Phylogenetic systematics of Schacontia Dyar with descriptions of eight new species (Lepidoptera, Crambidae)

Abstract The Neotropical genus Schacontia Dyar (1914) is reviewed and revised to include eleven species. Schacontia replica Dyar, 1914, syn. n. and Schacontia pfeifferi Amsel, 1956, syn. n. are synonymized with Schacontia chanesalis (Druce, 1899) and eight new species are described: Schacontia umbra,sp. n., Schacontia speciosa,sp. n., Schacontia themis, sp. n., Schacontia rasa, sp. n., Schacontia nyx,sp. n., Schacontia clotho, sp. n., Schacontia lachesis, sp. n., and Schacontia atropos, sp. n. Three species, Schacontia medalba, Schacontia chanesalis, and Schacontia ysticalis, are re-described. An analysis of 64 characters (56 binary, 8 multistate; 5 head, 13 thoracic, 13 abdominal, 25 male genitalic, and 8 female genitalic) scored for all Schacontia and three outgroup species (Eustixia pupula Hübner, 1823, Glaphyria sesquistrialis Hübner, 1823, and Hellula undalis (Fabricius, 1781)) retrieved 8 equally most parsimonious trees (L=102, CI=71, RI=84) of which the strict consensus is: [[[[medalba + umbra] + chanesalis] + speciosa] + [ysticalis + [rasa + themis + [atropos + lachesis + nyx + clotho]]]]. The relevance of male secondary sexual characters to the diagnosis of Schacontia species is discussed.

introduction Schacontia Dyar, 1914: 400 represents a small cluster of species recently transferred to the Glaphyriinae (Solis 2009) (Figs 1-12). Both the male genitalia and the external appearance of described Schacontia are atypical for pyraloids, so much so that the type species was originally described by Schaus as a member of the noctuid genus Acontia Ochsenheimer (as "Acontia? [sic] medalba"; Schaus, 1904: 163). The subsequent taxonomic history of these moths is one of taxonomic curiosity and nomenclatural flux. Schacontia caught the attention of taxonomists in part by virtue of its unusual male genitalic apparatus, which comprises a uniquely configured gnathal complex and reduced valvae. Schacontia was originally described in the Schoenobiinae, retained there by Amsel (1956), transferred to the Epipaschiinae (Pyralidae) by Munroe (1958), and then tentatively transferred to the Cybalomiinae (Munroe 1995). It was most recently transferred to the Glaphyriinae by Solis (2009) based in part on the external morphology and genitalia, but mostly based on the morphology of the tympanal organs. Solis (2009: 493) characterized the subfamily with the following combination of characters: chaetosemata absent; concavity on the costa of the forewing present; fovea between Rs 2+3 and Rs 4 present; forewing with Rs 4 in a non-apical position and a costal crescent present; and lateral indentations of Sternite 2 present (Luquet and Minet 1982).
With respect to their actual biology, Schacontia larvae have been variously associated with Capparaceae (Brassicales) and have been recently reported as parasites of cassidine chrysomelid beetles (Cuignet et al. 2008), but that latter report is unverified as Schacontia. Without more life history data and more taxon-rich analysis, it is not possible at this time to address life history evolution, the macroevolution of host plant associations, or larval feeding behaviors in Schacontia. It is in the widespread species (S. chanesalis, S. themis, S. ysticalis) that life history and larval data are most sorely needed rangewide. The association of some species with Capparaceae is not unusual for crambids (cf. Solis et al. 2009), and as such, Schacontia may provide a forum for exploring its origin(s). The evolution of glucosinolates in the Brassicales (e.g., Mithen and Marquez 2010) may bear on the origins of these moths' specialized feeding habits, including the gall-forming behavior and internal feeding reported by Solis et al. (in prep.).
In the present work, we treat newly assembled historical and recent material from the Western Hemisphere. Our purpose is to refine the circumscription and composition of Schacontia by identifying and describing new species and presenting a phylogenetic analysis of their relationships. Recent collecting and rearing work, including the efforts in Costa Rica by D. Janzen and W. Hallwachs, have generated life history information, most importantly the association of Schacontia with capparaceous plants. Those potentially allied with Schacontia on the basis of wing venation and features of the gnathos and tegumen, comprise eight undescribed species ranging from Mexico through Central and South America and the Caribbean, some narrowly endemic, others widespread.

Materials and methods
Pinned specimens were examined with an incandescent light source (reflected light). Male and female genitalic preparations varied, some of those pre-dating this study having accumulated from several sources. Most were prepared following Clarke (1941), using chlorazol black and in some cases mercurochrome as staining agents; Eosin-Y was used in some preparations [those originating with Dr. V.O . Becker]. The more recent dissections were made following a hot soak in supersaturated sodium hydroxide, and held in glycerine caps or temporary slides for character scrutiny. Some older preparations of wings were prepared following Borror, Triplehorn, and Johnson (1989): soaked in bleach, stained with Eosin-Y, and slide mounted in Canada balsam. Slide preparations were examined with dissecting and compound microscopes. Photographs were made using the Microptics and Visionary Digital imaging systems and images manipulated with the Gnu Image Manipulation Program (The GIMP Team, gimp. org) and, when appropriate, retouched with Adobe Photoshop® (Adobe Systems, Mountain View, CA). All measurements were made with the aid of an ocular micrometer. Forewing length was measured from the center of the axillary area up to the apex of the forewing (FW). Terminology follows Wooton (1979), Klots (1970), Maes (1985Maes ( , 1995, Yoshiyasu (1985), Phillips and Solis (1996), Solis and Maes (2002), and Mally and Nuss (2011), except where noted (Figs 63,64). The terminology of Maes (1985Maes ( , 1995 is adhered to strongly with respect to the tympanal organs; Mally and Nuss (2011) are consulted as a more recent reference with respect to coding the female genitalic characters.

Material examined
This work drew in part from an effort to treat taxa with taxonomic and nomenclatural problems identified during preliminary surveys of pyraloids in the extensive Costa Rican collection of D. Janzen and W. Hallwachs. Because the genitalic characters of Schacontia species had not been adequately explored, specimens of all known Costa Rican species were initially dissected to survey diagnostic characters of each species and putative synapomorphies for the genus. In order to determine the nomenclatural status of Costa Rican species, types of all neotropical species at the Zoologische Staatssammlung München, Munich, Germany (ZSM), Naturhistoriches Museum, Vienna (NHMV), The Natural History Museum, London (BMNH), and the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM) were examined. Following this preliminary work, a more expansive series of material (all the specimens of Schacontia we could locate) was examined, most recently including Bolivian and Puerto Rican material housed at the Carnegie Museum of Natural History (CMNH) as well as at Cornell University Insect Collection (CUIC), and all the available material at USNM, including material from the V.O. Becker collection (VOB). Specimens are listed for each species with all attendant label data, including genitalic dissection slide numbers and record numbers from the database of Janzen and file and then as a TNT file, using a text editor to ensure rooting at the first terminal encountered. Exhaustive searches (implicit enumeration) were then run in TNT (Willi Hennig Society edition; Goloboff et al. 2008). Bremer values (Bremer 1988) were calculated in TNT from exhaustive searches of progressively longer suboptimal trees (increments of 1 step). Synapomorphies were mapped with Winclada.

Taxonomic scope and outgroup selection
The scope of our treatment of described Schacontia is based on Munroe (1995); type material is deposited at USNM except where designated in text. We treat the taxonomic and nomenclatural issues in the subfamilial placement of the genus only insofar as they pertain to outgroup selection and rooting.
Initial examination of specimens tentatively identified as Schacontia revealed, first, a cohesive group of species comparable to the type species [S. medalba (Schaus, 1904)] unified by a uniquely hood-like or mucronate uncus, reduced male valvae, a divided tegumen with a prominent medial sulcus, and a gnathos with a unique, four-armed configuration. A somewhat more variable group, including S. ysticalis (Dyar, 1925) and several undescribed species, appeared to bear similarities to Schacontia in forewing pattern and, in modified form, features of the male valva, tegumen, and gnathos. Bearing in mind that member species of Schacontia have been placed in several subfamilies prior to the genus' transfer to Glaphyriinae by Solis (2009), and in the interest of being thorough, type species of all 33 known glaphyriine genera were examined as outgroup candidates. We also surveyed types of Cybalomia Lederer, 1863 and a range of crambid subfamilies in order to mine specific character systems for putative synapomorphies of Schacontia and to ensure proper character polarization. We selected three glaphyriine outgroup taxa, all type species of their respective genera: Eustixia pupula Hübner, 1823, Hellula undalis Fabricius, 1781, and Glaphyria sesquistrialis Hübner, 1823, at which our tree is rooted. These outgroups were chosen both on the basis of their status as name-bearers and on the basis of what we estimated to be comparable suites of observable similarities (putative homologies) with S. medalba. The rooting at G. sesquistrialis was implied by the current classification, but more importantly was based on a preliminary screening of male and female genitalia and tympanal structures. These include the configuration of the saccus tympani and corpus bursae.

Phylogenetic relationships
We were unable to discern consistently different characters among S. chanesalis, S. pfeifferi, and S. replica, but in view of there being extremely limited material of S. pfeifferi in particular, and despite Amsel's description's being the only detailed and wellfigured one to date, we elected to synonymize S. replica and S. pfeifferi with S. chanesalis based in large part on a lack of discernable discrete variation in the male genitalia.
From cladistic analysis eight most parsimonious trees obtain (L=102, CI=71, RI=84), the strict consensus of which (L=108, CI=67, RI=81) is presented ( Fig. 66 The monophyly of Schacontia is supported by synapomorphies enumerated in the diagnosis of the genus (below). Two primary groupings appear in the strict consensus (character numbers given parenthetically). The first comprises the type S. medalba, S. chanesalis and the newly described species S. umbra and S. speciosa; these are united by the absence of ocelli (0); reduced proboscis (1); a compound, non-uniform ground color that does not sharply delineate the medial area (8); a distinct hindwing postmedial line approaching or reaching the inner margin (10); robust, broad fornix (27); and wide, gently tapered venulae secundae (28). In these species the outer margin of the valva is also continuous, the valva highly reduced in all but S. speciosa (40), and without a conspicuous saccular bend; phallus simple, without cornuti (55); attachment of the ductus bursae basal (63). Three of these species (excluding S. speciosa) are characterized by having the saccus tympani deep, with a posterior ridge, but not invaginated posteriad (19) and a tipped, mucronate uncus that is not conspicuously obovoid and is longer than its width at the base. As will be discussed, 4 of 5 described species -all but one of which fall within this group -were described on the basis of female types. The morphology of the female genitalia is rather uniform among the species in this group; their putative association with males is based on a combination of wing pattern and geographical proximity.
The second major intrageneric grouping, the ysticalis-themis group comprises S. ysticalis and six newly described species, whose association with Schacontia had been hypothesized initially. This group is united by a forewing pattern that is either essentially unicolorous excepting the antemedial and postmedial lines and orbicular spot, or two toned, but with the basal area unbroken and the medial area contrasting with the basal and apical areas (8); an inconspicuous hindwing postmedial line not nearing the inner margin (10); the saccus tympani a capacious ovate chamber with a conspicuous broad lip, comparable to that of Cybalomia Lederer (19); the dorsal ridge of the tegumen cruciate, crossing near the base of the uncus (33); the uncus either gently tapered and bluntly rounded, wider at the base than long, or variously nippled, trefoiled, and/ or conspicuously obovoid (35, 36); the outer margin of the valva with upper and lower lobes, not with a continuous edge (39), but with a saccular bend or elbow either at its midpoint or proximal to the vinculum (44, 45); ventro-marginal setae present and well-developed (48); costal setae present, sometimes arranged in a recurved, fish-hookshaped cluster (52); phallus with cornuti (54); attachment of the ductus bursae subbasally, creating the shouldered appearance on the corpus bursae (62).
Morphologically, this second, perhaps more enigmatic species-group, is less homogeneous than that surrounding the type species of Schacontia. Its most basal member (S. ysticalis) retains numerous features common to the latter group, viz. concentration of white scales apical to the postmedial line (character 9), the narrow distance between the postmedial line and the wing terminus (character 11), the light wing lines in contrast with the dark ground color (character 12), the undifferentiated uncus (character 35), the configuration of the intra-saccular process (character 41), and the elongate corpus bursae (character 61). The remaining species, all hitherto undescribed, form a complex of species exhibiting a heterogeneous collection of male secondary sexual characters, including unusual metatarsal structures, tibial scales and spurs, and abdominal coremata. These appear somewhat homoplastically, such that their downweighting or removal results in a more decisively resolved topology, but we retain them in analysis to emphasize their relevance to future work. Schacontia Dyar, 1914: 400 http://species-id.net/wiki/Schacontia Type species. Acontia medalba Schaus, 1904: 163, by subsequent designation.
Type locality: Brazil. Etymology. "Schacontia" seems to be Dyar's contraction of Schaus and Acontia, the noctuid genus in which Schaus mistakenly attributed medalba and subsequently designated by Dyar as the type species of Schacontia.
Diagnosis. Schacontia species may be recognized by (character numbers listed parenthetically): Foreweing Rs 3 and Rs 4 stalked (5); M 1 and M 2 stalked (6); hindwing M 2 M 3 + CuA 1 stalked (13); bullae tympani invaginated in S2 (18); absence of puteoli (22); fornix heavily sclerotized and far removed from the edge of Ve1 (24); fornical angle a low arc > 90 degrees (25); presence of gnathos-ventrotergal rods complex (31), bearing a finger-like middle process (32); presence of teguminal sulcus (34); intrasaccular process a bump or flange towards base of valve or as a trigger-like process at margin of lower lobe of valve (41); pair of terminal black dots on abdominal dorsum of male (53); uncus hood-like, mucronate, or obovoid, with variously modified terminal nipple (35,36). In addition, the costal bulge in the FW postmedial line is frequently coupled with a color contrast between the FW medial area and the basal and terminal areas, often involving white scaling. Unlike the medalba group (for present purposes including S. speciosa), the proboscis is not reduced in the ysticalis-themis group, the labial palpi droop, the tympanal fornix is narrow, ribbonlike; venulae secundae tapered to form a "neck." Habitus. In the species most readily identifiable as Schacontia (by virtue of their similarity to the type species S. medalba), hereafter referred to as the medalba group, the forewings are gray with a metallic sheen and the antemedial and postmedial lines variously suffused with white, the exception being S. umbra, which may be almost uniformly shaded dark brown. Towards the costa, the postmedial line bulges outward; the hind wings are by and large nondescript in pattern beyond the presence of a faint postmedial line. The ysticalis-themis group including the S. themis-rasa sister pair and the S. nyx complex [S. nyx+S. clotho+S. lachesis+S. atropos], are distinguished from these in having ocelli present; frons with normal, convex contour, except in S. ysticalis; and labial palps porrect, extending beyond the clypeus.
Male genitalia. All Schacontia bear a modification of the intrasaccular region of the valva. In the case of those species surrounding the type species of Schacontia, this comprises a naked or denticled flange; the valvae are characteristically reduced, if not truncate, and the uncus prominent but unadorned, mucronate. The valvae become progressively more complex in the ysticalis-themis group, with the intrasaccular feature transposed laterally to form a sclerotized trigger-like structure. Also in the ysticalisthemis group: the dorsal ridges of the tegumen are cruciate, meeting near the uncus; the tegumen is much wider than the uncus such that the lateral edges of the tegumen appear to taper/fall away from the uncus gradually; the outer margin of the valva is complex, including a variously adorned subcostal process, the costa associated with a fleshy lobe at its terminus and at least one setal tuft; the sacculus bears a localized patch or cluster of setae ventrad; and a membranous area exists between the costa and the subcostal process.
Description. Head -In medalba group, ocelli and chaetosemata absent; proboscis reduced; frons conical; labial and maxillary palpi straight. In ysticalis-themis group, ocelli present; frons of normal, convex contour except in S. ysticalis; labial palps porrect, extending beyond clypeus. Thorax -In medalba group, pronotum, mesonotum, legs gray; hind leg of female with 1 pair of tibial spurs. Males of several members of ysticalis-themis group bear a flattened, hind tibial spur, specialized hind tibial scales, a shallow concave spoon-like metatarsal modification, and coremata on 4 th abdominal segment (on S. themis, S. nyx, S. clotho, S. lachesis, and S. atropos); in addition, epipleural setae may be present (in S. rasa, S. clotho, S. lachesis, and S. atropos); and female hind tibia usually bear two pair of spurs (a medial pair present) except in S. ysticalis and S. rasa. Forewing (FW) -Schacontia exhibit a characteristic curvature of postmedial line, outwardly bulging towards costa. In medalba group FW medial area partially suffused with white; in ysticalis-themis group, FW either unicolorous with basal and postmedial areas or polymorphic, with some specimens more darkly shaded. Rs 3 and Rs 4 stalked; M 1 and M 2 stalked. Hindwing -In medalba group, HW generally pale with few contrasting markings; female frenulum with a single seta; postmedial line sometimes present, conspicuous, but never in ysticalis-themis group. [M 2 M 3 ]+CuA 1 stalked. Abdomen -Scales arranged in two terminal black dorsal spots in males, more conspicuous in ysticalis-themis group. Tergites gray with dark-gray scaling in medalba group. Tympanal organs crambiform (tympanum and conjunctivum not co-planar, praecinctorium present, bullae tympani open anteromedially), but somewhat variable. In medalba group, bullae tympani broad, tympanal assemblage wider than long (cf. Solis 2009: 503); processi tympani present, towards antero-lateral end of fornix, prominent, lamellate, hemi-circular; processus spiniformis present; fornix tympani strongly sclerotized, broad, removed from edge of venula prima; fornical ulna gradually arched at approximately >90° angle; pons short, broad, V-shaped, length more or less equivalent to breadth of fornix; rami (posteromedial margins of sacci) weakly sclerotized, arcuate, not strongly angled medially; venulae secundae present, tapering gently such that posterior width only slightly less than anterior width; puteoli absent; posterior lip of saccus weakly sclerotized, saccus indistinct and grading into second sternite; posterior width of tympanal organs narrower than anterior width, but venulae secundae not tapering sharply to form a neck; bullae not conspicuously invaginated in S2. In ysticalis-themis group, tympanal assemblage less asymmetrical than in medalba group (i.e., not conspicuously wider than long); tergo-sternal sclerite robust, conspicuous; bullae tympani longer than wide, saccus or rim of bullae tympani sclerotized at base; processi tympani present, lamellate, thumb-like, towards antero-lateral end of fornix; fornix tympani sclerotized; angle of fornical ulna obtuse; pons elongate, comprising (in part) two parallel, elongate, sclerotized prongs, divergent only at anterior terminus (posteromedial margin of saccus appears delimited by sclerotized rami, extends and remains parallel to pons for most of its length, pons extending towards bottom of saccus); saccus deep, pronounced (cf. "poches ou dépressions tympaniques" of Minet 1985); venulae secundae prominent, tapering such that "partie libre" (sensu Minet) of second sternite forms a "neck" as in S. speciosa; puteoli absent; posterior width of tympanal organs roughly half of anterior width. Male genitalia (Figs 36-60, part). Medalba group: Uncus oblong, cuspidate or mucronate, terminal edge entire; tegumen robust, divided into two dihedral, di-trapezohedral, or hemispherical bubbles that meet for a length that varies across species such that its dorsal ridges appear cruciate; juncture may appear as an elongate strut that divides anterior to base of uncus, such that anterior margin of tegumen may appear moderately emarginate (as in S. chanesalis) or more deeply invaginate (as in S. medalba and S. umbra). A transparent, membranous or sub-sclerotized area within uncus overlies a finger-like process arising from within center of gnathos, configuration harness-like, comprising a plate suspended by four arms, one pair extending to and (apparently) articulating with base of uncus dorsocaudally; other subtergal pair extending ventrally to and articulating with vinculum; connection between gnathal plate and tegumen membranous. Lower arms of gnathos appear to represent a fusion with ventro-tergal rods (Cf. Yoshiyasu 1985). Characteristic reduced male valvae extend straight out at roughly a 90° angle, and with a localized patch or cluster of ventral, filiform saccular setae. Valvae either simple and rounded or broadly emarginate to bilobed; reduced, their most prominent feature a pair of intrasaccular processes (one in each valva) oriented dorsally and variously naked or adorned with spines or denticles. Ventro-marginal setae absent or rudimentary. Juxta U-shaped or broadly V-shaped, robust at base, vaguely taurean. Phallus simple, cornuti absent. Ysticalis-themis group: Uncus obovoid or superficially tridentate (appearing trefoilor spade tipped); tegumen robust, divided into two obliquely-oriented oval sections meeting caudally near base of uncus, but diverging widely cephalad such that anterior margin of tegumen appears deeply invaginated; gnathos comprising a suspended rectangular plate with arms arising from each corner and a small, nub-like process arising centrally; dorsal arms wrap around anal tube, a ventral pair extend to termini of vinculum, such that gnathos almost appears to articulate both with uncus-tegumen and with vinculum, which is variously U-shaped or horseshoe shaped with pronounced pockets at each terminus. Valvae complex, comprising regions and processes that are variously sclerotized, fleshy in appearance, and/or bearing tufts of setae: intrasaccular flange located towards latero-ventral edge and sclerotized to form a trigger-shaped process; robust, spine-like setae on valva; ventro-marginal setae present on valva, either distributed evenly along length of outer margin of sacculus or concentrated at ventro-saccular "ulna"; costa robust and joined to rest of valva by a narrow membranous area; valva with secondary outer fleshy setose lobe or process below costa; recurved/decumbent setal plume associated with terminus of costa. Juxta robust, V-shaped or broadly Ushaped, ventral tip curved outward forming a small chin-like platform in S. themis and S. rasa; a less robust, more open U-shape in S. nyx complex. Phallus with two cornuti. Female genitalia (Figs 38-63, part) -Medalba group: Papillae anales convex, partially appressed but separate, setose; posterior and anterior apophyses roughly equivalent in length, not especially robust; antrum may be conspicuous, chalice-like; ductus bursae short, not discretely circumscribed; corpus bursae membranous, elongate, without signa; ductus seminalis arising from posterior end of corpus bursae. Ysticalis-themis group: Papillae anales setose, rounded, not conspicuously dihedral (except in S. lachesis); colliculum, if present, a partial collar, sometimes shortened to form a narrow ring immediately outside corpus bursae, ductus bursae per se all but eliminated; note that in contrast to Udea Guenée (1845), for example, ductus bursae, if present, developed posterior to colliculum (cf. Mally and Nuss 2011: 63, Fig. 3), an elongate band or partial sleeve immediately occupying antrum, appearing as a sclerotized band on floor of ductus bursae; corpus bursae globular or ovoid (more elongate in S. ysticalis), without signa, one or two accessory bursae posteriad where ductus seminalis attached.
Species variation. Individual species variation with respect to wing polymorphism is especially acute in the S. nyx complex; of particular interest here are the male sec-ondary sexual characteristics, which covary imperfectly across species and are discussed below. Schacontia species may vary greatly in size (>100% wingspan).
Distribution. Collectively, Schacontia species are distributed across Mexico, south to Central America (Guatemala, Costa Rica, Panama) and South America (Bolivia, Brazil, Ecuador, Venezuela) and the Caribbean (Puerto Rico, Cuba, Hispaniola). A single North American record of Schacontia themis is reported here from Sanibel Island, Florida (USA: Lee Co.).
Biology. Larvae are internal feeders that may induce galls, and pupate within the host. The only known host plant records are in Capparaceae: in Costa Rica, larvae have been reared from Podangrogyne decipiens (Triana & Planch.) Woodson (Solis, Nishida and Metz, in preparation); Cleome spinosa Jacq. has been reported as host for S. chanesalis; Capparis frondosa Jacq., and C. verrucosa Jacq. are reported for other Schacontia species.
Remarks. Schacontia was described by Dyar (1914) to accommodate three species, whose original descriptions were based primarily on wing pattern: the type species Schacontia medalba (Schaus, 1904; formerly Acontia medlba); S. chanesalis (Druce, 1899), formerly Pionea chanesalis; and Schacontia replica Dyar, 1914, the last of which accompanied the generic description (Druce 1899: 557, Schaus 1904: 163, Dyar 1914. Dyar (1925: 8) later described Thlecteria ysticalis from a female specimen, also on the basis of wing pattern, and this species was later removed to Schacontia by Munroe (1995: 42), who also recognized S. pfeifferi Amsel, 1956, raising the total number of species recognized in the genus to five. Amsel's (1956: 101-102) description of S. pfeifferi, which placed Schacontia in the Schoenobiinae, is the most complete description to date and one of only two works prior to the present to figure or characterize genitalia (the other being Solis 2009). Neither Schaus nor subsequent authors were explicit in their characterization of what makes Schacontia unique or in their rationale for describing and including new species in the genus.

Key to species of Schacontia
Key to species of Schacontia: Male Genitalia + Habitus + Female genitalia (part)  Diagnosis. Specimens of S. medalba are most readily diagnosed from those of S. chanesalis by male genitalia, specifically the reduced, unlobed valvae and the naked intrasaccular process, features they share with S. umbra.
Re-description (Fig. 1). Forewing length 6.5-1.0 mm. Head -Frons conical; labial palpi straight, extending as far as clypeus. Thorax -Female with one pair of hind tibial spurs (medial pair absent); legs uniform gray brown. Forewing. Basal area primarily gray brown, undivided; antemedial (am) line meets anal margin. Subterminal line interrupted by wing veins; medial area partially suffused with white, especially basad; white postmedial line appears shaded basally, interrupts/traverses dark shading between apical area and distal region of medial area; this "double" line faintly common to HW; FW fringe gray-brown. Hindwing. Postmedial line present, conspicuous (see above); terminal area lightly shaded, fringe white. Abdomen -Apical bands of pale scales on abdominal segments; terminal dots grayish brown, faint if present. Tympanal organs (Fig. 21). As for the medalba group, vide supra. Male genitalia (Figs 36, 37). Teguminal sulcus short, such that anterior margin of tegumen appears deeply invaginate; juxta U-shaped; valvae simple, reduced, rounded, not bilobed or emarginate; intrasaccular process a simple flange; intrasaccular process naked; phallus simple, cornuti absent. Female genitalia (Fig. 38)    Diagnosis. Specimens of S. chanesalis are best distinguished from those of S. medalba by the male genitalia, specifically a more sinuate valva and more denticled or rugose (as opposed to naked) intrasaccular process. The valvae are less conspicuously lobate than in S. umbra (below). Forewing pattern somewhat variable, as in S. medalba, but antemedial area more often traversed by white bar originating at scapula, enhancing the baso-costal patch.
Re-description (Fig. 2). Forewing length: 4.5-9.0 mm. Head -Ocelli and chaetosemata absent; proboscis reduced. Labial palpi porrect, extending slightly beyond clypeus. Frons conical; vertex and frons grayish brown, intermixed with white scales medially and along anterior bases of antennae. Thorax -Prothoracic collar light gray intermixed with gray-brown and white scales. Tegula and mesoscutum mostly gray, intermixed with light-gray and/or grayish-brown scales, the posterior apex of tegulae pale gray. Legs predominantly white, gray shading throughout foreleg; female with one pair of hind tibial spurs (medial pair absent). Forewing. Baso-costal triangle flanked by white scaling towards inner margin and in medial area, which is outwardly shaded brown (suffused with white basad). Postmedial area (between postmedial line and subterminal line) grayish brown. Subterminal line white; terminal line black, interrupted. Marginal scales brown. Basal area grayish brown traversed by a white band. Fringe scales light gray. Hindwing. Ground color white/very light gray, darker postmedially; postmedial lines grayish brown, white distally, conspicuous and common with FW. Subterminal area shaded darker brown; fringe white. Sc+R1 and Rs anastomosed slightly beyond dilated base of former. Male and female acanthae of frenulum fused from near base to apex to form one bristle. Abdomen -Ground color mostly dark gray intermixed with gray and light-gray scales above, white on undersurface. Tympanal organs (Fig. 22). As above for medalba group, vide supra. Male Genitalia (Figs 39, 40) -Tegumen divided dorsally into two dihedral or hemi-spherical "bubbles" that meet at a central sulcus, which divides anterior to base of uncus and forms a Y-shaped strut. Teguminal sulcus long, extending length of two teguminal lobes, such that anterior margin of tegumen appears emarginate, but not deeply invaginated. Uncus oblong, mucronate or miter-like, culminating in a distinct tip; concave or spatulate, setose on inner (ventral) surface. A membranous, more or less circular region at base of uncus positioned directly above (dorsal to) finger-like projection of gnathos, which also comprises a floating sub-tegumenal plate with four arms. Finger-like process arises from center of gnathos; dorsal pair of arms, which meet at juncture of uncus and tegumen, appearing to fulfill traditional description of gnathos by enveloping the anal tube, and the anterior pair extending ventrolaterally towards the vinculum, resembling a wishbone. Gnathos thus appears as a subtegumental (ventrad) suspension. Valvae reduced, broadly emarginate, bilobed; intrasaccular process a simple flange, denticled or rugose; subapical setal cluster near saccular margin. Costa robust, curved, appearing to arise near the respective vincular terminus. Juxta horseshoe-shaped, the base wider than the lateral "arms." Phallus simple, moderately sclerotized throughout; cornuti absent. Female Genitalia (Fig. 41) -Papillae anales appressed; antrum apparent, chalice-like; ductus bursae short; corpus bursae elongate, without signa, caeca, or appendix bursae; ductus seminalis originating from posterior portion of corpus bursae. Ostium bursae with membrane between seventh and eighth segments.
Distribution. Mexico, Guatemala; Costa Rica; Venezuela. Immature stages. Unknown. Variation. In size, with Mexican specimens appearing smaller in wingspan (FW length 4.5-7.0 mm) than Central American specimens.
Remarks. It is with some trepidation that we synonymize both replica and especially pfeifferi with chanesalis. Pfeifferi in particular was, until this work, the only Schacontia for which a detailed description had been published, and its continental separation from the type locality and primary distribution of chanesalis might suggest the potential for as yet unrecognized diagnosable species. Diagnosis. Habitus, male genitalia (Figs 3, 42). This species is most readily diagnosed by the darkly shaded forewings and by the male genitalia, which have the following features in common with medalba: anterior margin of tegumen deeply invaginate, outer margin of valva entire, intra-saccular process naked.
Immature stages. Unknown. Variation. Markings may be obscured in some specimens, rendering them more or less uniformly gray brown.
Etymology. The specific epithet refers to the dark wing shading of this species and is treated as a noun in apposition.
Biology. Unknown. Distribution. Central Ecuador.  (Figs 4, 44). The forewing pattern of this species makes it unmistakable; readily diagnosed by a combination of the frosted medial area common to other Schacontia and the interruption of the brown basal area to render a medio-basal patch encircled in white. Male genitalia diagnosed from those of other Schacontia species by the combination of expanded (not truncate) but inornate valvae, and reduced features associated with them, such as the inconspicuous intrasaccular flange; and a blunt, squarish, barely-tapering uncus.
Immature stages.  (Fig. 25). As for ysticalis-themis group, vide supra. Male genitalia (Figs 46, 47) -Teguminal sulcus short, such that anterior margin of tegumen appears deeply invaginate, the two oblong teguminal lobes joined obliquely. Uncus wider than long; terminal edge of uncus entire. Gnathos quadrate. Juxta an inverted triangular plate or robust "V", less sclerotized at center. Valvae complex; costa robust with recurved, elongate tufts of setae; subcostal lobe with petiolate scales, most arched towards dorsal articulation of valva; with secondary outer, oblong lobe or process below costa; with fleshy setose lobe associated with terminus of costa and located between distal portion of costa and lower portion. Intrasaccular process a simple flange, the inner surface of which bears chisel-shaped setae; with robust, spine-like setae at base; submarginal area of sacculus setose. Saccular margin angled close to vinculum, not at saccular mid-point; ventro-marginal setae concentrated at saccular ulna. Phallus moderately sclerotized; vesica with a small cornutus. Female genitalia (Fig. 48) (Fig. 6). Unlike many Schacontia, there is little to no contrast between the medial area and the rest of the forewing; although this holds for both S. rasa and S. clotho as well, those two are readily distinguished on other grounds. Male S. themis exhibit the full range of secondary sexual features known from the genus (flattened hind tibial spur, elongate hind tibial scales with embedded dark patch, epipleural setae, and concave metatarsal structure) as well as long abdominal coremata ( Fig. 19; Table 1); S. rasa (below), the putative sister species of S. themis, has none of these (but see remarks). Two other Schacontia species (S. clotho and S. lachesis) do share these features in part, but not the genitalic configuration that characterizes S. themis and S. rasa (see below). Male and female genitalia (Figs 49-51). As with the remaining Schacontia, males of this species are most readily diagnosed by a combination of genitalic and external secondary sexual characteristics. The male genitalia best distinguish this species and its putative sister, S. rasa (below), from other Schacontia: The uncus has a characteristic, expansive trefoil-shaped tip and lateral edges that appear swollen or re-enforced (as S. lachesis and S. atropos do), and a raised, pronounced medial ridge (as they do not). The intrasaccular flange is robust and forms a trigger-shaped process at the latero-ventral edge of the valva.
Description. Male. (Fig. 6). Forewing length: 5.3-10.0 mm. Head -Ocelli present; proboscis with pale basal scales in males, light brown basal scales in females. Vertex and frons yellowish white in males, intermixed with brownish-gray scales in females; frons of normal (convex) contour; maxillary palpi uniformly grayish brown; labial palpi grayish brown in males, fading to gray apically in females; labial and maxillary palpi porrect, extending well beyond clypeus; antennal scape and pedicel yellowish white, flagellomeres grayish brown. Thorax -Thoracic collar, tegula and mesoscutum yellowish white in males, brownish gray in females. Males with flattened hind tibial apical spur, hind tibial tuft of long black scales intermixed with pale yellowish-white scales, epipleural setae, and shallow concave metatarsal modification. Females with two pair of hind tibial spurs (medial pair present). Forewing ground color straw, with few contrasting mark-table 1. Species diagnoses for the S. nyx complex based on male genitalia and secondary sexual characters. Format inspired by Ferguson (1992: 259 ings other than jagged chocolate-brown antemedial and postmedial lines, postmedial line outwardly bulging only slightly, towards costa. Medial area unicolorous with basal area and postmedial areas. Fringe darkened apically. Hindwing. Almost uniformly pale, shaded brown at subterminal area. Abdomen -Dorsal surface straw brown. Scales arranged in two terminal black dorsal spots in males. Lobe-like extensions resembling rudimentary coremata on 4 th abdominal sternum. Tympanal organs. (Fig. 26). As for ysticalis-themis group, vide supra. Male genitalia (Figs 49, 50) -Tegumen divided into two obliquely opposed oval sections meeting caudally near base of uncus and diverging widely anteriorly towards valvae. Teguminal sulcus short, such that anterior margin of tegumen appears deeply invaginate, two oblong teguminal lobes joined obliquely. Uncus with prominent medial ridge; uncal tip hastate, trefoil-like; lateral edges of uncus swollen, appearing re-enforced and lending a shovel-or scoop-like appearance to uncus. Gnathal plate narrowed to a transverse band with arms at each corner and a small, nublike process arising centrally; dorsal arms wrap around anal tube while a ventral pair extend to termini of vinculum. Vinculum variously U-shaped or horseshoe shaped with pronounced pockets or eyelets at each terminus. Juxta robust, V-shaped; valvae complex, robust costa arising near vincular terminus, extending almost length of valva, and taper-ing to a point. A serrate, trigger-like ventral spine arises from ventral margin of sacculus; robust, spine-like setae at base of valva; saccular margin angled close to vinculum, not at saccular mid-point; ventro-marginal setae concentrated at venter or saccular ulna; valva with secondary fleshy subcostal setose lobe, setal plume recurved/decumbent. Elongate saccular process at outer saccular margin; saccular margin with several stout setae. Inner surface of valvae dorsal to saccular area with a small, circular setal cluster. Phallus moderately sclerotized; vesica with two large cornuti. Female genitalia (Fig. 51) -Papillae anales separate, not especially swollen; anterior and posterior apophyses threadlike, approximately equivalent in length; antrum/ductus bursae conspicuously sclerotized with a partial ventral collar or sleeve ventrally anterior to colliculum; a sharp constriction between ductus bursae and colliculum; colliculum a short sclerotized ring immediately posterior to corpus bursae with a narrow differentially sclerotized band around its center; corpus bursae membranous, more or less round, without signa or conspicuous appendices except single posterio-dorsal lobe; ductus seminalis appears to originate from antrum. Immature stages. Unknown.
Variation. This species varies most obviously in size (5.3 mm-10.0 mm in male forewing length), and based on the examination of several anomalous specimens from the British Virgin Islands, the presence of male secondary sexual characteristics (tibial hair pencils and abdominal coremata) do not perfectly covary: Hair-penciled males with and without coremata are noted from Guana Island, examples annotated and/or segregated in "Material examined" section above; see also Discussion.
Etymology. The specific epithet refers to the Greek Titaness and embodiment of divine order and is treated as a noun in apposition.
Distribution. Mexico, Cuba, Dominican Republic (essentially, vic. Gulf of Mexico). Remarks. Schacontia rasa is evidently the sister species of S. themis. Were it not for the characters associated with the forewing ground color, female hind tibia, and male genitalia and given the homoplastic nature of certain male secondary sexual characters comparable to the system described by Ohno (2000) [see discussion], S. rasa would be a more obvious candidate for conspecificity with S. themis. We have treated anomalous specimens with "chimeric" distributions of male secondary sexual chacters under S. themis (above), and considered only those lacking both tibial hair pencils and abdominal coremata (in addition to genitalic features) to be unambiguously S. rasa, recognizing the need for future molecular work to evaluate the degree to which these character systems are functionally and genetically linked. DNA barcode data (meeting the Barcode Data Standard of Genbank, noted in Benson et al. 2012) are limited to three Dominican Republic specimens and do decisively unite two specimens of S. themis to the exclusion of S. rasa. Not enough specimens are sampled to test the variable sites as diagnostic characters and enable their use in the species' diagnoses (Goldstein and DeSalle 2011), but the data corroborate (albeit by a distance measure of >7%) the reliability of the morphological characters as consistent with two distinct species.
It was suggested by V.O. Becker (pers. comm., following the submission of this work) that the name Dichogama fernaldi Möschler, 1890, the type material of which has apparently been lost from MNHU, might refer to this species (see Becker & Miller, in prep., for discussion) and that it should be placed in the now monotypic genus Dichochroma, whose description is, in turn, based on a single female (and only known specimen) of the type species, D. muralis Forbes, 1944. This attribution of the specimens we consider to fall within S. themis (or S. rasa, below) to D. fernaldi is, however, not well corroborated by any character identified in the original description by Möschler, but only by process of elimination and the report of its being reared on Capparis by Wolcott (1950-1951: 658, cited in Becker andMiller, in prep.). While eliminating a nomen dubium is desirable and the process of elimination by which such an attribution might be reached intriguing, the recognition of two similar co-occurring species (S. themis and S. rasa) described here, corroborated at least indirectly by DNA barcode data, precludes any specific attribution. We therefore retain the description of Schacontia themis and S. rasa as such. Further, notwithstanding the superficial similarity of certain female Schacontia genitalia to those of the only known specimen Dichochroma muralis, both phylogenetic placement described in this work and priority of Schacontia would dictate that Dichochroma be sunk were it determined that these species were congeneric, even if male Dichochroma muralis were discovered and or more compelling character data were brought to bear.
The Schacontia nyx complex: Some of these species are not readily diagnosed by a single character; each, rather, is characterized by either an absence of characters (as in S. atropos) or by combinations of characters, all of them male, both genitalic and external, the latter presumably secondary sexual features. Diagnosis. Habitus (Fig. 8). Although more readily diagnosed by the male genitalia, nyx can be differentiated on the basis of wing pattern. Nyx shares with other members of the genus (and in particular of the complex of sibling species to which it belongs) the configuration of the medial area, with its outward subcostal bulge, but its more mottled appearance and less uniformly contrasting ground coloration between the medial and both the antemedial and postmedial areas. Male S. nyx does not bear the epipleural setae shared by most other members of the complex; nor do they exhibit a dark patch embedded within the elongate hind tibial scales (Table 1). Genitalia (Figs 55, 56). Male specimens of S. nyx are most readily diagnosed by the obovate uncus, which is without pronounced medial ridges or lateral swellings. The subcostal processes are more conspicuous and elongate than in S. themis or S. rasa, but less narrow than in other species in the complex (S. clotho, S. lachesis, S. atropos, or S. androgyne).
Immature stages. Unknown Etymology. Nyx, the primordial goddess of the night who according to myth stood at the beginning of creation, refers to the first of five closely related species in this complex. The specific epithet is treated as a noun in apposition.
Biology  (1♂, 1♀), the latter accompanied by "Genitalia Slide ♀ by JAL." Diagnosis. Habitus (Fig. 9). This species superficially resembles S. rasa in coloration and maculation; it is smaller and the male bears all of the secondary sexual characters, including coremata, known to occur within Schacontia (Table 1). Genitalia (Figs 58-60). The male genitalia of S. clotho place it unambiguously within the S. nyx complex as opposed to with S. themis or S. rasa. Moreover the subcostal lobe of the valva is elongate.  Description. Male. (Fig. 9). Forewing length: 6.9-7.0 mm (n=3) (Female 6.8 mm). Head -Ocelli present; proboscis normal; frons of normal contour; labial palpi porrect, extending beyond clypeus. Thorax -Prothoracic collar and tegulae an admixture of brown and mouse-gray scales. Flattened hind tibial spur, specialized hind tibial scales, epipleural setae present, and dark patch amidst male hind tibial scales all present. Female with two pairs of hind tibial spurs (medial pair present); shallow concave metatarsal modification present. Forewing. More lanceolate than in other Schacontia species. More or less uniform mouse gray, with very light dusting of very pale gray in medial and postmedial areas; medial area more darkly shaded than basal area and postmedial areas; FW fringe brown; subterminal line unbroken. Hindwing. Nearly translucent; postmedial line absent; fringe pale yellowish; subterminal line unbroken. Abdomen -Scales arranged in two terminal black dorsal spots in males; elongate coremata on 4 th abdominal segment (Fig. 34). Tympanal organs (Fig. 29). As for ysticalis-themis group, vide supra. Male genitalia (Figs 58, 59) -Uncus trefoil-shaped tip reduced to a small, more or less rhomboid nipple; lateral edges of uncus simple, undifferentiated; juxta broadly V-shaped, comparable in shape to an avian furcula, arms not robust; valvae complex, intrasaccular flange at latero-ventral edge and sclerotized to form a trigger-shaped process; robust, spine-like setae at base of valva; saccular margin rounded at mid-point; prominent setal comb at ventro-medial margin of valva; ventro-marginal setae concentrated at saccular ulna; costal bar diverges from subcostal lobe towards base of costa (isolation of costa >75% along length, character 49); valva with pronounced, elongate secondary outer lobe or process below costa; recurved/decumbent setal plume associated with terminus of costa; sharply hooked setal cluster prominent. Phallus moderately sclerotized; vesica with two large cornuti. Female genitalia (Fig. 60) -Papillae anales separate, not swollen; antrum/ductus bursae elongate (not chalice-like), faintly sclerotized posterior to colliculum, separated from colliculum by a sharp constriction; colliculum a short ring (not an elongate collar), with faintly sclerotized band, immediately posterior to corpus bursae; corpus more or less globular, surface complex; ductus seminalis originates at posterior end of corpus bursae. Immature stages. Unknown. Etymology. The specific epithet refers to the youngest of the three fates in Greek mythology, responsible for spinning the thread of human life, and is treated as a noun in apposition.
Biology. Unknown. Adults December. Distribution. Southern Ecuador. cus trefoil-shaped tip reduced to a small, more or less rhomboid nipple; juxta U-shaped, tapered ventrally; valvae complex, intrasaccular flange transposed towards latero-ventral edge and sclerotized to form a trigger-shaped process; robust, spine-like setae at base of valva; saccular margin sharply angled at saccular mid-point; ventro-marginal setae distrib-uted along length of outer margin of sacculus; valva with pronounced, elongate secondary outer lobe or process below costa; fleshy setose lobe and recurved/decumbent setal plume associated with terminus of costa; sharply hooked setal cluster prominent. Phallus moderately sclerotized; vesica with two large cornuti. Female genitalia (Fig. 63) -Papillae anales separate, unswollen; colliculum present, short, sclerotized, immediately posterior to corpus bursae, with narrow sclerotized band around center, sometimes separated from bursa by a sharp constriction; ductus bursae present, conspicuously sclerotized ventrally, entering corpus bursae dorsally; appendix bursae ventral, superficially complex; ductus seminalis attached at posterior end of ventral corpular out-pocketing. Immature stages. Unknown. Etymology. In Greek mythology, Lachesis is the middle sister of the three fates, the personification of destiny responsible for measuring the duration of human life. The specific epithet is treated as a noun in apposition.

Schacontia lachesis
Biology. Unknown. Adults in Brazil active January, April, November, December; adults in Bolivia active October-December.
Distribution. Central Brazil (Rondonia east to Bahia, Ceara and Rio de Janeiro), Bolivia (Santa Cruz). Diagnosis. Habitus (Fig. 12). Overlaps in appearance with S. lachesis, but males lack all secondary sexual features; hindwing uniformly pale throughout. Genitalia (Figs 64,65). Male genitalic features place it squarely in the nyx complex, from whose other member species it may be distinguished by the combination of the angled ventral edge of the saccus (ulna) and unmodified edges of the uncus.
Immature stages. Unknown. Etymology. The specific epithet refers to the third of the three fates. Treated as a noun in apposition.
Biology. Unknown. Distribution. Northern Venezuela. Remarks. Given the phenomenon described by Ohno (2000) it is well within the realm of possibility that this species represents a synonym of S. lachesis.

Discussion
The moths treated in this paper compise a range of geographically widespread and potentially localized cryptic species united by a range of synapomorphies. It is not possible to infer an unambiguous center of origin for Schacontia; members of both the typical medalba group and the ysticalis-themis group are distributed from Mexico to South America. While we note at least three trans-isthmian species (S. chanesalis, S. ysticalis, and S. themis), several species -including the entire S. nyx complex -are restricted to South America, while Schacontia rasa is known only from Mexico and the Caribbean. Among the more intriguing features of Schacontia that warrant further study is their gall-forming habits in association with capparaceous plants, and the enormous variation in size, which may be a function of indeterminate instar number.
There likely exist undiscovered Schacontia species, cryptic and otherwise. Schacontia chanesalis in particular may represent a complex of Mayrian species that are difficult to diagnose without more extensive molecular data. However, available data are such that obvious breaks in the continuum of variation are not obvious, and rather than allowing the weakly articulated epithet S. replica to persist as valid we have elected to synonymize it, along with S. pfeifferi. Although its presumptive range differs geographically from that of S. chanesalis, we saw little point in retaining S. pfeifferi given a lack of apparent distinguishing characters. Although its presumptive range differs geographically from that of S. chanesalis, we saw little point in retaining S. pfeifferi given a lack of apparent distinguishing characters. We took a less conservative approach within the S. nyx complex, nominating species on the basis of characters we suspect may be more labile than our limited collection of specimens suggests. We attribute the lack of phylogenetic decisiveness, particularly in the nyx complex, to homoplasy among characters associated with the male secondary sexual features.
By far the richest source of phylogenetic signal in our matrix are the male genitalia, accounting for almost half the characters included in our analysis. This is not unusual for species-level studies of Lepidoptera (or insects generally), and there may be a growing consensus that despite their likely being subjected to sexual selection and thus potentially evolving quite rapidly (Fisherian runaway), male genitalic characters nonetheless contain valuable information at multiple hierarchic levels (Simonsen and Roe 2009;Song and Bucheli 2010), a situation analogous to that of third positions or transitions in molecular phylogenetics (e.g. Källersjö et al. 1998). A noteworthy exception is the work of Solis and Maes (2002), who concluded that male genitalic characters were not especially useful at the subfamily level within the Crambidae.
In contrast, it is of particular relevance to the taxonomy of Schacontia that relatively little complex variation is observed in the female genitalia, particularly given that four of the five species described prior to this work -two of which are synomized herein -have female holotypes. Solis et al. (2009) discussed the roles of secondary sexual structures, including scent-producing structures and their associated modified scales (reviewed by Hallberg and Poppy 2003), in lepidopteran diversification broadly and in Pyraloidea specifically. They highlighted the historical differences of opinion between taxonomists who discounted the importance of such structures and those who viewed them as invaluable in lepidopteran classification (e.g., Janse 1931) and, following Solis (1993) and Simonsen and Roe (2009), in phylogenetic reconstruction in spite of empirical demonstrations of homoplasy. Based on the present work, male secondary sexual characters were among the more homoplastic we analyzed, at least insofar as their removal contributes resolution to the phylogenetic hypothesis generated. The intraspecific lability of characters such as hind tibial hair pencils or the abdominal coremata remains to be studied in detail. Other pyraloid examples of this kind have been discussed by Solis et al. (2009), and Hayden (2010, 2011 has since added examples of independently derived and readily reversed structures from the odontiine crambids: some species of Cliniodes bearing tufts on the prothoracic femora and 7 th sternite have sister species in which these structures are reduced or absent. In the genus Dicepolia, the two most commonly encountered species share a prothoracic tibial tuft but are otherwise unrelated; meanwhile an S7 tuft is an unreversed synapomorphy of Neotropical members of the genus. Apropos of species diagnosis, we recall that as compelling as are the raw diversity of secondary sexual characters and the demonstrations of their phylogenetic lability, there have been suggestions that the expression of such structures may be underlain by rather simple genetic systems. Following Ohno's (2000) observation of withinbrood polymorphism with respect to mid-tibial tufts in Ostrinia (Crambidae), Frolov et al. (2007) suggested that this polymorphism may depend on two di-allelic loci unrelated to reproductive isolation. Although Frolov et al. speculated somewhat with respect to the putative roles of such structures in sympatric speciation, it is clear that even seemingly complex structures may be subject to the simple rules of Mendelian inheritance, character fixation and extinction. If such is the case, then even "important" characters such those involved in courtship, lekking, or mate recognition might be polymorphic, locally fixed, or even frequency-dependent, and might not necessarily serve to diagnose species. Apparent polymorphism in features such as tibial hair pencils (as in S. lachesis) or abdominal coremata (as in S. themis) speak to the possibility that species pairs such as S. lachesis-atropos or S. themis-rasa might be conspecific. Our limited DNA barcode data suggest otherwise for the latter species pair, but can not corroborate the diagnostic power of the morphological characters themselves without more extensive sampling.
format in post-review. In particular, we wish to thank V.O. Becker for providing material that formed the basis for several new species descriptions, and to S. Miller and V.O. Becker for making certain material from the British Virgin Islands available to this study. DNA barcode sequences were provided by the University of Guelph under the iBOL project funded by Genome Canada. Paul Goldstein was supported a Specific Cooperative Agreement, USDA with Charles Mitter, Dept. of Entomology, University of Maryland.