer Biologische Beiträge 45(1): 673–679.
  • Florencourt Chassot K (1796) Verzeichniss der Insecten Gottnischer Gegend. In: Meyer FAA (Ed. ) Zoologisches Archiv (Leipzig) Band 1: 197–244.
  • Fuente JM (1910) Datos para la fauna de la provincial de Ciudad Real. XXI. Boletín de la Real Sociedad Española de Historia Natural 10: 444–449.
  • Fürsch H (1961) Revision der afrikanischen Arten um Exochomus flavipes Thunb. Col. Cocc. Entomologische Arbeiten aus dem Museum G. Frey. Tutzingbei München 12: 68–92.
  • Fürsch H (1987) Neue Coccinelliden aus Ethiopien. Folia Entomologica Hungarica 48: 39–44.
  • Gautam RD (1990) Mass-multiplication technique of coccinellid predator, ladybird beetle (Brumoides suturalis). Indian Journal of Agricultural Sciences 60: 747–750.
  • Gebler FA von (1830) Bemerkungen über die Insecten Sibiriens, vorzüglich des Altai (Part 3). In: Ledebour CF (Ed.) Reise durch Altai-Gebirge und die songorische Kirgisen-Steppe. Auf Kosten der Keiserlichen Universität Dorpat im Jahre 1826 in Begleitung der Herren D. Carl Anton Miecherund, D. Alexander von Bunge. Zweiter Theil. G. Reimer, Berlin, 1–228.
  • Gebler FA (1841) Characteristik mehrerer neuen sibirischen Coleopteren. Bulletin Scientifique de l’Académie Impériale des Sciences St. Péterbourg 8: 369–376.
  • Geoffroy EL (1785) [new taxa]. In: Fourcroy AF de (Ed.) Entomologia parisiensis; sive Catalogus Insectorum quae in Agro Parisiensi reperiuntur; Secundum methodum Geoffraeanam in sections, genera et species distributus: cui addita sunt nomina trivialia et fere trecentae novae species Parisiis, Serpentineis. Pars 1: i–viii + 1–231; pars 2, 233–544.
  • Ghanbari A, Sadeghi SE, Ladan Moghadam AR, Fakhredini M (2012) An Investigation on predaceous coccinellid`s fauna, their distribution and determining dominant species on shading trees and shrubs in green spaces and parks of 16th Tehran municipal zone. Proceedings of the 20th Iranian Plant Protection Congress (Shiraz), 179.
  • Giorgi JA, Vandenberg NJ, McHugh JV, Forrester JA, Ślipiński SA, Miller KB, Shapiro LR, Whiting MF (2009) The evolution of food preferences in Coccinellidae. Biological Control 51: 215–231. https://doi.org/10.1016/j.biocontrol.2009.05.019
  • Giorgi JA, Lima MS, Vandenberg NJ (2014) The first record of Brumoides foudrasii (Mulsant) (Coleoptera: Coccinellidae: Chilocorini) from South America, with notes on its biology. The Coleopterists Bulletin 68(2): 336–338.
  • Gmelin JF (1790) Caroli a Linné Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Edito decimatertia, aucta, reformata.Tomus 1. Pars IV. Classis V. Insecta. Georg Emanuel Beer, Lipsiae, 1517–2224.
  • Goeze JAE (1777) Entomologische Beiträgezu des Ritter Linné, zwölften Ausgabe des Natursystems. Leipzig: Weidmanns Erben und Reich 1st Ed., 736 pp.
  • Gordon RD (1985) The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society 93: 1–912.
  • Hajizadeh J, Jalili Sendi J, Peyrovi H (2003) Introducing a part of the coccinellids (Col., Coccinellidae) fauna of Guilanprovince. Journal of Agricultural Sciences and Natural Resources 9: 99–111.
  • Inayatullah C (1984) Sugarcane aleurodids, Aleurolobus barodensis (Maskell) and Neomaskellia andropogonis Corbett (Hom.: Aleyrodidae), and their natural enemies in Pakistan. International Journal of Tropical Insect Science 5(4): 279–282. https://doi.org/10.1017/S1742758400001570
  • Jafari R, Kamali K (2007) Faunistic study of ladybird (Col.: Coccinellidae) in Lorestan province and report of new records in Iran. New Findings in Agriculture 4: 349–359.
  • Kaydan MB, Kilincer N, Uygun N, Japosvilli G, Gaimari S (2006) Parasitoids and predators of Pseudococcidae (Homoptera: Coccoidea) in Ankara, Turkey. Phytoparasitica 34(4): 331–337. https://doi.org/10.1007/BF02981018
  • Kaydan MB, Atlihan R, Uygun N, Şenal D (2012) Coccinellid (Coleoptera: Coccinellidae) species feeding on coccoids (Hemiptera: Coccoidea) in Van Lake Basin, Turkey. Türkiye Biyolojik Mücadele Derneği 3(1): 37–46.
  • Kovář I (1995) Revision of the genera Brumus Muls. and Exochomus Redtb.(Coleoptera, Coccinellidae) of the Palaearctic Region. Part I. Acta Entomologica Musei Nationalis Pragae 44: 5–124.
  • Kovář I (2007) Coccinellidae. In: Löbl I, Smetana A (Eds) Catalogue of Palaearctic Coleoptera. Volume 4. Elateroidea, Derodontoidea, Bostrichoidea, Lymexyloidea, Cleroidea, Cucujoidea. Apollo Books, Stentrup, Denmark, 71–74, 568–630.
  • Kraatz G (1873) Revision der europäischen Arten der Coccinelliden-Gattung Exochomus Redtb. Berliner Entomologische Zeitschrift 17: 191–194.
  • Küster HC (1848) Die Käfer Europa’s. Nach der Natur beschrieben. Mit Beiträgen mehrerer Entomologen. 13. Heft. Bauer and Raspe, Nürnberg, 100 pp. [2 pls]
  • Küster HC (1849) Die Kafer Europa’s. Nach der Naturbeschrieben. Mit Beiträgen mehrerer Entomologen. 17. Heft. Bauer and Rapse, Nürnberg, 100 pp. [2 pls]
  • Latreille PA (1807) Genera crustaceorum et insectorum secundum ordinem naturalem in familias disposita, inconibus exemplisque plurimis explicata. Tomus Tertius. A. Koenig, Parisiset Argentorati.
  • Leach WE (1815) Entomology. In: Brewster D (ed) The Edinburgh Encyclopedia. Balfour, Edinburgh 9: 572–172.
  • Li W, Huo L, Wang X, Chen X, Ren S (2015) The genera Exochomus Redtenbacher, 1843 and Parexochomus Barovsky, 1922 (Coleoptera: Coccinellidae: Chilocorini) from China, with descriptions of two new species. The Pan-Pacific Entomologist 91(4): 291–304. https://doi.org/10.3956/2015-91.4.291
  • Li W, Huo L, Chen X, Ren S, Wang X (2017) A new species of the genus Phaenochilus Weise from China (Coleoptera, Coccinellidae, Chilocorini). ZooKeys 644: 33–41. https://doi.org/10.3897/zookeys.644.9825
  • Linnaeus C (1758) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species cum Characteribus, Differentiis, Synonymis, Locis. Holmiae: Laurentii Salvii. Editio Decima (reformata) Edn., 823 pp.
  • Linnaeus C (1767) Systema Naturae, per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tom I. Pars I. Editio Decima tertia, ad Editionem duodecimam reformatam Holmiensem.Vindobonae, Typis Ioannis Thomae nob. de Trattern, 533–1364.
  • Łączyński P, Tomaszewska W (2012) Chapinaria, new genus of Chilocorini for Endochilus meridionalis Sicard from Africa (Coleoptera: Coccinellidae). Annales Zoologici 62: 1–9. https://doi.org/10.3161/000345412X633658
  • Mahghari H, Ostovan H (2006) Predator arthropods, fauna of whiteflies (Homoptera: Aleyrodidae) in Mazandaran and Golestan provinces and their feeding efficiency. Journal of Agricultural Sciences and Natural Resources 12(6): 171–180.
  • Mader L (1955) Evidenz der palaarktisehen Coceinelliden und ihrer Aberrationen in Wort and Bild. Entomologische Arbeiten aus dem Museum G. Frey Tutzing bei München 6: 764–1035.
  • Magro A, Lecompte E, Magné F, Hemptinne JL, Crouau-Roy B (2010) Phylogeny of ladybirds (Coleoptera: Coccinellidae): are the subfamilies monophyletic? Molecular Phylogenetics and Evolution 54(3): 833–848. https://doi.org/10.1016/j.ympev.2009.10.022
  • Miyatake M (1970) The East-Asian coccinellid-beetles preserved in the California Academy of Sciences, tribe Chilocorini. Memoirs of the College of Agriculture, Ehime University 14: 19–56.
  • Moddarres-Awal M (2012) List of agricultural pests and their natural enemies in Iran. Third ed. Ferdowsi University Press, Mashhad, Iran, 759 pp.
  • Montazeri MM, Mossadegh MS (1995) The coccinellids (Coleoptera) fauna of Gorgan plain and Gonbad Kavus. Proceedings of the 12th Iranian Plant Protection Congress (Karaj), 325.
  • Moodi S, Mossadegh MS (1995) The coccinellids (Coleoptera) of southeast Khorasan province, Iran. Proceedings of the 12th Iranian Plant Protection Congress (Karaj), 326.
  • Motschulsky V (1837) Description de quelques Coléoptères recueillis dans un voyage au Caucase et dans les provinces transcaucasiennes russes en 1834 et 1835. Nouveaux Memoirés de la Société Impériale des Naturalistes de Moscou 5: 413–425. [pl. 16]
  • Motschulsky V (1840) Insectes du Caucase et des provinces transcaucasiennes recueillis decritis (Continuation). Bulletin de la Société Impériale des Naturalistes de Moscou 13: 169–180. [pl. 4]
  • Motschulsky V (1849) Coléoptères reçus d’un voyage de M. Handshuch dans le Midi de l’Espagne, énumérés et suivis de notes. Bulletin de la Société Imperiale des Naturalistes de Moscou 22(3): 52–163.
  • Mulsant E (1846) Histoire Naturelle des Coléoptères de France. Sulcicolles-Sécuripalpes. Maison, Paris, xxiv + 26 pp + 280 pp + 1 pl.
  • Mulsant E (1850) Species des Coléoptères Trimères Sécuripalpes. Annales des Sciencies Physiques et Naturelles, d’Agriculture et d’Industrie, publiées par la Société nationale d’Agriculture, etc. , de Lyon, Deuxième Série, 1104 pp.
  • Mulsant E (1853) Supplement a la Monographie des Coleopteres Trimeres Securipalpes. Annales de la Société Linnéenne de Lyon (Nouvelle Série) 1: 129–333. https://doi.org/10.5962/bhl.title.60609
  • Müller OF (1776) Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae indigenarum characteres, nomina et synonyma imprimis popularium. Hallageriis, Hafniae, xxxii + 282 pp. https://doi.org/10.5962/bhl.title.13268
  • Nedvěd O, Kovář I (2012) Phylogeny and classification. In: Hodek I, Van Emden HF, Honek A (Eds) Ecology and Behaviour of the Ladybird Beetles (Coccinellidae). Blackwell Publishing Ltd., Chichester, UK, 1–12. https://doi.org/10.1002/9781118223208.ch1
  • Ponsonby DJ, Copland JW (1997) Predators. Coccinellidae and other Coleoptera. In: Ben-Dov Y, Hodgson CJ (Eds) Soft Scale Insects-their biology, natural enemies and control, Vol. 7B. Elsevier, Amsterdam, 29–60. https://doi.org/10.1016/S1572-4379(97)80076-0
  • Raimundo AAC, van Harten A (2000) An annotated checklist of the Coccinellidae (Insecta: Coleoptera) of Yemen. Fauna of Arabia 18: 211–243.
  • Raimundo AC, Fürsch H, Van Harten A (2008) Order Coleoptera, family Coccinellidae. Arthropod fauna of the UAE 1: 217–239.
  • Redtenbacher L (1843) Tetamen dispositionis generum et specierum Coleopterorum Pseudotrimeorum Archiducatus Austriae.Vindobone: Disert. Inaug., 32 pp.
  • Robertson J, Ślipiński A, Moulton M, Shockley FW, Giorgi A, Lord NP, McKenna DD, Tomaszewska W, Forrester J, Miller KB, Whiting MF, McHugh JV (2015) Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinelloidea (Coleoptera: Cucujiformia). Systematic Entomology 40: 745–778. https://doi.org/10.1111/syen.12138
  • Roubal J (1926) Seieben neue Coleopteren aus der Paläarktischen Region. Coleopterologisches Centrablatt 1: 244–249
  • Roubal J (1927) Sex nova Coleoptera palaearctica. Boletín de la Real Sociedad Española de Historia Natural 27: 134–136
  • Sahlberg JR (1903) Messis hiemalis Coleopterorum Corcyraeorum. Enumeratio Coleopterorum mensibus Novembri-Februario 1895–1896 et 1898–1899 nec non primo vere 1896 in insula Corcyra collectorum. Öfversigtaf Finska Vetenskaps-Societetens Förhandlingar 45(11) [1902–1903]: 1–87.
  • Salehi T, Pashaei Rad SH, Mehrnejad MR, Shokri MR (2011) Ladybirds associated with pistachio trees in part of Kerman Province, Iran (Coleoptera: Coccinellidae). Iranian Journal of Animal Biosystematics 7(2): 157–169.
  • Sasaji H (2005) Additional revision of the tribe Chilocorini (Coleoptera: Coccinellidae) of Japan. Elytra 33(1): 61–68.
  • Say T (1835) Descriptions of new North American Coleopterous insects, and observations on some already described. Boston Journal of Natural History 1: 151–203.
  • Schaufuss LW (1862) Beschreibungen von neuen Käferarten. Sitzungsberichte und Abhandlungen der Naturwissenschaflichen Gesselschaft Isis zu Dresden 1861: 47–51.
  • Scriba LG (1791) Beiträgezu der Insecten-Geschichte. Zweites Heft. Frankfurt: Varrentrapp and Wenner, 69–194. [pls 7–12]
  • Schneider WG (1881) Einigeneue Varietäten von Coccinellidaen. Zeitschriftfür Entomologie (N.F. ) (Breslau) 8: 10–16.
  • Schrank F von Paula (1798) Fauna Boica. Durchgedachte Geschichte der in Baierneinheimischen und zahmen Thiere. Erster Band, zweyte Abtheilung. Nürnberg: Stein’sche Buchhgandlung, 293–720.
  • Seago AE, Giorgi JA, Li J, Ślipiński A (2011) Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetics and Evolution 60: 137–151. https://doi.org/10.1016/j. ympev. 2011.03.015
  • Sicard A (1929) Description de deux espèces nouvelles de coccinellides paléarctiques. Bulletin de la Societéd Histoire Naturelle de l’Afrique du Nord 20: 60–62.
  • Stansly PA (1984) Introduction and evaluation of Chilocorus bipustulatus (Col.: Coccinellidae) for control of Parlatoria blanchardi (Hom.: Diaspididae) in date groves of Niger. Entomophaga 29(1): 29–39. https://doi.org/10.1007/BF02372206
  • Stephens JF (1832) Illustrations of British entomology or, a synopsis of indigenous insects: containing their generic and specific distinctions; with an account of their metamorphoses, times of appearance, localities, food, and economy, as far as practicable. Mandibulata. Vol. 4. Baldwin & Cardock, London, 367–413.
  • Szawaryn K, Bocak L, Ślipiński A, Escalona HE, Tomaszewska W (2015) Phylogeny and evolution of phytophagous ladybird beetles (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera. Systematic Entomology 40: 547–569. Https://doi.org/10.1111/syen.12121
  • Ślipiński A, Giorgi JA (2006) Revision of the Australian Coccinellidae (Coleoptera). Part 6. Tribe Chilocorini. Annales Zoologici 56: 265–304.
  • Ślipiński A (2007) Australian ladybird beetles (Coleoptera: Coccinellidae) their biology and classification. ABRS, Canberra, 286 pp.
  • Tavakoli M, Ansari pour A, Pirhadi A, Pirozi F (2014) Study of fauna ladybirds (Col., Coccinellidae) on pastures Astragalus adscendens (Fabaceae) in Lorestan province. Journal of Entomological Research 6: 287–297.
  • Thomson GG (1859) Skandinaviens Coleoptera, Synoptiskt Bearbetade. Tom I. Berlingska Boktryckeriet, Lund, 290 pp.
  • Thunberg CP (1781) Dissertatio entomologica novas Insectorum species, sisten scujus partem primam. J. Edman, Upsaliae, 28 pp.
  • Thunberg CP (1792) Description esinsectorumsvecicorum. Nova Acta Regiae Societatis Scientiarum Upsaliensis 5: 85–119.
  • Tomaszewska W, Szawaryn K (2016) Epilachnini (Coleoptera: Coccinellidae). A Revision of the World Genera. Journal of Insect Science 16(1): 101; 1–91. https://doi.org/10.1093/jisesa/iew082
  • Uygun N (1981) Taxonomic research on Coccinellidae (Coleoptera) fauna of Turkey. Çukurova University, Faculty of Agriculture, Publications No: 157, Adana, Turkey 48: 110.
  • Ülgentürk S, Toros S (2001) Natural enemies of the oak scale insect, Eulecanium ciliatum (Douglas) (Hemiptera: Coccidae) in Turkey. Entomologica 33: 219–224.
  • Weise J (1878) [New taxa]. In: Schneider O, Leder H (Eds) Beiträge zur Kenntnis der kaukasischen Käferfauna. Verhandlungen des Naturforschenden Vereins in Brunn, 16[1877]: 1–359. [3 pls]
  • Weise J (1879) Bestimmungs-Tabellen der europaischen Coleopteren II. Coccinellidae. Zeitschrift fur Entomologie (N.F. ), Breslau, 7: 88–156.
  • Weise J (1885) Coccinellidae. Bestimmungs-Tabellen der europäischen Coleopteren. II. Auflge mit Berücksichtung der Arten aus dem Nördlichen Asien. Mödling, 83 pp.
  • Weise J (1895) Neue Coccinelliden, sowie bemerkungen zu bekannten Arten. Annales de la Société Entomologique de Belgique 39: 120–146.
  • Yaghmaei F, Kharrazi Pakdel A (1995) A faunistic survey of Coccinellids in Mashhad region Proceedings of the 12th Iranian Plant Protection Congress, Karaj, 307 pp.
  • Yigit A (1992) Serangium parcesetosum Sicard (Col., Coccinellidae), new record as a citrus whitefly predatory ladybird in Turkey. Türkiye Entomologi Dergisi 16: 163–167.
  • Zubkov B (1833) Nouveaux coléoptères recueillis en Turcménie. Bulletin de la Société Impériale des Naturalistes de Moscou 6: 310–340.
  • Zare Khormizi M, Ostovan H, Fallahzadeh M (2016) A contribution to the fauna of ladybird beetles (Coleoptera: Coccinellidae) from Yazd Province, Iran. Proceedings of the 22th Iranian Plant Protection Congress, Karaj, 448 pp.