ZooKeys 149: 5 (67 (201 1) A peer-revi iewed open-access journa I doi: 10.3897/zookeys. 149.1519 RESEARCH ARTICLE #7Z,00Ke y

www.zookeys.o rg Launched to accelerate biodiversity research

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of Elasmia mandela (Druce) from Texas and Oklahoma (Lepidoptera, Notodontidae, Nystaleinae)

Eric H. Metzler!*, Edward C. Knudson**

| Adjunct Curator of Lepidoptera, Michigan State University, PO. Box 45, Alamogordo, NM 88311-0045 USA 2. Texas Lepidoptera Survey, 8517 Burkhart Rd., Houston, TX 77055-7517 USA

T urn:lsid:zoobank. org:author:BBF4EAD8-74F4-455D-B28E-EFA8015EC34C * urn:lsid:zoobank. org:author:84095B59-A263-4E 1A-8FF2-987A8 14E3210

Corresponding author: Eric H. Metzler (metzlere@msu.edu)

Academic editor: Donald Lafontaine | Received 13 May 2011 | Accepted 16 August 2011 | Published 24 November 2011 urn:lsid:zoobank. org:pub:D1914106-7804-49A0-9AA0-9C9288 98 E44F

Citation: Metzler EH, Knudson EC (2011) A new species of Elasmia Méschler from New Mexico and Texas, and a new subspecies of Elasmia mandela (Druce) from Texas and Oklahoma (Lepidoptera, Notodontidae, Nystaleinae). In: Schmidt BC, Lafontaine JD (Eds) Contributions to the systematics of New World macro-moths II. ZooKeys 149: 51-67. doi: 10.3897/zookeys. 149.1519

Abstract

Hippia packardii (Morrison) and Hippia insularis (Grote) are moved to the genus Elasmia Méschler as comb. n. Elasmia cave Metzler, sp. n. is described from New Mexico and Texas, and Elasmia mandela san- taana Metzler & Knudson, ssp. n. is described from Texas and Oklahoma. A key to the species of Elasmia of southwestern U.S. is provided. Adult male and female moths of Elasmia from southwestern U.S. and

their genitalia are illustrated.

Keywords Lepidoptera, Notodontidae, Nystaleinae, Arizona, Oklahoma, New Mexico, Texas, Kansas, Hippia, Elas- mia, Carlsbad Caverns National Park, Santa Ana National Wildlife Refuge

Copyright E.H. Metzler, E.C. Knudson. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

52 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

Introduction

Lafontaine and Schmidt (2010) listed two species of Hippia Méschler, 1878 (Noto- dontidae: Nystaleinae) for North America north of Mexico: H. packardii (Morrison, 1875), described from Texas; and H. insularis (Grote, 1866), described from Cuba. They listed no species of Elasmia Moschler, 1886 (Notodontidae: Nystaleinae). Our investigations show that 1. packardii and H. insularis belong in the genus Elasmia, and that 7. insularis, reported from Texas (Knudson and Bordelon 1999), is an error. United States specimens thought to be H. insularis instead represent an undescribed species described here as F. cave. Our investigations further show an undescribed sub- species of Elasmia mandela (Druce, 1887) from the United States. Elasmia mandela santaana is described here from Texas and Oklahoma.

Methods

Adult moths were collected in U.S.D.A. type black-light traps and at black light and sheet as described in Covell (1984).

Genitalia were examined following procedures outlined in Clarke (1941), Hard- wick (1950), Lafontaine (2004), and Pogue (2002). Abdomens were removed, wetted in 95% ethyl alcohol, and soaked in 10% KOH for 1.5 hours at 50°C. Genitalia were dissected in 5% ethyl alcohol, stained with Safranin O in ethyl alcohol and Chlorozol Black in water, dehydrated in 100% ethyl alcohol, cleared in oil of cloves, rinsed in xylene, and slide mounted in Canada balsam.

The aedeagus of species of Elasmia is held firmly in place by membranes within the genital capsule, and the aedeagus is nearly always broken into two pieces during the process of removal. The anterior portion is short and abruptly flared out. The posterior portion with the everted vesica is illustrated in this paper (Figs 13, 16, 19).

Wing pattern terminology came from Lafontaine (1987, 2004) and Mikkola et al. (2009). Morphological structure terminology came from Common (1990) and Forbes (1954), Genital structure terminology came from Lafontaine (1987, 2004), Franclemont (1946), Forbes (1954), Klots (1970), Miller (1991), and Weller (1995). Forewing lengths, from the base to the apex excluding fringe, were measured to the nearest mm, using a stereo-microscope. Nearly all specimens from New Mexico were collected as part of a long-term faunal study of Lepidoptera at Carlsbad Caverns National Park.

Specimens of Lepidoptera from this study are deposited in the following collec- tions:

AMNH American Museum of Natural History, New York, New York BMNH Natural History Museum, London, England CMNH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 53

CUIC EHM JBS JBWw MSU

MCZ ORU OSU TAM TLSRC UFL UNM

USNM

Results

Cornell University, Ithaca, New York

Eric H. Metzler, Alamogordo, New Mexico, for subsequent transfer to MSU J. Bolling Sullivan, Beaufort, North Carolina

J. Bruce Walsh, Tucson, Arizona

Albert J. Cook Arthropod Research Collection, Department of Entomology, Michigan State University, East Lansing, Michigan

Museum of Comparative Zoology, Harvard University, Cambridge, Massa- chusetts

Oral Roberts University, Tulsa, Oklahoma

Oklahoma State University, Stillwater, Oklahoma

Texas A&M University, College Station, Texas

Edward C. Knudson, Texas Lepidoptera Survey, Houston, Texas

McGuire Center for Lepidoptera and Biodiversity, University of Florida, Gainesville, Florida

Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico

United States Museum of Natural History (Smithsonian Institution) Wash- ington, DC

Key to the species of Elasmia in Arizona, Kansas, New Mexico, Oklahoma, and

Texas

Forewing gray and/or blue gray, sides of uncus convex, evenly curved (Fig. 11), forewing length: VST ANY o ceya spss ibsviccaics sumed Myitiea he shonyd eat sy adhe packardii Forewing brown or gray, sides of uncus flared outward (Figs 14, 17), forewing lenient WAS EG ity casan ca teeavaseimn ee cpertnaned a oanienrntneavahadeneta danas duaninenneent eenaadttedt 2 Forewing brown gray (Fig. 5), apex of costulae swollen and bent........... cave Forewing gray brown (Fig. 9), apex of costulae straight and not bent............ nah cahad ea ec led eattbed a cad aida ites daetaeh Aaah bse decade RelA oases mandela santaana

In south central Texas, larvae of the genus Elasmia (not identified to species) feed on Ungnadia speciosa Endl. (Mexican buckeye) and Sapindus saponaria var. drummon- dit (Hook. & Arn.) L. Benson (soapberry tree) (both Sapindaceae) (Val Bugh pers. comm. 2010).

54 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

Systematics

Hippia Méschler, 1878

Discussion. We examined the illustration of the type and the male and female genita- lia of Hippia mumetes (Cramer, [1775]), the type species of Hippia Méschler (1878). Those examinations show that the North American species, placed in Hippia, are not congeneric with H. mumetes.

Elasmia Méschler, 1886

Discussion. We examined the illustration of the type of Elasmia lignosa Moschler, 1886, the type species of Elasmia. The North American species, previously placed in Hippia, are determined to be congeneric with Elasmia

Elasmia insularis (Grote, 1866), comb. n.

Discussion. ‘The adult and genitalia of E. insularis were illustrated in Torre and Alayo (1959). We examined male specimens, and their genitalia, of E. insularis from Cuba. Elasmia insularis is not known to occur in Florida (Heppner 2003), and it is doubtful that it occurs in the U.S. Inclusion of E. insularis in Lafontaine and Schmidt (2010) was based on erroneous reports from Texas (Knudson and Bordelon 1999).

Elasmia packardii (Morrison, 1875), comb. n. http://species-id.net/wiki/Elasmia_packardii Figs 1-4, 11-13, 20, 23, 26

Description. Overall color light gray blue to gray with obscure transverse forewing markings, sometimes showing slight brownish shadings over reniform spot and in postmedial and subterminal areas. Males and females similar in appearance; male an- tenna narrowly bipectinate in basal 34, with dense setae on ventral surface. Female antenna filiform for entire length, with sparse setae. Apex of forewing marked with a diagonal white and dark shade. Forewing length in males 12-15 mm (mean = 14 mm, n = 72), and in females 13-16 mm (mean = 14 mm, n = 25). Male genitalia (Figs 11-13) distinguished by a helmet-shaped uncus with gradually widening sides. Female genitalia (Fig. 20) with membranous papilla anales partially hidden from view. Ductus bursae broad and short, dorso-ventrally compressed; corpus bursae, round in profile, with a single shark tooth-shaped signum, also with a heavily sclerotized, perpendicular, thumb-like projection ventrally and a sclerotized finger-like pocket appressed to corpus bursae dorsally. Deciduous cornuti from male vesica may be found in corpus bursae.

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 55

PEERETECTEPP TEP EP aT

Figures |-6. Elasmia adults. | FE. packardii female holotype 2 E. packardii holotype labels 3 E. packardii male 4 Elasmia packardii female 5 Elasmia cave male holotype 6 Elasmia cave female paratype.

Remarks. Morrison (1875) described Elasmia packardii from Waco, Bosque County, Texas (Fig. 2) based on a single female specimen (Fig. 1). Adults are on the wing from April through early October.

Distribution and Biology. Elasmia packardii occurs in Texas, Arizona, New Mex- ico, Oklahoma, and Kansas (Fig. 26); it is common at Carlsbad Caverns National Park. Its distribution in Mexico is unknown. ‘The larvae feed on Ungnadia speciosa Endl. (Mexican buckeye) (R.O. Kendall specimens in TAM) and Sapindus saponaria var. drummondii (Hook. & Arn.) L. Benson (soapberry tree) (both Sapindaceae) (R.O. Kendall specimens in AMNH and TAM).

56 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

Figures 7-10. Elasmia adults. 7 E. mandela male 8 E. mandela female 9 E. m. santaana male holotype 10 £. m. santaana female paratype.

Elasmia cave Metzler, sp. n. urn:lsid:zoobank.org:act:93E887C7-757F-4EC6-B5B5-5E4012D9822A http://species-id.net/wiki/Elasmia_cave

Figs 5—6, 14-16, 21, 24, 27

Type material. Holotype male: “USA: NM: Eddy Co. Carlsbad Caverns N[ational] Park], riparian habitat, 32°06.566' N 104°28.257' W, 29 August 2006, Eric H. Metzler, CCNP4, uv trp Accsn #: CAVE - 02263", “HOLOTYPE USNM Elasmia cave Metzler” [red handwritten label] (USNM). Paratypes: 19 males; 14 females: NEW MEXICO: USA: NM: Eddy Co. Carlsbad Caverns NP, arroyo habitat 32°05.98'N 104°33.57'W, 5 September 2010, Eric H. Metzler, CCNP2 uv trp Accssn #: CAVE - 02263. TEXAS: Alpine, Tex., 1-7 May 1926, 8-14 May 1926, 1-7 July 1926, 8-14 July 1926, 15-21 July 1926, 1-7 Aug. 1926, 15-21 Aug. 1926, O.C. Poling, Coll[ector]. Barnes Col- lection (USNM), Texas, Uvalde Co. Concan, 12-V-90, leg. E.C. Knudson. 14-X-93, Concan, Uvalde Co., TX, Coll C. Bordelon. TX: Brewster Co., Big Bend N.P., Green Gulch/5400' 5—7-V-97/ECK. Big Bend, Tex. Brewster Co., 6—7000 ft., Poling, F John- son donor, 8-1-26. USA: Texas: Jeff Davis Co. Davis Mountains, Limpia Canyon, elev: 4920', 30°30.0'N 103°52.5'W, 8 August 1991, Eric H. Metzler. TEXAS: Jeff Davis Co., 25-V1-81, Davis Mt. St. Pk., Jeff Davis Co. TX: Ft. Davis, 3-x-94, leg. E. Knud- son. 24 Aug 1995. Jeff Davis Co. Texas, 5-V-78, Kokernaut Creek, leg. E.C. Knudson.

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 57

costulae

Figures | 1-19. Elasmia male genitalia. 11 E. packardii male genitalia genital capsule slide E.H.M. 343 12 £. packardii male genitalia detail of terminus of costulae slide E.H.M. 343 13 E. packardii male geni- talia aedeagus slide E.H.M. 343 14 E. cave male genitalia paratype genital capsule slide E-H.M. 355 15 £. cave male genitalia detail of terminus of costulae slide E.H.M. 355 16 £. cave male genitalia paratype aedeagus slide E.H.M. 355 17 E. m. santaana male genitalia paratype genital capsule slide E.H.M. 359 18 £. m. santaana male genitalia detail of terminus of costulae slide E.H.M. 359 19 E£. m. santaana male genitalia paratype aedeagus slide E.H.M. 359.

58 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

ny | |

24

Figures 20-25. Elasmia female genitalia and male eighth sternites. 20 E. packardii female genitalia slide E.H.M. 347 21 E. cave female genitalia paratype slide EH.M. 409 22 E. m. santaana female genitalia paratype slide E.H.M. 410 23 Eighth sternite of male of E. packardii slide E.H.M. 343 24 Eighth sternite of male of E. cave paratype slide E.H.M. 355 25 Eighth sternite of male of E. m. santaana paratype slide Ele. Mio 59:

Jeff Davis Co., TX, Ft. Davis, 10,11-[X-10 Bordelon & Knudson coll. Jeff Davis Co., TX, Ft. Davis, 24-26-V-07 Bordelon & Knudson coll. TX: Culberson Co., Guadalupe Mts. N.P., Lamar Cyn., Coll. C. Bordelon. TEXAS: Culberson Co., Guadalupe Mts. N.P., Pine Spring, 6-8-IX-91, leg. E.C. Knudson. Green Gulch 5400' Big Bend Natl. Park Brewster Co., Texas 4 May 1972 J. G. Franclemont ¢ Genitalia slide 6419 J. G. Franclemont. Alpine, Brewster Co. Texas 15-21 Aug. 1926 O.C. Poling CS Genitalia slide 2535 J. G. Franclemont. (EHM, MSU, CUC, TLSRC, USNM).

Etymology. CAVE is the acronym, used by the U.S. National Park Service, for Carlsbad Caverns National Park. The specific name of this species, cave, treated as a noun in apposition, refers to the type locality, Carlsbad Caverns National Park.

Diagnosis. Elasmia cave is a dark brown-gray moth with obscure transverse markings. Elasmia cave looks like brown example of E. mandela; E. mandela is dark

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 59

gray brown. The brown color of the imago and its genitalia will separate E. cave from E. packardii, which is gray blue to gray. The uncus of £. packardii (Fig. 11), narrow at its apex, gradually widens with evenly curved sides. The uncus of E. cave (Fig. 14) is wide, like a manta ray, and narrows immediately before the apex. The distal end of the costulae of F. insularis, n=3, are narrow, straight or slightly sinuous, and without bend or swelling apically (illustrated by Torre and Alayo 1959); the costulae of E. mandela, n=3, are nearly identical to E. insularis. In comparison to E. insularis and E. mandela, the costulae of E. cave (Fig. 15) are broader, and they are abruptly bent upward and swollen at the distal end.

Description. Adult male (Fig. 5): Head: smoky gray, scales strap-like, erect, a fuscous line between eyes below antennae. Labial palpus erect, extending upward to slightly beyond base of antenna, smoky brown gray with a fuscous lateral stripe, extends to slightly beyond base of antenna, ventral scales on Ist and 2nd segments long, not shaggy, 3rd segment closely scaled. Haustellum coiled between labial palpi. Antenna narrowly bipectinate for basal 3/4, each ramus tipped with long setae, api- cal 1/4 ciliate with short setae, dorsal surface alternating fuscous and smoky, close- ly scaled, ventral surface naked, brown. Thorax: collar black, sometimes preceded by brown, dorsum smoky brown gray, longitudinal narrow black lines anteriorly, posteriorly, and laterally, tegula smoky brown gray, scales strap-like; underside smoky dark gray brown, laterally smoky, scales erect long hair-like or narrowly forked. Legs: dark smoky gray brown, closely scaled, except lateral margin with shaggy scales, tar- somere apex yellow. Forewing: length 14-18 mm, mean 16 mm, n = 16; dorsal surface ground color smoky gray brown, sometimes slightly hoary; antemedial line obscure, pale basally, black mesally, sinuous; postmedial line vague, sinuous, black basally outer element pale; subterminal line a series of fuscous bars; terminal line narrow, black; orbicular spot absent or vaguely pale; reniform spot inconspicuous, dark with pale outline; costa brown except white shade at apex; subreniform spot fuscous, contrasting; dark line from apex running obliquely toward reniform spot; fringe smoky gray. Ventral surface: smoky dark gray black, apical markings similar to dorsal surface, fringe concolorous. Hindwing dorsal surface: dark smoky gray, slightly paler basally, markings absent, fringe pale smoky. Ventral surface: apex to tornus dark smoky gray, tornus pale gray along inner margin, base pale gray, mark- ings absent, fringe pale gray. Abdomen: dorsum smoky gray, with fuscous tufts on first and second segments, elsewhere closely scaled, underside pale smoky gray. Genitalia (Fig. 14): Uncus broad, flattened, setose, apex bluntly pointed, dorsally with nar- row ridge, ventrally with two short cornutus-like spines; socii broad, setose, bent at approximately 90°, with one ear-like dorsal projection; tegumen flattened; saccus short, broadly U-shaped; juxta shield shaped, dorsal margin a half circular cutout; diaphragma bearing two sclerotized processes (costulae) near bases of valvae costa, bent at 90°, bent and swollen club-like apically (Fig. 15), valve setose, dorsally scle- rotized, ventrally membranous, Barth’s Organ large, with many chevron-shaped par- allel pleats, cucullus not well differentiated, with three narrow, curved ridges, corona with weak, mesally-directed curved setae. Aedeagus (Fig. 16) straight, anterior end

60 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

abruptly flared out, posteriorly flattened, spoon-shaped; vesica lightly sclerotized, subbasal diverticulum with a nipple-shaped cornutus; a patch of deciduous (may be dislodged during mating) stellate (like a starfish) spicule-shaped cornuti; basal diver- ticula lightly sclerotized, with two finger-like subbasal diverticulae.

Adult female (Fig. 6). Similar to male except; antenna filiform, without long se- tae, top of head yellow to orange, collar to disc of thorax yellow to orange. Forewing length 15-18 mm, mean 17 mm, n = 11. Genitalia (Fig. 21). Papilla anales mem- branous, setose, partially hidden from view between sclerotized extensions of ninth abdominal segment. Posterior apophyses slender. Anterior apophyses slender. Ductus bursae short, broad. Corpus bursae round, with a single shark tooth-shaped signum; sclerotized ventral wall forming a thumb-like extension with bulbous terminus; scle- rotized dorsal wall with a pock-marked, finger-like extension appressed to surface of corpus bursae.

Remarks. Elasmia cave was mistakenly identified in the U.S. as E. insularis. The costulae of the male genitalia, Figured in (Torre and Alayo 1959) from Cuba and noted in the diagnosis, separate the species. Elasmia cave is placed in the genus Flas- mia Moschler, 1886, because the imago is closely similar to E. /ignosa, and the male genitalia are closely similar to those of E. mandela.

Distribution and biology. Elasmia cave occurs in the U.S. in New Mexico and Texas; its distribution in Mexico is not known. Three specimens were collected in riparian habitats in Texas and New Mexico. Two specimens from Alpine, Texas and one from Big Bend, Texas, leg. Poling, have additional handwritten labels that say “Buckeye” or “bred Buckeye” respectively. The type locality was selected because it will be protected by the U.S. National Park Service into perpetuity.

Elasmia mandela (Druce, 1887) http://species-id.net/wiki/Elasmia_mandela Figs 7, 8

Description. Overall color dark gray brown with obscure transverse forewing mark- ings. Males and females similar in appearance. Male antenna narrowly bipectinate in basal 34, with dense setae on ventral surface. Female antenna filiform for entire length, with sparse setae. Apex of forewing with a diagonal white mark. Reniform spot outlined with pale-orange scales, not contrasting. Forewing length in males 17.0-18.0 mm (mean = 17.2 mm, n = 5), and in females 18.0—20.0 mm (mean = 19.3 mm, n = 7). Male genitalia distinguished by uncus with abruptly widening sides, like a manta ray, and robust saccular area (Barth’s Organ). Female genitalia with membranous papilla anales that are partially hidden from view. Ductus bursae broad and short, dorso-ventrally compressed; corpus bursae round in profile, with a single shark tooth shaped signum, also with a heavily-sclerotized, perpendicular, thumb-like projection ventrally and a sclerotized finger-like pocket appressed to cor- pus bursae dorsally.

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 61

Remarks. Druce (1887) described Elasmia mandela from Presidio, Mexico, based ona single female specimen. We examined a photograph of the type and it’s genitalia. We also examined specimens from Vera Cruz and Yucatan, Mexico (AMNH), and from Costa Rica (JBS).

Distribution and biology. Elasmia mandela occurs in Mexico and Costa Rica. Its distribution in other Central American countries is unknown. The larval hosts in Costa Rica are one species of Rhamnaceae and 22 species of Sapindaceae (Janzen and

Hallwachs 2009).

Elasmia mandela santaana Metzler & Knudson, subsp. n. urn:lsid:zoobank.org:act:9B30138B-9B92-4B8 1-9797-82B59ABE417F http://species-id.net/wiki/Elasmia_mandela_santaana

Figs 9, 10, 17-19, 22, 25, 28

Type material. Holotype male: “Hidalgo Co. Texas 31-X-83 Santa Ana Refuge leg. E.C. Knudson” “HOLOTYPE USNM Elasmia mandela santaana Metzler & Knud- son” [red handwritten label] (USNM). Paratypes: 13 males; 10 females: TEXAS: Harris Co: TX. Houston, Leg. E.C. Knudson, 9-VIII-75. Hidalgo Co. TX. Santa Ana NWR, 6-IX-92, leg. E.C. Knudson. Texas: Uvalde Co. Concan, 15-V-10, B/K. Tarrant Co. Texas Benbrook, 30-IV-78, leg. E.C. Knudson. Terrel Co. Tex. Sanderson, 25-IV-81, leg. E.C. Knudson. Kerrville. Texas, Barnes Collection. Kerrville, Texas. VIII 1904. Kerrville, TX. H. Lacy Collector. Kerrville, 4-23-08, TX. EC. Pratt Collector. Texas, San Patricio Co. Welder Wildlife Refuge near Sinton, Texas, 14-16-V1-85, leg. E.C. Knudson. Hidalgo Co., TX, Bentsen State Park, 6-VHI-94, E. Knudson coll. Mont- gomery Co. Tex. Sawdust Rd. & LS. 45, leg. E.C. Knudson, 20-VI-75. Brownsville, Tex HI-10-29, RH. Benjamin collr, Barnes Collection. Alpine, TX. 8-14 May, 8-14 July, 15-21 Aug. 1926, O.C. Poling, Coll[ector]. TEXAS: Smith Cany., Guadalupe Mountains, Culberson Co., 5750’ May 22, 1973, Douglas C. Ferguson. OK: Caddo Co. Methodist Youth Camp 1 October 1994 J.M. Nelson Coll. OK: Tulsa Co. Sand Springs 145" & W. 19" St. Aug 27 - Sept 1, 2008 J.F. Fisher, Collr. at black light. OK: Tulsa Co. Sand Springs 145 & W. 19" St. September 12, 2008 J.F. Fisher, Collr. at black light. (TLSRC, ORU, USNM).

Etymology. The name of this subspecies, santaana, refers to its type locality, Santa Ana National Wildlife Refuge in Texas. The name is treated as a noun in ap- position.

Diagnosis. Elasmia mandela santaana is gray overall. The adult resembles a large example of E. packardii; E. m. santaana has a contrasting dark scale patch in the re- niform/subreniform area. Elasmia m. santaana (mean forewing length = 16 mm) is larger than E. packardii (mean forewing length = 14 mm) and E£. m. santaana’s Barth's Organ is relatively larger. The lateral margin of the uncus in E. packardii has a slight shoulder immediately below the apex, whereas in E. m. santaana the lateral margin of the uncus is flared outward. Elasmia m. santaana is a gray moth, and E. cave is a

62 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

brown moth. The male and female genitalia of E. m. santaana are similar to those of E. cave. The costulae of £. cave are abruptly bent and swollen apically (Fig. 15); the costulae of E. m. santaana may be slightly swollen but not bent apically (Fig. 18).

Description. Adult male (Fig. 9): Head: smoky gray, scales strap-like, erect, a vague fuscous shade between eyes below antennae, a vague fuscous shade behind antennae. Labial palpus erect, extending to base of antenna, smoky brown gray with two dark-brown lateral lines, ventral scales on 1st and 2nd segments long, not shaggy, 3rd segment closely scaled. Haustellum coiled between labial palpi. Antenna nar- rowly bipectinate for basal 3/4, each ramus tipped with long setae, apical 1/4 ciliate, with short setae, dorsal surface smoky, closely scaled, ventral surface naked. Thorax: pale brown behind head; collar narrow, black, dorsum smoky with blackish brown- tipped scales on disc; tegula pale smoky, edged with black scales, scales strap-like; underside dark smoky gray with pale-tipped scales, smoky laterally, hair-like. Legs: smoky dark gray brown, closely scaled, each segment and each tarsomere apex ringed with pale. Forewing: length 15-17 mm, mean 16 mm, n = 7. Dorsal surface ground color smoky gray; basal line pale at costa, finely lined with black; antemedial line pale, sinuous, finely lined with black; postmedial line sinuous, pale, finely lined with black; subterminal line a series of fuscous black bars; terminal line a fine fuscous line; orbicular spot inconspicuous; reniform spot a black bar outlined with pale, dark blackish shade in lower part; subreniform spot contrasting black and fuscous; costa apex pale gray tan to white; dark line with dark shade from apex oblique to subreni- form spot. Ventral surface smoky; terminal line black; fringe smoky. Hind wing. Dorsal surface ground color smoky gray, darker distally; fringe pale. Ventral surface ground color smoky, with scattered fuscous scales; fringe smoky. Abdomen: smoky, basal tuft blackish, underside smoky. Genitalia (Fig. 17): Uncus flattened, flared out- ward laterally, with narrow shoulders immediately below apex, apex setose, pointed; socii large, setose, bent at approximately 90°, each arm with an ear-like ridge; tegu- men flattened; saccus U-shaped, short; juxta shield shaped, dorsal margin a half cir- cular cutout; diaphragma bearing two sclerotized processes (costulae) near bases of valvae costa, bent at 90°, apex slightly swollen (Fig. 18); valve setose, sclerotized dor- sally, membranous ventrally, Barth’s Organ robust, with numerous chevron-shaped parallel pleats, cucullus poorly defined with three narrow curved ridges, corona with weak, mesally-directed, curved setae. Aedeagus (Fig. 19) straight, abruptly flared out anteriorly, flattened, spoon shaped; vesica lightly sclerotized, with a patch of de- ciduous stellate (like a starfish) spicule-shaped cornuti; subbasal diverticulum with a nipple-shaped cornutus; apex lightly sclerotized, one large basal diverticulum with two subbasal lobes.

Adult female (Fig. 10). Similar to male except; antenna filiform without long setae; top of head yellow to orange; collar to disc of thorax yellow to orange. Fore-

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 63

wing length = 16-18 mm, mean 17 mm, n = 9. Genitalia (Fig. 22). Papillae anales membranous, setose, hidden from view between sclerotized extensions of ninth ab- dominal segment; posterior apophyses slender; anterior apophyses slender; ductus bursae short, broad; corpus bursae round, with a single shark tooth-shaped signum; dorsal wall of corpus bursae sclerotized, forming a thumb-like extension, without bulbous terminus; sclerotized ventral wall with a pock-marked, finger-like extension appressed to surface of corpus bursae.

Remarks. We make this a subspecies of FE. mandela because the color of the forewings is different from E. m. mandela, it is slightly smaller, and it is geographi- cally separated from E. mandela. The male and female genitalia, however, are indis- tinguishable from those of E. m. mandela. Some specimens from Oklahoma were previously misidentified as EF. insularis.

Distribution and biology. In the U.S., E. m. santaana has been recorded from Texas and Oklahoma; its distribution in Mexico is unknown. A larval host (R. O. Kendall specimens in TAM) is Unganadia speciosa Endl. (Mexican buckeye) (Sapin- daceae). The type locality was selected because the U.S. Fish and Wildlife Service will protect it into perpetuity.

Figure 26. Distribution map for Elasmia packardii in United States.

64 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

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Figure 28. Distribution map for Elasmia mandela santaana in United States.

A new species of Elasmia Moschler from New Mexico and Texas, and a new subspecies of... 65

Discussion

For all characters, except those we noted in the key and diagnoses, the species are closely similar in appearance.

The details of the shape of the costulae, in combination with the color and size of the adults, is important in defining the species. The costulae can be slightly variable within a species, thus all the characters should be consulted in making an identifica- tion.

The female genitalia of E. m. mandela, E. m. santaana, E. insularis, E. packardii, and FE. cave are nearly identical. The male genitalia of E. packardii are distinct. The differences between the male genitalia of E. mandela, E. insularis, and E. cave are more subtle; the most reliable character we found was the shape of the terminal por- tion of the costulae (Figs 12, 15, 18). The costulae of EF. insularis and E. mandela are closely similar; the superficial appearance of the adults are different. The costulae of E. m. santaana and E. cave are similar (see the key and Figs 15 and 18 for differences); most adults can be identified by external appearance, but a few specimens require examination of the male genitalia for positive identification.

The specimens from Carlsbad Caverns National Park were collected by Metzler as part of a 10-year study of the Lepidoptera of the Park initiated by the Park in 2006. This is the second in a series of papers (Metzler et al. 2010) detailing the moths of Carlsbad Caverns National Park.

Acknowledgments

The National Parks Foundation, Washington, DC, the Adopt-A-Bat program, admin- istered by the Carlsbad Caverns-Guadalupe Mountain Association, Carlsbad, NM, and the Chihuahuan Desert Conservation Alliance, Carlsbad, NM all contributed travel funding for the study of Lepidoptera at Carlsbad Caverns National Park. We are grateful for their financial support. Renée West, Supervisory Biologist, Carlsbad Caverns National Park was especially helpful with logistics. Several persons made spec- imens or photographs or data available for study from their institutions or personal collections; James K. Adams, Don C. Arnold, Val Bugh, Charles V. Covell, Jr., Mark Dreiling, John F Fisher, E. Richard Hoebeke, Alan Jenkins, Hugo L. Kons, Jr., Jon A. Lewis, James K. Liebherr, John M. Nelson, John D. Oswald, Michael G. Pogue, Suzanne Rab Green, John E. Rawlins, Edward G. Riley, J. Bolling Sullivan, David L. Wagner, Andrew D. Warren, and Catherine Weisel. Philip D. Perkins, MCZ, made the type of Elasmia packardii available for study. The Museum of Comparative Zool- ogy, Harvard University, provided the illustration of the type of E. packardii for use. Alessandro Giusti, Natural History Museum, London, provided a photograph of the genitalia of the type, a female, of E. mandela. Patricia A. Metzler faithfully accompa- nied Metzler on his many collecting trips to Carlsbad Caverns National Park, and she accompanied him on several long driving trips to Washington, DC and other locations

66 Eric H. Metzler & Edward C. Knudson / ZooKeys 149: 51-67 (2011)

for the purpose of identifying specimens. She contributed financially to the study. Maurice Klapwald, New York City Library, J. Donald Lafontaine, and Michael Pogue helped with literature searches. Conversations with, and a thorough review by, James S. Miller were especially helpful. We thank two anonymous reviewers for reading the paper and offering valuable suggestions.

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