The North American species of Charadra Walker , with a revision of the Charadra pata ( Druce ) group ( Noctuidae , Pantheinae )

Th e North American species of the genus Charadra Walker are reviewed, and the species of the yellowhindwing (C. pata) group are revised. Four species of the pata group are described as new: Charadra franclemonti sp. n. (Arizona), C. tapa sp. n. (Arizona), C. cakulha sp. n. (Mexico), and C. coyopa (Mexico) sp. n. A new species related to C. deridens (previously treated as C. ingenua) is described from Arizona / New Mexico / Texas, and Charadra ingenua syn. n. is synonomized with Charadra deridens. Th e types of Charadra pata, C. oligarchia, C. patafex and C. ingenua are illustrated.


Introduction
Th e genus Charadra Walker as currently understood is comprised of three yellow-hindwinged species (C.pata (Druce), C. oligarchia Dyar, and C. patafex Dyar) and fi ve greyhindwinged species (C.deridens (Guenée), C. ingenua J.B. Smith, C. dispulsa Morrison, C. nigracreta H. Edwards and C. nitens Schaus).Charadra is restricted to the New World with the greatest species diversity found from the southwestern United States to Central America.Th e widespread and variable Charadra deridens is a well-known species found across most of the continental United States and southern Canada.Based on phenotype and genitalic structure, the genus is arranged into three groups: 1) the pata group (yellow hindwing; male vesica with two simple thorn-like cornuti; clasper not reaching apex of valve; saccus V-shaped; female antevaginal plate bilobed or double pronged; female ductus bursae lacking lateral twisted fl anges); includes C. pata, C. oligarchia, C. patafex, and four additional species described herein (C.franclemonti sp.n., C. tapa sp.n., C. cakulha sp.n., C. coyopa sp.n.).Th is group is primarily Mexican -Central American in distribution, with two species reaching southeastern Arizona.
2) the C. deridens group (male cornuti multi-spined, basal cornutus crest-like; clasper reaching apex of valve; saccus broadly U-shaped; female ductus bursae with lateral, ventrally twisted fl anges); includes C. deridens, C. moneta sp.n., C. dispulsa, C. nitens, and several undescribed Central American species (BCS, in prep.); as a whole, this group occurs from temperate North America southward to at least Costa Rica.3) the nigracreta group, with only C. nigracreta, is characterized by a simple valve that lacks a clasper; a massive plate-like transtilla; absence of cornuti on the vesica, and presence of a sclerite between the ductus and corpus bursae.Th ese groups may warrant recognition as separate genera, but a review of other pantheine genera, notably the closely related Colocasia Ochsenheimer and Pseudopanthea McDunnough, is needed before generic limits can be revised and is beyond the scope of this work.
With the exception of Arizona material, the pata group is very poorly represented in collections.It appears to be quite diverse in Mexico, where more species will likely be discovered.J. G. Franclemont collected and reared series of the yellow-winged species in southern Arizona in the 1960's; this material consists of two species, neither has an available name.Th ese two Arizona taxa and two new Mexican species are described herein.Th e western North American taxon C. ingenua has been misunderstood and is here synonymized under C. deridens syn.n., and a new name is provided for the taxon previously treated as C. ingenua.

Methods and materials
Adult genitalia were prepared following the methods detailed by Lafontaine (2004).Cleaned, stained genitalia were stored and examined in 30% ethanol, and slidemounted in Euparal before being photographed.Molecular variation for some species was assessed based on the 658 base-pair 'barcode' region of the fi rst subunit of the cytochrome oxidase (cox1) gene (Hebert et al. 2003).DNA was extracted from one leg removed from a dried specimen, and processed at the Canadian Centre for DNA Barcoding, Guelph, Ontario.DNA extraction, amplifi cation and sequencing protocols for the Barcode of Life initiative are given in Hebert et al. (2003).Haplotypes of all barcode sequences were compared with phylograms constructed using the neighbourjoining method in PAUP 4.0*b10 (Altivec) (Swoff ord 2002).Phyletic distances were calculated using the Kimura-2-Parameter (K2P) distance model.Data for molecular voucher specimens, including trace fi les and photographs, are available at http://barcodinglife.com (project: Lepidoptera of NA Phase II: "Charadra revision" under the "Published Projects" tab).Molecular sequences have been submitted to GenBank, but accession numbers were not available at the time of publication.
Th e generic placement of Charadra in relation to other pantheine genera, especially Pseudopanthea and Colocasia, is in need of review, and more than one genus for the species currently placed in Charadra may need to be recognized.Th e morphology of the genus is quite heterogeneous, but important shared genital characters include the following: apex of uncus with slight to very pronounced medial notch; clasper parallel to ventral valve margin (perpendicular in Panthea Hübner) but absent in C. nitens; dorsal tegumen lacking process (lobed in Panthea, prong-like or fl ange-like in Colocasia); basal costal process absent or highly developed; vesica with two strongly sclerotized cornuti (absent in C. nitens); female ductus bursae sclerotized laterally; corpus bursae lacking signa.Some of the character states treated as autapomorphic for Panthea by Fibiger et al. (2009) are also present in Charadra, including a dorsally membranous aedeagus and the presence of cornuti on the vesica.(Druce) Fig. 1 Trisulodes pata Druce, 1894: 362.Charadra patens J. B. Smith, 1908;misspelling.‡Charadra basifl ava J. B. Smith, 1908; unavailable name.Type material.Charadra pata -Guatemala, Guatemala City.Holotype female.BMNH; examined.Charadra basifl ava -this taxon was listed as a synonym of pata by Franclemont & Todd (1983).Todd (1982) questioned the validity of Smith's description, which consists of: "Trisuloides patens [sic] Druce, is a Charadra which I had named basifl ava before Dr. Barnes called my attention to the fi gure in the Biologia (II, 509, p. 96).It has been taken at Palmerlee, Cochise Co., Arizona, and is no doubt a member of our fauna."(Smith 1908).Th e question is then what Smith meant when he said "had named."Since there is no earlier published mention of basifl ava, we take this statement to mean that Smith had determined and labeled a specimen as a new species and intended to describe it as basifl ava.Todd (1982) also concluded this was the most likely meaning of Smith's statement, and designated as lectotype a specimen labeled "Charadra basifl ava Smith Type".However, Smith's (1908) statement does not qualify as a valid description under the provision of Article 12 of the ICZN (1999), and basifl ava is therefore an unavailable name (a conclusion apparently also reached by Poole 1989, as the taxon is not included in his publication).Th e lectotype designated by Todd (1982) is therefore not a true "type."Even if Smith's description is deemed to be valid, the type specimen is the illustration of C. pata in Druce (1894), not the specimen designated as lectotype by Todd (1982).

Charadra pata
Diagnosis.Th e wing markings of the female holotype, the only known specimen of this species, are most similar to those of C. oligarchia (only known from two males) and C. patafex.Compared to C. oligarchia, C. pata has a darker grey-brown forewing subterminal area with a contrasting white reniform area, but lacks any outline of a reniform (reniform outlined in oligarchia); the orbicular spot is slightly larger and more oblong in C. oligarchia.Compared to C. patafex, the forewing medial area of C. pata is contrastingly darker (concolorous with basal area in C. patafex) and the reniform area is white (brownish grey in C. patafex); also the hindwing marginal band is darker and narrower in C. pata than in C. patafex.
Distribution and biology.Known only from the type locality, Guatemala City, Guatemala.Nothing is known of the biology, although the larvae possibly feed on oak, as do those of C. tapa and C. franclemonti.
Remarks.After studying the type specimen of C. pata, we have come to the conclusion that this is not the same species as the Arizona taxon that has gone under this name, and belongs to a southern Mexican / Central American group of species consisting of C. pata, C. oligarchia, C. cakulha sp.n. and coyopa sp.n., here termed the oligarchia subgroup.Th e holotype female of C. pata (Fig. 1) diff ers from the tapa subgroup (C.franclemonti sp.n. and C. tapa sp.n.) in several key characters, namely the white, almost completely unmarked reniform area (grey and well marked with the usual markings in the tapa subgroup), a prominent and thick, well-defi ned black terminus of the subterminal line near the anal angle characteristic of the oligarchia subgroup (thinner, diff use and poorly defi ned in tapa subgroup), browner tone of the forewing ground colour (grey in tapa subgroup).Th e genitalic structure of the type female of C. pata diff ers from that of both C. franclemonti and C. tapa in that the antevaginal plate has short lobes, like C. tapa (long and prong-like in C. franclemonti), but with a more fl ared-out basal region than in either C. tapa or C. franclemonti, and the sclerotized lateral margins of the ductus bursae are nearly symmetrical, lacking the pronounced ventral twist of the right lateral margin of C. tapa (also nearly symmetrical in C. franclemonti).
Based on the brownish ground colour, prominent black mark of the anal angle and whitish reniform area, we place C. pata in the oligarchia subgroup.Th e lack of associated specimens of corresponding sexes is problematic, as it leaves the possibility that C. pata is the same species as C. oligarchia, C. patafex, C. cakulha or C. coyopa; a correlation in the structure of the male and female genitalia in this group is of some help, since asymmetrical placement or size of the male vesica cornuti corresponds to asymmetry in the shape and sclerotization of the ductus bursa (where the cornuti are presumably positioned during copulation: for example, in C. tapa, males have both cornuti positioned on the right, while females have a more heavily sclerotized, twisted right lateral margin of the ductus bursae).Th e nearly symmetrical ductus bursae of C. pata suggests a similar symmetrical placement and size of male cornuti, which would rule out C. oligarchia and C. patafex, (Figs 2, 3), a conclusion that also is supported by diff erences in wing markings.C. cakulha has both symmetrical placement and size of cornuti (Fig. 24), but diff ers markedly in wing markings (Fig. 7), as does C. coyopa (Fig. 10).
Diagnosis.Th e relatively uniform grayish-brown forewing pattern is similar to that of C. tapa and C. franclemonti, but the latter two species lack the bold, thick anal terminus of the subterminal line.Th e huge costal process of the male valve are unique in the yellow-winged Charadra species.
Distribution and biology.Known only from the type locality of Guerrero Mill, State of Hidalgo, Mexico at 9000 feet elevation.Possibly associated with dry oak woodlands at higher elevations, as are C. tapa and C. franclemonti.
Diagnosis.Superfi cially similar to C. cakulha and C. pata.Th e contrastingly pale outer third of the forewing is similar only to C. cakulha, but C. oligarchia is darker overall; internally, the male vesica has the two cornuti placed at the base of the vesica, one massive, one small, whereas C. cakulha has two lateral, symmetrical cornuti that are fused to the aedeagus.See also 'Diagnosis" of C. pata.
Distribution and biology.Known only from Guerrero Mill, Mexico.Possibly associated with dry oak woodlands at higher elevations, as in C. tapa and C. franclemonti.
Remarks.As discussed in the diagnosis of C. pata, C. oligarchia and C. pata may be the same species, but lack of specimens of each taxon of the corresponding sex prevents further comparison; however, the pata holotype diff ers slightly in wing markings, and the type localities of the two species are in diff erent mountain ranges separated by several hundred kilometers, so we treat both as distinct taxa until more study material becomes available.Etymology.We name this species after the late John G. Franclemont who collected and reared this species and recognized that there were two distinct species in southern Arizona.

Charadra franclemonti
Diagnosis.Superfi cially very similar to Charadra tapa, but can be separated by the forewing pattern and in particular by genital characters in both sexes.Th e outer half of the forewing in both sexes is darker and more patterned, with the outer forewing more even and paler grey in C. franclemonti than in C. tapa.Females can be separated without dissection by brushing the underside of the terminus of the abdomen and examining the sterigma and in particular the lamella antevaginalis, which extends caudally in two long prongs in C. franclemonti (often visible with the naked eye); in C. tapa these prongs are reduced to two blunt lobes.In males, the vesica of C. franclemonti, when infl ated, expands into a pouch as it exits the aedeagus, and the two cornuti are on opposite sides of the vesica; in C. tapa the vesica is not signifi cantly expanded where it exits the aedeagus and the two cornuti are together on one side.Th e valves of C. franclemonti are more truncated and squared off than those of C. tapa, which are slightly more produced and tapered.Although there is some overlap in fl ight periods, C. franclemonti primarily fl ies in July and August, whereas C. tapa fl ies in September and October.
Description.Sexes externally alike, except females slightly larger than males.FW length averaging 18 mm in males, 19 mm in females.Head -palps short, covered in stiff grey, black and white hair-like scales; proboscis well developed; eyes large, globular; frons with short grey and black hair; antennae broadly bipectinate, with longest rami about 7 times as long as width of shaft.Th orax -clothed in long dark-grey, black, and white scales; tegulae mostly white anteriorly, black and grey banded on posterior; Forewing dark grey with a brown-black medial band.Basal area a mix of black and white scales, appearing light grey, with white scales bleeding into medial area below orbicular producing a small pale streak.Antemedial line narrow, black, erratic and dragged outward below orbicular to meet or almost meet postmedial line midway.Orbicular spot prominent, oval, fi lled with dark blackish-brown scales and narrowly outlined in black.Medial band broad, brownish black, narrower on upper side and extending to upper margin above orbicular, but extending almost to wing base on lower half.Postmedial line black, narrow, erratic, pulled inward to meet or almost meet antemedial line midway.Postmedial line erratic, poorly defi ned, bordered outwardly with white scales that expand into a diff use whitish-grey reniform spot.Subterminal line poorly defi ned by dark scales, except more prominently lined with black at lower margin of wing and where it bends basad before meeting upper margin of wing.A diffuse patch of white scales at anal angle.Terminal line narrow, black, broken at veins.Fringe dark grey and black, faintly checked with lighter grey at veins.Veins beyond medial area narrowly lined with black.Hindwing pale yellow with an orange tint on basal half, sharply divided from grey black outer half.Fringe grey on inner half, white on outer half and lightly checkered with black between veins.Abdomen -clothed in lead grey hair-like scales mixed with numerous white scales at terminus, with a series of 3-4 small dark-grey tufts midway along dorsal centerline.Legs grey, banded with black at the joints.Male genitalia (Fig. 23) -Valves simple, subquadrate apically with a slightly pointed dorsal apex; costal process absent; clasper a simple scoop-shaped process about 1/10 length of dorsal valve margin, with broad surface directed dorso-cephalad; sacculus unmodifi ed; uncus with a wide base and narrowest medially, apex broadly  squared with a medial notch giving a slightly bifi d appearance; tegumen greatly expanded dorsally; saccus quadrate with broad U-shaped medial dorsal notch; aedeagus 5x longer than diameter, with a very wide, dorsally directed opening to vesica, opening about 1/3 total length of aedeagus; bulbous base of vesica directed at about 90 degrees to aedeagus; vesica base with two large, equal-sized, thorn-like cornuti positioned opposite each other, one ventral and one dorsal; one medial and one distal diverticulum of similar size, oriented dorsally.Female genitalia (Fig. 30 Description.Sexes externally alike, except females slightly larger than males.FW length averages 18 mm in males, 19 mm in females.Head, thorax, abdomen -as for C. franclemonti, with following diff erences in wing markings: Forewings slightly paler grey overall, particularly distal third; medial dark area slightly paler; hindwing with slightly paler yellowish base, dark marginal band on average slightly wider than in C. franclemonti.Male genitalia (Fig. 24) -structurally very similar to that of C. franclemonti, with following diff erences: valve tip slightly more triangular (more squared in C. franclemonti); base of vesica with two large, thorn-like cornuti positioned adjacent to each other on left-sublateral area.Female genitalia (Fig. 31) -as in C. franclemonti, with following diff erences: antevaginal plate with broad Ushaped medial notch and two short, broadly-rounded triangular lobes, not extending beyond margin of sternum; ductus bursae with a more pronounced ventral twist of right lateral margin.
Distribution and biology.Th e known distribution of C. tapa is limited to the Chiricahua, Huachuca, and Santa Rita Mountains of southeastern Arizona, although the species probably occurs in adjacent parts of Mexico.Th e main fl ight period is from September to October; a single specimen from early May indicates there may be spring fl ight.Larvae have been reared on Gambel Oak (Quercus gambelii).
Remarks.Although previously treated as such, neither C. tapa nor C. franclemonti is referable to C. pata, as pointed out in the 'remarks' section of C. pata.Etymology.A deity from Mayan mythology, Cakulha is the ruler of the lesser lightning bolts, and brother of Coyopa.It is a noun in apposition.

Charadra cakulha
Diagnosis.Externally recognizable by the very pale greyish-white outer third of the forewing and thorax, unique among the pata group.In addition to the wing markings, C. cakulha is distinguishable from the similar C. oligarchia by the lateral, symmetrical cornuti that are fused to the aedeagus (in C. oligarchia they are at the base of the vesica with one cornutus massive and one small).
Description.Female unknown.Forewing length 19.7 mm.Head, thorax and abdomen -as in C. franclemonti, diff ering in the following characters: vestiture much paler grey, nearly white; thorax with margin of tegulae and patagia bordered with black scales.Forewing light grey, distal third powdery whitish grey; reniform spot not discernible, reniform area entirely whitish; orbicular broadly oval, whitish with brown diffuse central area; subterminal line poorly defi ned medially by irregular white line, black scales at costal and anal margin, expanded to a thick, prominent line at anal margin.Male genitalia (Fig. 25) -Valves simple, subquadrate apically with a strongly incurved, scoop-shaped dorsocaudal apex; costal process absent; clasper a simple scoop-shaped process about 1/6 length of dorsal valve margin, with broad surface directed dorso- cephalad; sacculus unmodifi ed; uncus strongly constricted medially and apex bifi d, with an overall Y-shaped appearance; tegumen expanded dorsally; saccus V-shaped; juxta triangular; subscaphium strongly sclerotized; aedeagus 3 × longer than wide, with a very wide, dorsally directed opening to vesica, opening about 1/3 total length of aedeagus; vesica directed at about 90 degrees to aedeagus; cornuti usually placed on base of vesica and fused laterally to aedeagus, so aedeagus armed distolaterally with two stout, laterally projecting spines, left one placed slightly more apically; one medial and one distal diverticulum of similar size, oriented dorsally.
Distribution and biology.Known only from the two specimens of the type series, collected in mid-June at San Cristobal de las Casas, Chiapas, Mexico.Etymology.Th e name is derived from Mayan mythology.Coyopa is the ruler of the sound of thunder, and the brother of Cakulha.It is a noun in apposition.

Charadra coyopa
Diagnosis.Charadra coyopa is superfi cially most similar to C. oligarchia, but with an oblong rather than a round orbicular, and with a prominent pale patch (absent in C. oligarchia) below the orbicular.Th e male vesica has a single massive cornutus, unique in Charadra.
Description.Female unknown; forewing length 18.9 mm.Head, thorax and abdomen -as for C. franclemonti, diff ering in the following characters: prothoracic collar pale brown with a whitish-grey border; vestiture paler grey brown; forewings with basal two-thirds grey brown, distal third (beyond postmedial line) grey, terminal area grey brown; reniform spot yellowish white with oblong, hollow-centered medial line; orbicular elongate oval, yellowish white with diff use brown central scales; postmedial line distinct and serrate at veins; subterminal line diff use and brown, expanded to a thick black line at costal and anal margins, two apical black dashes along veins.Male genitalia (Fig. 24) -Valves simple, subquadrate apically with an incurved, scoop-shaped dorsocaudal apex, which appears irregular and somewhat spinulose; costal process absent; clasper a simple scoop-shaped process about 1/6 length of dorsal valve margin, with broad surface directed dorso-cephalad; sacculus unmodifi ed; uncus strongly constricted medially and apex bifi d, Y-shaped; tegumen expanded dorsally; saccus V-shaped; juxta triangular; subscaphium sclerotized and minutely scobinate; aedeagus 4.5 × longer than wide, with a very wide, dorsally directed opening to vesica, opening about 1/3 total length of aedeagus; vesica directed at about 90 degrees to aedeagus; a single, massive rose-thorn like cornutus positioned near base on left side, directed towards vesica base.
dens.Specimens from the Guadalupe Mountains of west Texas reported as C. ingenua by Blanchard and Franclemont (1981) are C. moneta sp.n. (see below).
Diagnosis.C. deridens can be identifi ed by it being the only Charadra species throughout its range, and is characterized by the typically monochromatic colouring of the forewing (lacking brown shades), and the black scaling in the orbicular spot (diff use and brownish in C. moneta).Charadra deridens may prove to be sympatric with C. moneta in New Mexico and western Texas, but the two can be distinguished by the more contrasting 'pupil' of the orbicular spot in deridens, and the white-grey rather than brownish forewing colour of moneta, as well as by the genital diff erences presented in the diagnosis of C. moneta.
Distribution and biology.Widely distributed, from Nova Scotia to British Columbia (not yet recorded from Alberta), south to Florida, Texas (Knudson and Bordelon 2004) and northeastern New Mexico (Raton, Colfax Co.).Th e larva, illustrated by Wagner (2005), feeds primarily on beech, oak, and white birch.
Remarks.Although quite consistent in appearance within a given geographic region, there is a moderate amount of variation in phenotypes across its range, and a melanic form (Fig. 16) occurs in the northeastern part of its range (Klots 1968).Th e palest specimens are from Atlantic Canada (Fig. 11, 15), particularly Nova Scotia.Kentucky specimens are on average smaller with a slight brown cast and reduced white scaling (Figs 14,18), whereas Colorado specimens are paler overall with less contrasting forewing markings (ingenua, Fig. 12).
Four specimens of C. deridens from Ontario and Kansas expressed three haplotypes, diff ering by less than 1%, and at least 2.5% divergent from three C. moneta samples.Etymology.During a discussion regarding the distinctness of this taxon compared to C. deridens, BCS bet GGA ten dollars that the DNA barcodes of C. moneta and C. deridens would be more than 1% divergent.Moneta is the Latin term for money.
Distribution and biology.Recorded from central and eastern Arizona (Coconino and Apache Cos.), the San Mateo Mountains of New Mexico, the Guadalupe Mountains of New Mexico and Texas, and the Big Bend region of Texas; south to the Sierra Madre in Nuevo Leon, northern Mexico.Collection dates range from March to June (Arizona, Texas, and New Mexico) and September (Mexico), possibly indicating two or more broods.Much of the type series was reared on Quercus gambelii.
Remarks.Specimens from New Mexico and Texas have a more smoothly-marked forewing and a less contrasting hindwing marginal band than those from Arizona, but are indistinguishable internally; the type series is therefore restricted to Arizona specimens.
Th ree barcoded specimens from NM exhibited a single haplotype, at least 2.5% divergent from the sampled C. deridens haplotypes.Diagnosis.A relatively small, pale silvery-grey species with few forewing markings that are largely confi ned to a narrow, black antemedial line and a prominent partial medial band from the costa to mid wing.Th e small size, pale colour, reduced markings, lack of central orbicular black scales, and black medial dash connecting the AM and PM lines allow this species to easily be distinguished.

Charadra dispulsa Morrison
Distribution and biology.Occurs from Texas southward and westward to at least San Luis Potosi, Mexico.Collection records range from March through May and July for southern Texas.Nothing is known of the larval stages or food plants, but larvae are possibly oak feeders.
Remarks.Barcode sequence for one specimen of C. dispulsa was available, which diff ered about 3.5% from C. deridens, 4% from C. moneta, and 6% from C. franclemonti.

Institutional collections are abbreviated as follows: AMNH American
Museum of Natural History, New York, New York, USA.BMNH Th e Natural History Museum (formerly British Museum [Natural History]), London, United Kingdom.
CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada.CUIC Cornell University Insect Collection, Ithaca, New York, USA.MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, USA.USNM National Museum of Natural History (formerly United States National Museum), Washington, D.C., USA UASM University of Alberta Strickland Museum, Edmonton, Alberta, Canada.