Review of the Tribe Hyperaspidini Mulsant (Coleoptera: Coccinellidae) from Iran

Zootaxa 4236 (2): 311–326 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4236.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:01F6A715-AA19-4A2A-AD79-F379372ACC65 Review of the tribe Hyperaspidini Mulsant (Coleoptera: Coccinellidae) from Iran

AMIR BIRANVAND1, WIOLETTA TOMASZEWSKA2,11, OLDŘICH NEDVĚD3,4, MEHDI ZARE KHORMIZI5, VINCENT NICOLAS6, CLAUDIO CANEPARI7, JAHANSHIR SHAKARAMI8, LIDA FEKRAT9 & HELMUT FÜRSCH10 1Young Researchers and Elite Club, Khorramabad Branch, Islamic Azad University, Khorramabad, Iran. E-mail: amir.beiran@gmail 2Museum and Institute of Zoology, Polish Academy of Sciences, Warszawa, Poland. E-mail: [email protected] 3Faculty of Science, University of South Bohemia, Branišovská 1760, CZ-37005 České Budějovice, Czech Republic. E-mail: [email protected] 4Institute of Entomology, Biology Centre, Branišovská 31, 37005 České Budějovice, Czech Republic. 5Department of Entomology, College of Agricultural Sciences, Shiraz Branch, Islamic Azad University, Shiraz, Iran. E-mail: [email protected] 627 Glane, 87200 Saint-Junien, France. E-mail: [email protected] 7Via Venezia 1, I-20097 San Donato Milanese, Milan, Italy. E-mail: [email protected] 8Plant Protection Department, Lorestan University, Agricultural faculty, Khorramabad, Iran. E-mail: [email protected] 9Department of Plant Protection, Faculty of Agriculture, Ferdowsi University of Mashhad, Mashhad, Iran. E-mail: [email protected] 10University Passau, Didactics of Biology, Germany. E-mail: [email protected] 11Corresponding author. E-mail: [email protected]

Abstract

The Iranian species of the tribe Hyperaspidini Mulsant, 1846 (Coleoptera: Coccinellidae) are reviewed. The current list includes 12 species, all placed in a single genus Hyperaspis Chevrolat, 1836. Hyperapsis asiatica Lewis, 1896 and H. pumila Mulsant, 1850 are excluded from the Iranian list of Coccinellidae. Diagnoses of the tribe Hyperaspidini and the genus Hyperaspis are given. Images of adult beetles and diagnostic characters of the male genitalia of all species distributed in Iran are shown. A key to identification of the species is presented. Distribution records are provided for each species along with information on host plants and prey species when available.

Key words: Coccinelloidea, Hyperaspis, checklist, key, lady beetles

Introduction

Superfamily Coccinelloidea (Coleoptera, Polyphaga) comprises taxa formerly placed in the Cerylonid Series of the superfamily Cucujoidea. It includes 15 families: Bothrideridae, Teredidae, Euxestidae, Murmidiidae, Discolomatidae, Cerylonidae, Latridiidae, Akalyptoischiidae, Alexiidae, Corylophidae, Anamorphidae, Endomychidae, Mycetaeidae, Eupsilobiidae and Coccinellidae (Robertson et al. 2015). The family Coccinellidae, with 360 genera and over 6,000 species (Vandenberg 2002; Ślipiński & Tomaszewska 2010; Nedvěd & Kovář 2012), is extremely diverse both morphologically and ecologically. Since the establishment of the family name by Latreille (1807), a vast number of systematic studies of Coccinellidae have proposed various subfamily and tribe-level classifications (Mulsant 1846, 1850; Crotch 1874; Casey 1899; Sasaji 1968, 1971; Gordon 1994). Kovář (1996) recognized 7 subfamilies including Coccinellinae, Coccidulinae, Chilocorinae, Epilachninae, Scymninae, Sticholotidinae and Ortaliinae. Ślipiński (2007) found these classifications phylogenetically unacceptable and argued the basal split of Coccinellidae into two subfamilies Microweiseinae and Coccinellinae. This classification was confirmed by subsequent morphological and molecular

Accepted by M. Gimmel: 19 Dec. 2016; published: 23 Feb. 2017 311 studies (Robertson et al. 2008; Giorgi et al. 2009; Seago et al. 2011; Robertson et al. 2015). Nedvěd & Kovář (2012), however, proposed another classification including 9 subfamilies (Coccinellinae, Coccidulinae, Scymninae, Chilocorinae, Epilachninae, Sticholotidinae, Ortaliinae, Exoplectrinae and Microweiseinae) and 42 tribes for this family. Considering food preferences, Coccinellidae can be divided into two major groups: predaceous and phytophagous. Predaceous Coccinellidae attack a wide range of insects including whiteflies (Aleyrodoidea), aphids (Aphidoidea), chrysomelid beetle larvae (Chrysomelidae), mealybugs (Coccoidea), psyllids (Psylloidea) (Hodek & Honěk 2009; Obrycki et al. 2009; Hodek & Evans 2012) and ants (Formicidae) (Harris 1921; Pope & Lawrence 1990; Samways et al. 1997; Majerus et al. 2007). Stethorini are specialist predators on tetranychid mites (Biddinger et al. 2009) while numerous and diverse Epilachnini (Szawaryn et al. 2015) are phytophagous (Gordon 1985; Giorgi et al. 2009) and preferably feed on plants from the families Solanaceae and Cucurbitaceae (Raimundo & van Harten 2000; Canepari 2011). Mycophagy is observed in former tribes Tytthaspidini and Halyziini (Sutherland & Parrella 2009). Some species of Coccinellini visit flowers and feed on pollen and nectar, usually as a supplement to a predatory diet (Ślipiński 2007). The tribe Hyperaspidini Mulsant is represented in the Palearctic by a single genus (Hyperaspis Chevrolat, 1836) but is much more diverse in the New World with 6 genera (Blaisdelliana Gordon, 1970; Brachiacantha Dejean, 1837; Hyperaspidius Crotch, 1873; Thalassa Mulsant, 1850; Helesius Casey, 1899 and Hyperaspis) known from North America including Mexico and 8 genera (Hyperaspis; Tenuisvalva Duverger, 1989; Thalassa; Menoscelis Mulsant, 1850; Diazonema Weise, 1926; Clypeaspis Gordon & Canepari, 2008; Prognataspis Gordon & Canepari, 2008 and Peruaspis Gordon & Canepari, 2008) from South America (Gordon 1985; Gordon & Canepari 2008). Hyperaspis is a large genus with 48 species recorded from the Palearctic region (Kovář 2007; Raimundo et al. 2006; Bogaert et al. 2012), 103 species from North America and Mexico (Gordon 1985; Gordon & Canepari 2008) and 68 species from South America (Gordon & Canepari 2008). Larvae and adults of Hyperaspis are voracious predators feeding on all development stages of scale insects (e.g., Coccidae and Pseudococcidae) and on aphids (Gordon & Canepari 2008). Specimens of Hyperaspis chevrolati Canepari, 1985 have also been observed in a field full of nests of ants of the genus Messor (Canepari, pers. obs.), which may suggest myrmecophile habits like those known in some other Hyperaspidini (Brachyacantha ursina (Fabricius, 1787), and B. quadripunctata (Melsheimer, 1847), Menoscelis saginata (Mulsant, 1850)) (Berti et al. 1983; Chapin 1966; Orivel et al. 2004). Duverger (1983) reported 6 species of the genus Hyperaspis from Iran, i.e. H. persica Duverger, 1983, H. polita Weise, 1885, H. pumila Mulsant, 1850, H. quadrimaculata Redtenbacher, 1843, H. syriaca Weise, 1885 and H. transversoguttata Weise, 1878. Subsequently, some other records of this genus were reported by Yazdani (1990), Moodi & Mossadegh (1995), Bagheri & Mossadegh (1997), Yaghmaei & Kharazi Pakdel (1995), Kovář (2007), Moddarres-Awal (2012), Sadat Alizadeh et al. (2013) and Zare Khormizi et al. (2014). In this paper, we attempt to complete previous studies on Iranian Hyperaspidini, correct previous possible misidentifications and update the information on Iranian species of the tribe.

Material and methods

Coccinellid samples were collected by sweep net from different localities in Iran between 2013 and 2015 (Fig. 28). Moreover, specimens of Coccinellidae preserved in Hayk Mirzayans Insect Museum of the Iranian Research Institute of Plant Protection and Insect Museum of Islamic Azad University of Shiraz were also examined. External morphology of ladybirds was observed with a stereo microscope (Olympus SZ- ST). Male and female genitalia were dissected, boiled for 10 minutes in 10% KOH and examined with an Olympus CX21 compound microscope. Images of genitalia were first photographed with a digital camera, then transferred to the Adobe Photoshop CS5.1 software and finally, all genital pictures were prepared by turning images into drawings (Adobe Photoshop CS5.1). The beetles were identified to the species level with the help of available keys and resources (Capra 1929; Mader 1955; Iablokoff-Khnzorian 1971; Duverger 1983; Canepari et al. 1985; Fürsch 1996; Raimundo et al. 2008). Specimens collected and examined during this study are deposited in the Department of Entomology, College

312 · Zootaxa 4236 (2) © 2017 Magnolia Press BIRANVAND ET AL. of Agricultural Sciences, Shiraz Branch, Islamic Azad University, Shiraz, Iran, and in Hayk Mirzayans Insect Museum of the Iranian Research Institute of Plant Protection and Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran. In this paper we follow the higher classification of Coccinellidae by Seago et al. (2011). Taxonomy of the species level agrees with Kovář (2007) except for Hyperaspis inexpectata Günther, 1959, which we consider as a valid species and not a junior synonym of Hyperapsis concolor Suffrian, 1843 (Canepari 2009). Although Kovář (2007) listed H. inexpectata as a synonym of H. concolor Suffrian, 1843, Canepari et al. (1985) considered this species totally different from H. concolor. We could not examine the holotype of H. inexpectata. These species need a thorough revision to clarify their status (Canepari 2009). Species are arranged alphabetically for convenience. The geographical distribution and host plants are given for all species based on literature, labels of the museum specimens examined by the first author and on personal observations of other authors. The prey species are given when available. The geographical distribution here is arranged first according to the year of record publication and thence in alphabetical order. For some species, such as H. concolor, H. duvergeri Fürsch, 1985 and H. polita, that we could not find in our sampling nor in museums, literature descriptions were used in order to prepare the identification key. Summaries of diagnostic characters of the tribe Hyperaspidini and the genus Hyperaspis are given after Gordon & Canepari (2008). Specific terminology used in morphology of Coccinellidae follows Ślipiński & Tomaszewska (2010).

Results

A checklist of the species of the tribe Hyperaspidini from Iran is presented, with information on host plant data and distribution. The list includes 12 species, all of them belonging to the genus Hyperaspis. This list represents approximately 25 percent of the species diversity of Hyperaspis in the Palearctic region. Re-examination of specimens reported by Yazdani (1990) as H. asiatica Lewis and comparing them with H. reppensis (Herbst) revealed that they were misidentified and actually belong to H. reppensis; therefore, H. asiatica is excluded from the Iranian list of coccinellids. Re-investigation of H. pumila Mulsant which was reported by Duverger (1983) and also Yazdani (1990) and comparing it with H. polita Weise revealed that these are probably erroneous records and the specimens most likely are H. polita. Hyperaspis pumila is a rare species previously reported from North of Africa (Duverger 1983) and some countries like Iraq (Duverger 1983), Yemen (Raimundo et al. 2006) and Saudi Arabia (Kovář 2007). As this species is morphologically very similar in general body shape and pattern to H. polita, hence these two species are often confused. We could not find enough evidence to confirm the existence of this species in Iran, so we excluded H. pumila from the list of Iranian coccinellids.

Subfamily Coccinellinae Latreille, 1807

Tribe Hyperaspidini Mulsant, 1846

Diagnosis. Hyperaspidini can be separated from other tribes of the subfamily Coccinellinae by the following combination of characters: body elongate oval, depressed to rounded, convex; dorsal surface glabrous (except in Blaisdelliana Gordon); antenna short, composed of 9 to 11 antennomeres, club elongate, fusiform, terminal antennomere small, recessed in preceding antennomere, antennal insertion exposed or concealed; eye large, not emarginate by an eye canthus, finely faceted, without pubescence; maxillary palpus with apical palpomere securiform; scutellar shield usually large; elytral epipleuron narrow, usually excavated for reception of femoral apex (except Hyperaspidius Crotch and Blaisdelliana Gordon); legs short, femur grooved or flattened for reception of tibia, protibia slender, unmodified, or expanded, tarsus cryptotetramerous; abdomen with 6 ventrites in both sexes; male genitalia with penis guide asymmetrical, toothed, penis guide rooted in phallobase; female genitalia with coxites usually short, transverse, styli reduced or absent (Gordon 1985; Gordon & Canepari 2008).

Syn: Hyperaspis Redtenbacher, 1843; Hyperaspis Chevrolat in Dejean, 1833

Diagnosis. The genus Hyperaspis can be differentiated from other genera within the tribe Hyperaspidini by the following combination of characters: body oval or rounded, convex, dorsum glabrous; head usually yellow in male, at least partially brown or black in female; elytron black or with pale maculation on dark background or dark maculation on pale background; antenna with 10 or 11 antennomeres, scape longer than wide, antennal insertion exposed; terminal maxillary palpomere slightly emarginate apically; mandibular basal tooth strongly curved, apex rounded; clypeus usually long, abruptly or obliquely, smoothly joined to frons laterally, distinctly emarginate apically, often concealing all or most of labrum; labrum short, narrow, weakly sclerotized, yellow or yellowish brown; scutellar shield large, wider than long; elytral epipleuron narrow, usually medially grooved, excavated for reception of middle and hind femoral apices; prosternal process usually slightly convex, with two carinae of varying lengths; posterior margin of metaventrite abruptly descending between coxa and lateral margin; metendosternite with anterior tendons of fork shape, curved toward middle or not; protibia usually slender, rarely slightly flanged; tarsal claw with or without basal tooth; postcoxal line on first abdominal ventrite incomplete, of Scymnus s.str. type; in male apical abdominal ventrite weakly modified; apex of female abdominal ventrite 6 rounded; female genitalia with coxites basically transverse (Gordon & Canepari 2008).

Key to the Iranian species of Hyperaspis

1. Pronotum black in the middle, with yellow, orange or red lateral, sometimes also front margins ...... 2 - Pronotum orange or brown with contrasting pattern ...... 13 2. Elytra completely black, without spots (Fig. 14); pronotum black with red lateral margins; body length 2.8–3.0 mm; penis guide almost symmetrical, pointed (Fig.1) ...... H. concolor Suffrian - Elytra black with yellow, orange or red markings ...... 3 3. Single pair of red spots in posterior half of elytra (preapical) ...... 4 - More than one pair of yellow, orange or red markings ...... 5 4. Body not uniformly oval, wider posteriorly; pronotum with broad red lateral margins; each elytron with reddish spot (Fig. 23); penis guide with truncate apex and large median tooth (Figs. 9, 10); body length 3.1–5.0 mm ...... H. reppensis (Herbst) - Body uniformly oval; pronotum with narrow orange lateral margins; each elytron with orange spot (Fig.15); penis guide broad, shorter than parameres, with oblique apex and small median tooth (Fig. 2); body length 3.2–4.0 mm . . . H. duvergeri Fürsch 5. Each elytron with two rounded spots, one (discal) in anterior half, one (preapical) in posterior half ...... 6 - Elytra with more complex pattern ...... 10 6. Each elytron with two yellowish spots of almost same size (Fig. 20); penis guide with asymmetric apex and small median tooth (Fig. 6); body length 2.8 mm ...... H. persica Duverger (male) - Discal spot smaller than preapical one, orange or red ...... 7 7. Penis guide symmetrical, slender, with rounded apex, without median tooth (Fig. 3) ...... H. femorata (Motschulsky) - Penis guide asymmetrical, with straight or oblique apex ...... 8 8. Penis guide broad with squarely truncate apex and large blunt median tooth (Fig. 8); elytron with discal spot moderately large (Fig. 22) ...... H. quadrimaculata Redtenbacher - Penis guide with obliquely truncate apex and small median tooth; elytron with discal spot very small or absent ...... 9 9. Penis guide much shorter than parameres (Fig. 4); spots on elytra orange (Fig. 17) ...... H. histeroides (Faldermann) - Penis guide almost as long as parameres (Fig. 11); spots on elytra red (Fig. 24) ...... H. syriaca Weise 10. Each elytron with three spots (anterior, discal, preapical) and narrow lateral stripe joined with at least anterior spots, yellow or orange; penis guide long, slender, with sharp median tooth ...... 11 - Each elytron with large irregular, red patch in the middle, often reaching lateral margin, sometimes with small black spot in center of patch; head black; body length 3.0 mm (Fig. 19) ...... H. persica Duverger (female) 11. Head brown; pronotum black with anterior and lateral margins yellowish orange; preapical spot on elytron not confluent with lateral stripe (Fig. 21); body length 2.8–3.0 mm; penis guide as long as parameres and much wider than parameres (Fig. 7) ...... H. polita Weise - Head black with yellow or yellowish brown part anteriorly and sometimes also laterally; pronotum black with yellowish area laterally; preapical spot on elytron confluent with lateral stripe; penis guide much longer and hardly wider than parameres ...... 12 12. Head black except for yellowish brown anterior and lateral margins; elytron with discal spot separate from anterior spot/stripe (Fig. 18); body length 2.0–3.0 mm; basal part of penis guide twice as wide as distal part (Fig. 5) ...... H. marmottani (Fairmaire) - Head black, yellow anteriorly; elytron with discal spot connected to anterior spot/stripe (Fig. 26); body length 1.8–3.2 mm . . .

314 · Zootaxa 4236 (2) © 2017 Magnolia Press BIRANVAND ET AL...... H. vinciguerrae Capra (female) 13. Elytra brown, with basal, discal, lateral and apical creamy brown spots which are variously confluent and irregularly scattered; pronotum brown with creamy brown apical and lateral margins, with an anchor-like spot basomedially which is connected to the anterior margin (Fig. 25); body length 2.5 mm ...... H. transversoguttata Weise (female) - Elytra whitish, with black stripe along suture and each elytron with black, elongate discal, comma-like stripe; pronotum orange, darker mediobasally (Fig. 27); body length 1.2–2.7 mm; penis guide slender, rounded apically, with small median tooth (Figs. 12, 13) ...... H. vinciguerrae Capra (male)

Updated list of Iranian species of Hyperaspis

Hyperaspis concolor Suffrian, 1843 (Figs. 1, 14)

Hyperaspis concolor Suffrian 1843: 93.

Distribution in Iran. Chaharmahal and Bakhtiari (Bagheri & Mossadegh 1997). General distribution. Germany (Klausnitzer 1985), Iran (Bagheri & Mossadegh 1997), Austria, Belgium, Bulgaria, Croatia, Czech Republic, France, Greece, Hungary, Italy, Poland, Portugal, Romania, Slovakia, Spain, Sweden, Switzerland (Canepari 2004). Host plants in Iran. This species has been collected from Echinops sp., Astragalus sp. and Quercus sp. (Bagheri & Mossadegh 1997). Remarks. We used the species descriptions and photographs by Bagheri & Mossadegh (1997) and Canepari et al. (1985) with some modifications.

Hyperaspis duvergeri Fürsch, 1985 (Figs. 2, 15)

Hyperaspis duvergeri Fürsch 1985: 233.

Distribution in Iran. Chaharmahal and Bakhtiari (Bagheri & Mossadegh 1997). General distribution. Algeria, Croatia, France, Hungary, Italy, Morocco, Portugal, Spain (Kovář 2007), Iran (Moddarres-Awal 2012). Host plants in Iran. This species has been collected from Echinops sp. and Astragalus sp. (Bagheri & Mossadegh 1997). Remarks. We used the species descriptions and photographs by Bagheri & Mossadegh (1997) and Canepari et al. (1985) with some modifications.

Hyperaspis femorata (Motschulsky, 1837) (Figs. 3, 16)

Coccinella femorata Motschulsky 1837: 421. Hyperaspis inaudax Mulsant 1853: 231. Hyperaspis collaris Fleischer 1900: 119.

Material examined. 1♂: Iran, Lorestan province, Chenar (33°42'40.8"N, 48°09'11.8"E), Amygdalus sp., VII.2015, lgt. Elham Shakarami, det. Biranvand. 3♂, 1♀: Iran, Semnan province, Clouds forest (36º44ʹ24.7ʹʹN; 55º02ʹ17.9ʹʹE), Quercus sp., V.2015, lgt. Mino Toozandejani, det. Biranvand. Distribution in Iran. Alborz (Borumand 2000), Lorestan, Semnan (current study). General distribution. Armenia, Azerbaijan, Bulgaria, Russia, Turkmenistan, Turkey, Ukraine (Kovář 2007), Iran (Moddarres-Awal 2012). Host plants in Iran. This species has been collected from Medicago sp. (Borumand 2000), and recently from Amygdalus sp. and Quercus sp. (current study).

REVIEW OF HYPERASPIDINI OF IRAN Zootaxa 4236 (2) © 2017 Magnolia Press · 315 Remarks. Specimens examined are deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

FIGURES 1–8. Tegmen of the male genitalia of Hyperaspis species. 1. H. concolor Suffrian. 2. H. duvergeri Fürsch. 3. H. femorata (Motschulsky). 4. H. histeroides (Faldermann). 5. H. marmottani (Fairmaire). 6. H. persica Duverger. 7. H. polita Weise. 8. H. quadrimaculata Redtenbacher.

316 · Zootaxa 4236 (2) © 2017 Magnolia Press BIRANVAND ET AL. FIGURES 9–13. Tegmen of the male genitalia of Hyperaspis species. 9, 10. H. reppensis (Herbst). 11. H. syriaca Weise. 12, 13. H. vinciguerrae Capra.

Hyperaspis histeroides (Faldermann, 1837) (Figs. 4, 17)

Scymnus histeroides Faldermann 1837: 408. Hyperaspis vicaria Zaslavskij 1964: 153.

Material examined. 2♂: Iran, Yazd province, Khatam (30º02ʹ23.9ʹʹN; 54º18ʹ30.4ʹʹE), Amygdalus scoparia L, Atraphaxis sp, VI.2013, lgt. Mehdi Zare Khormizi, det. Canepari. Distribution in Iran. Iran (Kovář 2007), Yazd (current study). General distribution. Armenia, Azerbaijan, Iran, Kazakhstan (Kovář 2007), Turkey (Kaydan et al. 2012). Host plants in Iran. This species has been collected from Amygdalus scoparia L. and Atraphaxis sp. (current study). Remarks. Specimens examined are deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

Coccinella marmottani Fairmaire, 1868: 501. Hyperaspis albidiceps Walker 1871: 19. Hyperaspis camillae Bedel 1885: 90.

Material examined. 1♂: Iran, coll. Museum of Islamic Azad University of Shiraz, det. Canepari and Fürsch. Distribution in Iran. Fars (Yazdani 1990), Khorasan (Yaghmaei & Kharazi Pakdel 1995) General distribution. Nigeria (Umeh 1982), Algeria, Egypt, Iran, Libya, Tunisia (Kovář 2007). Remarks. Specimen examined is deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

Hyperaspis persica Duverger, 1983 (Figs. 6, 19, 20)

Hyperaspis persica Duverger 1983: 79.

Material examined. 4 specimens: Iran, coll. Hayk Mirzayans Insect Museum, det. Fürsch. Distribution in Iran. Kerman (Duverger 1983), South Khorasan (Moodi & Mossadegh 1995). General distribution. This species has been reported only from Iran (Kovář 2007). Host plants in Iran. This species has been collected from Tamarix sp. (Moddarres-Awal 2012). Remarks. Specimens examined are deposited in Hayk Mirzayans Insect Museum of the Iranian Research Institute of Plant Protection, Iran.

Hyperaspis polita Weise, 1885 (Figs. 7, 21)

Hyperaspis polita Weise 1885: 60. Hyperaspis decemguttata Fleischer 1900: 119.

Distribution in Iran. Deeh-Bala region, Ramine region (Duverger 1983), Fras (Yazdani 1990), Golestan (Montazeri & Mossadegh 1995), Kermanshah (Jalilvand et al. 2014), Kerman, Khuzestan, Mazandaran, Zanjan (Moddarres-Awal 2012). General distribution. Egypt, Ethiopia, Iran, Kenya, Libya, Palestine, Somalia, Sudan, Yemen (Duverger 1983), Iraq (Ali et al. 1990), Afghanistan, Iran, Israel, Syria, Tajikistan, Turkmenistan, Turkey (Kovář 2007). Host plants and prey species in Iran. This species has been collected from Ficus sp., Citrus sp., Morus sp. as a predator of Maconellicoccus hirsutus Green and Nipaecoccus viridis (Newstead) (Hemiptera: Pseudococcidae) (Yazdani 1990; Moddarres-Awal 2012). Remarks. We used the species descriptions and photographs by Capra (1929), Mader (1955) and Iablokoff- Khnzorian (1971) with some modifications.

Hyperaspis quadrimaculata Redtenbacher, 1843 (Figs. 8, 22)

Hyperaspis quadrimaculata Redtenbacher 1843: 16.

Material examined. 1♂, 1♀: Iran, Lorestan province, Rig Sephid (33º29ʹ56.7ʹʹN; 48º27ʹ46.1ʹʹE), Medicago sp. , VI.2014, lgt. Amir Ansari pour, det. Canepari and Fürsch. Distribution in Iran. Isfahan (Daran region), Lorestan, Vannae region, Sagdar region (Duverger 1983), Fars (Yazdani 1990).

318 · Zootaxa 4236 (2) © 2017 Magnolia Press BIRANVAND ET AL. General distribution. Iran (Duverger 1983), Albania, Austria, Bulgaria, Czech Republic, Greece, Hungary, Lebanon, Romania, Syria, Turkey (Kovář 2007). Host plants in Iran. This species has been collected from Medicago sp. (Yazdani 1990). Remarks. Specimens examined are deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

FIGURES 14–22. Dorsal habitus of Hyperaspis species. 14. H. concolor Suffrian. 15. H. duvergeri Fürsch. 16. H. femorata (Motschulsky). 17. H. histeroides (Faldermann). 18. H. marmottani (Fairmaire). 19. H. persica Duverger (female). 20. H. persica Duverger (male). 21. H. polita Weise. 22. H. quadrimaculata Redtenbacher.

Coccinella reppensis Herbst 1783: 48. Chrysomela bipustulata Thunberg 1784: 13. Dermestes xanthocephalus Quensel 1790: 10. Dermestes marginellus Quensel 1790: 9. Coccinella reppensis Thunberg 1795: 111. [Homonym] Hyperaspis subconcolor Weise 1879: 136. Hyperaspis occidentalis Fürsch 1967: 261. [Homonym] Hyperaspis galliae Duverger 1989: 144. [Replacement name]

Material examined. 2♂, 1♀: Iran, Yazd province, Khatam (30º02ʹ23.9ʹʹN; 54º18ʹ30.4ʹʹE), Atraphaxis sp., Prunus sp., Pistacia sp., VI.2013, lgt. Mehdi Zare Khormizi, det. Canepari; 2♂: Iran, Yazd province, Khatam (30º02ʹ23.9ʹʹN; 54º18ʹ30.4ʹʹE), Amygdalus sp., IX.2013, lgt. Mehdi Zare Khormizi, det. Canepari. Distribution in Iran. Yazd (Zare Khormizi et al. 2014), Lorestan (Tavakoli et al. 2014). General distribution. Iraq (Ali et al. 1990), Israel (Halperin et al. 1995), Austria, Belgium, Belarus, Czech Republic, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Kazakhstan, Poland, Romania, Russia, Sweden, Switzerland, Tunisia, Ukraine (Kovář 2007), Morocco (Smaili et al. 2010). Host plants in Iran. This species has been collected from Astragalus sp. (Tavakoli et al. 2014), and recently from Medicago sp., Atraphaxis sp., Amygdalus sp., Punica sp., Prunus sp. and Pistacia sp. Remarks. Specimens examined are deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

Hyperaspis syriaca Weise, 1885 (Figs. 11, 24)

Hyperaspis syriaca Weise 1885: 57.

Material examined. 1♂: Iran, Yazd province, Khatam (30°02'23.9"N, 54°18'30.4"E), Atraphaxis sp., VI.2014, lgt. Mehdi Zare Khormizi, det. Canepari and Fürsch. Distribution in Iran. Vannae region (Duverger 1983), Chaharmahal and Bakhtiari (Bagheri & Mossadegh 1997), Yazd (current study). General distribution. Israel, Lebanon, Syria, Turkey (Kovář 2007). Host plants in Iran. This species has been collected from Amygdalus sp., Zea sp., Astragalus sp. (Bagheri & Mossadegh 1997) and recently from Atraphaxis sp. Remarks. Specimen examined is deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran.

Hyperaspis transversoguttata Weise, 1878 (Fig. 25)

Hyperaspis transversoguttata Weise 1878: 94. Oxynychus alexandrae Weise 1890: 488.

Material examined. 1♀: Iran, coll. Hayk Mirzayans Insect Museum, det. Canepari and Fürsch. Distribution in Iran. Ramine region, Khuzestan (Duverger 1983). General distribution. Iran (Duverger 1983), Iraq (Ali et al. 1990), Armenia, Kyrgyzstan, Kazakhstan, Mongolia, Tajikistan, Turkmenistan, Uzbekistan (Kovář 2007), Turkey (Kaydan et al. 2012). Host plants in Iran. This species has been collected from Tamarix sp. (Duverger 1983). Remarks. Specimen examined is deposited in Hayk Mirzayans Insect Museum of the Iranian Research Institute of Plant Protection, Iran.

320 · Zootaxa 4236 (2) © 2017 Magnolia Press BIRANVAND ET AL. FIGURES 23–27. Dorsal habitus of Hyperaspis species. 23. H. reppensis (Herbst). 24. H. syriaca Weise. 25. H. transversoguttata Weise (female). 26. H. vinciguerrae Capra (female). 27. H. vinciguerrae Capra (male).

Hyperaspis vinciguerrae Capra, 1929 (Figs. 12, 13, 26, 27)

Hyperaspis vinciguerrae Capra 1929: 241.

Material examined. 1♀: Iran, Yazd province, Mehriz (31°34'36.1"N, 54°25'16.0"E), Medicago sp., VIII.2015, lgt. Mehdi Zare Khormizi, det. Canepari and Fürsch. Distribution in Iran. Khuzestan (Sadat Alizadeh et al. 2013), Yazd (current study). General distribution. Libya, Saudi Arabia, Yemen (Kovář 2007), United Arab Emirates (Raimundo et al. 2008). Host plants species in Iran. This species has been collected from Medicago sp. (current study). Comments. Capra (1929) described Hyperaspis vinciguerrae from North Africa. This description was published in July, four months earlier than a very similar species H. maindroni described from India by Sicard (1929). According to C. Canepari these names are probably synonyms (Canepari, unpublished). Remarks. Specimen examined is deposited in Plant Protection Department, Lorestan University, Agricultural Faculty, Khorramabad, Iran. We used the species descriptions and photographs by Capra (1929) and Raimundo et al. (2008) with some modifications for male genitalia. Figures 12 and 13 both show the same genitalia which were drawn from different angles.

According to the studies hitherto conducted in Iran, 12 species of Hyperaspis (Hyperaspidini) have been reported. They have been recorded from 15 out of 31 provinces of Iran (Fig. 28). Because of the existence of so many localities of the same name in various provinces of the country, we could not determine the exact localities of four regions mentioned in Duverger’s paper, i.e. Deeh-Bala, Ramine, Vannae and Sagdar.

FIGURE 28. Distribution of Hyperaspis species in Iran. Abbreviations used: Hyperapsis concolor: Cn, H. duvergeri: Dv, H. femorata: Fm, H. quadrimaculata: Qd, H. histeroides: Ht, H.marmottani: Mm, H. persica: Ps, H. polita: Po, H. reppensis: Rp, H. syriaca: Sy, H. transversoguttata: Ts, H. vinciguerrae: Vc.

Most of the Iranian species are distributed in the western Palearctic region. Hyperaspis marmottani and H. polita are distributed in North Africa and Asia (south-western parts of Asia and middle Asia) and there is no evidence of their presence in Europe. Hyperapsis concolor, H. femorata, H. histeroides, H. quadrimaculata, and H. transversoguttata are widely distributed in Asia and Europe and do not extend to North Africa. Hyperaspis syriaca is distributed in Middle East and there is no evidence of its presence in Europe, North of Africa. The report of H. persica from Iran is the only record of this species in the world; furthermore, the reports of H. concolor, H. duvergeri and H. marmottani from Iran are the only records of these species in Asia. We recorded 15 host plants for these 12 Iranian species; most of these plants are wild. Four Hyperaspis species were collected on Astragalus and Medicago, three species on Amygdalus and Atraphaxis, two species on Echinops, Quercus and Tamarix, and only one species was found on each of the other host plant species (genera).

The authors would like to thank Dr. Ebrahim Ebrahimi (Hayk Mirzayans Insect Museum of the Iranian Research Institute of Plant Protection) for a loan of several specimens of Hyperaspis used in this paper. The authors also would like to thank Dr. Saeed Moodi (Department of Plant Protection, College of Agriculture, Birjand University, Birjand, Iran) for many helpful comments and suggestions. Adam Ślipiński (Australian National Insect Collection, CSIRO, Canberra, Australia) read this manuscript providing valuable suggestions and corrections. Two anonymous reviewers are acknowledged for their constructive criticism. ON was supported by grant GA JU 152/2016/P provided by University of South Bohemia.

References

Ali, H.A., Abdul-Rassoul, M.S. & Swail, M.A. (1990) Systematic list of Coccinellidae recorded for Iraq. Bulletin of the Iraq Natural History Museum, 8 (3), 45–51. Bagheri, M.R. & Mossadegh, M.S. (1997) Fauna of coccinellid beetles (Col.: Coccinellidae) in Chaharmahal and Bakhtiari province. Second part of subfamilies: Scymninae and Sticholotidinae. Journal of Agricultural Sciences Islamic Azad University, 3 (9, 10), 97–108. Bedel, L. (1885) Recherches sur les coléoptères du nord de l’Afrique. Recherche synonymiques (1). Annales de la Societé Entomologique de France, Series 6, 5, 85–90. Berti, N., Boulard, M. & Duverger, C. (1983) Fourmis et Coccinelles. Revue bibliographique et observations nouvelles. Bulletin de la Societé Entomologique de France, 88, 271–279. Biddinger, D.J, Weber, D.C. & Hull, L.A. (2009) Coccinellidae as predators of mites: Stethorini in biological control. Biological Control, 51, 268–283. https://doi.org/10.1016/j.biocontrol.2009.05.014 Bogaert, J., Adriaens, T., Constan, J., Lock, K. & Canepari, C. (2012) Hyperaspis ladybirds in Belgium, with the description of H. magnopustulata sp. nov. and faunistic notes (Coleoptera, Coccinellidae). Bulletin de la Société royale belge d’Entomologie/Bulletin van de Koninklijke Belgische Vereniging voor Entomologie, 148, 34–41. Borumand, H. (2000) Insects of Iran: The List of Coleoptera in the Insect Collection of Plant Pests and Disesases Research Institute, Coleoptera (XXIV): Cucujoidea: Fam. (120): Coccinellidae. Plant Pests & Diseases Research Institute, Teheran, Iran, 44 pp. Canepari, C., Fürsch, H. & Kreissl, E. (1985) Die Hyperaspis-Arten von Mittel, West- und Südeuropa. Systematik und verbreitung (Coleoptera Coccinellidae). Giornale Italiano di Entomologia, 9 (2), 223–252. Canepari, C. (2004) Fauna Europaea: Coccinellidae. In: Audisio, P. (Ed.), Fauna Europaea: Coleoptera 2, Beetles. Fauna Europaea version 1.1. Available from: http://www.faunaeur.org (accessed 29 August 2013) [as version 2.6.2] Canepari, C. (2009) New data on some Coccinellidae (Coleoptera) from the Mediterranean Region. Zootaxa, 2318, 394–399. Canepari, C. (2011) Contribution to the knowledge of the Coccinellidae of Sardinia (Coleoptera). Conservazione Habitat Invertebrati, 5, 501–516. Capra, F. (1929) Sulle forme affini all’ Hyperaspis polita WSE. Bollettino della Società Entomologica Italiana, 66 (7), 101– 108. Casey, T.L. (1899) A revision of the American Coccinellidae. Journal of the New York Entomological Society, 7, 71–169. Chapin, E.A. (1966) A new species of myrmecophilous Coccinellidae with notes on other Hyperaspini (Coleoptera). Psyche, 73, 278–283. https://doi.org/10.1155/1966/73641 Chevrolat, L.A.A. (1836) [new taxa]. In: Dejean, P.F.M.A. (Ed.), Catalogue des Coléoptères de la collection de M. le Comte Dejean. Méguignon-Marvis, Paris, pp. 1–443. Crotch, R.G. (1873) Revision of the Coccinellidae of the United States. Transactions of the American Entomological Society, 4, 363–382. Crotch, G.R. (1874) A Revision of the Coleopterous Family Coccinellidae. E.W. Janson, London, 311 pp. https://doi.org/10.5962/bhl.title.8975 Dejean, P.F.M.A. (1833) Catalogue de la Collection des Coléptères de M. le Comte Dejean. Deuxième Edition. Livraison 1–2. Méguignon-Marvis Père et Fils, Paris, 176 pp. [Livraison 1: pp. 1–96, Livraison 2: pp. 97–176] Dejean, P.F.M.A. (1837) Catalogue des Coléptères de la Collection de M. le Comte Dejean. Troisieme Edition, revue, corrigée et augmentée, livraisons 1–4. Mequisiiion-Marvis Père et Fils., Paris, 468 pp. Duverger, C. (1983) Contribution à la connaissance des Coccinellidae d'Iran. Nouvelle Revue d'Entomologie, Paris, Nouvelle Serie, 13 (1), 73–93. Duverger, C. (1989) Contribution à l'étude des Hyperaspinae. 1ère Note (Coleoptera, Coccinellidae) Bulletin de la Sociétè Linnéenne de Bordeaux, 17 (3), 143–157.

REVIEW OF HYPERASPIDINI OF IRAN Zootaxa 4236 (2) © 2017 Magnolia Press · 323 Fairmaire, L. (1868) Essai sur les Coléoptères de Barbarie, Cinquieme partie. Annales de la Société Entomologique de France, Série 4, 8, 471–502. Faldermann, F. (1837) Fauna entomologica Trans–Caucasica. Coleoptera. Pars II. Nouveaux Mémoires de la Société Impériale des Naturalistes de Moscou, 5, 1–433. Fleischer, A. (1900) Neue Coccinelliden aus der Sammlung des Kais. Rathes Herrn Edmund Reitter. Wiener Entomologische Zeitung, 19, 116–120. https://doi.org/10.5962/bhl.part.3442 Fürsch, H. (1967) Coccinellidae. In: Freude, H., Harde, K. & Lohse, G.A. (Eds.), Die Käfer Mitteleuropas. Band 7 Clavicornia. Krefeld, Goecke & Evers, pp. 227–228. Fürsch, H. (1985) [new taxa]. In: Canepari, C., Fürsch, H. & Kreissl, E. (Eds.), Die Hyperaspis-Arten Mittel-, West- und Südeuropa. Systematic und Verbreitung (Coleoptera, Coccinellidae). Giornale Italiano di Entomologia, 2, 223–252. Fürsch, H. (1996) Coccinellid collections from Iran. Coccinella, Dresden, 6, 24–25. Giorgi, J.A., Vandenberg, N.J., McHugh, J.V., Forrester, J.A., Ślipiński, S.A., Miller, K.B., Shapiro, L.R. & Whiting, M.F. (2009) The evolution of food preferences in Coccinellidae. Biological Control, 51, 215–231. https://doi.org/10.1016/j.biocontrol.2009.05.019 Gordon, R.D. (1970) New genera and species of Coccinellidae from the western United States (Coleoptera). Proceedings of the Entomological Society of Washington, 72, 42–50. Gordon, R.D. (1985) The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society, 93 (1), 1–912. Gordon, R.D. (1994) South American Coccinellidae (Coleoptera). Part III. Definition of Exoplectrinae Crotch, Azyinae Mulsant, and Coccidulinae Crotch; a taxonomic revision of Coccidulini. Revista Brasileira de Entomologia, 38, 681–775. Gordon, R.D. & Canepari, C. (2008) South American Coccinellidae (Coleoptera), Part XI: a systematic revision of Hyperaspidini (Hyperaspidinae). Annali del Museo Civico di Storia Naturale “G. Doria”, 99, 245–512. Günther, V. (1959) Vertreter des Tribus Hyperaspini (Col., Coccinellidae) aus der Tschechoslowakei. Časopis Československé Společnosti Entomologické, 56, 225–264. Halperin, J., Merkl, O. & Kehat, M. (1995) An annotated list of the Coccinellidae (Coleoptera) of Israel and adjacent areas. Phytoparasitica, 23, 127–137. https://doi.org/10.1007/BF02980972 Harris, R.H.T.P. (1921) A note on Ortalia pallens Mulsant. South African Journal of Science, 18, 170–171. Herbst, J.F.W. (1783) Kritisches Verzeichniss meiner Insektensammlung. Archiv der Insectengeschichte, 4, 1–72. Hodek, I. & Honěk, A. (2009) Scale insects, mealybugs, whiteflies and psyllids (Hemiptera, Sternorrhyncha) as prey of ladybirds. Biological Control, 51, 232–243. https://doi.org/10.1016/j.biocontrol.2009.05.018 Hodek, I. & Evans, E.W. (2012). Food relationships. In: Hodek, I., van Emden, H. & Honěk, A. (Eds.), Ecology and Behaviour of the Ladybird Beetles (Coccinellidae). Blackwell Publishing Ltd., Chichester, pp. 141–274. https://doi.org/10.1002/9781118223208.ch5 Iablokoff-Khnzorian, S.M. (1971) Synopsis des Hyperaspis Paléartiques (Coleoptera Coccinellidae). Annales de la Société entomologique de France, 7, 163–200. Jalilvand, K., Shirazi, M., Fallahzadeh, M., Vahedi, M.H.A., Samih, M.A. & Moeini Naghadeh, N. (2014) Survey of natural enemies of mealybug species (Hemiptera, Pseudococcidae) in Kermanshah Province, western Iran to inform biological control research. Journal of the Entomological Research Society, 16 (3), 1–10. Kaydan, M.B., Atlihan, R., Uygun, N. & Şenal, D. (2012) Coccinellid (Coleoptera: Coccinellidae) species feeding on coccoids (Hemiptera: Coccoidea) in Van Lake Basin, Turkey. Türkiye Biyolojik Mücadele Dergisi, 2012, 3 (1), 37–46. Klausnitzer, B. (1985) Zur Kenntnis der Hyperaspis-Arten der DDR. Entomologische Nachrichten und Berichte, 29 (6), 271– 274. Kovář, I. (1996) Phylogeny. In: Hodek, I. & Honek, A. (Eds.), Ecology of Coccinellidae. Kluwer Academic Publishers, Dordrecht, pp. 19–31. https://doi.org/10.1007/978-94-017-1349-8_2 Kovář, I. (2007) Coccinellidae. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 4. Elateroidea, Derodontoidea, Bostrichoidea, Lymexyloidea, Cleroidea, Cucujoidea. Apollo Books, Stentrup, pp. 71–74, 568–630. Latreille, P.A. (1807) Genera Crustaceorum et Insectorum Secundum Ordinem Naturalem in Familias Disposita, Iconibus Exemplisque Plurimis Explicata. Tomus tertius. A. Koenig, Paris, 258 pp. Lewis, G. (1896) Notes on Japanese Coccinellidae. The Annals and Magazine of Natural History, Series 6, 17, 22–41. https://doi.org/10.1080/00222939608680319 Mader, L. (1955) Evidenz der palaarktisehen Coccinelliden und ihrer Aberrationen in Wort and Bild. Entomologische Arbeiten aus dem Museum G. Frey Tutzing bei München, 6, 765–1035. Majerus, M., Sloggett, J., Godeau, J. & Hemptinne, J.L. (2007) Interactions between ants and aphidophagous and coccidophagous ladybirds. Population Ecology, 49, 15–27. https://doi.org/10.1007/s10144-006-0021-5 Moddarres-Awal, M. (2012) List of Agricultural Pests and Their Natural Enemies in Iran. Fourth print, Third edition. Ferdowsi University Press, Mashhad, 759 pp.

324 · Zootaxa 4236 (2) © 2017 Magnolia Press BIRANVAND ET AL. Montazeri, M.M. & Mossadegh, M.S. (1995) The coccinellids (Coleoptera) fauna of Gorgan plain and Gonbad Kavus. In: Abai, M. (Ed.), Proceedings of the 12th Iranian Plant Protection Congress (Karaj), 1995, pp. 325. Moodi, S. & Mossadegh, M.S. (1995) The coccinellids (Coleoptera) of southeast Khorasan province, Iran. In: Abai, M. (Ed.), Proceedings of the 12th Iranian Plant Protection Congress (Karaj), 1995, pp. 326. Motschulsky, T.V. (1837) Description de quelques Coléoptères, recueillis dans un voyage au Caucase et dans les provinces Transcaucasiennes Russes en 1834 et 1835. Fungicoles. Nouveaux Mémoires de la Société Impériale des Naturalistes de Moscou, 5, 413–425. Mulsant, M.E. (1846) Histoire Naturelle des Coléoptères de Frances. Sulcicolles–Sécuripalpes. Maison, Paris, 280 pp. Mulsant, E. (1850) Species des Coléoptères Trimères Sécuripalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, publiées par la Société nationale d’Agriculture, ect., de Lyon, Deuxième Série, 2, 1–1104. Mulsant, E. (1853) Supplément a la Monographie des Coléoptères Trimères Sécuripalpes. Annales de la Société Linneenne de Lyon, Nouvelle Série, 1, 129–333. [1852–1853] https://doi.org/10.5962/bhl.title.60609 Nedvěd, O. & Kovář, I. (2012) Phylogeny and classification. In: Hodek, I., Van Emden, H. & Honěk, A. (Eds.), Ecology and Behaviour of the Ladybird Beetles (Coccinellidae). Blackwell Publishing Ltd., Chichester, pp. 1–12. https://doi.org/10.1002/9781118223208.ch1 Obrycki, J.J., Harwood, J.D., Kring, T.J. & O’Neil, R.J. (2009) Aphidophagy by Coccinellidae: application of biological control in agroecosystems. Biological Control, 51, 244–254. https://doi.org/10.1016/j.biocontrol.2009.05.009 Orivel, J., Servigne, P., Cerdan, P., Dejean, A., Corbara, B. (2004) The ladybird Thalassa saginata, an obligatory myrmecophile of Dolichoderus bidens ant colonies. Naturwissenschaften, 91, 97–100. https://doi.org/10.1007/s00114-003-0499-z Pope, R.D. & Lawrence, J.F. (1990) A review of Scymnodes Blackburn, with the description of a new Australian species and its larva (Coleoptera: Coccinellidae). Systematic Entomology, 15, 241–252. https://doi.org/10.1111/j.1365-3113.1990.tb00315.x Quensel, C. (1790) Dissertatio historico-naturalis ignotas insectorum species continens, cujus partem primam, consent. ampliss. facult. philosoph. Public examine modeste subjiciunt praeses Conrad. Quensel, philosophiae magister, et respondens C.P. Lundgård. Typis Berlingianis, Lundae, 20 pp. Raimundo, A.A.C. & van Harten, A. (2000) An annotated checklist of the Coccinellidae (Insecta: Coleoptera) of Yemen. Fauna of Arabia, 18, 211–243. Raimundo, A.C., Fürsch, H. & van Harten, A. (2006) Notes on the ladybird beetles (Coleoptera: Coccinellidae) of Yemen, with the description of two new species. Fauna of Arabia, 21, 217–245. Raimundo, A.C., Fürsch, H. & van Harten, A. (2008) Order Coleoptera, family Coccinellidae. Arthropod fauna of the UAE, 1, 217–239. Redtenbacher, L. (1843) Tentamen dispositionis generum et specierum Coleopterorum Pseudotrimerorum Archiductus Austriae. Carolus Ueberreuter, Vindobona, 32 pp. Robertson, J.A., Whiting, M.F. & McHugh, J.V. (2008) Searching for natural lineages within the Cerylonid Series (Coleoptera: Cucujoidea). Molecular Phylogenetics and Evolution, 46, 193–205. https://doi.org/10.1016/j.ympev.2007.09.017 Robertson, J., Ślipiński, A., Moulton, M., Shockley, F.W., Giorgi, A., Lord, N.P., McKenna, D.D., Tomaszewska, W., Forrester, J., Miller, K.B., Whiting, M.F. & McHugh, J.V. (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 Sadat Alizadeh, M., Mossadegh, M.S. & Esfandiari, M. (2013) Natural enemies of Maconellicoccus hirsutus (Green) (Hemiptera: Pseudococcidae) and their population fluctuations in Ahvaz, southwest of Iran. Journal of Crop Protection, 2 (1), 13–21. Samways, M.J., Osborn, R. & Saunders, T.L. (1997) Mandible form relative to the main food type in ladybirds (Coleoptera: Coccinellidae). Biocontrol Science and Technology, 7, 275–286. https://doi.org/10.1080/09583159730974 Sasaji, H. (1968) Phylogeny of the family Coccinellidae (Coleoptera). Etizenia, 35, 1–37. Sasaji, H. (1971) Fauna Japonica. Coccinellidae (Insecta: Coleoptera). Academic Press of Japan, Tokyo, 340 pp. Seago, A.E., Giorgi, J.A., 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) Descriptions de quelque espèces nouvelles de Coccinellides de la faune Indo-malaise. Annals and Magazine of Natural History, Serie 10, 3, 179–184. https://doi.org/10.1080/00222932908672954 Ślipiński, A. (2007) Australian Ladybird Beetles (Coleoptera: Coccinellidae). Their Biology and Classification. ABRS, Canberra, 286 pp. Ślipiński, A. & Tomaszewska, W. (2010) Coccinellidae Latreille, 1802. In: Leschen, R.A.B., Beutel, R.G & Lawrence, J.F.

REVIEW OF HYPERASPIDINI OF IRAN Zootaxa 4236 (2) © 2017 Magnolia Press · 325 (Eds.), Handbook of Zoology. Vol. 2. Coleoptera. Walter de Gruyter, Berlin/New York, pp. 454–472. Smaili, M.C., Blenzar, A. & Fürsch, H. (2010) First record of new species and phenotypes of ladybird (Coleoptera: Coccinellidae) in citrus orchards in Morocco. Entomologie faunistique – Faunistic Entomology, 62 (3), 103–107. Suffrian, C.W.L.E. (1843) Entomologische Bemerkungen. Entomologische Zeitung (Stettin), 4, 91–95. Sutherland, A.M. & Parrella, M.P. (2009) Mycophagy in Coccinellidae: review and synthesis. Biological Control, 51 (Special Issue: SI), 284–293. https://doi.org/10.1016/j.biocontrol.2009.05.012 Szawaryn, K., Bocak, L., Ślipiński, A., Escalona, H.E. & 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 Tavakoli, M., Ansari pour, A., Pirhadi, A. & Pirozi, F. (2014) Study of fauna ladybirds (Col., Coccinellidae) on pastures Astragalus adscendes (Fabaceae) in Lorestan province. Journal of Entomological Research, 6, 287–297. Thunberg, C.P. (1784) Novae insectorum species. Nova Acta Regiae Societatis Scientarum Upsaliensis, 4, 1–28, 1 pl. Thunberg, C.P (1795) Dissertatio entomologica, sistens Insecta Suecica. IX. Exam. Steno Edvardus Westman, Upsaliae, pp. 105–113. Umeh, E.D.N.N. (1982) Biological studies on Hyperaspis marmottani Fairm. (Col., Coccinellidae), a predator of the cassava mealybug Phenacoccus manihoti Mat-Ferr. (Hom., Pseudococcidae). Journal of Applied Entomology, Berlin, 94 (1–5), 530–532. https://doi.org/10.1111/j.1439-0418.1982.tb02600.x Vandenberg, N.J. (2002) Family 93. Coccinellidae Latreille 1807. In: Arnett, R.H., Thomas, M.C., Skelley, P.E. & Frank, J.H. (Eds.), American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press LLC, Boca Raton, pp. 371–389. Walker, F. (1871) List of Coleoptera Collected by J.K. Lord, Esq. in Egypt, Arabia and Near the African Shore of the Red Sea. With Characters of the Undescribed Species. E.W. Janson, London, 19 pp. Weise, J. (1878) [new taxa]. In: Schneider, O. & Leder, H. (Eds.), Beiträge zur Kenntniss der kaukasichen Käferfauna. Verhandlungen des Naturforschenden Vereins in Brünn, 16, 1–359, 3 pls. [1977] Weise, J. (1879) Bestimmungs-Tabellen der europaischen Coleopteren II. Coccinellidae. Zeitschrift für Entomologie, New Series, 7, 88–156. Weise, J. (1885) Coccinellidae. II. Auflage. Mit Berücksichtigung der Artenaus dem nördlichen Asien. In: Reitter, E. (Ed.), Bestimmungs–Tabellen der europäischen Coleopteren. II. Heft. Büsing, Mödling, 83 pp. Weise, J. (1890) Insecta, a CL. G.N. Potanin in China et in Mongolia novissime lecta. XVI Chrysomelidae et Coccinellidae. Horae Societatis Entomologicae Rossicae, 24 [1889–1890], 477–492. Weise, J. (1926) Uber bekannte und new Chrysomeliden und Coccinelliden aus dem Reich museum zu Stockholm. Arkiv för Zoologi, 18, 34, 1–34. Yazdani, A. (1990) The Coccinellids (Col.; Coccinellidae) Fauna of Fars Province. Unpublished Thesis, M. Sc, Shiraz University, Shiraz, 145 pp. Yaghmaei, F. & Kharazi Pakdelm, A. (1995) A faunistic survey of coccinellids in Mashhad region. In: Abai, M. (Ed.), Proceedings of the 12th Iranian Plant Protection Congress (Karaj), 1995, p 307. Zare Khormizi, M., Ostovan, H., Fallahzadeh, M. & Mossadegh, M. (2014) Report of three ladybird beetles (Coleoptera: Coccinellidae) from Iran. In: Panah, R. (Ed.), Proceedings of the 21th Iranian Plant Protection congress (Urmia), 2014, pp. 490. Zaslavskij, V.A. (1964) Novye i maloizvestnye Hyperaspis (Coleoptera, Coccinellidae) iz Kazakhstana i Srednej Azii. Trudy Zoologichekogo Instituta (Leningrad), 34, 152–154.