Agrotis Ochsenheimer (Lepidoptera, Noctuidae): a systematic analysis of South American species Author Blas, Germán San text Zootaxa 2014 3771 1 1 64 journal article 36891 10.11646/zootaxa.3771.1.1 9f2ddd99-87fb-478b-938c-99e60937a1b7 1175-5326 285634 7C4129A9-DE4F-4CAE-AD88-EE14195A3E64 Agrotis Ochsenheimer, 1816 Agrotis Hübner, 1806 : 1 . Work rejected for nomenclatural purposes by International Commission on Zoological Nomenclature, 1926, Smithsonian Miscellaneous Collections 73 (4) Opinion 97: 19. Also idem, 1954, Opinions and Declarations rendered by the International Commission on Zoological Nomenclature 6 Opinion 278: 140. Only included species: Agrotis segetis . Agrotis Hübner, 1808 : 4 . Work rejected for nomenclatural purposes by International Commission on Zoological Nomenclature 1966, Bulletin of Zoological Nomenclature 23 Opinion 789: 213. Placed on the Official Index of Rejected and Invalid Generic Names in Zoology: Name No. 1826. Only included species: Agrotis grata Hübner (1808) . Agrotis Ochsenheimer, 1816 : 66 . Type species: Noctua segetum [ Denis & Schiffermüller], 1775 . Designated by Curtis, 1827 : 165 . Agronoma Hübner, 1821 : 227 . Type species: Noctua valligera [ Denis & Schiffermüller], 1775 = Phalaena vestigialis Hufnagel, 1766a . Designated by Grote, 1895 : 64 . Georyx Hübner, 1821 : 227 . Type species: Noctua segetum [ Denis & Schiffermüller], 1775 . Designated by Hampson, 1903 : 153 . Scotia Hübner, 1821 : 226 . Type species: Noctua cinerea [ Denis & Schiffermüller], 1775 . Designated by Hampson, 1903 : 153 . Noctua Boisduval, 1829 : 63 . Type species: Phalaena exclamationis Linnaeus, 1758 . Designated by Duponchel, 1829 : 71 . Note: Noctua Boisduval (1829) is pre-occupied by Noctua Linnaeus (1758) , it is therefore a junior homonym of the latter. Psammophila Stephens, 1850 : 72 . Type species: Noctua ripae Hübner, 1823 . By monotypy. Note: Psammophila Stephens (1850) is pre-occupied by Psammophila Brown (1827) , it is therefore a junior homonym of the latter. Tetrapyrgia Walker, 1865b : 711 . Type species: Tetrapyrgia graphiphorides Walker, 1865b = Agrotis porphyricollis Guenée , in Boisduval & Guenée, 1852 . By monotypy. Elegarda Walker, 1865b : 712 . Type species: Agrotis dorsicinis Walker, 1858 = Agrotis porphyricollis Guenée , in Boisduval & Guenée, 1852 . Designated by Nye, 1975 : 173 . Comophorus Alphéraky, 1887 : 168 . Type species: Comophorus villosus Alphéraky, 1887 . By monotypy. Note: Comophorus Alphéraky (1887) is pre-occupied by Comophorus Agassiz (1848) , it is therefore a junior homonym of the latter. Objective replacement name is Lycophorus Staudinger. Porosagrotis Smith, 1890 : 11 . Type species: Agrotis muraenula Grote & Robinson, 1868 = Agrotis vetusta Walker, 1856 . By original designation. Lycophorus Staudinger , in Staudinger & Rebel, 1901 : 154 . Type species: Comophorus villosus Alphéraky, 1887 . By monotypy. Note: Lycophorus Staudinger is the objective replacement name for Comophorus Alphéraky (1887) . Onychagrotis Hampson, 1903 : 465 . Type species: Agrotis rileyana Morrison, 1875 . By original designation. Neosema Rebel, 1907 : 55 . Type species: Neosema sesamioides Rebel, 1907 . By monotypy. Powellinia Oberthür, 1912 : 330 . Type species: Luperina lasserrei Oberthür, 1881 . By monotypy. Hermonassoides Strand, 1915 : 157 . Type species: Agrotis problematica Strand, 1915 . By monotypy. Brachypteragrotis Viette, 1959 : 25 . Type species: Brachypteragrotis patricei Viette, 1959 . By original designation. Crassagrotis Beck, 1992 : 180 . Type species: Noctua crassa Hübner, 1803 . By original designation. Putagrotis Beck, 1992 : 181 . Type species: Noctua puta Hübner, 1803 . By original designation. Leucagrotis Beck, 1992 : 181 . Type species: Agrotis graslinii Rambur, 1848 . By original designation. Militagrotis Beck, 1992 : 181 . Type species: Agrotis militaris Staudinger, 1888 . By original designation. Agrotis subgenus Striagrotis Beck, 1996 : 91 . Type species: Noctua fatidica Hübner, 1824 . By original designation. Agrotis subgenus Exagrotis Beck, 1996 : 91 . Type species: Phalaena exclamationis Linnaeus, 1758 . By original designation. Agrotis subgenus Ripagrotis Beck, 1996 : 92 . Type species: Noctua ripae Hübner, 1823 . By original designation. Agrotis subgenus Spinagrotis Beck, 1996 : 92 . Type species: Agrotis biconica Kollar , in Kollar & Redtenbacher, 1844 . By original designation. Agrotis subgenus Schawagrotis Beck, 1996 : 92 . Type species: Agrotis schawerdai Bytinsky-Salz, 1937 . By original designation. Type species. Noctua segetum [ Denis & Schiffermüller], 1775 . Designated by Curtis, 1827 : 165. Etymology. Agrotis derives from the Greek word Αγgότις that means from the land or from the soil. Thus named by Ochsenheimer (1816) because the larvae live in the soil. Diagnosis. Lafontaine (2004) identifies six character states that define adults of the genus: 1) vesica of male genitalia with presence of a spiny bar near base on left side (absent in some primitive South American species); 2) basal swelling present, usually with numerous lobes; 3) long looping vesica in male genitalia and appendix bursae of female genitalia; 4) absence of medial and apical diverticula; 5) apex of aedeagus hook-like shaped on right side; and 6) costa margin with a marked rounded pouch close to ampulla base. Diagnosis of South American species. The species of Agrotis can be differentiated from other South American Noctuidae genera by the characters listed above in addition to the following characters: 1) aedeagus with posterior half sclerotized and anterior half lightly sclerotized, almost membranous in some species, being confused with the bulbus ejaculatorius ( fausta -group); 2) aedeagus projected into the vesica in the following way: a dorsal strip with posterior half projected ventrolaterally through right margin, a right ventrolateral strip, and a strip close to the latter, like a band with posterior 1/3 forming a 1/4 of a spiral; 3) vesica with right basal diverticulum present, absent only in some species; 4) vesica with postbasal, medial, subapical, and apical diverticula absent; 5) vesica lacking cornuti; and 6) tergum 8 sclerotized like a longitudinal rectangle, slightly narrowed anteriorly, with an anterior membranous area, and an anterior band slightly sclerotized and projected laterally. Other characters that can help to identify the species of this genus are: 1) male antenna never doubly-biserrate; 2) forewing costal band concolorous with forewing ground color or darker, never lighter; 3) forewing postmedial line with no strong basal projections on posterior half. Generic redescription of South American species. Head. Frons smooth, with central pointed projection, or with a circular or subrectangular raised edge. When a raised edge is present, anterior surface rugous and can have a slightly pointed anterior projection. Antenna concolorous with forewing ground color, bifasciculate, biserrate or bipectinate in males, filiform in females. Labial palp with long hair-like scales on ventral margin of basal and medial segments, apical segment with short and wide scales. Thorax . Concolorous with forewing ground color; patagium concolorous with thorax, darker, or distally lighter, with transverse lines; tegulum concolorous with thorax, lighter, or darker, basal and marginal dark lines can be present. Legs with whitish rings on segment joints and on base and apex of tibial spurs. Forewing length in males: 10.6–20.9 mm and in females: 9.2–21.1 mm; veins R2, R3, and R4 stalked; R5 from anterior corner of areole; M1 from posterior corner of areole; M2, M3, and CuA1 adjacent, from posterior corner of discal cell; CuA2 from apical 1/3 of discal cell. Hind wing with vein Sc+R1 from basal 1/3 of discal cell; Rs and M1 stalked, from anterior corner of discal cell; M2 weak; M3 and CuA1 adjacent, from posterior corner of discal cell; CuA2 from apical 1/3 of discal cell. Under side of both wings with neither generic nor specific characters, the color pattern is composed of dark shadows not constant even in different specimens of the same species. Abdomen . Tergum 8 sclerotized like a longitudinal rectangle, slightly narrowed anteriorly, with an anterior membranous area, and an anterior band slightly sclerotized and projected laterally ( Fig. 2 ). Sternum 8 sclerotized like a transverse rectangle, with a slightly sclerotized anterior oval area ( Fig. 2 ). Male genitalia . Uncus uniformly curved over its entire length or sinuous, tapered apically, apex generally rounded; in Agrotis ipsilon apex very thin, like a spine; with no spine-like setae. Tegumen with strong or marked “shoulders,” arms extended to valve costa, pleurite like a vinculum contiguous structure, elongate and narrow, obliquely joined to end of tegumen arms. Juxta subrectangular, ventral 1/3 of lateral margins subquadrate and projected, ventral margin projected as a strongly sclerotized spine-like triangle. Anal tube with two ventrolateral bands with posterior half less sclerotized. Clavus varying from an undifferentiated area of hair-like scales to a long cylindrical projection, 4 × as long as wide. Valve with anterior margin convex near ampulla apex, posterior margin convex in dorsal half of valve, shape variable as follows: 1) subrectangular, neither narrowed nor widened in any part, 2) subrectangular, curved, elongate, very narrow, and 3) subrectangular, basal half narrow, then widened; cucullus apex strongly or slightly projected anterodorsally (some species with no projection at all); sacculus strongly sclerotized, with two possible shapes: 1) 3/5 × as wide as valve or 2) between 4/6–9/10 × as wide as valve; costa margin with a rounded pouch close to clasper base; digitus and editum absent; clasper proper absent; ampulla uniformly curved inward, 1/4–1/6 × as long as valve, teardrop shaped basally, with a flat ventral margin adjacent to valve, twisting 1/4 of a turn to apex, flat-ended; saccus subtriangular or hemispherical, ventrally projected as a spine, some species with a dorsal notch. Aedeagus completely sclerotized, or posterior half sclerotized and anterior half lightly sclerotized (anterior half almost membranous in some species), being confused with the bulbus ejaculatorius; aedeagus projected onto base of vesica in following way: a dorsal strip with posterior half projected ventrolaterally through right margin, another right ventrolateral strip, and a third strip close to the latter, like a band with posterior 1/3 making a 1/4 turn; vesica 2–12 × as long as aedeagus, and consisting of a variable number of wide loops, basal swelling present, right basal diverticulum subtriangular, subcylindrical, or absent, no other diverticula present, basal spined band present on left side, absent in some species, vesica same diameter through entire length or gradually swelling toward apex. Female genitalia . Anal papillae slightly sclerotized, on lateral view 2 × as long as wide, with hair-like setae; posterior apophysis 1–2 × as long as anterior apophysis; ostium bursae membranous; ductus bursae 1.5–3 × as long as anterior apophysis, membranous; corpus bursae 3–12 × as long as anterior apophysis, with or without one or two signa, apex globose or subtriangular; appendix bursae 1.5– 14 × as long as corpus bursae, consisting of a variable number of wide loops, apex globose or subtriangular; ductus seminalis originating at corpus bursae apex or laterally, near its apex. Biology. Species of Agrotis are usually found in open xeric areas. In western North America , most Agrotini species avoid flying in summer, larvae feed during spring and aestivate during summer, with the adults emerging at the end of summer when temperatures are lower ( Lafontaine 2004 ). According to material checked for this study, the flight period of Agrotis species in South America is mainly between late spring and autumn (from November to April), but some specimens are collected throughout the year, even in winter ( Table 1 ). Larvae of this genus belong to the “true cutworms,” because they cut stems and leaves of young plants and carry them to holes in the soil where they feed. In some cases, larvae also feed on plant roots. The combination of larval habits being subterranean, nocturnal, and living in desert areas (generally unoccupied by people), make studying immature stages difficult and result in poor biological knowledge of almost all species. Economically important species, such as Agrotis ipsilon (Hufnagel) and A. malefida Guenée , are best known biologically, but there are no detailed studies on host preferences or larval habits. Biology of the better-known species (e.g., A. ipsilon and A. malefida ) indicates that egg-laying begins in spring and continues throughout the year. Each female lays up to 2,000 eggs. Larvae live buried in the ground, where they construct a protection cell. At dusk and at night they leave the cell to feed on stems and leaves of young plants. Larvae can spend the summer in diapause within the cell. Pupation occurs inside the cell. Adults can emerge all year long, but most emergences occur in autumn. Winter can be spent as larvae or pupae ( Artigas 1994 ). In Argentina , depending on the species, there can be between two and six generations a year ( Igarzábal et al . 1994 ). Their life cycle is related to temperature, humidity, and food availability, generally taking between 40 and 60 days to complete. Economic importance and hosts. Numerous species of Agrotis are economically important in all areas the world, but only a few occur on more than one continent. Some species can damage 100% of seedlings ( Artigas 1994 ). In Argentina , Agrotis species are not major pests, but one larva can cut between 10 and 15 young plants at ground level, causing the death of the plants ( Igarzábal et al . 1994 ). Larvae can attack many different crops: alfalfa, corn, soybeans, tomatoes, peppers, potatoes, cabbage, melon, etc. ( Chiesa Molinari 1942 ; Angulo & Weigert 1975a ; Artigas 1994 ; Igarzábal et al . 1994 ; Pastrana 2004 ; Angulo & Olivares 2005 ). They also attack ornamental plants and weeds ( Artigas 1994 ; Pastrana 2004 ). Bibliographical information about economically important hosts in Argentina was presented by Pastrana (2004) . Biological control. Biological control has only being studied for species of economic importance. There are numerous works about parasitoids species of Agrotis from all over the world. More common parasitoid families include: Braconidae (Hymenoptera) , Ichneumonidae (Hymenoptera) , and Tachinidae (Diptera) . Migrations. There are several migratory species including: Agrotis segetum and A. exclamationis in Europe ( Wood et al . 2009 ), A. infusa Boisduval in Australia ( Common 1954 ), and A. ipsilon in New Zealand ( Greenslade et al . 1999 ), Israel ( Pedgley & Yathom 1993 ), and North America ( Showers et al . 1993 ). In Northern Hemisphere countries, adults fly northward in spring and southward in fall. Showers et al . (1993) released A. ipsilon in Iowa, USA , and among the recaptured moths they found one live adult in Texas 11 days later and 1900 kilometers from the release site. In the Southern Hemisphere, the flight pattern is opposite; flying southward in spring and northward in fall. In Australia , A. infusa flies southward when the weather gets warm. Adults fly to the southern mountains in spring and return to breed in the North in fall ( Common 1954 ). Migration is carried out with the help of prevailing winds and they can fly as high as 1500 meters ( Pedgley & Yathom 1993 ). In South America there are no studies of noctuid migration. It is likely that A. ipsilon presents migratory behaviors similar to those displayed in the Southern Hemisphere. TABLE 1. Number of specimens of Agrotis captured by species and months in South America. Species\Month Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. Specimen s/species Agrotis propriens ( Dyar, 1 4 1 6 1913 )

Agrotis acronyctoides Angulo & Olivares, 2006	1	1
Agrotis caliginosa Angulo & Olivares, 2006	1	1
Agrotis steniptera (Dognin, 1916)	0

Agrotis bistrigata 2 1 3 Maassen, 1890 Agrotis dispar Köhler 0 (1958) 1959 Agrotis peruviana 2 4 1 4 2 2 2 17 (Hampson, 1909) Agrotis elegans (Köhler, 17 1 1 2 21 1945 ) Agrotis benitezi León , 10 3 13 2010 Agrotis leonoides Poole, 6 45 51 1989 Agrotis edmondsi Butler , 12 24 28 6 7 39 32 148 1882 Agrotis leucovenata San 1 13 14 Blas & Gentili, 2011 Agrotis experta (Walker, 1 3 4 1 3 12 1869) Agrotis fausta (Köhler, 2 10 2 14 1958) Agrotis malefida Guenée , 1 6 21 4 7 3 10 8 19 1 80 1852

Agrotis canities (Grote, 1902)	1	1	2	1	2	1	3	5	3	19
Agrotis schreiteri (Köhler, 1945)	4	4
Agrotis araucaria (Hampson, 1903)	37	55	33	13	2	16	24	16	196

Agrotis ipsilon (Hufnagel, 2 3 4 4 1 1 4 3 8 3 33 1766) Agrotis robusta 120 53 35 21 50 2 2 8 42 333 ( Blanchard, 1852 ) Specimens/month 205 150 158 104 66 3 17 3 19 34 106 101 966 Species/month 12 10 13 10 6 1 4 3 5 7 7 8 Distribution. The main distribution of the genus comprises United States , southern Canada and Europe, southern South America , and South Africa , with fewer species in northern North America , northern South America , Mexico , North Africa, and India ( Lafontaine 2004 ) . In South America , the main distribution of Agrotis includes Chilean and Argentinean Patagonia extending northward along the Andes Mountains to Colombia , across the central and northern extra-Andean mountains of Argentina and Paraguay , and northeastwards as far as the coasts of Argentina , Uruguay , and Brazil . Some species are present in the Falklands and Juan Fernandez Islands. As pointed out by Lafontaine (2004) , diversity of the genus decreases to the north of South America , with few species in the Andes Mountains north of Argentina and Chile and in coastal regions of Uruguay and Brazil . Species groups. Three species groups are proposed: edmondsi -group, fausta -group, and robusta -group based on differences seen on South American species. fausta - and robusta -groups share cucullus apex strongly projected anterodorsally, sacculus 3/5 × as wide as valve, and saccus notched dorsally. robusta -group is characterized by valve subrectangular, basal half narrow, then widened and fausta -group by valve subrectangular, curved, elongate, and very narrow and aedeagus posterior half sclerotized and anterior half lightly sclerotized. Characters of fausta - group are shared with Agrotis ceramophaea Meyrick from Hawaii, even it is the only species seen of Hawaii it is likely other Agrotis from Hawaii belong to this group. edmondsi -group is characterized by valve subrectangular, neither narrowed nor widened in any part, cucullus apex slightly projected anterodorsally, sacculus between 4/6–9/ 10 × as wide as valve, and saccus without dorsal notch. edmondsi -group characters are shared with species from other continents, e.g., A. cinerea (Europe and Asia) and A. exclamationis (Europe) , but combination of all this characters is unique for the group.