A revision of the genus Antepione Packard with description of the new genus Pionenta Ferris (Lepidoptera, Geometridae, Ennominae).

Abstract Based on genitalic studies, the new genus Pionenta is established for two taxa formerly placed under Antepione. The taxa hewesata and ochreata (and previously associated synonyms) are now synonomized as Pionenta ochreata. Three species of Antepione are now recognized: Antepione thisoaria, Antepione imitata, Antepione tiselaaria with the taxa comstocki, constans, and indiscretata synonomized under Antepione imitata. No new species are described. Adults and genitalia are illustrated, including type specimens.


Introduction
A genitalic study of the eight species recognized by Parsons et al. (1999) associated with genus Antepione uncovered two distinct and quite diff erent forms in both the male and female genitalia. Th is situation was alluded to by Pitkin (2002: 283), who excluded Antepione ochreata (Hulst) in her treatment of the genus. In Antepione, as restricted herein, the DHW Dorsal hindwing.

FWL
Forewing length, measured along costa from base to apex. MB medial band = area between DFW AML and PML.

TL
Type locality.

Key to genera
(based on DFW pattern and genitalia) 1 DFW triangular costal dark patch present; male genitalia lack furca; female genitalia lack colliculum and signum .. Type species: Epione depontanata Grote, 1864. Location of type unknown; originally placed in ANSP. Described from Maryland, USA.
Description. Adults. Sexually dimorphic and sexes polyphenic; FWL 13-21 mm. Antenna simple, more slender in females. Head -Dark ochreous speckled with darker scales, concolorous collar; labial palpi broad, barely extending beyond frons, ochreous speckled with darker scales. Th orax, abdomen, legs -Ochreous or pale tan as in wings with widely scattered small brown scales. Wings -FW outer margin arcuate at vein M 3 and HW; DFW apex acute to falcate. Usually obscure narrow dark DFW submarginal band; small dark discal spots both wings. Males. Dorsal color varies from gray, medium ochreous to medium brown. DFW AML and PML variable from pronounced and dark to broken and indistinct; medial band concolorous with remainder of wing, or paler and yellowish; a dark triangular patch with blunted or acute apex, with or without pale oblong spot, located along costa distad of PML. DHW with dark narrow medial band varying individually from dark to indistinct. Ventrally paler with dorsal maculation repeated, usually with less intensity. Females. Dorsal color varies from yellow through pale ochreous to medium ochreous and gray. Crosslines usually indistinct. DFW triangular patch as in males, PML above inner margin expanded into two large oblong brown spots. Ventrally paler with dorsal maculation repeated, usually with less intensity. Male genitalia -Uncus stout, slightly decurved, tapering to a rounded tip; medial gnathos with a few small teeth; valva rounded at apex; anellus with small spines; aedeagus truncate with one large oblong cornutus near base of vesica. Female genitalia -Apophyses long, slender; posterior apophyses ca. 1.8 × anterior apophyses; colliculum absent; ductus bursae ridged, short, partially sclerotized at posterior end; corpus bursae without signum, oblong with membranous anterior sac; ductus seminalis originates at top of ductus bursae.

Systematics
Fixation of type locality. Th e Central American taxa were not recognized and described until 1892 (azonax) and 1912 (rhomboidaria). On this basis, I infer that specimens from this region were not available to Guenée in 1857 when he described thisoaria, and that the holotype was collected in eastern North America. In habitus, the HT matches exactly female specimens of the sulphuraria/sulphurata form. Th e HT was most probably collected in the Middle Atlantic region. I hereby fi x the type locality as eastern North America. Based on my research, it appears that Forbes (1948) was the fi rst to use the name thisoaria in the North American fauna.
Other material examined. 84 specimens (a few by photograph) from Alabama, Indiana, Kentucky, Mexico, Michigan, New Jersey, New York, Nova Scotia, Pennsylvania, Quebec, Tennessee, Virginia. Additional distribution records were obtained from individuals and several museums, including 439 from the Carnegie Museum of Natural History, Pittsburgh, PA.
Diagnosis. Antepione thisoaria is most easily separated from A. imitata based on geography. It does not occur west of the 95th parallel, while A. imitata extends eastward only to west Texas, and is not recorded from Central America. In the male genitalia, the apical region of the valva lacks spines, which are present in the valva of imitata. In the female genitalia, the corpus bursae is initially swollen while not so in A. imitata.
Biology and distribution (Fig. 59). McGuffi n (1987: 88-89) described the early stages and cited three specifi c larval hosts: Alnus rugosa (Du Roi) Spreng; Physocarpus opulifolius (L.) Maxim; Prunus serotina Ehrh. Various additional larval hosts are report-ed in the literature in the families Aceraceae, Anacardiaceae, Betulaceae, Ebenaceae, and Rosaceae. Th e last instar larva was illustrated by Wagner et al. (2001, p. 155) and Wagner (2005, p. 195). Adults fl y April-May with an occasional mid-March and mid-June record, July-August with occasional September to mid-October records. Th ere is (Eugonobapta) constans HT, adult, pin labels (AMNH photo) and male genitalia. Th e balsam embedding medium has fogged with age producing the apparent lack of focus in the genitalia photo. one generation in Canada, and at least two southward. Th e distribution map (Fig. 59) represents the data that I was able to locate. Th e heavy distribution in Pennsylvania refl ects intensive collecting in that state by CMNH personnel and volunteers. Undoubtedly similar eff orts in neighboring areas should produce additional records. Th e overall range of this species is: in CANADA from Nova Scotia to Manitoba; in the UNITED STATES (county records in parentheses) then south and west to the Gulf states to the 95th parallel, including Alabama ( (Pitkin et al. 1996). Covell (1984) stated the westward range of the species to Texas. I have been unable to confi rm Texas from museum records. Forbes (1948) stated: ". . . varieties in Colorado, Texas, and Arizona." His "varieties" are assumed to be A. imitata. Although the distribution map (Fig. 59) suggests occurrence of thisoaria in Florida, Vermont and Wisconsin, no records were found.
Remarks. Th e gray spring form of the moth (Figs 11, 15) was described by Packard as the species furciferata. Th e male (Fig. 14) represents the summer form arcasaria, and the female (Fig. 16) represents the summer form sulphuraria = sulphurata. Packard (1876) redescribed Heterolocha sulphuraria Packard, 1873 as Antepione sulphurata. Once barcoding data are available, the disjunct distributions of Mexican and Central American populations may ultimately prove to be separate species, in which case the name azonax Druce, 1892 (Costa Rica, Guatemala) is available and has date priority over rhomboidaria Oberthür, 1912 (Costa Rica) and rivulata Warren, 1897 (Costa Rica). Th e two female specimens in the CNC from Tuxpan, Michoacan, Mexico are exact matches for the sulphuraria/sulphurata phenotype and were collected in early August, 1959. Edwards, 1884 Figs 2-5, 20-33, 59 Antepione comstocki Sperry 1939, syn Antepione comstocki male HT (Fig. 5), Arizona, [Pima Co.], Baboquivari Mts., 26 April, 1938 [CNC]. Tetracis indiscretata female HT (Fig. 3), New Mexico, [San Miguel Co.], Las Vegas, August, 1882 [SEMC]. Eugonobapta constans male HT (Fig. 4) Diagnosis. Antepione imitata is most easily separated from A. thisoaria based on geography. It does not occur east of west Texas and is not recorded from Central America, while A. thisoaria extends west only to the 95th parallel. In the male genitalia, the apical region of the valva exhibits 3 long robust spines and additional fi ne setae, which are not present in the valva of thisoaria. In the female genitalia, the corpus bursae is not initially swollen as in A. thisoaria.
Remarks. One male specimen (Fig. 25) of the comstocki phenotype examined from Las Animas Co., Colorado lacks the characteristic DFW costal triangular patch, caus-ing it to resemble superfi cially the ligata form of Pionenta ochreata. Th e male genitalia, however, are typical of A. imitata.
Biology and distribution (Fig. 59). Noel McFarland (Hereford, AZ) reared the species on Ribes aureum Push. from ova from an adult female of the nearly uniformly brownish-ochreous April-May generation; adults emerged June-July. Th e  Th is locality is essentially due south of the western Texas records, and one might anticipate that with further collecting A. imitata will prove to be widespread in northern Mexico. It is generally associated with riparian canyons up to 6000' (1830m).
Discussion. As is also the case with Antepione thisoaria, most spring individuals of A. imitata are rather drab in appearance with lightly maculated brownish males (the comstocki phenotype) and pale creamy colored or ochreous females. Th e strongly maculated males and yellow females appear in the later generations in company with the rather drab early-season phenotypes. In his original descriptions of imitata and indiscretata, Edwards provided no insight as to why he assigned imitata to Antepione and indiscretata to Tetracis. Both taxa are described on the same page with the description of imitata preceding that of indiscretata. He characterized the color of imitata as similar to the yellow sulphurata phenotype of thisoaria, and indiscretata as "Ochraceus drab." Over the years the type specimens have faded to some extent so that they now appear nearly identical in color, the only diff erence being the extent of the dark maculation. Th e name constans appears to have been applied to the heavily maculated male phenotype, as best can be determined from the poor condition of the HT.

Antepione tiselaaria (Dyar, 1912) Figs 6-10
Paragonia tiselaaria Dyar 1912 Type material. Male HT (Fig. 6), Mexico, Minerale de Zacualpan, January, 1911 [USNM]. Comment: Dyar (1912: 87) stated the type locality only as "Zacualpan" and not "Minerale de Zacualpan" as shown on the specimen label. I interpret the label to mean the Zacualpan mining region located in the state of Morelos south of Mexico City, today still an active silver mining district.
Diagnosis. Females not known to the author. Mexican specimens of Antepione tiselaaria males are most easily separated from A. imitata based on geography, since the latter species does not penetrate south to central Mexico. In Costa Rica, where A. thisoaria is also  reported, A. tiselaaria manifests a more orange-brown overall color than the drab ochreousgray form of thisoaria. In the male genitalia, the apical region of the valva is covered with multiple short slender translucent spines over most of the surface except toward the base; spines are absent in the valva of A. thisoaria, and 3 long robust spines occur in A. imitata.
Description. Adults. Only males were available for examination. As described above for the genus, other than the wings. FWL 17-18 mm. Wings -FW outer margin arcuate (roundly produced about) vein M 3 and HW; DFW apex sightly acute, not fal-cate. Dorsal color pale orange-brown-ochreous with darker maculation. AML a narrow band centrally with a few paler scales, PML an interrupted band with irregular edges and centrally paler, widening substantially approaching inner margin; MB not clearly defi ned with splotchy brown maculation over paler ground color; a dark triangular patch with blunted or acute apex, with or without pale oblong spot, located along costa distad of PML; small dark discal spots FW and HW. Ventrally paler with dorsal maculation repeated with slightly less intensity. Male genitalia. Figs 9-10. Dissection 1m. Uncus stout, slightly decurved, tapering to a rounded tip; gnathos with unjoined slender arms, medial gnathos with a few very small dark teeth; valva rounded at apex with multiple short slender translucent spines over most of the surface excepting toward the base, produced ventral ridge forming one large and one short projection; anellus with two sclerotized spinose lobes; aedeagus truncate with one large (equal to diameter of aedeagus shaft) oblong triangular cornutus near base of vesica; fully everted vesica initially spherical becoming a tapered tube.
Biology and distribution. Early stages unknown. Current distribution records are for the Mexican states of Morelos and Puebla, and Costa Rica.
Pionenta Ferris, gen. n. urn:lsid:zoobank.org:act:95BF80CA-F55F-40D9-B877-A25252635770 Type species: Antepione ochreata Hulst, 1898. Etymology. Pionenta is a masculine anagram of Antepione. Diagnosis. Pionenta ochreata lacks the DFW triangular costal patch found in all species of Antepione. Th e well-developed DFW AML and PML form a wedge-shaped medial band absent in Antepione. Th e robust centered furca in the male genitalia and large stellate signum in the female genitalia of Pionenta are absent in Antepione.
Description. Adults. Sexually dimorphic and both sexes are polyphenic; FWL 14-19 mm. Antenna simple. Head -Uniformly ochreous, collar concolorous; labial palpi relatively narrow, slightly upcurved, ochreous, barely extending beyond frons. Th orax, abdomen, legs -Uniformly colored as in ground color of wings with a few widely scattered small brown scales on legs. Wings -Outer margin arcuate FW (about M3) and HW; DFW apex normally sightly falcate. Wing color variable from pale creamy white to ochreous tan. AML and PML narrow and brown (occasionally reddish-brown), PML continues on DHW as medial line; AML with narrow pale shading basad, PML with narrow pale shading distad. MB trapezoidal tapering inward from costa to inner margin. Small dark discal spots present FW and HW. Scattered dark patches may be present basally and submarginally on DFW, and submarginally on DWH. Ventrally paler with dorsal maculation only weakly repeated. Male genitalia (7 dissections by author, additional museum slides examined) -Uncus stout, slightly decurved, tapering at apex to a rounded tip; gnathos v-shaped with well-sclerotized edges, medially a sharp upcurved tip with numerous very small teeth; valva rounded at apex, but with blunt triangular projection at end of sclerotized costa; anellus membranous without spines or setae, with central robust cylindrical furca covered by numerous short spines on rounded apex; aedeagus truncate with two long sclerotized pointed extensions from apical margin and a variable patch of apparently deciduous dark setae near base of otherwise membranous short cylindrical vesica. Female genitalia (6 dissections) -Posterior apophyses short, anterior apophyses much reduced ca. 0.4 × posterior apophyses; sterigma well sclerotized; posterior margin of lamella antevaginalis rounded at ends with central depression; well-defi ned colliculum; partially ridged short ductus bursae opens into ovoid membranous corpus bursae; one large centrally located oval stellate signum; ductus seminalis originates immediately below colliculum.   Hulst, 1898 Figs 34-58, 60 Antepione hewesata Sperry 1948, syn. n. Sabulodes arizonata Taylor 1905, syn. rev. Sabulodes dyari Grossbeck 1908, syn. rev. Sabulodes ligata Grossbeck 1908 Type material. Male HT (Fig. 34), Arizona, Senator [probably Senator Mine, Yavapai Co.] [AMNH].
Biology and distribution (Fig. 60). Early stages unknown. Adults from mid-May to August in riparian canyons and dry coniferous forest to 8400' (2560m); probably more than one generation. Collection records include Arizona (Cochise, Coconino, Pima, Santa. Cruz, Yavapai), New Mexico (Grant).
Discussion. Based on the male and female genitalia, Pionenta is closely related to Tetracis. Th e male genitalia of both genera possess a well defi ned central furca. Th e female genitalia of both genera possess a well defi ned colliculum and prominent single signum. Th e gnathos in Pionenta does not have a quadrate dorso-caudal margin with two to four (occasionally fi ve) widely separated, dorsally projecting teeth as found in Tetracis (Ferris & Schmidt, 2010). Once barcoding of the North America geometrid genera has been completed, the relationship of Pionenta can be established.