Neominois carmen A. Warren, Austin, Llorente, Luis & Vargas

Warren, Andrew D., Austin, George T., Llorente-Bousquets, Jorge E., Luis-Martínez, Armando & Vargas-Fernández, Isabel, 2008, A new species of Neominois from northeastern Mexico (Lepidoptera: Nymphalidae: Satyrinae), Zootaxa 1896, pp. 31-44 : 32-39

Neominois carmen A. Warren, Austin, Llorente, Luis & Vargas , NEW SPECIES

( Figs. 1 a–h, 3a–d, 4a–b, 5a–b)

Description. Male ( Figs. 1 a–d, 3a–b (right)): mean forewing length = 22.3 mm (range 20.0–23.6 mm, n=27); forewing costa slightly curved, apex rounded, termen straight to CuA2, then rounded to nearly straight anal margin; termen of hindwing convex; venation as illustrated for genus ( Holland 1916, Miller 1968); dorsum of both wings pale tan; forewing with medial and submedial area orange-tan; base and costa and outer margins heavily overscaled with brown; medial area may be lightly overscaled with brown; veins brown; submargin with elongate pale tan macules divided by brown veins in R5-M1, M1-M2, anterior 2/3 M2-M3, posterior 2/3 M3-CuA1, and CuA1-CuA2, shorter and more quadrate macule (may be divided horizontally by thin brown line) in CuA2-2A, all macules indistinctly outlined with brown, space between macules in M2-M3 and M3- CuA1 brown as outer margin; large and white-pupiled black ocellus within submarginal macules in M1-M2 and CuA1-CuA2, supernumerary ocelli with or without white pupils often within submarginal macule in M3-CuA1 (in R5-M1 on one specimen); terminal line dark brown; fringes brown. Hindwing heavily overscaled with brown extending distad to submarginal macules, may be less dense distad in discal cell and bases of M1-M2, M2-M3, and M3-CuA1, these regions may have slight tinge of orange; marginal area overscaled with brown, most dense apically; elongate submarginal macules contrastingly tan forming continuous band divided or not by brown veins from vein Rs to 2A and variably into 2A–3A without overscaling except lightly cephalad, distal margin of macules sharply angled cephalad, more rounded caudad, distinctly margined distad by dark brown; distal portion of submarginal macule in CuA1-CuA2 usually with black white-pupiled or not ocellus; terminal line dark brown; fringes brown.

Ventral forewing variably pale orange-tan proximad, becoming pale gray along costa and outer margin; gray-brown striations in base of discal cell and in its anterior 1/3 to 1/2 to its end, striations finer apically along costal and outer margins and in space between macules in M2-M3 and M3-CuA1; submarginal macules pale tan, all indistinctly outlined with brown, ocelli repeated from dorsum, but smaller; terminal line vaguely darker than ground color. Ventral hindwing pale tan; entire surface except submarginal area finely striated with gray-brown, may be denser in median area and especially proximad of submarginal macules; submarginal macules lightly overscaled or not, outlined distad with dark gray-brown, no ocellus; terminal line vaguely darker than ground color.

Dorsal head, thorax, and abdomen pale gray-brown; head white behind eyes; long tan scales on abdomen cephalad; eyes reddish; antennae gray above, whitish on venter with vague and thin black at segments, nudum pale reddish tan; palpi pale gray; ventral thorax and abdomen pale gray.

Genitalia ( Fig. 4 a–b): uncus in lateral view relatively broad, decurved slightly to a pointed and slightly hooked caudal end, tapering in dorsal view; tegumen broad in both lateral and dorsal views; appendix angularis prominent; subuncus thin, curved dorsad, tapering to sharply pointed caudal end; combined ventral arm of tegumen and dorsal arm of saccus thin, slightly sinuate; anterior arm of saccus broad in both lateral and ventral views, variable in length, often slightly curved to right in ventral view; valva tapering caudad to blunt caudal end, upcurved, variable fine serrations on dorsal and ventral edges caudad; aedeagus long (1/ 5X length of valva), thin, nearly straight; vesica with lightly sclerotized leaf-shaped cornutus.

Female ( Figs. 1 e–h, 3c–d (left)): mean forewing length = 26.2 mm (range 24.7–27.6 mm, n=7); wings broader and more rounded than on male, costa gradually curved, more so than on male, apex rounded, termen convex, curving evenly to straight anal margin, termen of hindwing strongly convex; venation similar to male; dorsum of both wings tawny-orange; forewing with base overscaled with gray-brown, diminishing in intensity distad; costal margin gray-brown exhibiting vague darker striations on basal half; outer margin overscaled with gray-brown cephalad of vein CuA1 or CuA2; submargin with pale and elongate ochreous-orange macules (paler cephalad) divided by brown veins in R4-R5 (may be absent), R5-M1, M1-M2, anterior 2/3 M2-M3, posterior 1/2 to 2/3 M3-CuA1, and CuA1-CuA2, shorter and more quadrate macule in CuA2-2A, all macules individually finely outlined with brown, most prominent on distal edges, space between macules in M2-M3 and M3- CuA1 overscaled with gray-brown; large and white-pupiled oval black ocellus within submarginal macules in M1-M2 and CuA1-CuA2; terminal line prominent, dark gray-brown extending thinly along full length of anal margin; fringes pale gray-brown. Hindwing overscaled extensively with pale gray-brown extending distad to submarginal macules, least dense distad in discal cell and bases of M1-M2, M2-M3, and M3-CuA1; marginal area overscaled with pale gray-brown, most densely adjacent to submarginal macules; elongate submarginal macules pale ochreous-orange as on forewing forming continuous band divided by brown veins from vein Rs to 2A and vaguely into 2A-3A, distal margin of macules sharply angled cephalad, more rounded caudad; inner margin of macules variably shaped; band outlined narrowly with brown both proximad and distad; distal portion of submarginal macule in CuA1-CuA2 with or without black white-pupiled or not ocellus; terminal line prominent, dark gray-brown; fringes pale gray-brown.

Ventral forewing tawny-orange, becoming pale gray along costal and outer margins; gray-brown striations in base of discal cell and in its anterior 1/3 to its end, along costal margin becoming finer apically, along outer margin, and in space between macules in M2-M3 and M3-CuA1; submarginal macules as on dorsum, those in R5-M1, M1-M2, and M2-M3 tan or with slight tinge of orange; macules all indistinctly outlined with brown; ocelli repeated from dorsum but smaller, terminal line distinct, brown. Ventral hindwing pale gray to pale tan with indistinct tinges of orange especially submarginally; entire surface striated by pale gray-brown densest as outline of often otherwise nearly indistinguishable submarginal macules; no ocellus; terminal line distinct, brown.

Dorsal head, thorax, and abdomen brown to gray-brown; head with white behind eye; long pale tan scales cephalad on abdomen; eyes reddish; antennae, palpi, and ventral thorax and abdomen as on male.

Genitalia ( Fig. 5 a–b): sterigma broad in ventral view (width about 2X length); largely lightly sclerotized or membranous; lamella antevaginalis with well-sclerotized central lobe, variably notched on caudal edge, (which is vertical in lateral view); antrum long, broad, lightly sclerotized, strongly folded on ventral surface; ductus bursae broad, membranous, expanding at junction with globular corpus bursae; no signa.

Types. Holotype male ( Figs. 1 a–b) with the following labels: / MEXICO: COAHUILA: / Mpio. Acuña: Maderas / del Carmen . Los Cohos [sic = Cojos] / Ridge. 7000–7400’ / N 28°56’21.5’’ W 102° / 36’01.8’’. 18- VI-2007 / Jim P. Brock /, to which is attached a red, printed label - / HOLOTYPE / Neominois carmen / A. Warren, Austin, Llorente, Luis & Vargas /. Allotype female ( Figs. 1 e–f) with the following labels: / MX, Coahulia, / Maderas del Carmen , / Los Cohos [sic = Cojos] Ridge, / 7000–7400’ / N28 51’ 21.5” / W102 36’ 01.8” / 18 June 2007 / leg. Jim P. Brock /, to which is attached a red, printed label - / ALLOTYPE / Neominois carmen / A. Warren, Austin, Llorente, Luis & Vargas /. Paratypes: same locality, date, and collector as holotype, 18 males, 8 females; MEXICO: COAHUILA: Mpio. Acuña: Maderas del Carmen : Vista Hermosa, 16- VI-2005, Bonnie Reynolds McKinney & Jonás Delgadillo Villalobos (1 male); MEXICO: COAHUILA: Mpio. Acuña: Maderas del Carmen : Casa Negro, 30-VI-2005, Bonnie Reynolds McKinney & Jonás Delgadillo Villalobos (1 male); MEXICO: COAHUILA: Mpio. Acuña: Maderas del Carmen, 2279m , UTM 733798 x 3203239, 6 -VI-2007, Bonnie Reynolds McKinney & Jonás Delgadillo Villalobos (7 males); MEXICO: COAHUILA: Mpio. Acuña: Maderas del Carmen, 2238m , UTM 733510 x 3203060, 6 -VI-2007 (1 male); MEXICO: COAHUILA: Mpio. Acuña: Maderas del Carmen, 2176m , UTM 733388 x 3202774, 17 -VI-2007, Bonnie Reynolds McKinney & Jonás Delgadillo Villalobos (1 male, 1 female); MEXICO: NUEVO LEÓN: 9 km E San Roberto, 1850m., 23-VII-2006, D. C. Robacker (1 female); MEXICO: NUEVO LEÓN: San Roberto, 6500’, 14-22-VII-2007, D. C. Robacker (1 male, 1 female). The holotype, allotype and most paratypes are deposited at the Museo de Zoología “Alfonso L. Herrera”, Facultad de Ciencias, Universidad Nacional Autónoma de México, México D.F., México; a pair of paratypes are at the McGuire Center for Lepidoptera and Biodiversity, and an additional pair temporarily remains with ADW.

Type locality. MEXICO: COAHUILA: Mpio. Acuña: Maderas del Carmen : Los Cojos Ridge, 7000– 7400’ (2134–2256 m), 28°56’21.5’’N 102°36’01.8’’W ( Fig. 3 e–f). Los Cojos Ridge is a high igneous ridge managed by CEMEX-Proyecto El Carmen .

Distribution and phenology. The type series of Neominois carmen suggests that the species occupies a rather extensive range in northeastern Mexico, at least from the Maderas del Carmen in northwestern Coahuila, to the area west of Linares in southern Nuevo León; these areas are separated by about 480 km. It is probable that N. carmen inhabits regions between northern Coahuila and southern Nuevo León, and perhaps even in southwestern Texas, although its overall range remains to be elucidated. Dates of capture of the type specimens indicate a single generation annually, from early June to late July.

Habitat. The Sierra Maderas del Carmen , a sky island with elevations from 600 to 2700 m encircled by low elevational Chihuahuan Desert, represents the major forested portion of the Sierra del Carmen . Its location in northwestern Coahuila, Mexico, is adjacent to the Big Bend region of southern Texas and only 60 km south of Big Bend National Park. The region encompasses a series of biotic associations from desert scrub at lower elevations to mesic montane forest, including Pseudotsuga Carr. and Abies P. Mill. , at the higher elevations ( Jiménez-G & Zuñiga-R 1991). The climate is temperate with average monthly temperatures ranging from 10C in winter to 32C in summer. Annual rainfall varies from negligible at low elevations to perhaps as much as 60 cm at higher elevations, principally from mid-summer to early autumn.

The part of Los Cojos Ridge in the Maderas del Carmen where N. carmen is most abundant is comprised of open areas along a dirt road at the beginning of the pine-oak woodland habitat ( Fig. 3 e–f). This area is dominated by the following plants: Quercus mohriana Buckl. ex Rydb. (Fagaceae) , Pinus cembroides Zucc (Pinaceae) , Juniperus deppeana Steud. (Cupressaceae) , Muhlenbergia emersleyi Va ss e y, Piptochaetium pringlei (Beal) Parodi (both Poaceae ), Lepidium montanum Nutt. (Brassicaceae) , Nolina erumpens (Torr.) S. Wats. , Agave spp. (both Agavaceae ), and Opuntia polyacantha var. polyacantha Haw. (Cataceae) . Other sites in the Maderas del Carmen inhabited by N. carmen include Casa Negro (UTM 734154 x 3203755), 2470m, and Vista Hermosa (UTM 731220 x 3198759), 1398m. The former site, Casa Negro, is at a slightly higher elevation than Los Cojos Ridge, and is dominated by Quercus spp. ( Fagaceae ), Juniperus monosperma (Engelm.) Sarg. , Juniperus flaccida Schltdl. (both Cupressaceae ), Pinus cembroides , Muhlenbergia emersleyi , and Panicum hallii var. hallii Vasey (latter two Poaceae ). The latter site, however, is significantly lower in elevation, at the beginning of the transition zone from low desert to foothill grasslands; Vista Hermosa is dominated by Yucca sp., Dasylirion leiophyllum Engelm. ex Trel. , Agave lechuguilla Torr. (all Agavaceae ), Opuntia engelmannii Salm-Dyck ex Engelm. (Cataceae) , Bouteloua curtipendula (Michx.) Torr. var. caespitosa Gould & Kapadic (Poaceae) , and Viguiera stenoloba Blake (Asteraceae) . The habitat for N. carmen east of San Roberto, Nuevo León, is dominated by “Pinyon pine, Juniper, cactus and grasses” (David Robacker, pers. comm. 2006).

Biogeography. Various authors have discussed biogeographic attributes of taxa occurring in the Sierra del Carmen (e.g., Miller 1955, Encina & Villareal 2002), as well as diverse aspects of their ecology and bioconservation (e.g., Ceballos et al. 1998, Hellgren et al. 2005, McCormack et al. 2005, Villalobos et al. 2005, Bhagabati & Horvath 2006, Poulus et al. 2007). Studies based on birds ( Miller 1955) and oaks ( Encina & Villareal 2002) have reached similar conclusions regarding biogeographic affinities of the faunal elements. While the Sierra del Carmen can be considered an extreme northern geological or biogeographical province of the Sierra Madre Oriental ( Villaseñor & Téllez-Valdés 2004), mesic habitats above 1500m represent a relictual, southern distributional limit for a number of nearctic taxa (including two mammals Sorex milleri Jackson (Soricidae) and Ammospermophilus interpres (Merriam) (Sciuridae) , and subspecies of another mammal Erethizon dorsatum L. ( Erethizontidae ) and a butterfly Limenitis weidemeyerii W. H. Edwards ( Nymphalidae )), and have strong faunal affinities to various Trans-Pecos ranges in western Texas ( Miller 1955, Encina & Villareal 2002) and higher ranges in northern Coahuila (e.g., Sierra de la Madera, Sierra Cruces).

A number of endemic or quasiendemic animals are known from the Sierra del Carmen , including the mammal Scalopus montanus Baker (Talpidae) , subspecies of two additional mammals, Odocoileus virginianus Zimmermann (Cervidae) and Tamias dorsalis Baird (Sciuridae) , and two snakes Natrix erythrogaster (Forster in Bossu) and Coluber constrictor L. (both Colubridae ) ( Kobelkowsky 2003). Encina & Villareal (2002) noted the Sierra del Carmen to be the most diverse region for Quercus in Coahuila, with 16 of 30 species known from the state. Thus, the Sierra del Carmen can be considered a biological island, a component of the “sierras coahuilenses,” which has served various taxa as a relictual distributional outpost.

While the butterfly genera Neominois and Gyrocheilus have been cited as relictual elements of the fauna of the Sierra del Carmen ( Kobelkowsky 2003) , both genera are also distributed far to the southeast in the northwestern Sierra Madre Oriental (data on northeastern Mexican Gyrocheilus will be presented in a subsequent study). The Lepidoptera of the entire region between the known distributional limits of Neominois carmen has been poorly studied, and more endemic taxa could well be found here with continued fieldwork and collections (especially between April and October).

Larval foodplant. Adults of Neominois carmen were found in openings dominated by Piptochaetium pringlei ( Fig. 3 f–h) at both Los Cojos Ridge and Casa Negro. While no oviposition events were witnessed, the close association of adults with this particular grass on Los Cojos Ridge strongly suggests that it is the local larval foodplant. No Piptochaetium was encountered at Vista Hermosa. Neominois was seen only once at this latter locale, indicating a possible stray from higher elevations, but if it breeds in the area around Vista Hermosa, it is possible that the species may use the locally common Bouteloua as a larval foodplant. The early stages are unknown.

Etymology. Neominois carmen is named for the Maderas del Carmen of northwestern Coahuila. This species has been known by the common name Joboni Satyr (see Warren et al. 2008), after Jonás A. Delgadillo Villalobos and Bonnie Reynolds McKinney, who collected part of the type series.

Figures

FIGURE 1. Types of Neominois carmen. Holotype male, dorsal (a) and ventral (b), from Mexico: Coahuila: Mpio. Acuña: Maderas del Carmen: Los Cojos Ridge, 7000 - 7400 ’, 18 - VI- 2007. Paratype male, dorsal (c) and ventral (d), from same locality and date as holotype. Allotype female, dorsal (e) and ventral (f), from same locality and date as holotype. Paratype female, dorsal (g) and ventral (h), from same locality and date as holotype.

FIGURE 3. Live adults and habitats of Neominois carmen. (a) Dorsal and (b) ventral view of male; (c) dorsal and (d) ventral view of female (same individual as Figs. 1 g-h), with male at right; habitat (e) from a distance, and (f) close up, showing Piptochaetium pringlei, a grass that probably serves as a larval foodplant; all from Mexico: Coahuila: Mpio. Acuña: Maderas del Carmen: Los Cojos Ridge, 7000 - 7400 ’, 18 - VI- 2007, by Jim P. Brock; (g) and (h) show additional views of Piptochaetium pringlei in the Maderas del Carmen, by Bonnie McKinney.

FIGURE 4. Male genitalia of Neominois. a) N. carmen — MEXICO: Coahuila; Maderas del Carmen (ADW # 08 - 21); b) N. carmen — MEXICO: Coahuila; Maderas del Carmen (ADW # 08 - 19); c) N. ridingsii — CANADA: Saskatchewan (GTA # 11579); d) N. ridingsii — USA: Nevada; Elko County (GTA # 11576).

FIGURE 5. Female genitalia of Neominois. a) N. carmen — MEXICO: Coahuila; Maderas del Carmen (GTA # 14022), ventral view; b) same, lateral view c) N. ridingsii — USA: Nevada; Esmeralda County (GTA # 11598), ventral view; d) same, lateral view.