A Survey of the Agrotis of Iran

Abstract The present study reviews the genus Agrotis Ochsenheimer, 1816 (Lepidoptera: Noctuidae: Noctuinae) in Iran from a taxonomic and faunistic point of view. An identification key of external features is presented for 16 Iranian species and subspecies. A description of each taxon is presented based on external male and female genital characteristics. Diagnostic features and comparisons with the closest relatives are given for each species. Original combination and citation with the synonymy of each species or subspecies are expounded as well as their distribution and bionomy. Adult moths and male genitalia are illustrated.


Introduction
The territory of Iran primarily encompasses semi-desert plains. However, it also is covered by two mountain chains (i.e., Elburz and Zagros) with xeromountain type (Varga 1996) vegetation. Despite the rather dry climate of the country, the northern line of the country comprises humid forests. Owing to its various geographic and climatologic diversity, relatively rich fauna and flora are expected in the country.
Cutworms, members of the genus Agrotis Ochsenheimer, 1816 (Noctuinae: Agrotini), feed on roots and foliage of their host plants, and many species are important pests of crops. Some of the larvae spend the day inside the soil and consume leaves at night. Agrotis spp. inhabit various geographical regions and prefer open habitats. The prominent feature of the genus is the pectinate antennae of the male as well as the configuration of male and female genitalia (described by Fibiger 1990), especially the elongated vesica (the longest in the subfamily) with basal and apical swellings and basal diverticulum with a basal scobinated belt. A very long appendix bursae (sometimes five to six times longer than corpus bursae), fitted with the vesica of the male, is a unique feature among Agrotis species. These particular structures encouraged Fibiger (1990) to propose a new method to delimit the species of the genus based on measuring the vesica and appendix bursae length of both male and female sexes.
Until now, many researchers have primarily studied this genus from the faunistic and taxonomic point of view (e.g., Wiltshire 1975Wiltshire , 1994Fibiger 1990Fibiger , 1993Fibiger , 1997Hacker 1990;Ivinskis and Miatleuski 1999;Kravchenko et al. 2006;Michael 2006;Lafontaine and Schmidt 2010). To date, 103 species of this genus have been described worldwide. Of those, 80 species exist in the Palaearctic region. Twenty-three Agrotis species of are endemic to North America in the Nearctic region and do not exist in the Palaearctic region (Lafontaine and Schmidt 2010). The European fauna, with 23 species, have been well studied by Fibiger (1990Fibiger ( , 1993Fibiger ( , 1997, who classified these species into 10 species groups: the fatidica, the cinerea, the graslini, the segetum, the endogaea, the vestigialis, the ripae, the trux, the crassa, and the biconica. The Near East and its neighboring areas contains around 38 Agrotis species (Hacker 1990;Wiltshire 1994;Ivinskis and Miatleuski 1999). Among them, Iran is represented by 16 species, Turkey by 19 species (Kemal et al. 2007;Koçak et al. 2008), Israel by 19 species (Kravchenko et al. 2006), Turkmenistan by six species (Ivinskis and Miatleuski 1999), and Cyprus by eight species (Fibiger et al. 1999).
The 16 Iranian species are arranged into six species groups: cinerea, segetum, endogaea, trux, bigramma, and spinifera (A. crassa ([Hübner], 1803) and A. biconica Kollar, 1844 are synonyms of A. bigramma (Esper, [1790]) and A. spinifera (Hübner, [1808]), respectively, so in this study their groups are treated as synonyms of the bigramma and spinifera species groups). The cinerea species group, with Southwestern Palaearctic distribution (Bond and Gittings 2008), is represented in Iran by one species, A. cinerea ([Denis & Schiffermüller], 1775), which is distributed in western Iran (Ebert and Hacker 2002). A. segetum ([Denis & Schiffermüller], 1775), a member of the segetum species group, lives in all geographical regions in Iran in spite of its other congroup, A. clavis (Hufnagel, 1766), which populates the temperate zones in the north and northeast parts of the country. In Iran, three species belonging to the endogaea species-group (A. herzogi Rebel, 1911, A. pu-ta (Hübner, 1803, and A. sardzeana Brandt, 1941) are dispersed in the warm and dry lands in the southern parts of the country (Ebert and Hacker 2002), the preferred habitat mentioned by Fibiger (1990). A. puta have also been reported from the European countries (Bond and Gittings 2008). The cosmopolitan species (Fibiger 1997) A. ipsilon (Hufnagel, 1766), A. exclamationis (L., 1758), and A. trux (Hübner, [1824) comprise the trux species group and are distributed in all climatic regions. Due to the lack of comprehensive faunistic investigations, the distribution pattern of some species is based on local surveys. The significant example is the bigramma species group, in which A. bifurca grossi Hacker & Kuhna, 1986 is reported from the east (Ebert and Hacker 2002) and A. obesa scytha Alphéraky, 1889 and A. bigramma occupy northern latitudes (Wieser and Stangelmaier 2005), although a few specimens of A. obesa scytha have been locally collected by the authors from the southern part of the country. The taxonomic placement of the sixth species group, spinifera, is dubious; therefore, this survey follows Fibiger (1997), in which A. spinifera, A. lasserrei (Oberthür, 1881), A. benigna (Corti, 1926) and A. psammocharis Boursin, 1950 are referred as members of the spinifera species group. All members are locally distributed in both semiarid and temperate regions (Ebert and Hacker 2002;Wieser and Stangelmaier 2005;Kravchenko et al. 2007).
The present paper reviews 16 species of the genus Agrotis in Iran. Citations of the original combination, synonymy, and distribution of each species are given. Adult moths and their genitalia are described; diagnostic features, bionomy, and notes on their distribution are discussed. An dentification key is presented based on the external characteristics. Adult moths and their genitalia, if they are presented, are illustrated.

Materials and Methods
Main data of the recorded noctuid moths were obtained from a bibliographic survey (e.g., Hacker 1990;Ebert and Hacker 2002). Furthermore, the results of recent expeditions carried out in different parts of the country (Kerman, Sistan va Balouchestan, Semnan, Markazi, Ardabil, Fars, Golestan, Khorasan, Hormozgan, and Esfahan provinces), were added. Distinctive regions included diverse vegetation, and geographical locations were chosen and sampled in separate provinces beginning in 2004. Two kinds of light traps (portable and constant) applied to collect material and were sampled regularly. Chloroform solution was used to kill the attracted moths inside the light traps. The collected moths were maintained with insect pins on spreading boards. In order precisely identify the species, the genitalia of moths were extracted and mounted using Euparal mounting medium. The adult and genitalia images were taken by a Canon Digital camera (model Power Shot A710, www.canon.com). Terminology for genital structures and wing morphology follows that of Fibiger (1977Fibiger ( , 1990. All collected and studied material (except those of A. lasserrei and A. cinerea, which are deposited in the Hungarian Natural History Museum) are deposited in the Collection of Entomology, Shahid Bahonar University of Kerman, Iran. [Synonymy: denticulatus (Haworth, 1803); tephrina Staudinger, 1901] General distribution. Southwestern Palaearctic (Bond and Gittings 2008). Caucasus, Armenia, Turkey (Hacker 1990), Iraq, Levante, Morocco, Tunisia, Libya, Southern and Central Europe (Kravchenko et al. 2007).
Bionomy. Multivoltine. The moths flying period is late winter to early fall. They are usually attracted to light and occur in every location from forest to desert habitats in varied altitudes. Larvae are the most important pests in cereals and vegetables.
Taxonomic notes. This species can be confused with A. trux, but antennae of the male are less pectinated and apical sign on the costal margin of the forewing is more distinct in A. trux.  (Fibiger 1990).
Description. Male (Figure 4). Antennae slightly pectinated. Wingspan 32-43 mm; forewings light brown to gray, noctuid maculation complete, claviform and orbicular stigmata elongated, claviform stigma light brown surrounded by black line, orbicular stigma cream, reniform stigma elongated, surrounded by black line, cross lines distinguishable; hindwing pure white without discal spot. Female similar to male but larger than male; antennae filiform. (Figure 19). Valve elongated, distal one-third relatively narrow, slightly pointed, costal and ventral margin curved, clasper in moderate size and approximately slender. Aedeagus basally and terminally narrow, medially broader. Vesica narrow, long, nearly about five times as long as the aedeagus, basal swelling with a small bulge, scobinated bar short, apical swelling cylindrical.

Male genitalia
Female genitalia (Figure 32). Ovipositor of medium size, papilla anales pointed posterior and anterior apophyses in moderate size, posterior apophyses longer than anterior ones; appendix bursae, curved, very long, more than three times longer than corpus bursae, both drop-shaped.
Bionomy. Univoltine. Late winter or early spring flyers, from October to April. Species attracted to light (Kravchenko et al. 2006). Widespread in desert and semi-desert habitats. The early stages and host plants are unknown.
Taxonomic notes. It can be confused with A. puta, but the basal area of the forewing in A. puta is dark colored, as is that of A. syricola. The aedeagus in A. puta is absolutely tubular and the papilla anales are short and not acute. A. herzogi also might be confused with A. sardzeana, but the forewings of A. sardzeana are light yellowish brown and its clasper is pointed.  (Fibiger 1990); Cyprus (Fibiger et al. 1999); Israel (Kravchenko et al. 2007); Austria, France, Belgium, Southurn Germany, Hungary, Denmark, Poland, Britain (Bond and Gittings 2008).

Male genitalia
Female genitalia (Figure 33). Ovipositor short, posterior apophyses longer than anterior ones; appendix bursae twice as long as the corpus bursae, with torsion.

Bionomy.
Univoltine. Adults are active in mid-autumn and usually live in arid and semiarid habitats but sometimes exist in shrubs and forest vegetation. The early stages and host plants are undescribed. General distribution. Palaearctic region. Widespread through the entire Palaearctic region except Iceland. In North Africa and the Near and Middle East, this species is limited in semi-desert and desert zones (Kravchenko et al. 2006). In Iraq (Hacker 1990); Mongolia (Gyulai and Ronkay 1999); Lebanon, Caucasus, Russia, Turkey, Armenia, Afghanistan, Turkmenistan (Ivinskis and Miatleuski 1999); Cyprus (Hacker 2001).
Description. Male (Figure 7). Antennae weakly pectinated. Patagia with black line. Wingspan 36-39 mm; forewings bright to dark brown, forewings ground color, almost unicolor except the terminal area is darker, noctuid maculation present, reniform and claviform stigmata distinct, black, orbicular stigma as ground color in some specimens a little darker, antemedian and postmedian lines more or less distinct, sometimes weak, the former more distinct; hindwings white to light cream. Female similar to male, antennae filiform. (Figure 22). Valve long, acute, costal margin slightly arched, clasper slightly pointed. Vesica long, more than five times as long as the aedeagus, basal swelling elliptical, broad, with basal bulge, scobinated bar long, turned, apical swelling distinct. Fibiger (1997, Figure 77). Ovipositor short, posterior apophyses a little longer than anterior ones; ductus bursae short, narrower at distal half, sclerotized bands present; appendix bursae and corpus bursae tubular, globularly widened; appendix bursae long, curved, about three times as long as the corpus bursae.

Female genitalia. Described by
Bionomy. Multivoltine. The reported flight period in Israel is November (Kravchenko et al. 2006), but it has been recorded in May to July in some countries, like Ireland (Bond and Gittings 2008). In Iran, it has been collected in May to October and usually occurs in open spaces, especially in farming lands, in all altitudes. Adults are attracted to light and feed on nectar of flowers. Larvae are known as a serious pest of agricultural crops, feeding on roots of herbaceous plants, specifically plants from the Gramineae family.

Male genitalia
Bionomy. Probably multivoltine. Flight period in October, March and June in Iran. July to mid-August in Ireland, July and August in Britain (Bond and Gittings 2008), September to April in Israel (Kravchenko et al. 2006). The adults are attracted to light, sugar, and flowers (Fibiger 1990) and inhabit various areas. The larvae feed on grassy plants and have been described by several authors.
Description. Male (Figure 9). Antennae pectinated. Wingspan 36-43 mm; forewings light brown to dark brown, noctuid maculation complete, reniform and orbicular stigmata centered with black, a black dart shaped patch present in the ostium of the reniform stigma as well as the two small dart shaped patches in the subterminal area, these features are missing in other species of the genus Agrotis. Black band presented between the reniform and orbicular stigmata, terminal and subterminal areas bright; hindwing white and crystalline but darker in terminal area. Female as male, antenna filiform. (Figure 24). Valve elongated, basal one-third narrow, widened; clasper pointed. Aedeagus broader basally; vesica elongated, tubular, with small sclerotized grooves, six times as long as the aedeagus, basal swelling similar to cylinder, everted laterally, scobinated bar long, apical swelling weakly broader than the vesical tube.

Male genitalia
Female genitalia (Figure 34). Ovipositor small, both the apophyses short, posterior apophyses longer than the anterior ones; ductus bursae in the same length of the ovipositor, appendix bursae tubular, less than one-half times as long as the corpus bursae; corpus bursae pyriform.
Bionomy. Usually multivoltine. Active throughout the year throughout Iran. Moths attracted to flowers, light, and sugar (Fibiger 1990) with heavy migration. Larvae feed on various plants belonging to many families and are one of the most important pests on herbaceous plants, feeding on the roots.

Distribution in Iran.
Description. Male (Figure 10). Antennae strongly pectinated. Wingspan 36-45 mm; forewings bright brown to bright gray, all three spots distinct, reniform and claviform stigmata darker than forewing ground color, a dark patch between reniform and orbicular stigmata, antemedian and postmedian lines not distinct, subterminal line complete, white streak extended from the upper edge of reniform stigma to the cilia, veins distinct surrounded by white scales; hindwing bright, sometimes the outer margin light brownish. Female as male but antennae filiform and outer margin of the hindwing gray to brown. (Figure 25). Valve slightly arched at the costal margin, clasper sickleshaped. Aedeagus nearly tubular; vesica tubular, four times longer than the aedeagus, basal swelling with almost large rounded dorsal hump, scobinated bar relatively short. Female genitalia. Described by Fibiger (1997, Figure 80). Ovipositor large, posterior and anterior apophyses in the same length, posterior apophyses slender; ductus bursae nearly in the same length of ovipositor; appendix bursae short, slightly longer than corpus bursae; corpus bursae elliptical.

Male genitalia
Bionomy. Univoltine, flying generally from August to October, adults attracted to sugar and light. In Israel, a univoltine autumnal species flying from September to October (Kravchenko et al. 2006). This species is mostly found in open habitats along with ex-tending vegetation. The early stages feed on sprouts and aerial parts of fresh plants.

Female genitalia. Described by
Bionomy. Univoltine in Israel and Europe (Fibiger 1990;Kravchenko et al. 2007). Adults are on the wing in September to December in Israel and in August and September in Europe. Imagines attracted to flowers, sugar, and artificial light. Occurs in steppe regions with sparse vegetation (Hacker 2001). Larvae feed on Poaceae species and low density vegetation (Hacker 2001 (Kravchenko et al. 2006).
Description. Male ( Figure 13). Antennae weakly pectinated. Wingspan 28-40 mm; forewings brown to gray, three stigmata complete, orbicular stigma elongated, claviform stigma sharply defined, pointed, pure black, a black band present between the reniform and orbicular stigmata, antemedian and postmedian lines missing, subterminal line distinct as an incomplete waved line, terminal line distinct; hindwings white to bright cream. Female as male, antennae filiform. (Figure 28). Valve of moderate size, costal margin approximately straight, ventral margins arched, clasper short and thickened. Aedeagus broader basally, two subbasal spine fields at the apex of aedeagus; vesica tubular, four times as long as the aedeagus, basal swelling pyriform, scobinated bar of medium size, spiny.

Male genitalia
Female genitalia (Figure 35). Ovipositor relatively large, posterior apophyses a little longer than anterior ones; ostium bursae sclerotized. Both appendix and corpus bursae tubular; corpus bursae medially tubular; appendix bursae slightly longer than the corpus bursae.
Bionomy. Multivoltine. Active throughout the year (Kravchenko et al. 2006). Attracted to light, and appears in various geographical regions with various altitudes. The early stages have been described by several authors. The caterpillars feed on the leaves and roots of herbaceous plants, especially on Poaceae family.
Taxonomic notes. This species can be distinguished from its other Iranian congeners by a very long, black claviform stigma and elongated orbicular stigma. General distribution. South of Palaearctic region especially in droughty places (Kravchenko et al. 2006); Lebanon, Syria, Israel, Iraq, Jordan, Arabian Peninsula, Egypt (Hacker 1990); In Europe, in Spain and Malta; from the West to Mauretania, West Sahara in North Africa and the Canary Isles, from the East to Turkmenistan and Iran (Hacker 2001).
Bionomy. Univoltine, with autumnal flight period (Kravchenko et al. 2006). This species is widespread in arid and semi-arid regions. Early stages and host plants are as yet unknown. General distribution. Russia, Turkey, Iraq (Hacker 1990).

Bionomy.
Univoltine. This species flies in October and is attracted to light. The early stages and host plants are yet undisclosed.  (Kravchenko et al. 2007).
Description. Male (Figure 16). Antennae strongly pectinated. Wingspan 33-41 mm; forewing ground color light brown to gray, reniform and orbicular stigmata present as a two pale gray signs, claviform stigma absent, antemedian and postmedian lines distinguishable; hindwing white with an elongated discal spot. Female as male, antennae filiform. (Figure 30). Costal and ventral margin of the valve straight, cucullus rounded; clasper almost sharp. Aedeagus tubular, nearly as long as the vesica; vesica short, basal swelling nearly two times as broad as the aedeagus, scobinated belt in moderate size, vesica narrower to the end.

Male genitalia
Female genitalia. Female specimens were not collected in this survey.

Bionomy.
Univoltine. Adults commonly fly in fall. They are attracted to light, sugar, and flowers and are present in areas with poor vegetation and dry climate. The early stages and host plants are unknown.
Taxonomic notes. External characteristics of this species are close to A. benigna but can be recognized by absence of claviform stigma in A. psammocharis and in the male genitalia by thickened clasper and longer vesica, which is more than three times longer than aedeagus in the latter species.