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Theory on plasticity driving speciation, as applied to insect–plant interactions (the oscillation hypothesis), predicts more species in clades with higher diversity of host use, all else being equal. Previous support comes mainly from specialized herbivores such as butterflies, and plasticity theory suggests that there may be an upper host range limit where host diversity no longer promotes diversification. The tussock moths (Erebidae: Lymantriinae) are known for extreme levels of polyphagy. We demonstrate that this system is also very different from butterflies in terms of phylogenetic signal for polyphagy and for use of specific host orders. Yet we found support for the generality of the oscillation hypothesis, in that clades with higher diversity of host use were found to contain more species. These clades also consistently contained the most polyphagous single species. Comparing host use in Lymantriinae with related taxa shows that the taxon indeed stands out in terms of the frequency of polyphagous species. Comparative evidence suggests that this is most probably due to its nonfeeding adults, with polyphagy being part of a resulting life history syndrome. Our results indicate that even high levels of plasticity can drive diversification, at least when the levels oscillate over time.

Both community composition changes due to species redistribution and within-species size shifts may alter body size structures under climate warming. Here we assess the relative contribution of these processes in community-level body size changes in tropical moth assemblages that moved uphill during a period of warming. Based on resurvey data for seven assemblages (>8000 individuals) on Mt. Kinabalu, Borneo in 1965 and 2007, we show significant wing-length reduction (mean shrinkage of 1.3% per species). Range shifts explain most size re-structuring, due to uphill shifts of relatively small species, especially at high elevations. Overall, mean forewing length shrank by ca. 5%, much of which accounted for by species range boundary shifts (3.9%), followed by within-boundary distribution changes (0.5%), and within-species size shrinkage (0.6%). We conclude that the effects of range shifting predominate, but considering species physiological responses is also important for understanding community size reorganization under climate warming.

The phylogeny of geometrids sampled in this study and used for analysis.

The phylogeny of plant hosts sampled in this study for geometrids or pyraloids.

Plots of fitted quasibinomial regressions for both oxidative activity (mg/g) and protein precipitation (mg/g). Plots i) and ii) include only the significant phylogenetic axes common to both moth families, while plots ii) and v) include only the chemical traits, plots iii) and vi) include both phylogenetic axes and chemical traits.

Literature review and summary of Geometroidea, Pyraloidea and Thyrididae host use in the oriental region.

A histogram of mean oxidative activity (mg/g) (± one s.e.) for all 88 species analysed.

A histogram of mean protein precipitation (mg/g) (± one s.e.) for all 88 species analysed.

A schematic diagram of our analytical steps for predicting network structure.

A list of individuals sequenced for each gene included in this study, including from published studies (Haines and Rubinoff, 2012; Sihvonen et al., 2011). Red text indicates individuals where close relatives were used as BOLD extracts were not available for the exact specimen.

Correlations between Euclidean distance matrices derived from observed and predicted models.

A heatmap of geometrid abundance, oxidative activity (mg/g) and predicted occurrence.

Detailed methods and results section.

A description of taxonomic updates published since Novotny et al. (2010) or resulting from this study.

Plots of different fitted quasibinomial regressions for both oxidative activity (mg/g) and protein precipitation (mg/g).

Symondson, William O.C., Holloway, Jeremy D., Goossens, Benoit, Müller, Carsten T. (2014): Bornean caterpillar (Lepidoptera) constructs cocoon from Vatica rassak (Dipterocarpaceae) resin containing multiple deterrent compounds. Journal of Natural History 49 (9): 553-560, DOI: 10.1080/00222933.2014.939731

Pyrrhia umbra (Hufnagel, 1766) Pl. 2, fig. 11; male genitalia Pl. 5, fig. 29; female genitalia Pl. 9, fig. 45. Phalaena umbra Hufnagel, 1766, Berl. Mag. 3: 294, Taf. 51: 6 (TL.: Germany: Berlin). Synonymy: Noctua rutilago [Denis & Schifferm��ller], 1775, Noctua marginata Fabricius, 1775, Phalaena (Noctua) conspicua Borkhausen, 1792, Phalaena (Noctua) umbrago Esper, [1796], Noctua marginago Haworth, 1809, Heliothis cilisca Guenee, 1852, Hydroecia tibetana Moore, 1878, Chariclea vexilliger Christoph, 1893. References: Christoph 1877 (Chariclea umbra); Hampson 1910; Hacker 1990; Ebert & Hacker 2002 (Pyrrhia umbra). Distribution: Transpalaearctic. Europe, Caucasus, Transcaucasia, Kazakhstan, Central Asia, south Siberia, Far East, China (including Tibet), north India, Nepal, Australia (introduced). ��� In Iran (Pl. 11, fig. 59) occurs in provinces West Azerbaijan, Guilan, Mazandaran, Golestan and Tehran. Bionomics: Bivoltine. Moths flying from May to October. The species inhabits steppe habitats on lowmedium elevations from 0 to 1800 m. Larvae are polyphagous, feed on 33 species of herbaceous plants, shrubs and trees of 16 botanical families, prefer Fabaceae and Scrophulariaceae. Material examined: 61 specimens from provinces West Azerbaijan, Guilan, Mazandaran, Golestan and Tehran, collected between 14.V to 29.X on elevations from 0 to 1800 m., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on pages 14-15, DOI: 10.5281/zenodo.181966

Aedophron venosa Christoph, 1887 Pl. 2, fig. 13; male genitalia Pl. 6, fig. 31; female genitalia Pl. 9, fig. 47. Aedophron venosa Christoph, 1887, Stett. Ent. Z. 48: 165 (TL.: [Turkmenistan]: Ashkhabad). References: Wiltshire 1951 (Aedophron venosa); Kalali 1976 (Aedophron phlebophora ��� misidentification). Bionomics: Univoltine. Moths flying from May to July. The species inhabits dry steppe biotopes on medium and moderate altitudes (1150 ��� 2950 m). Larval host-plants are unknown. Distribution: Irano-Turkestanian. Central Asia (Turkmenistan). ��� In Iran (Pl. 11, fig. 61) occurs in provinces Golestan, Khorasan and Semnan. Material examined: Golestan: Sulgerd, Park-e-Melli Golestan, 1150 m, 16-18.VII. 1975, leg. Pazuki A. Khorasan: Aladagh, Khosravieh, 1600 m, 16.VI. 1974, leg. Rajabi & Pazuki; Allahakbar Mount., Kopeh Dagh Mount., 2950 m, 16.VI. 1974, leg. Rajabi & Pazuki; Almeh, Park-e-Melli-Golestan, 1590 m, 17- 25.V. 1988, leg. Pazuki A.; Almeh, Park-e-Melli-Golestan, 1600 m, 26-29.V. 1986, leg. Pazuki A.; Assadly, Bojnurd, 30 km S, 1970 m, 17.VI. 1977, leg. Pazuki & Abai, Eizman, Bojnurd, 17.V. 2005, leg. Falsafi & Nematian; Yakhtikalan, Park-e-Melli Golestan, 1650 m, 02.VI. 1986, leg. Pazuki A. Semnan: Kalate-Amirieh, Shahrud, 1550 m, 30.V. 1982, leg. Hashemi; Kamare Cheheldokhtar, Shahrud, 1700 m, 03.VI. 1982, leg. Hashemi. PLATE 4. Figures 22 ���25. 22, Heliothis nubigera, slide 443 (genitalia), Hormozgan, and slide 221 (aedeagus), Tehran; 23, H. peltigera, slide 60 (genitalia), Tehran (Karadj), and slide 446 (aedeagus), Semnan; 24, H. incarnata, slide 449, Fars; 25, Helicoverpa armigera, slide 258, Esfahan (Karkas Kuh). PLATE 5. Figures 26 ���29. 26, Schinia scutosa, slide 461, Golestan; 27, Periphanes delphinii, slide 344, Golestan (Gorgan); 28, P. victorina, slide 341 (genitalia), Tehran (Damavand), and slide 462 (aedeagus), Tehran (Damavand); 29, Pyrrhia umbra, slide 456, Mazandaran (Tonekabon). PLATE 6. Figures 30 ���33. 30, Aedophron phlebophora, slide 339, Fars (Didegan); 31, A. venosa, slide 454, Golestan; 32, A. rhodites, slide 277, Kerman; 33, A. sumorita, slide 279, Fars. PLATE 7. Figures 34 ���36. 34, Heliocheilus confertissima, slide 451 (genitalia), and slide 343 (aedeagus), Baluchestan; 35, Masalia albida, slide 342, Kerman; 36, M. perstriata fuscostriata, slide 345, Hormozgan. PLATE 8. Figures 37 ���42. 37, Heliothis viriplaca, slide 274, Fars; 38, H. maritima, slide 442, Tehran; 39, H. peltigera, slide 445, Ardebil; 40, H. incarnata, slide 448, Fars; 41, Helicoverpa armigera, slide 447, Hormozgan; 42, Schinia scutosa, slide 460, Golestan. PLATE 9. Figures 43 ���48. 43, Periphanes delphinii, slide 459, Fars; 44, P. victorina, slide 469, Tehran (Damavand); 45, Pyrrhia umbra, slide 457, Mazandaran; 46, Aedophron phlebophora, slide 452, Kordestan; 47, A. venosa, Golestan, slide 455; 48, Masalia philbyi, Hormozgan, SMNK., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on pages 15-21, DOI: 10.5281/zenodo.181966

Genus Schinia H��bner, 1818 Type species: Noctua scutosa [Denis & Schifferm��ller], 1775 by original designation Synonymy: Euclidia H��bner, 1808; Melicleptria H��bner, [1823]; Anthoecia Boisduval, 1840; Alaria [Duncan & Westwood], 1841; Trypana Guen��e, 1841; Oria Guen��e, 1852; Rhodophora Guen��e, 1852; Tamila Guen��e, 1852; Euleucyptera Grote, 1865; Lygranthoecia Grote & Robinson, 1873; Tricopis Grote, 1874; Adonisea Grote, 1875; Heliophana Grote, 1875; Oxylos Grote, 1875; Pippona Harvey, 1875; Porrima Grote, 1877; Rhododipsa Grote, 1877; Bessula Grote, 1881; Dasyspoudaea Smith, 1882; Pseudotamila Smith, 1883; Canidia Grote, 1890; Eupanychis Grote, 1890; Trichosellus Grote, 1890; Thyreion Smith, 1891; Incita Grote, 1895; Palada Smith, 1900; Chlorocleptria Hampson, 1903; Heliothis Hampson, 1903; Tricraterifrontia Berio, 1940; Trilenca Neave, 1940; Uollega Berio, 1945; Protoschinia Hardwick, 1970; Purpurschinia Beck, 1996., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 12, DOI: 10.5281/zenodo.181966

Genus Heliocheilus Grote, 1865 Type species: Heliocheilus paradoxus Grote, 1865 Synonymy: Raghuva Moore, 1881, Canthylidia Butler, 1886, Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 22, DOI: 10.5281/zenodo.181966

Genus Heliothis Ochsenheimer, 1816 Type species: Phalaena dipsacea Linnaeus, 1767 (Syst. Nat. (Ed. 12) 1: 856) by subsequent designation by Samouelle, 1819: 252. Synonymy: Heliothisa Meigen, 1832; Heliotis Sodoffsky, 1837; Chloridea Duncan & [Westwood], 1841; Asphila Guen��e, 1852; Heliocheilus Grote, 1865; Dorika Moore, 1881; Rhodosea Grote, 1883; Disocnemis Grote, 1883; Dysocnemis Grote, 1890; Neocleptria Hampson, 1903; Nubiothis Beck, 1996; Peltothis Beck, 1996. Heliothis viriplaca (Hufnagel, 1766) Pl. 1, fig. 1; male genitalia Pl. 3, fig. 20; female genitalia Pl. 8, fig. 37. Phalaena viriplaca Hufnagel, 1766, Berlinisches Mag. 3 (4): 406 (TL.: [Germany]: Berlin). Synonymy: Phalaena (Noctua) dipsacea Linnaeus, 1767 References: Christoph 1873, 1877 (Heliothis dipsaceus); Schwingenschuss 1938; Barou 1967 (Chloridea dipsacea); Kalali 1976; Modarres Awal 1994, 1997 (Chloridea viriplaca); Zahedi 1983 (Heliothis dipsacea); Hacker & Kautt 1999); Hacker & Meineke 2001; Hacker 2001; Ebert & Hacker 2002 (Heliothis viriplaca). Bionomics: Bivoltine, probably multivoltine (Hacker 2001), univoltine in Israel (Kravchenko et al. 2005). Moth in flight from March to September. The early stages have been described by Hampson (1903), Spuler (1908), Forster & Wohlfahrt (1971), Bretherton et al. (1979) and Skou (1991). The species flies by day as well as at night. The species inhabits steppe-like habitas, usually at medium altitude up to 2900 m. Larvae are polyphagous, feed on 70 species of herbaceous plants of 22 botanical families (prefer Caryophyllaceae, Fabaceae, Lamiaceae and Asteraceae). Distribution: West Palaearctic. Europe, North Africa, Near East, Caucasus, Transcaucasia, Central Asia, Kazakhstan, south Siberia (including Transbaikalia), China and north India. ��� In Iran (Pl. 10, fig. 49) distributed almost everywhere except eastern and some south-eastern provinces. Material examined: 351 specimens from provinces West Azerbaijan, East Azerbaijan, Ardebil, Guilan, Mazandaran, Golestan, Khorasan, Semnan, Tehran, Qazvin, Qom, Markazi, Zanjan, Kermanshah, Kordestan, Esfahan, Kohkiluyeh va Boyer-Ahmad, Chaharmahal va Bakhtiari, Lorestan, Fars and Kerman, collected between 10.III to 22.IX on elevations from 0 to 2900 m., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 6, DOI: 10.5281/zenodo.181966

Heliothis maritima Graslin, 1855 Pl. 1, fig. 2; male genitalia Pl. 3, fig. 31; female genitalia Pl. 8, fig. 38. Heliothis maritima Graslin, 1855, Annls. Soc. Entomol. France, Serie 3, 3: 65-74 (TL.: France: Vendee). Synonymy: Heliothis adaucta Butler, 1878; Chloridea maritima subsp. angarensis Draudt, 1938; Chloridea maritima subsp. bulgarica Draudt, 1938; Chloridea maritima subsp. centralasiae Draudt, 1938; Heliothis maritima subsp. septentrionalis Hoffmeyer, 1939; Chloridea maritima subsp. hungarica Kovacs, 1950; Chloridea maritima subsp. warneckei Boursin, 1963. References: Modarres Awal 1999 (Heliothis maritima). Bionomics: Bivoltine. Moths flying in two overlapping generations from June to September. Inhabits steppe habitats on elevation 830-2040 m. Larvae are polyphagous, feed on 28 species of herbaceous plants and shrubs of 13 botanical families, prefer Fabaceae, Asteraceae, Chenopodiaceae, Ericaceae and Cucurbitaceae. Distribution: Transpalaearctic. Europe, Near East, Caucasus, Transcaucasia, Central Asia, Kazakhstan, south Siberia, Far East, Korea, Japan, China and north India. ��� In Iran it is known from Khorasan, Tehran and Lorestan (Pl. 10, fig. 50). Material examined: Khorasan: Gonabad, 36 km N, 830 m., 7.VI. 1977 (Safavi, Pazuki & Abai). Tehran: Damavand, 1910 m., 7-25.VIII. 1986 (Rajabi); Evin, Tehran, 1600 m., 28.VI. 1974; 13.VII. 1974: 2.VIII. 1974 (Light trap); Malard, Karaj, 20.IV. 1971 (Sabzevari); Vanak, 15 km N Tehran, 1600 m., 1- 10.VII. 1962 (Vartian E.). Lorestan: Kamanda, N, Oshtoran Kuh, 2040 m., 22-24.VII. 1971 (Pazuki & Borumand). Schahrud, 1887, leg. O. Herz, 2 females (ZISP)., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on pages 6-7, DOI: 10.5281/zenodo.181966

Aedophron sumorita Ronkay, 2002 Bionomics: Univoltine. Moths flying from June to July. The biotope was described by Ronkay (2002) as a relatively high part of a long, deep rocky valley, near a small stream bordered by sparse stands of willow. The species inhabits medium and high elevations from 1650 to 2700 m. Larval host-plants are unknown. Distribution: Iranian. In Iran (Pl. 11, fig. 63) occurs in provinces West Azerbaijan, Kordestan, Hamedan, Zanjan, Lorestan, Esfahan and Fars. Material examined: West Azerbaijan: Ghasemlu valley, 1650 m, 19.VII. 2005, leg. Zahiri & Khiaban. Zanjan: Baba Sorkheh, 10 km W Bijar, 2180 m, 29.VI. 1975, leg. Pazuki; Zanjan, 50 km SW Zanjan-Bijar, 1700 m, 28.VI. 1975, leg. Pazuki. Kordestan: Stra��e Zanjan-Bijar, 53 km S Zanjan, 1700 m, 28-29.VI. 1975; leg. Ebert & Falkner; Ariz, 27 km W Sanandaj, 2200 m., 10.VII. 1975, leg. Ebert & Falkner. Hamadan: Khakadan, 25 km W, Hamedan, 2200 m, 21.VI. 2000, leg. Fabian (paratype, coll. L. Sz��cs��nyi, Budapest). Lorestan: Dorud, 06- 17.VI. 1978, leg. Parvin. Esfahan: Esfahan, Park-e-Ghamishlu, 2000 m, 16-17.VI. 1993, leg. Mirzayans & Badii; Shibak, 10 km S, Esfahan, 2700 m, 10.VI. 2000, leg. Fabian. (holotype, HMNH). Fars: Kamfiruz (Kam-Firuz), 1900 m, 26.VI. 1975, leg. Abai M., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 22, DOI: 10.5281/zenodo.181966

Genus Periphanes H��BNER, [1821] Type species: Phalaena Noctua delphinii Linnaeus, 1758 Synonymy: Chariclea Curtis, 1825, Chariclea Stephens, 1829, Philareta Moore, 1881, Calocharia Beck, 1996, Helivictoria Beck, 1996., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 13, DOI: 10.5281/zenodo.181966

Masalia perstriata fuscostriata (Brandt, 1941) Pl. 3, fig. 19; male genitalia Pl. 7, fig. 36. Timora fuscostriata Brandt, 1941, Mitt. M��nch. Ent. Ges. 31: 854, fig. 43 (TL.: Iran: Tahte Malek). References: Ebert & Hacker 2002 (Masalia perstriata fuscostriata). Bionomics: Univoltine. Moths flying from February to April. The species inhabits deserts and semideserts on low elevations (550-850 m). Larval host-plants are unknown. Distribution: Irano-Sindian. Pakistan, India. ��� In Iran (Pl. 12, fig. 67) occurs in provinces Hormozgan and Sistan va Baluchestan. PLATE 12. Figures 65 ���71. 65, Distribution of Masalia albida in Iran; 66, Distribution of M. philbyi in Iran; 67, Distribution of M. perstriata fuscostriata in Iran; 68, Provinces of Iran (1, West Azerbaijan; 2, East Azerbaijan; 3, Ardebil; 4, Guilan; 5, Mazandaran; 6, Golestan; 7, Khorasan [Khorasan has recently divided by 3 provinces: 7 (1): North Knorassan; 7 (2): Razavi Khorasan; 7 (3): South Khorasan]; 8, Semnan; 9, Tehran; 10, Qazvin; 11, Zanjan; 12, Kordestan; 13, Kermanshah; 14, Hamedan; 15, Markazi; 16, Qom; 17, Esfehan; 18, Yazd; 19, Lorestan; 20, Ilam; 21, Khuzestan; 22, Chahar Mahal va Bakhtiari; 23, Kohkiluye va Boyer-Ahmad; 24, Fars; 25, Bushehr; 26, Kerman; 27, Hormozgan; 28, Sistan va Balouchestan); 69, Climatic zones of Iran; 70, Geographical regions of Iran (1, borders of regions; 2, borders of districts), 1, North Iranian mountain region (1 a, North-Western Iran; 1 b, South Caspian mountains; 1 c, Turkmen-Khorasan mountains); 2, Caspian wet forest region; 3, South-West Iranian mountain region (3 a, Poshte-Kuh or Kabir-Kuh mountains; 3 b, Zagros mountains; 3 c, Karun Plain); 4, Central Iranian mountain region (4 a, Ghohrud mountains; 4 b, Kuhbenan mountains; 4 c, Gavkhan hollow); 5, South Iranian mountain region (5 a, mountains of Fars; 5 b, Kuh-e-Furgun mountains; 5 c, eastern part of the South Iranian mountains); 6, East Iranian mountain region (6 a, Dzham mountains; 6 b, Kayen mountains; 6 c, Pelengan mountains; 6 d, Serhed tableland); 7, high plains of the Iranian highland (7 a, Dasht-e- Kavir desert; 7 b, Dasht-e-Lut desert; 7 c, Sistan hollow, 7 d, Namakzar hollow); 71, Division of the Iranian territory into studied sites, abbreviations as in Table 1. TABLE 1. The two-way table of presence-absence data for Heliothinae species and areas in Iran. Geographical territories are abbreviated as: NNW���north part of the North-West Iran, the Araxes River basin, Talysh Mountains and Arasbaran Forest; NW���North-West Iran, excluding the Araxes River basin, NW of Zagros Mountains including West Azerbaijan, Kordestan and North of Kermanshah; C���the Caspian wet forest region; G ��� Golestan Forest; E���southern macroslope of the Alburs (=Elburs) Mountains including North of Tehran and Qazvin; NE���the Turkmen-Khorasan Mountains including Binalud and Kopet Dagh Mountains; TS���the Torkman-Sahra Transcaspian plain and other plains North-East Iran; Z���South-West Iran Mountain region, mainly Zagros Mountains including S Kermanshah, Lorestan, North of Khuzestan, Chaharmahal va Bakhtiari, Kohkiluye Boyer-Ahmad, Ilam, Eat of Esfahan, Markazi, Hamedan, excluding the part in Fars; ZF���Zagros Mts. in Fars; NZK���North Iran provinces Zanjan and Qazvin; NTQ��� North Iranian high plains, mainly Dasht-e-Kavir within the provinces of Tehran, Qom, Semnan, North of Esfahan; Gh���the Central Iranian mountains, mainly Ghohrud Mountains; SKh���the mountains of Southern Khorasan; NB���the mountains of Northern Baluchestan; Hmt ��� the mountains of Hormozgan including Genu Mountains; Bmt ��� the mountains of Southern Baluchestan; KzB ��� the plains of South Khuzestan and the coast of Bushehr; Oc ��� the coast of the Strait of Persian Gulf and the Oman Sea; I ��� the islands of the Strait of Persian Gulf. Material examined: Unknown place: Keine Angaben zum Fundort, Quli Kush, leg. Brandt (paratype, SMNK). Hormozgan: Geno M., Issin, 300 m, 11.III. 1991, leg. Mirzayans & Badii (HMIM); Stra��e nach Sirjan, km 107, Bandar-e-Abbas, 850 m, 07.III. 1973, leg. Ebert (SMNK); Kuh-e-Geno, 550-650 m, 01- 05.III. 1973, leg. Ebert (SMNK); Bander-Abbas, 40 km n��rdl, 04.IV. 1974, leg. Vartian (NHMW); Bander- Abbas, 40 km n��rdl, 31.III. 1972, leg. Exped. Mus. Vind. (NHMW); Bander-Abbas, 75 km n��rdl., 03.IV. 1974, leg. Vartian (NHMW). Sistan va Balouchestan: Bandar-e-Chabahar, 15.II. 1938, leg. Brandt (paratype, NHMW); Bandar-e-Chabahar, 15.II. 1938, leg. Brandt (NHMW)., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on pages 27-31, DOI: 10.5281/zenodo.181966

Genus Aedophron Lederer, 1857 Type species: Heliothis rhodites Eversmann, 1851, Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 15, DOI: 10.5281/zenodo.181966

Helicoverpa armigera (H��bner, [1808]) Pl. 1, fig. 6; male genitalia Pl. 4, fig. 25; female genitalia Pl. 8, fig. 41. Noctua armigera H��bner, [1808], Samml. Eur. Schmett. 4: pl. 79, fig. 370 (TL.: [Europe]). Synonymy: Noctua obsoleta sensu auct., nec Fabricius, 1793; Heliothis obsoleta Fabricius, 1775; Noctua barbara Fabricius, 1794; Heliothis pulverosa Walker, 1857; Heliothis conferta Walker, 1857; Heliothis uniformis Wallengren, 1860; Heliothis armigera fusca Cockerell, 1889; Heliothis guidellii Constantini, 1922; Helicoverpa armigera subsp. commoni Hardwick, 1965; Heliothis rama Bhattacherjee & Gupta, 1972. References: Bienert 1870; Lederer 1871 (Heliothis Armigera); Christoph 1877 (Heliothis armiger); Barou 1967 (Chloridea obsoleta); Kalali 1976 (Chloridea armigera); Modarres Awal 1994, 1997 (Heliothis (= Chloridea) obsoleta); Modarres Awal 1999 (Helicoverpa (= Heliothis) armigera); Hacker & Kautt 1999; Hacker & Meineke 2001; Hacker 2001; Ebert & Hacker 2002 (Helicoverpa armigera). Bionomics: Multivoltine, continuously brooded throughout the year. Occurs in various habitats from sea level to 3300m. Moths flying from February to December. Larvae have been recorded from a wide range of herbaceous plants, shrubs and trees from 38 botanical families. The crops attacked by the larvae most often in Near and Middle East are cotton and tomato. Taxonomic notes: Hardwick (1965) recognised three subspecies. The typical race flies throughout most of the range, ssp. conferta Walker in Australia and the Pacific, and ssp. commoni Hardwick on southeast China (Holloway 1989). Distribution: Cosmopolitan (Palaeotropic-Palaeosubtropical). Europe, Africa (including Madagascar and R��union), Near East, Caucasus, Transcaucasia, Kazakhstan, Central Asia, India, South-East Asia, Australia, Oceania, New Zealand. ��� In Iran (Pl. 10, fig. 54) occurs everywhere except the eastern provinces. Material examined: 648 specimens from provinces West Azerbaijan, East Azerbaijan, Ardebil, Mazandaran, Guilan, Golestan, Khorasan, Semnan, Tehran, Qom, Markazi, Qazvin, Hamedan, Zanjan, Kordestan, Kermanshah, Ilam, Lorestan, Esfahan, Charmahal va Bakhtiari, Kohkiluyeh va Boyer-Ahmad, Khuzestan, Fars, Yazd, Kerman, Bushehr, Hormozgan and Sistan va Baluchestan, collected between 16.II to 21.XII on elevations from 0 to 3300 m., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 12, DOI: 10.5281/zenodo.181966

Heliothis incarnata Freyer, 1838 Pl. 1, fig. 5; male genitalia Pl. 4, fig. 24; female genitalia Pl. 8, fig. 40. Heliothis incarnata Freyer, 1838, Neuere Beitr. Schmetterlingskde 3: 91, Tab. 256, Fig. 4 (TL.: [Turkey]: [Istanbul]). Synonymy: Heliothis boisduvalii Boisduval, 1840; Rhodocleptria incarnata f. bernaldezina Fernandez, 1951. References: Modarres Awal 1999 (Rhodocleptria incarnata), Gutleb & Wiesser 2001 (Chazaria incarnata), Hacker 2001; Ebert & Hacker 2002 (Heliothis incarnata). Systematic notes: The forewing upperside varies from reddish-brown, through sandy-brown, to pale yellowgreen, with all possible intergrades (Hacker 2001). Bionomics: Bivoltine (but univoltine according to Hacker 2001). Moths flying from February to October. The species inhabits dry rocky and chalky steppes and semi-deserts in the plains on elevation up to 2600 m. The early stages have been described by Hampson (1903) and Spuler (1908). Larvae feed on Gypsophila juzepczukii, Silene vespertina, S. viscosa, S. volgensis. Distribution: West Palaearctic. Central and South Europe, North Africa, Near East, Caucasus, Transcaucasia, Central Asia, Kazakhstan. ��� In Iran (Pl. 10, fig. 53) occurs in northern, western and southern provinces. Material examined: 207 specimens from provinces West Azerbaijan, East Azerbaijan, Golestan, Khorasan, Tehran, Qom, Hamedan, Zanjan, Lorestan, Khuzestan, Fars, Kerman, Bushehr, Hormozgan and Sistan va Baluchestan, collected between 28.II to 2.X on elevations from 30 to 2600 m., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 8, DOI: 10.5281/zenodo.181966

Periphanes victorina (Sodoffsky, 1849) Pl. 2, fig. 10; male genitalia Pl. 5, fig. 28; female genitalia Pl. 9, fig. 44. Heliothis victorina Sodoffsky, 1849, Stett. Ent. Ztg. 10: 130 (TL.: [Caucasus]). Synonymy: Chariclea prazanoffskyi Guen��e, 1852. References: Schwingenschuss 1938; Modarres Awal 1994, 1997 (Pyrrhia victorina). Bionomics: Bivoltine. Moths in flight from March to August. The species inhabits steppe habitats on lowmedium elevations from 100 to 1900 m. Larvae feed on Dictamnus albus and Salvia. Distribution: Mediterranean-Asiatic. South Europe, Caucasus, Transcaucasia. ��� In Iran (Pl. 11, fig. 58) occurs in provinces East Azerbaijan, Mazandaran and Tehran. Material examined: Azerbaijan: Khalatpushan, Tabriz, 26.III. 1954. Mazandaran: Amol, 23.VII. 1969, leg. Vartian E. (NHMW). Tehran: Damavand, 1900 m, 08- 25.VIII. 1976, leg. Rajabi; Damavand, 1900 m, 05- 11.VI. 1976, leg. Rajabi; Damavand, 1900, 21 - 27.VI. 1976, leg. Rajabi: Damavand, 1900 m, 28.V. 1976, leg. Rajabi; Damavand, 1900 m, 28.V-04.VI. 1976, leg. Rajabi; Absard, Damavand, 1900 m, 3-7.VII. 1978, leg. Pazuki & Sabzevari., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 14, DOI: 10.5281/zenodo.181966

Masalia philbyi (Brandt, 1941) Pl. 3, fig. 18; female genitalia Pl. 9, fig. 48. Timora philbyi Brandt, 1941, Mitt. M��nch. Ent. Ges. 31: 853, fig. 44 (TL.: Iran: Tahte Malek). Synonymy: Timora philbyi nuristana Boursin, 1960, Timora philbyi arabica Boursin, 1960. References: Ebert & Hacker 2002 (Masalia philbyi). Bionomics: Univoltine. Moths flying from March to April. The species inhabits deserts and semideserts on low elevations (30-850 m). Larval host-plants are unknown. Distribution: Iranian-Arabian. Saudi Arabia. ��� In Iran (Pl. 12, fig. 66) occurs in province Hormozgan. Material examined: Unknown place: Keine Angaben zum Fundort, Quli Kush, leg. Brandt (paratype, SMNK). Hormozgan: Stra��e nach Sirjan, km 107, Bandar-e-Abbas, 850 m, 07.III. 1983, leg. Ebert (SMNK); Kuh-e-Geno, 550-650 m, 01-05.III. 1973, leg. Ebert; Minab, 10 km N, 30 m, 03.III. 1973, leg. Ebert (SMNK); Bander-Abbas, 75 km n��rdl. 03.IV. 1974, leg. Vartian (NHMW); Bander-Abbas, 40 km n��rdl, 04.IV. 1974, leg. Vartian (NHMW); Rudan Sarzeh, 21 km W, 200 m, 3-4.III. 1978, leg. Pazuki (HMIM)., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 27, DOI: 10.5281/zenodo.181966

Periphanes delphinii (Linnaeus, 1758) Pl. 1, fig. 8; male genitalia Pl. 5, fig. 27; female genitalia Pl. 9, fig. 43. Phalaena (Noctua) delphinii Linnaeus, 1758, Syst. Nat. Ed. 10, 1: 518 (TL.: [Europe]). Synonymy: Chariclea darollesi Oberth��r, 1876 References: Bienert 1870; Barou 1967 (Chariclea Delphinii); Kalali 1976; Modarres Awal 1994, 1997 (Periphanes delphinii darallesi); Hacker 1990 (Periphanes delphinii delphinii); Hacker 2001; Ebert & Hacker 2002 (Periphanes delphinii). Systematic notes: Represented in Iran by subspecies tekke L.Ronkay, Varga & Hreblay, 1998. Bionomics: Bivoltine; univoltine according to Hacker (2001) and Kravchenko et al. (2005). Flight period: April to August The species inhabits vegetation-rich steppe habitats on elevations from sea level up to 2500 m. The early stages have been described by Hampson (1903), Spuler (1908), Forster & Wohlfahrt (1971) and Bretherton et al. (1979). Larvae feed on flowers and seeds of species of Consolida ajacis and Aconitum napellus. Distribution: Mediterranean-Asiatic. Central and South Europe, North Africa, Near East, Caucasus, Transcaucasia, Kazakhstan, Central Asia. ��� In Iran (Pl. 10, fig. 56) occurs in provinces West Azerbaijan, East Azerbaijan, Ardebil, Mazandaran, Golestan, Tehran, Hamedan and Fars. Material examined: 22 specimens from provinces West Azerbaijan, East Azerbaijan, Ardebil, Mazandaran, Golestan, Tehran, Hamedan and Fars, collected between 12.IV to 20.VIII on elevations from 0 to 2500 m, Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on page 13, DOI: 10.5281/zenodo.181966

Heliocheilus confertissima (Walker, 1865) Pl. 2, fig. 16; male genitalia Pl. 7, fig. 34. Leucania confertissima Walker, 1865, List. Spec. Lep. Ins. Coll. Br. Mus. 32: 625 (TL.: [India]: South Indostan). Synonymy: Heliocheilus hyalosticta Hampson, 1894, Raghuva perdentata Hampson, 1903, Heliocheilus mekrana Brandt, 1941, Heliocheilus designata Brandt, 1941, Raghuva ennedica Herbulot & Viette, 1952. References: Brandt 1941 (Heliocheilus mekrana, Heliocheilus designata); Ebert & Hacker 2002 (Heliocheilus confertissima). Bionomics: Univoltine. Moths flying from February to April. The species inhabits deserts and semi-deserts on lowlands and medium elevations (200 ��� 1100 m). Larval host-plants are unknown. Distribution: Afrotropical-Asiatic. East Africa, Iran, Turkmenistan, Pakistan, India, Myanmar. ��� In Iran (Pl. 11, fig. 64) occurs in provinces Qom, Hormozgan and Sistan va Baluchestan. Material examined: Qom: Qom, 20 km S dl., 1100 m, 28.IV. 1975, leg. Thomas (SMNK). Hormozgan: Sarkhoun, 08.III. 1972, leg. Mirzayans & Borumand; Sirik, 30 m, 19.III. 1991, leg. Mirzayans & Badii; Kuh-e- Geno, 550-650 m, 01-05.III. 1973, leg. Ebert; Shirin Rud, 45 km NE, Bandar-e-Abbas, 20 m, 02.III. 1973, leg. Ebert (SMNK); Stra��e nach Sirjan, km 107, Bandar-e-Abbas, 850 m, 07.III. 1973, leg. Ebert (SMNK); Bandar-Abbas (D��den), 17 km ��stl., 15.IV. 1972, leg. Exped. Mus. Vind. (NHMW); Bandar-Abbas (D��den), 17 km ��stl., 05.IV. 1974, leg. Vartian (NHMW). Sistan va Balouchestan: Bandar-e-Chabahar, 1938, leg. Brandt; Bandar-e-Chabahar, 21.II. 1938, leg. Brandt (paratype, SMNK). Sharistan: Kuche Makran, 6 [19].03.1901, leg. Zarudnyj (ZISP)., Published as part of Matov, Alexej, Zahiri, Reza & Holloway, Jeremy D., 2008, The Heliothinae of Iran (Lepidoptera: Noctuidae), pp. 1-37 in Zootaxa 1763 on pages 22-23, DOI: 10.5281/zenodo.181966