Agrotis ipsilon Hufnagel 1766
Agrotis ipsilon (Hufnagel, 1766) (Figs 42, 43, 62, 79, 92, 110) Phalaena ipsilon Hufnagel, 1766 b: 416. Type: [Alemania] Berlín region. Destroyed. Phalaena ypsilon (Hufnagel) typo: Rottenburg, 1776: 141. Phalaena ypsilon Rottenburg typo and authorship error: Treitschke, 1825: 152 (= A. suffusa ). Ph...
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Summary: | Agrotis ipsilon (Hufnagel, 1766) (Figs 42, 43, 62, 79, 92, 110) Phalaena ipsilon Hufnagel, 1766 b: 416. Type: [Alemania] Berlín region. Destroyed. Phalaena ypsilon (Hufnagel) typo: Rottenburg, 1776: 141. Phalaena ypsilon Rottenburg typo and authorship error: Treitschke, 1825: 152 (= A. suffusa ). Phalaena-Noctua upsilon (Hufnagel) typo: Stephens, 1829: 66 (= A. suffusa ). Agrotis ypsilon (Rottenburg) typo and authorship error: Staudinger, 1871: 421; Anonymous a, 1875: 67; Berg, 1882: 280; Meyrick, 1886: 32; Edwards, 1889: 85; Leech, 1889 a: 132; Beutenmüller, 1890: 212 (hosts); Butler, 1890: 691; Druce, 1881 –1900: 281; Berg, 1892: 236 (larval cannibalism); Tutt, 1892: 7 (systematic notes); Smith, 1893: 66 (synonymies); Hampson, 1894: 182 (diagnosis); Harvey & Knight, 1897: 79; Coquillett, 1897: 13, 15, 18, 21, 23 (Diptera and Tachinidae parasitoid species); Hudson, 1898: 30; Meyrick, 1899: 143 (collected at Hawaii Islands); Forbes, 1900: 104 (biology); Walsingham & Durrant, 1900: 8; Dyar, 1901: 453 (collected at Florida, USA); Verrill, 1902: 769, fig. 131 a, b; Needham, 1903: 25 (sipping nectar from “Acebo común,” Cephalanthus occidentalis L.); Hampson, 1903: 368; Dyar, 1904: 821 (collected at British Columbia, Canada); Hampson, 1905 a: 702 (synonymies); Forbes, 1905: 17, 21, 22, 240 (biology); Gibson, 1905: 107; Swezey, 1907: 118 (on Sonchus sp.); Froggatt, 1909: 52 (clover and cotton as hosts); Smith, 1909: 780 –781 (Diptera and Tachinidae parasitoid species); Henninger, 1910: 238; Swezey, 1910: 132 (collected at Hawaii, sugarcane and snuff as hosts); Foaden & Fletcher, 1910: 446 (cotton as host); King, 1911: 95, 132–134 (biology and damages); Davis, 1912: 17; Perkins, 1913: 145 –146 ( Agrotis Hawaiian checklist); Swezey, 1913: 234 (collected at Hawaii); Titus, 1914: 151 (damages and treatment); Rothschild, 1914: 319; Holland, 1903: 182, fig. 103; Rothschild, 1915: 230; Anonymous b, 1916: 145 (captured in the cabin of a ship); Jones, 1918: 5 (damages at Louisiana); McAlpine, 1918: 20 (attracted to molasses); Rothschild, 1920: 118; Moore, 1922: 17; Thomson, 1922: 524 (as exotic species at New Zealand); Illingworth, 1923: 278; Ripley, 1923: 179, 280, 281, 285, 302, 314, 317, 320, 321 (larval development and postembryology); Aldrich & Webber, 1924: 24, 88 (Diptera parasitoid species); Anonymous c, 1924: 353 ( Berecyntus sp. as parasitoid species); Muesebeck, 1924: 32 (Ichneumonidae parasitoid species); Seitz, 1924: 56, pl. 9, row d (diagnosis); Williams, 1930: 283, 300, 401 (migration and economic importance); Williams, 1941: 62 (relationship between spermatophore and female genitalia); Chiesa Molinari, 1942: 328 –329 (diagnosis and control); Sellers, 1943: 44 (parasitoids); Köhler, 1945: 101 –102, pl. II, figs a, b (male genitalia); Quintanilla, 1946: 404 –405 (diagnosis and control); Hedges, 1949: 75; Pepper et al. , 1954: 12 (beetroot as host); Roeder, 1963: 203, figs 2, 4 (ultrasound produced by adult in flight); Artigas, 1972: 7, 11–14 (biology); Moulding & Madenjian, 1979: 138 (sampling in USA oak forests). Agrotis ypsilon (Hufnagel) typo: Grote, 1875 b: 313; Pagenstecher, 1902: 278; Holloway, 1967: 41 (migration). Peridroma ypsilon (Rottenburg): Butler, 1889: 380 (new combination); Butler, 1890: 691 (corrects the error, mentions that it was a lapsus calami ). Rhyacia ipsilon (Hufnagel): Franssen, 1935: 109 –137 (biology). Agrotis ypsilon Linnaeus authorship error: Forbes, 1954: 48. Agrotis ipsilon (Hufnagel): Linsley & Usinger, 1966: 159 (collected at Galápagos Islands); Hayward, 1969: 42 (hosts); Angulo & Weigert, 1975 a: 69 –70, 98, 129, 133, figs 118, 119, 154, 175 (egg, larvae, and pupae); Angulo & Weigert, 1975 b: 173 (mimicry of aggression in larvae); Angulo & Quezada, 1975: 117 –124, figs 1, 3, 5, 7 (description of all stages and differences with Agrotis malefida Guenée auct . nec. Agrotis robusta Blanchard); Hayes, 1975: 163, figs 31–32 (diagnosis and collected at Galápagos Islands); Zewadski, 1976: 154 –155 (biology); Linsley, 1977: 29 (collected at Galápagos Islands); Angulo, 1978: 15 –16 (larval behavior); Krombein & Hurd, 1979: 259, 260, 282, 284, 650, 651, 676, 698 (parasitoids); Pautler et al. , 1979: 3 –7 (virgin female trap); Kogan & Kuhlman, 1982: 31 (biology); Troester et al. , 1982: 1 –33 (USA attack prediction model); Fisher, 1983: 24 (photoperiod related to copulatory changes); Angulo & Jana-Saenz, 1984: 77, 82 (larval integument); Parra et al. , 1986: 82, 89–90, 98, 100, figs 41, 80–82 (diagnosis, male genitalia, and key); Godfrey, 1987: 567, Fig. 26.404 (larvae); Carrillo et al. , 1988: 34 –35 (biology and collecting curve); Olivares & Angulo, 1989: 24, 25 (head and larval integument); Jana-Sáenz, 1989: 63, 66–68, figs 2, 7, 8, 11 (diagnosis, male genitalia, and key); Poole, 1989: 43 (world noctuid checklist); López, et al. , 1990: 3479 –3491 (use of different species pheromone traps at the same time); Salama et al. , 1990: 147 –151 (control with Bacillus thuringiensis Berliner); Pedgley & Yathom, 1993: 67 –71 (migration in Israel); Showers et al ., 1993: 2303 –2314 (migration in USA); Artigas, 1994: 567 –569, pl. 27, fig. 7 (diagnosis, life cycle, biological control, damages, hosts, economic importance, distribution, and international implications); Igarzábal et al. , 1994: 103 –104, 123, 125, figs 1, 19, 37, 55, 73, 97–99 (damage, biology, and larval key); McNeil et al ., 1995: 282, 284, 286, 287, 296, 298 (migration); Gadenne et al. , 1997 (morphology and hybrid communication system of Agrotis ipsilon and A. segetum ); Greenslade et al ., 1999: 1162, 1163, 1166 (migration to Subantarctic Island); Beshkov & Goater, 2000: 42 (attracted by aphids secretions); Koch & Waterhouse, 2000: 24, 51, 53, 55, 58, 60, 62, 64, 66, 68, 71, 76, 86, 98, 119, 121, 123, 125, 128, 130, 132, 134, 136, 138, 141, 146, 158, 172, 196, 198, 201, 224 (agricultural and forestry importance); Angulo & Olivares, 2001: 56 (pupa key); Carrillo et al. , 2001: 28 (winter larvae at Valdivia, Chile); Fullard, 2001: 1376, 1378 (auditory system response to a Hawaiian species of bat); Silvegren, 2003: 24, 27, 28, 31–33 (pheromone production); Lafontaine, 2004: 250 –251, fig. 136, pl. L, figs 36–38, pl. 34, fig. 6, lam 50, fig. 4 (larval and adult diagnosis); Pastrana, 2004: 156 –157 (hosts); Specht et al. , 2004: 400; Angulo & Olivares, 2005: 138 (diagnosis, genitalia of both sexes, and hosts); Specht et al. , 2005: 136 (collected at Rio Grande do Sul, Brazil); Angulo et al. , 2006: 556 (immature stages on a high altitude salar); Capinera, 2006: 1 –5 (diagnosis, life cycle, and control); Pogue, 2006: 3, 13, 19, 20, 67, fig. 9, map 6 (diagnosis and distribution); Carrero & Planes, 2008: 284 (biology); Wood et al ., 2009: 532 (migration); San Blas & Barrionuevo, 2013: 1157, figs 3 A– 3 D (differences with A. malefida and A. robusta ). Agrostis ypsilon (Rottenburg) generic and specific typo and authorship error: Ortiz Romero & Zanabria, 1979: 122 (pest); Giménez, 1988: 81 (sunflower as host). Agrostis ipsilon (Hufnagel) generic typo: Reyes Vaca, 1988: 106 (sunflower as host). Noctua suffusa [Denis & Schiffermüller], 1775: 80. Type: [Austria] Región de Viena. Destroyed. Agrotis suffusa ([Denis & Schiffermüller]): Treitschke, 1825: 152; Stephens, 1829: 66; Boisduval & Guenée, 1852: 268 –269; Stephens, 1856: 65; Staudinger, 1871: 421 (= A. ypsilon Rottenburg); Butler, 1874: 33 (collected at New Zealand); Grote, 1874: 135; Grote, 1875 a: 193; Butler, 1877: 383; Butler, 1882: 126; Butler, 1883: 160 (collected at India); Meyrick, 1886: 32; Cotes & Swinhoe, 1888: 309 (collected at India); Leech, 1889 a: 132; Leech, 1889 b: 499 (collected at Japan and Korea); Poole, 1989: 57 (= A. ipsilon Hufnagel). Phalaena Bombyx idonea Cramer, 1780: 150, pl. 275, fig. h; Cotes & Swinhoe, 1888: 309 (= A. suffusa ). Type: [USA] New York. Unknown location. Agrotis idonea (Cramer): Boisduval & Guenée, 1852: 269 (as a variety of A. suffusa ); Poole, 1989: 49 (= A. ipsilon Hufnagel). Bombyx spinula Esper, 1786: pl. 63, figs 6, 7; Treitschke, 1825: 152 (= A. suffusa ); Stephens, 1829: 66 (= A. suffusa ); Godart et al. , 1837: 255 (= N. suffusa ); Druce, 1881 –1900: 281 (= A. ypsilon Rottenburg); Poole, 1989: 56 (= A. ipsilon Hufnagel). Type: type designation not mentioned. Phalaena Bombyx spinifera Villers, 1789: 174; Stephens, 1829: 66 (= A. suffusa ); Godart et al. , 1837: 255 (= N. suffusa ). Type: France. Unknown location. Noctua spinifera (Villers): Treitschke, 1825: 152 (= A. suffusa ); Stephens, 1829: 66 (= A. suffusa ). Agrotis spinifera (Villers): Treitschke, 1827: 382 –383; Stephens, 1829: 66 (= A. suffusa ); Poole, 1989: 56 (= A. ipsilon Hufnagel). Phalaena spinula Donovan, 1801: 52, pl. 345, figs 2–3; Poole, 1989: 56 (new homonym of Bombyx spinula Esper). Junior secondary homonym of Bombyx spinula Esper. Type: type designation not mentioned. Agrotis telifera Harris, 1841: 323; Riley, 1869: 80 (redescription, stages, and economic importance); Grote, 1874: 135 (= A. suffusa ); Grote, 1875 a: 193 (= A. suffusa ); Grote, 1875 b: 313 (= A. ypsilon Hufnagel); Saunders et al. , 1880: 40 (diagnosis of larvae and adult); Treat, 1882: 65 (damages); Druce, 1881 –1900: 281 (= A. ypsilon Rottenburg). Poole, 1989: 57 (= A. ipsilon Hufnagel, junior secondary homonym of Agrotis telifera Donzel, 1837). Type: [USA] Massachusetts. Lafontaine (2004) mentions that it is likely to be found at MCZ, but it has not yet been found. Noctua robusta of authors, not Blanchard, 1852: 75; Berg, 1882: 280 (= A. ypsilon Rottenburg); Butler, 1882: 126 (= A. suffusa ). See Agrotis robusta . Agrotis bipars Walker, (1857) 1856: 334; Leech, 1889 a: 378; Druce, 1881 –1900: 281 (= A. ypsilon Rottenburg); Poole, 1989: 45 (= A. ipsilon Hufnagel). Holotype: ♀ Venezuela (BMNH). Image examined. Agrotis consueta Walker, (1857) 1856: 334; Butler, 1882: 126 (= A. hostilis Walker in part and = A. bipars Walker in part ); Butler, 1889: 378 (= Agrotis bipars ); Hampson, 1903: 353 (= Feltia malefida (Guenée)). See Agrotis malefida. Agrotis frivola Wallengren, 1860: 169; Berg, 1882: 280 (= A. ypsilon Rottenburg); Poole, 1989: 48 (= A. ipsilon Hufnagel). Type: ♂ [Uruguay] Montevideo (NRS). Not examined. Noctua aneituma Walker, 1865 a: 701. Type: ♂ Nuevas Hébridas (BMNH). Image examined. Agrotis aneituma (Walker): Butler, 1889: 376 (= Agrotis munda Walker); Walsingham & Durrant, 1900: 7 –8 (= Agrotis munda Walker); Hampson, 1903: 164 (= Euxoa infusa (Boisduval)); Poole, 1989: 43 (world noctuid checklist); Lafontaine, 2004: 250 –251 (= A. ipsilon Hufnagel). Agrotis ipsilon aneituma (Walker): Holloway, 1977: 55, 60, 143, 157, 177, 201, 220, 208, 209, 214,217, 218, 242, 274 (diagnosis, biology, and migration); Common, 1990: 55, 64, 467, 485 (brief diagnosis, ectoparasites, and hosts). Agrotis suffusa var. pepoli Bertoloni, 1874: 139; Hampson, 1903: 368 (= A. ypsilon Rottenburg); Poole, 1989: 53 (= A. ipsilon Hufnagel). Type: Bondeno, Italy. Unknown location. Diagnosis. Agrotis ipsilon differs from other South American species of Agrotis by the following combination of characters: 1) male antenna bipectinate, widest segment 3 × as wide as central shaft, abruptly tapered at 3 / 4 its length; 2) forewing area between wing base and postmedial line darker than ground color, in some specimens as dark diffuse flecks; 3) reniform spot distal margin with a sharp streak extending between M 1 –M 2 veins, reaching postmedial line; 4) in male genitalia uncus slightly curved, narrowed to the apex; 5) vesica with right basal diverticulum absent; 6) in female genitalia appendix bursae 1.5– 2 × as long as corpus bursae; and 7) corpus bursae with one signum. Redescription. Male (Fig. 42). Head . Palpus whitish ventrally; frons smooth, central projection and raised edge absent. Antenna basal 2 / 3 bipectinate, widest at 1 / 5 its length, abruptly tapered at 3 / 4 its length, widest segment 3 × as wide as central shaft, anterior process 4 × as wide as posterior process. Thorax . Patagium with brown postbasal line and black medial line; tegulum without lines. Forewing length 16.2–19.5 mm; ground color grayish brown, area between wing base and postmedial line darker than ground color, in some specimens as dark diffuse flecks; subcostal band dark grayish brown; basal area undifferentiated; basal line undifferentiated; antemedial line black, double, convex between veins, extended as a short sharp tooth between 1 A+ 2 A vein and posterior margin, never close to medial line; claviform spot concolorous with ground color, black edged; orbicular spot strongly oval, extending toward reniform spot, rarely contiguous with reniform spot, grayish brown, black edged with grayish center, some specimens with orbicular spot concolorous with subcostal band, slightly differentiated by black edge; reniform spot concolorous with orbicular spot, distal margin with a sharp streak extending between M 1 –M 2 veins to postmedial line; discal cell concolorous with ground color, some specimens with a black streak of variable width joining both spots; medial line differentiated in light specimens as a dark thick waved band; postmedial line black, double, concave between veins; subterminal line light brown and black edged, strongly concave between veins, extended basally as arrows, longest ones placed between M 1 –M 2 –M 3 veins, in some specimens the first arrow joined with reniform streak like one continuous line; terminal line a series of darkish lunulae between veins; fringe concolorous with forewing ground color with dark transverse lines at veins apex. Hind wing iridescent, with light brown veins in lighter specimens and distal margin grayish brown on darker specimens; fringe iridescent, dark at apex of veins. Abdomen . Light grayish brown with dark dorsal line. Genitalia (Fig. 62). Uncus slightly curved, narrowed to the apex, spine-like ended. Clavus slightly sclerotized, cylindrical, short, 2 × as long as wide. Ampulla 1 / 5 × as long as valve, basal half expanded, then narrowed to half its widest diameter. Vesica (Fig. 79) 6 × as long as aedeagus, consisting of 1 1 / 2 wide loop, right basal diverticulum absent, basal spined band present, vesica swollen in apical 1 / 7, in some specimens it duplicates minor diameter. Female (Fig. 43). Differences from male. Forewing length: 15.4–20.6 mm; antenna filiform; forewing uniformly dark grayish brown between wing base and postmedial line; hind wing hyaline diffuse grayish brown on veins and margins. Genitalia (Fig. 92). Posterior apophysis 1.5 × as long as anterior apophysis; ductus bursae 2 × as long as anterior apophysis; corpus bursae 12 × as long as anterior apophysis, with one signum, apex globose; appendix bursae 1.5– 2 × as long as corpus bursae, consisting of one wide loop, apex globose; ductus seminalis originating laterally near corpus bursae apex. Biology. For Argentina and Chile there are several works that treat the biology of this species. Carrillo et al. (1988) describes the biology with temporal distribution and abundance for Chile. Angulo et al. (1990) and Angulo & Olivares (2005) give keys to larvae and adults. Igarzábal et al. (1994) do something similar but from the viewpoint of their economic importance to Argentina. Lafontaine (2004) makes a larval diagnosis. Parasitoids. The following list parasitoids for A. ipsilon : Harvey & Knight (1897), Smith (1909), Aldrich & Webber (1924), Muesebeck (1924), Sellers (1943), Krombein & Hurd (1979), and Common (1990). Artigas (1994) lists the following parasitoids: HYMENOPTERA. Braconidae: Apanteles bourquini Blanchard, Macrocentrus collaris Spinola, Meteorus rubens Nees. Ichneumonidae: Amblyteles sp., Ophion sp. DIPTERA. Tachinidae: Archytas cirphis Curran, Bonnetia comta (Fallen), Carcelia formosa (Aldrich & Webber), Chaetogaeclia montícola (Bigot), Eucelatoria armigera (Coquillet), Euphorocera claripennis (Macquart), Gonia longipulvilli Tothill, Gonia sequax Williston, Lespesia archippivora (Riley), Madremyia saundersii (Williston), Sisyropa eudryae (Townsend), Tachinomyia panaetius (Walker), Prosopochaeta fidelis (Reinhard). Hosts. Angulo et al. (1990) and Angulo & Olivares (2005) cite economically important hosts. Artigas (1994) and Pastrana (2004) give a list of cited hosts for Argentina. Some economically important hosts are: chard, peppers, alfalfa, cotton, celery, oats, broccoli, onions, cereals including corn, cauliflower, chrysanthemum, crucifers, dahlia, asparagus, raspberries, strawberries, lettuce, peanuts, melons, potatoes, paprika, pine (roots), beet, cabbage, watermelon, snuff, tomato, clover, wheat, carrots, squash, etc. (Artigas, 1994; Pastrana, 2004) There are several works on hosts for other parts of the World: Beutenmüller (1890), Froggatt (1909), Swezey (1910), Foaden & Fletcher (1910), King (1911), Pepper et al. (1954), Riley (1869), and Common (1990). Distribution. Worldwide distribution, except Poles (Fig. 110). Material examined. (28 ♂, 24 ♀). ARGENTINA: Córdoba. Córdoba, ♂ 23 -X- 1947 (F.P. López) (IMLA). Mendoza. Las Heras, Reserva Villavicencio, 1265m, 2 ♂ 3 ♀ 22 -XI- 2007 (GSB) (IADIZA); Santa Rosa, Reserva Ñacuñán, 561m, 2 ♀ 9 -IX- 2006 (GSB & E. Ruiz Manzanos) (IADIZA); Las Heras, Tambillos, 2462m, ♂ 11 -XII- 2007 (GSB & E. Scheibler) (IADIZA). Neuquén. Pucará, 707m, ♂ XII- 1959 (Orrow) (IADIZA). Río Negro. Cipolletti, El Cuy, ♂ 13 -XI- 1965 (IMLA). Salta. 2 km N Cafayate, 1585m, ♂ 24 -XI- 2006 (GSB & E. Ruiz Manzanos) (IADIZA); El Galpón, ♂ ♀ 15 -IX- 1966 (IMLA). Tucumán. Trancas, La Higuera, ♂ ♀ 8 -XI- 1966 (Köhler) (IMLA). BERMUDA: Botanical Gardens Paget, ♂ 8 -XI- 1987 (D.J. Hilburn) (USNM); Brighton Hill, Devonshire Park, ♀ 12 -III- 1988 (D.C. Ferguson) (USNM); Spittal Pond Nature Reserve, Smith : Published as part of Blas, Germán San, 2014, Agrotis Ochsenheimer (Lepidoptera, Noctuidae): a systematic analysis of South American species, pp. 1-64 in Zootaxa 3771 (1) on pages 47-51, DOI: 10.11646/zootaxa.3771.1.1, http://zenodo.org/record/285634 : {"references": ["Hufnagel, J. S. (1766 b) Forsetzung der tabelle von de nachtvogein. IV. Fortsetzung der dierten tabelle von de Insecten, besonders von denen so genannten nacheulen als der zwoten klasse. Der nachvogel hiesiger gegend. 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