Review of the species level taxonomy of the neotropical butterfly genus Oenomaus (Lycaenidae, Theclinae, Eumaeini)

Abstract Seven new species of the Neotropical hairstreak genus Oenomaus are described: Oenomaus mancha Busby & Faynel, sp. n. (type locality Ecuador); Oenomaus gwenish Robbins & Faynel, sp. n. (type locality Panama); Oenomaus lea Faynel & Robbins, sp. n. (type locality Ecuador); Oenomaus myrteana Busby, Robbins & Faynel, sp. n. (type locality Ecuador); Oenomaus mentirosa Faynel & Robbins, sp. n. (type locality Peru); Oenomaus andi Busby & Faynel, sp. n. (type locality Ecuador) and Oenomaus moseri Robbins & Faynel, sp. n. (type locality Brazil, Santa Catarina). For each new Oenomaus species, we present diagnostic characters and notes on its habitat and biology. We illustrate adults, genitalia, and distribution. New distributional and biological data are presented for 21 previously described Oenomaus species. Oenomaus melleus guyanensis Faynel, 2008 is treated as a new synonym of Oenomaus melleus melleus (Druce, 1907). Females are described and associated with males for ten species using a variety of factors, including mitochondrial COI DNA “barcode” sequences. We summarize the reasons why the number of recognized Oenomaus species has grown in the past decade from one species to 28 species. Finally, we overview the habitats that Oenomaus species occupy and note that the agricultural pest on Annonaceae, Oenomaus ortygnus, is the only Oenomaus species that regularly occurs in greatly disturbed habitats.

A close phylogenetic relationship between Oenomaus and Porthecla Robbins was suggested when Robbins and Duarte (2004) described the latter genus. However, the distinction between these two genera has been disputed because of different interpretations of male genitalic morphology, which has resulted in the uncertain generic placement for a few species (Faynel 2007;Faynel et al. 2011). The species level taxonomy of Porthecla has been treated (Faynel et al. 2011), but a similar overview for Oenomaus is lacking.
We present new species level taxonomic information for Oenomaus in this paper. We describe seven new Oenomaus species. Next, we update information on the distribution, habitat, variation, and biology of the 21 species that were previously described in or transferred to Oenomaus (Robbins 2004;Faynel 2007;Faynel et al. 2011). We also associate females with males for many species based on male-female pairs collected in copula or on similarity of ventral wing patterns, geographic distribution, and DNA 'barcode' sequences (the mitochondrial COI gene). The morphology of newly associated females is detailed. With the species level taxonomy of Porthecla recently reviewed (Faynel et al. 2011), the goal of this paper is do the same kind of review for Oenomaus. This information will serve as the foundation for a phylogenetic analysis of Oenomaus and Porthecla.

Materials and methods
Genitalic terms follow those in Klots (1970), as modified for the Eumaeini in Robbins (1991). Wing veins are named following Comstock (1918), and wing cells are named by the veins that border them. Otherwise, morphological terms follow Snodgrass (1935). Abbreviations used repeatedly in the text are FW: forewing, HW: hindwing, D: dorsal, V: ventral and SD: standard deviation. Brazilian states are noted by their standard two letter abbreviations.
Illustrated adults of Oenomaus are noted in the material examined sections, and each genitalia drawing is of the adult illustrated. The structure of the male genitalia valvae in Oenomaus is complex, for which reason we present them in ventral, lateral, and dorsal views.
Biogeographical zones follow Brown (1982), who partitioned the forested continental Neotropics into the Transandean Region, Andean Region, Amazon Region, and Atlantic Region. Larval food plant nomenclature follows the Tropicos database of the Missouri Botanical Garden (http://www.Tropicos.org, accessed April 2012). Following Holdridge (1967), we classify lowland forests as humid/wet (> 200 cm annual precipitation) or dry/deciduous (100-200 cm annual precipitation). Many eumaeines display male territorial behavior on hilltops (Nicolay 1971;Alcock and O'Neill 1987;Dahners 2006, 2009;Robbins et al. 2012). Males wait on hilltops for receptive females to fly through the territory and "defend" these areas by flying at other males that enter the territory. Recorded times from our fieldwork for hilltopping behavior are the standard time at that locality. Finally, traps baited with decaying fish attract some lycaenid species and not others. We note the gender for each species which has been collected using fish-baited traps.
The ventral wing pattern in Oenomaus is sexually monomorphic, so associating the sexes of species with distinct ventral wing patterns, such as O. ortygnus, is straightforward. However, a majority of Oenomaus species have a ventral wing pattern that is similar to that of O. atena (Hewitson). Among these, some can be associated because they have distinct ventral wing pattern elements, such as those of O. isabellae (Faynel 2008), or because a mating pair was collected in copula. In other cases, we associate females with males if at least three of the following four criteria are met: (1) females have a ventral wing pattern that is indistinguishable from that of males, (2) females have a geo-graphic distribution that is similar to that of males, (3) both sexes are found in a locality where other species with the same wing pattern are unrecorded, and (4) divergence of DNA "barcode" sequences between the sexes is less than 2% (see next paragraph).
The mitochondrial COI gene sequence (commonly called a DNA "barcode") has been useful, when combined with other characters, in distinguishing lepidopteran species in a single area (e.g., Hebert et al. 2004;Hajibabaei et al. 2006;Janzen et al. 2009). Because genitalic and wing pattern characters generally provide clear species boundaries in Oenomaus, our purpose in determining COI gene sequences was to aid in associating females with males, as noted.
We use the following acronyms for collections, following those for public institutions listed on the website hbs.bishopmuseum.org/codens/codens-inst.html (accessed April 2012(accessed April ): et al. 2011. However, six of the newly described species have non-triangular, bifurcate valvae in lateral aspect , which is characteristic of Oenomaus. The seventh species is known only from a female, but the similarities in its wing pattern (Figs 2-3) and genitalia  to two other Oenomaus species support its generic placement. For these reasons, the following new species are described in Oenomaus.  in Faynel 2006). In particular, the dorsal part of the valvae of the male genitalia in lateral aspect is shorter and has a more sharply tapered posterior end in ventral view. In the female, the bifid  posterior end of the lamella postvaginalis is less marked and the anterior end of the ductus bursae is curved more sharply. One paratype from Ecuador has been barcoded , and its sequence is 3.5% divergent from the sequences of two males of O. ortygnus (CF-LYC-147 from Peru and CF-LYC-146 from Mexico, see Table 1) while the two O. ortygnus sequences differ by only 1.5%. Oenomaus ortygnus and O. mancha are sympatric in eastern Ecuador in Napo Province at approximately 450 m.

Oenomaus mancha
Etymology. The name of this species is derived from the Spanish word "mancha", which means spot, referring to the very distinctive, elongated black spot in VHW cell Sc+R1-Rs. The name is a feminine noun in apposition.
Habitat and distribution. Oenomaus mancha occurs widely in wet forest in eastern Ecuador at elevations ranging from 400 to 1100 m (Fig. 46). Although it is sympatric with O. ortygnus in wet forest, it does not occur in the highly disturbed habitats in which O. ortygnus sometimes occurs. It is yet an open question whether O. mancha is a lowland or lower montane species.
Description, diagnosis and recognition as a distinct species. Female FW length: 20 mm (N = 1). Wing pattern ( We hesitated to describe this species because we cannot assess its intraspecific variation. However, the series of 10 females of O. mancha show little variation in the traits that distinguish them from the holotype of O. gwenish. For this reason, a hypothesis of specific distinctness is better supported than a hypothesis of geographical variation. Etymology. The holotype of O. gwenish is a unique and distinctive female, for which reason it gives us great pleasure to name this species for entomologist Dr. Jennifer (Gwen) Shlichta. The name is a feminine noun in apposition.
Paratypes (2) the dorsal part of the valvae in lateral aspect shorter than the ventral part, and not pointed at the posterior end, (3) a straight penis in lateral view, and (4) no tooth at the end of the penis. The eighth tergum shows no difference from that of O. atesa. Lastly, the divergence of "barcode" DNA sequence data between O. lea (CF-LYC-005) and O. atesa (CF-LYC-003) is more than 6% (Table 1). The female of O. lea is unknown.
Etymology. This species is named for Léa Faynel, daughter of Christophe Faynel. The name is a feminine noun in apposition.
Habitat and distribution. Oenomaus lea occurs in wet lowland forest up to 1200 m elevation in eastern Ecuador and eastern Peru (Fig. 47). The ventral wing pattern of O. myrteana is superficially similar to those of Enos myrtea (Hewitson) and Allosmaitia myrtusa (Hewitson), but in these genera, males lack a scent pad on the DFW. The genitalia of O. myrteana, as noted, are typical of Oenomaus.

Oenomaus myrteana
Etymology. The name O. myrteana is intended to highlight the striking resemblance between the ventral hindwing of this species and that of Enos myrtea (Hewitson). The name is a feminine noun in apposition.
Habitat and distribution. Oenomaus myrteana occurs in lowland wet forest from eastern Ecuador to western Brazil (Rondônia) (Fig. 46). Busby observed males in Ecuador low in the understory at 11:00 hours. This species and E. myrtea have been found at the same site. Type-locality. Peru: Madre De Dios, Río La Torre, Tambopata Res., 12°50'13"S, 69°17'35"W, 300 m. Tambopata is at the mouth of the Río La Torre. In 1986 there was a lodge and a network of trails through uncut wet lowland forest. The holotype was collected during the transition between the dry and wet seasons when butterfly abundance and diversity generally peak.
Paratypes Oenomaus mentirosa is the only known Oenomaus species with red scales at the base on the VHW. In addition, it has a distinctive white spot along the VFW costa in cell Sc-R1. This feature occurs in no other Eumaeini except Porthecla minyia (Hewitson) where there are two white markings placed side by side in the cell between the costa and Sc. In male Oenomaus species, the eighth tergum is generally rectangular, but the anterior and posterior edges may be modified. In O. mentirosa, the male eighth tergum has a slightly modified anterior edge which looks like a shallow "W". The female of O. mentirosa is unknown.
Etymology. The name of this species comes from the Spanish word 'mentirosa', which means a feminine liar. We picked this name because the underside wing pattern resembles that of Porthecla gemma (Druce) and P. minyia (Druce), but this resemblance appears to be a false indicator of relationship. We treat the name as a feminine noun in apposition.
Habitat and distribution. Oenomaus mentirosa is known from lowland wet forest in Amazonian Peru (Fig. 47).
Remarks. Resemblance of the ventral wing patterns of O. mentirosa and P. gemma/P. minyia was noted in the etymology. Adults of all three species fly in the same habitats at the same time of year in the vicinity of Puerto Maldonado, Peru.
Type-specimen. Holotype ♂ (Fig. 8)  Description, diagnosis and recognition as a distinct species. Male FW length: 16.3 mm (N = 1). Female FW length: 16.7 mm (SD = 0.8, N = 2). Wing pattern (Figs 8,9) and genitalia (Figs 24,29) illustrated. The ventral wing pattern of O. andi is similar to that of many other Oenomaus, but this species is distinguished by (1)  Etymology. This species is named for Andrea (Andi) Busby, wife of Robert Busby, in appreciation for her long standing support of his research. The name is a feminine noun in apposition.
Remarks. Valvae structure in O. andi is very similar to that found in O. gaia Faynel, suggesting that this new species belongs to the O. cortica subgroup (as characterized by Faynel and Moser 2008). Species in this subgroup have a modified 8th tergum (except for O. druceus Faynel & Moser, 2008). In the male of O. andi (Fig. 42), the posterior edge of the 8th tergum has a deep depression in the middle, while the anterior edge is shaped like a wide "W". In the female, the posterior edge is nearly straight but is split in the middle. The anterior edge is similar to that of the male, but is laterally sclerotized (Fig. 43) Habitat and distribution. Oenomaus andi is a species of montane forest (> 1300 m) that is recorded from Ecuador to Bolivia (Fig. 47).
Behavior. A male and two females were attracted to traps baited with rotting fish (vouchers in RCB).
Type-specimen. Holotype ♂ (Fig. 10) (Figs 10-12). The lack of geographical variation in the characters distinguishing O. moseri and O. cyanovenata argues against the hypothesis that the former is a geographical variant of the latter.
Etymology. It is with great pleasure that we name this distinctive species for our good friend and collaborator Alfred Moser. Alfred lives in Rio Grande do Sul and has made prodigious contributions to the knowledge of Lepidoptera from southern Brazil, including co-authoring papers on the taxonomy of Oenomaus and Porthecla Moser 2008, Faynel et al. 2011 and Hook. f. (Annonaceae), a plant of frequent occurrence in the cerrado. We identified the male from a digital image and from the locality where it was reared. However, it is possible that it is a male of O. morroensis, even though this species is not known to occur as far north as São Paulo.
Habitat and distribution. Oenomaus moseri occurs in lowland and lower montane forest in southern Brazil (Fig. 46).

New data for previously described species of Oenomaus
For each of the 21 previously described Oenomaus species, we give distribution, habitat, and remarks. We then note, where relevant, new information on taxonomy, intraspecific variation, behavior/biology, associated females, and COI DNA sequences. The species are treated in alphabetical order. Oenomaus curiosa and O. melleus are included in this section, even though their generic placement is yet unresolved (Faynel et al. 2011  Female. The female of this species was determined by a pair collected in copula and was illustrated by Faynel (2008, fig. 2).
COI DNA sequence. Three specimens of O. atena have been barcoded, including a male from Peru (LO) (CF-LYC-084) and two females from French Guiana (CF-LYC-054 and CF-LYC-057). The latter two have the same brown dorsal wing pattern, ventral wing pattern, and genitalia as the female of O. atena found in copula. The three barcodes show 0.4% divergence.

Oenomaus atesa (Hewitson)
http://species-id.net/wiki/Oenomaus_atesa Distribution, habitat, and remarks. Oenomaus atesa is a widespread species that has been recorded from Mexico, Panama, western Ecuador, French Guiana, Venezuela, Colombia, eastern Ecuador, Peru, and Brazil (AM, DF, MG, RJ, SP, SC). The vast majority of museum specimens were collected in the lowlands, but males have also been found at 1375-1700 m in western Ecuador and at 2200 m in western Colombia (Prieto and Dahners 2006). Intraspecific variation. Despite substantive geographical variation in O. atesa, we lack sufficient material to determine if this variation might represent more than one species. Females from Venezuela and western Ecuador have more extensive dorsal blue and a somewhat lighter color than females from Panama, French Guiana, eastern Ecuador, and Peru. In addition, males from western Ecuador have more blue on the dorsal forewings than males from eastern Ecuador. However, this variation is small compared to that between males of O. atesa and O. lea. For example, the forewing dorsal blue area never reaches the cells from vein R3 to Cu1 as it does in O. lea. Structure of the female genitalia also varies geographically. Females from Venezuela and Peru have two processes at the posterior end of the lamella postvaginalis while a female from French Guiana had none (see Faynel 2006, p. 29 (Fig. 14).
Female. We associate a female (Figs 14, 31) which has the same ventral wing pattern as the male, which occurs in French Guiana (as do the known males), and which has a very similar COI DNA sequence to that of the males.
COI DNA sequence. Divergence among the three known specimens is 0.2%.  Intraspecific variation. The male from Panama has the posterior edge of its 8th tergum more deeply incised than in others.

Oenomaus cortica (D'Abrera
Female. Unknown. A female paratype of O. cortica from Espírito Santo, Brazil was illustrated in D'Abrera (1995), but no definitive evidence was presented to support this identification.
COI DNA sequence. Two males from Brazil, Pará have been sequenced (CF-LYC-051 and CF-LYC-052) and show 0.6% divergence.  (Fig. 17) are uniformly brown on the dorsal wing surface while the female from Costa Rica (Fig. 16) has the basal parts of both wings blue. Their genitalia, however, are uniform. Additionally, their COI DNA sequences are similar. This geographic variability is similar to that in O. taua.

Intraspecific variation. Females from French Guiana and Brazil, Pará
Behavior/biology. Two females were reared in Costa Rica (97-SRNP-62841.1 and 97-SRNP-6283) from Guatteria verrucosa R.E. Fr. (Annonaceae) (adult vouchers in USNM). Details of the rearing records along with images of the immatures can be found in Janzen and Hallwachs (2012).
Female. Females of this species (Figs 16,17,32,33) have the same ventral wing pattern as males, occur at the same localities, and have similar COI DNA sequences.  A female paratype of O. cyanovenata from Pará, Brazil was designated and illustrated in D'Abrera (1995) without definitive supporting evidence. This female has a different dorsal wing pattern than the female from Pará that we have associated with the male. We are skeptical of the biological validity of this paratype designation.
COI DNA sequence. Four males and seven females from French Guiana and Brazil, Pará were barcoded. One male (CF-LYC-046) is 6.7% divergent from the other three males, but its sequence is identical with that from a male of O. magnus (CF-LYC-020). Potential explanations for this result range from contamination to biologically significant, but until we have additional information, we omit this male from the following results. Divergence among the 10 other specimens of O. cyanovenata was 0.1%. The reared females from Costa Rica, which were barcoded in another project, are 0.4% divergent from the South American specimens.

Oenomaus druceus Faynel & Moser
http://species-id.net/wiki/Oenomaus_druceus Distribution, habitat, and remarks. This species was described from one Brazilian (AM) male, which is the only known specimen. As noted, its genitalia are similar to those of O. mentirosa, but it has a distinctly different ventral wing pattern.
Female. Unknown. Female. Described by Faynel and Moser (2008). Female. Four males in the USNM were collected on hills on Los Ríos hill (approximately 9°00'32"N, 79°35'34"W) and in Cocolí (approximately 8°58'46"N, 79°35'59"W), Canal Area, Panama. These areas are drier (<2 m annual precipitation, Rand and Rand 1982) than the forest in which other Oenomaus with an "atena-like" wing pattern have been found in Panama. Four females from these two localities have the same ventral wing pattern as the males. Since no other males are known from these localities, we associate the sexes and illustrate the adult wing pattern and genitalia of one of these females (Figs 15, 34).

Oenomaus gaia
We also associate a female from Brazil, Pará (CF-LYC-072) with a male of O. gaia from French Guiana because they have the same ventral wing pattern and have similar barcode sequences (0.2%). Intraspecific variation. As noted by Faynel and Moser (2008), O. jauffreti is a variable species, especially ventrally. For example, the VHW basal spot in cell Sc+R1 is large and mostly white in French Guiana; is small, black with a white centered pupil in Brazil (MT), and is large, with black and white scales in Ecuador. The only element which seems to be stable is the presence of a white spot on the basal side of VHW cell Rs-M1.

Oenomaus magnus
Female. We associate females from French Guiana (Figs 13, 36), Peru, and Bolivia with this species. They have the same ventral wing pattern, a similar geographic range, and limited COI DNA sequences are the same.

Oenomaus melleus
Taxonomy. Faynel (2007Faynel ( , 2008 partitioned this species into a Transandean Region (terminology from Brown 1982) nominate subspecies and an Amazonian Region subspecies O. melleus guyanensis based on size and color of scales at the base of the VFW. The male genitalia of each taxon were the same. As noted in the next paragraph, the new material examined does not confirm this recognition of two taxa. For example, the Costa Rican specimens resemble the Amazonian ones. For this reason, we synonymize O. melleus guyanensis Faynel with O. m. melleus (Druce), new synonym.
Intraspecific variation. The wing pattern of O. melleus is highly variable. The type from Colombia and two specimens from Nicaragua and Venezuela are relatively large (male FW length = 16.8 mm, SD = 1.3, N = 3). They have a white spot on the basal part of VHW cell Rs-M1, no reddish scales on the basal part of ventral wing, and a black spot in VHW cell Cu1-Cu2. The specimens from French Guiana, Guyana, Brazil (PA), Venezuela and Peru (UC) are smaller (male FW length = 14.1 mm, SD = 0.4, N = 5). They have a white spot on the basal part of VHW cell Rs-M1, reddish scales on the basal part of ventral wing, and no black spot in VHW cell Cu1-Cu2. The males from Costa Rica are also relatively small (male FW length = 14.7 mm, SD = 1.6, N = 3). They have no white spot on the basal part of VHW cell Rs-M1, no reddish scales on the basal part of ventral wing and a black spot in VHW cell Cu1-Cu2.

Oenomaus ortygnus (Cramer)
http://species-id.net/wiki/Oenomaus_ortygnus Distribution, habitat, and remarks. This species occurs in many different habitats from sea level up to 1000 m. It is unique in the genus in that it is often found in highly disturbed habitats. It is the most common Oenomaus species in collections and has been recorded from the United States, Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, French Guiana, Surinam, Guyana, Trinidad, Venezuela, Colombia, Ecuador, Peru, and many states throughout Brazil. As noted in the introduction, this species is a well-known pest of commercial Annonaceae.
Intraspecific variation. The blacks spots on ventral wings vary in size and the blue on the dorsal wings vary from light cyan to dark purple. The "Thecla lauta Draudt" phenotype from western Mexico is smaller and duller than individuals from the remainder of its range.
Intraspecific variation. Females from Brazil and Ecuador (Fig. 19) are uniformly brown on the dorsal wing surface while the female from Panama (Fig. 18) has the basal parts of both wings blue. Their genitalia, however, are uniform. This geographic variability is similar to that in O. cyanovenata.
Behavior/ biology. Territorial behavior on a hilltop in Panama (Canal Area, Gamboa, Cerro Pelado) was observed in January and August at 15:00 hours (vouchers in USNM). Two mating pairs were also collected on the same hilltop in January and March at 15:00 hours (vouchers in USNM).
Female. We illustrate adult females that were collected in copula (Figs 18-19) and the genitalia of one (Fig. 37).
COI DNA sequence. A female from Brazil, Pará (CF-LYC-064), which has a wing pattern similar to the females collected in copula, is 3.1% divergent from a male from Peru (CF-LYC-085).

Taxonomy. A decade ago
Oenomaus was a monotypic genus, but it now consists of 28 described species (albeit, it is still unclear if O. melleus and O. curiosa belong to Oenomaus or Porthecla). Further, if a phylogenetic analysis shows that Porthecla is paraphyletic in terms of Oenomaus, which is possible because Porthecla was distinguished by character states that may be plesiomorphic, then Oenomaus will be one of the most species-rich eumaeine genera with 40 species (Robbins 2004).
There are three biological reasons why the diversity of Oenomaus was not recognized until recently. First, about ¾ of the species have an indistinguishable, or barely distinguishable, ventral wing pattern that is similar to that of O. atena (e.g. ,  Figs 12-19). Among species with this wing pattern, there is a great diversity of male genitalic forms that were first documented by Faynel (2006Faynel ( , 2008 and Faynel and Moser (2008). Second, the ventral wing pattern of a few species is different from that of O. atena (e.g. , Figs 1-7), but similar to that of sympatric species that are now considered to be distantly related. For example, Draudt (1919Draudt ( -1920 in Seitz grouped O. ortygnus (the type species of Oenomaus), now placed in the Panthiades Section, with Atlides rustan (Stoll), now placed in the Atlides Section (Robbins 2004). Similarly, he placed O. atesa in a group with Enos mazurka (Hewitson) in the Brangas Section. Oenomaus myrteana, which is described in this paper, closely resembles Enos myrtea while O. mentirosa, also newly described, has a ventral wing pattern that resembles species in Porthecla, Olynthus Hübner, Janthecla Robbins & Venables, and Atlides Hübner (documented in Faynel et al. 2011). Third, many Oenomaus species are exceedingly rare in collections. Indeed, three species are still known from only one individual each.
DNA barcoding. Thirty-eight Oenomaus males belonging to 19 species have been successfully "barcoded" (>200 bp) (extraction and sequencing methods given in Hajibabaei et al. 2006). For those nine species for which there is more than one barcode (Table 1), intraspecific divergence calculated on the Bold web site (http:// www.boldsystems.org/views/login.php) using the Kimura 2 parameter with sequences aligned by BOLD varied from 0% to 1.6%. Interspecific divergence (Table 1) varied from 0.8% to 9.7% (672 comparisons, mean distance: 6.1%). It was usually greater than 4% except in the O. cortica species group (O. gaia, O. cortica, O. morroensis), where it was about 1%. Similarity in COI sequences among closely related species is well-established (e.g., Burns et al. 2007).
Male-female associations. Associating males and females in Oenomaus is sometimes very difficult. Only eight of the 28 recognized species previously had the sexes associated. In this paper we associate the sexes of another ten species based on mating pairs collected in copula and on similarity of ventral wing patterns, habitats, geographic distributions, and mitochondrial COI DNA sequences. The DNA "barcodes" have great potential (e.g., Janzen et al 2009), especially if there are large samples from geographically diverse sites.
Biology. Oenomaus ortygnus is a well-known pest of cultivated soursop (also called guanábana, Annona muricata L., Annonaceae) and relatives (e.g., Dampf 1929, Fennah 1937, Ballou 1945, Guagliumi 1965, Araque 1967, d'Araújo e Silva et al. 1967-1968, Leal 1970, Kendall 1975, Domínguez 1978, Peña et al. 2002, Castañeda-Vildózola et al. 2011. As noted in the results of this paper, two other Oenomaus species have now been reared, and Annonaceae (Duguetia, Guatteria) is a food plant for each. Although data are yet too scanty to ask why O. ortygnus is the only Oenomaus species that has been recorded as a pest on cultivated Annonaceae, we note that it is also the only Oenomaus species that is regularly found in disturbed habitats.
Most Oenomaus species inhabit relatively undisturbed lowland wet forest, but some species seem to be restricted to other habitats. Oenomaus andi is montane, being found so far only above 1450 m, while O. geba is known only from lower montane forest. Oenomaus morroensis occurs so far only in subtropical and lower montane forest. Oenomaus druceus has been found only in scrubby deciduous forest. A number of other Oenomaus species have broader habitat requirements. Some wet lowland species also occur in montane habitats, for which O. atesa and O. moseri are representative examples. Others, such as O. floreus, O. gaia, and O. isabellae, may inhabit dry deciduous forest. As previously noted, O. ortygnus is the only Oenomaus species that is regularly found in both undisturbed and disturbed habitats.