Review of the New World genera of the subfamily Acontiinae (Lepidoptera, Noctuidae)

Th e taxonomic status of the 138 species of Acontiinae are reviewed and assigned to seven genera, Ponometia Herrich-Schäff er, Tarache Hübner, Acontia Ochsenheimer, Eusceptis Hübner, Pseudalypia H. Edwards, Spragueia Grote, and Trogotorna Hampson. A key to the genera, diagnoses of the genera, species groups of Tarache, illustrations of adults and genitalia of representatives of the seven genera, and a check list of the New World species are included.


Introduction
Th e purpose of this paper is to propose a new classifi cation for the New World species of the noctuid subfamily Acontiinae.
Th e subfamily "Acontiinae" has a checkered history.It has been treated as a receptacle for a large number of unrelated small genera of the Noctuidae.Th e family as treated in the last check list of the moths of North America (Franclemont and Todd 1983) included the tribes Acontiini, Eustrotiini, Bagisarini, Cydosiini, and Eublemmini.Th ese tribes have been disassociated and given subfamily ranking following Crumb (1956), Poole (1995), Kitching and Rawlins (1998), Fibiger and Lafontaine (2005), Lafontaine and Fibiger (2006), and Lafontaine and Schmidt (in press).
Th e subfamily Acontiinae as currently classifi ed by Fibiger and Lafontaine (2005) includes four tribes: Acontiini, Armadini, Aediini, and Hypercalymniini.Of the four, only the tribe Acontiini occurs in the New World and is the same group of genera listed as Acontiini in Franclemont and Todd (1983).
Th e Old World Acontiini was recently revised by Hacker et al. (2008).In this classifi cation the world fauna of the Acontiini is placed in a single genus (Acontia Ochsenheimer) and the species arranged in seven subgenera: one restricted to the New World (subg.Euseptis Hübner), two Holarctic (Acontia and Emmelia Hübner) and four Palearctic.One would expect that a tribe with a single genus with more than 300 species would be very conservative structurally, but the situation is the opposite.Th e "genus" is so structurally diverse that the authors have restricted the diagnosis of the tribe and the genus to the type species, Acontia lucida Hufnagel, stating that the other members of the genus share "the same autapomorphic characters as those of the typespecies, though sometimes in diff erent states, and some of which might have been lost" (Hacker et al. 2008: 10).A number of the more striking apomorphic characters, such as the hair tufts on the scaphium and the characters of the tympanic area, apply to the subfamily.Th e larval characters apply to the tribe, and most of the other characters listed (bird-dropping appearance, shape of the valves and vesica, asymmetry of the valves, sacculi, and claspers) are so varied throughout the "genus" that they are of little use in diagnosing any of the "subgenera" except a few small subgenera segregated out because of a particularly unusual character state, in one case the forewing maculation.Th e derived character states that support the monophyly of the subfamily Acontiinae, and those that characterize the tribe Acontiini, are combined and applied to a single genus by Hacker et al. (2008) as a way of dealing with the huge amount of structural diversity among the several hundred species in the tribe Acontiini.Th e result is that the genus Acontia (s.l.) is very well defi ned, being equivalent to the tribe and largely to the subfamily.Molecular data and a phylogenetic analysis of genital characters may be useful in arranging the Old World fauna into smaller better defi ned monophyletic genera.
Th e classifi cation that we present here is the result of the fusion of two separate research eff orts.An online draft of the Acontiinae of North America by the junior author (R.W. Poole) intended as the basis for an online identifi cation manual of the subfamily arranged the 100 or so species into six genera, mainly on the basis of genital structure.Th e senior author (J.D. Lafontaine), working on the classifi cation mainly with DNA data from a molecular gene sequence of 658 base-pairs of the cytochrome c oxidase 1 (COI) mitochondrial gene, commonly called "barcodes," arrived at the same six lineages that we treat as genera.Th e New World fauna of the Acontiinae was arranged in 14 genera by Poole (1989).We arrange the fauna in seven genera with the addition of the Neotropical genus Trogotorna Hampson to the six North American genera.A list of New World Acontiinae is given in the Appendix.Our classifi cation based on morphology is similar in most respects to the phenetic clustering-produced neighbor-joining tree analysis of the 658 base-pair sequences of the COI barcodes of the available 68 species of New World Acontiinae (Fig. 49).Although the COI DNA sequence analysis shows many relationships supported by genital characters, the placement of the genera Spragueia and Trogotorna are not consistent with their placement according to genital characters Genital terminology.Terms for genital structures and wing markings follow Lafontaine (2004).
Key to genera of North American Acontiinae (male genitalia) 1.
Ventral Ponometia is a large genus of New World Acontiinae with 47 described species.We list it fi rst because of the bilaterally symmetrical male genitalia, a character likely to be primitive and apparently absent in Old World Acontiinae (Hacker et al. 2008).Unlike Tarache and Acontia, only a few species have a moth that resembles a bird dropping.Th e genus is most easily characterized by the relatively small size of the moth (forewing length: 6-16 mm, but most species 8-12 mm), and the male and female genitalia.Male genitalia .Th e valves are bilaterally symmetrical with only a few species showing slight diff erences between the valves (e.g., P. albitermen, P. binocula, P. tortricina).Th e clasper is a long spine-like process on the ventral margin of the valve that turns upward, often abruptly so, to project onto the inner surface of the valve.Th e diagnostic feature of Ponometia is the vesica; there are one or two long diverticula, each with a longitudinally-ridged apical cornutus, and there is a comb-like row of small cornuti on the apical part of the vesica.Female genitalia (Fig. 39).Th e inner surface of the corpus bursae is covered posteriorly, sometimes entirely, with large patches of long spines 5-10 × as long as wide.

Tarache Hübner, [1823]
Figs 5-9, 23-30, 40-43 Th e genus Tarache contains 46 species, the majority of the species previously assigned to Acontia in the New World.Th e genus is heterogeneous in morphology, but can be arranged in three relatively well-defi ned and more homogeneous species-groups, and two of these can be divided further into subgroups.It may be desirable at some point in the future to subdivide the genus, but overlap in the presence of diagnostic character states among the species groups has encouraged us to adopt a larger defi nition of the genus.Included in Tarache are species previously placed in Th erasea and Hemispragueia.Male genitalia .Valves usually bilaterally asymmetrical, often markedly so; vesica with a dense rasp-like patch or patches of short spines at or near apex of vesica or on a diverticulum.Female genitalia .Inner surface of corpus with patches of spines 2-4 × as long as wide; appendix bursae sclerotized, most frequently forming a posterior lobe of corpus bursae but wrapping around and partially or completely fused to right side of corpus bursae in T. aprica group and in T. augustipennis and T. cora, so that ductus seminalis arises at or near anterior end of corpus bursae.
Tarache aprica species-group.Th is species-group contains the type species of Tarache (T.aprica) and 22 other species (Tarache abdominalis, T. apela, T. ardoris, T. assimilis, T. dacia, T. cratina, T. delecta, T. destricta, T. fl avipennis, T. isolata, T. knowltoni, T. lagunae, T. lactipennis, T. morides, T. parana T. phrygionis, T. quadriplaga, T. rufescens, T. sutor, T. tenuicola, T. terminimaculata and T. tetragona).Th e vesica of the aedeagus is the most distinctive feature of the T. aprica species-group; it tends to be globular with small subbasal diverticula and a massive subapical diverticulum, almost as long as the aedeagus, with its apex covered by a dense patch of spines.Species of the T. aprica subgroup have a distinctive subbasal lobe on the vesica armed with a row of 2-8 (depending on the species) large spike-like cornuti (Fig. 23).Th is row of cornuti is absent in the other three subgroups: T. terminimaculata (with T. dacia, T. cratina, T. phrygionis, and T. isolata), T. tetragona (with T. quadriplaga), and T. ardoris (with T. morides, T. parana, and T. rufescens).A diagnostic feature of species in the T. aprica, T.  24).Th e South American Tarache ardoris subgroup lacks the clavus on the right valve.In the female, the appendix bursae is very long and is fused to the right wall of the corpus bursae and extends 0.5-0.8 of the distance to the anterior end of the corpus bursae (Fig. 40).A similar form of bursa copulatrix is in T. augustipennis (Fig. 41) and T. cora, but in the former the fusion is less complete and in the latter there is no distinction between the corpus bursae and appendix bursae, and the ductus seminalis is at the anterior end of the bursa.Tarache bilimeki species-group.Th is species-group contains 21 species found in North and Central America.Like the species of the T. aprica species-group, the moths and male valve structure varies greatly, and the moth of the majority of the species resembles a bird-dropping when at rest.Th e species associated with the T. bilimeki species-group are best characterized by the form of the vesica .Th e apical part of the vesica has two, sometimes partially merged, fi elds of dense spines that form a rasp-like area; there are several sub-basal diverticula, some of which have peculiar cornuti that arise obliquely, sometimes to the degree that they appear to be on their side.In the female genitalia (Fig. 41) the posterior part of the ductus bursae forms a sclerotized, pouch-like ostium bursae, and the appendix bursae is sclerotized, mostly posteriorly, but sometimes partially or completely fused with the right side of the corpus bursae, as in the T. aprica species-group.Species in the T. bilimeki species-group can be arranged in four subgroups.1) Tarache areli subgroup (with T. albifusa, T. areletta, T. areloides, T. geminocula, and T. toddi) characterized by a forewing divided into a mainly pale basal half and a mainly dark outer half, except for a contrasting white preapical patch and a contrasting reniform spot, usually with some blue scaling (Fig. 7).Th e group was recently revised by Ferris and Lafontaine (2009).2) Tarache expolita subgroup (with T. arida, T. bella, T. cora, T. phaenna) with dark and light shading forming a patchwork parttern (T.arida, T. bella), or a dark streak through the middle and lower part of the wing that curves up to the forewing apex (T.cora, T. expolita) or almost entirely dark (T.phaenna).3) Tarache bilimeki subgroup (with T. acerba, T. augustipennis, T. axendra, T. lanceolata, T. major, T. mizteca, and T. sedata) characterized by elongated forewings with the costal half mainly pale and the posterior half mainly dark (Fig. 8).4) Tarache idella subgroup that includes a single species with a white forewing with narrow black transverse bands and a clear yellow-orange hindwing (Fig. 9).
Tarache lucasi species-group.Th is species group includes only two species (T.lucasi and T. vittamargo).Th e males resemble some species in the T. bilimeki subgroup and females are like some in the T. aprica subgroup.Th e male genitalia (Fig. 29), how- Acontia Ochsenheimer, 1816 [30][31][32][33][34]44 Acontia is the only New World genus also represented in Old World as well.Th ere are fi ve described species in the genus in North America and six in Central and South America.Th e moths diff er greatly in appearance and size; however most species tend to be some combination of white and gray, the white tending to be shiny.Th e forewing tends to be broad, often with a rounded apex.Male genitalia .All of the species have a well-developed, setose ampulla on the clasper and the clasper is usually weakly sclerotized except for a spinelike apex.In many species the costal part of the valve is more heavily sclerotized than the ventral part.Th e vesica in the New World species is elongated and without diverticula or spiny areas, except for a long, tapered, horn-like subapical diverticulum that is covered with minute denticles so that it appears to function as an enlarged cornutus; Acontia lucida, the type-species from western Eurasia, has a spinulose subbasal diverticulum with a spiny apex, as well as the false cornutus.Female genitalia (Fig. 44).Th ese consist of an elongate, sclerotized, ostium bursae, a tubular, membranous ductus bursae, and an oval membranous corpus bursae with the ductus seminalis at the anterior end.In Old World Acontia there are separate sclerotized plates in the ostium and ductus bursae and the ductus seminalis is at the end of a sclerotized appendix bursae, which is on the posterior left side of the corpus bursae.
Food plants.Th e food plants are recorded in the New World only for an undescribed species related to A. cretata that occurs in Texas and northeastern Mexico; it has been reared from Abutilon pedunculare Kunth (Malvaceae).In the Old World, Acontia lucida feeds primarily, but not exclusively, on species of Malvaceae.Th is genus includes 11 species, mainly of the Neotropical Region, but one species extends northward to Texas.Th e relatively large moths (forewing length 9-17 mm) with their broad boldly-patterned forewings, are more reminiscent of some species of Eulepidotis Hübner than an acontiine.One species (E.obscura) is drably colored with more rounded wings, but the genitalia are typical for Eusceptis.Male genitalia (Fig. 35).Th e rows of stiff setae on the ventral surface of the uncus are diagnostic for the genus.Th e valves are broad and apically rounded, widest near the apex; they are highly asymmetrical in almost all species (less so in E. irretita) and in most the right valve is more structurally complex than the left one.In all species except two the corona is concentrated in a small cluster at the dorsal-apical corner of the valve at the apex of a rod-like thickening of the costal margin of the valve.Female genitalia (Fig. 45).Th ese are similar to those of Acontia.Th e ostial area is sclerotized but the ductus bursae and corpus bursae are elongated and membranous; the appendix bursae is on a short membranous lobe on the left posterior margin of the corpus bursae.Th e apophyses and abdominal segment eight are very long and suggest a diff erent mode of oviposition than other acontiines.
Food plant.Malvaviscus arboreus Cav.(Malvaceae); reared by D. Janzen and W. Hallwachs in Costa Rica.Pseudalypia H. Edwards, 1874 Figs 16,36,46 Th is only species known in the genus Pseudalypia is P. crotchii and is so diff erent from other acontiines that it was described in the Agaristinae.It was fi rst recognized as an acontiine by Crumb (1956) from the larval characters.Th e moth is unmistakable (Fig. 16) and is reminiscent of an arctiine because of the contrasting orange prothoracic collar.Male genitalia (Fig. 36).Th ese are most similar to those of Spragueia, but the valves are almost bilaterally symmetrical, except for the larger ampulla on the right valve.Th e vesica has two large subapical diverticula covered with spines.Female genitalia (Fig. 46).Th ese are similar to those of Tarache and have a large sclerotized appendix bursae that extends anteriorly part way down the right side of the corpus bursae.

Spragueia Grote, 1875
Figs 17,37,47 Th e genus Spragueia contains 21 species of small, colorful moths found from southern Canada to the American tropics.Th e genus has traditionally been associated with the group of genera now amalgamated into Ponometia, probably because of small size and bold forewing pattern, compared with the bird-dropping look of most species of Tarache and Acontia.Th e male genitalia, however, are asymmetrical, and the ampulla well developed, as in Acontia, Pseudalypia, and Eusceptis, so we associate Spragueia with these genera.Male genitalia (Fig. 37).Th e clasper of each valve is a broad plate arising from the ventral margin of the valve with a well developed ampulla on each side with the setae enlarged, so the ampulla resembles a mace.Th e right clasper ends in a heavily sclerotized spine-like process that is absent on the left valve.Th e vesica has four lobes covered with spicules.A strong corona is usually present.Female genitalia (Fig. 47).Th e ostium is deeply invaginated and heavily sclerotized with the sclerotization commonly extended into the corpus bursae.Th e corpus bursae is usually globular with its anterior half weakly sclerotized.Food plants.Species have been reared from plant species in the families Malvaceae, Sterculiaceae, Asteraceae, Convolvulaceae, and Poaceae.
Trogotorna Hampson, 1910 Figs 18, 38, 48 Th is genus has only recently been confi rmed as an acontiine though the research of J. B. Sullivan.It currently contains only the type species, Trogotorna persecta, but several undescribed species are under study (J. B. Sullivan, pers. comm.).Th e moth (Fig. 18) does not at all look like it would belong to the Acontiinae, but the scaphium has the characteristic pair of setose patches, the enlarged alula over the tympanum, a vestigial tympanal hood, and asymmetrical male sacculi and claspers.Also, the CO1 barcode consistently places Trogotorna among the acontiine genera.Male genitalia (Fig. 38).Th ese would hardly be recognizable as belonging to the Acontiinae were it not for the characteristic setose pouch on each side of the scaphium.Th e valve tapers apically, unlike most acontiines, and the clasper is short leaf-shaped.Th ere is no corona, ampulla, or saccular extension.Th e valves are slightly asymmetrical in that the sacculus and clasper on the left valve are smaller than those on the right valve.Th e aedeagus is unusually short and wide for an acontiine.Th e vesica is short and bulbous, except for a large subapical diverticulum with a dense patch of spines at the apex.Female genitalia (Fig. 48).Th e apophyses, abdominal segment eight, and the ostium and ductus bursae are very short, whereas the corpus bursae is very large, 10 × as long as the ductus bursae.
Food plants.Unknown.
Barcoding, Biodiversity Institute of Ontario, University of Guelph, Guelph, Canada, provided data and information from the Barcode of Life Data (BOLD) system.Bo Sullivan (Beaufort, North Carolina) generously shared data, specimens, and insights from his study of the Acontiinae in Central America.Chris Schmidt and James Adams reviewed the manuscript and made many useful suggestions.