Noctuidae, Pantheinae)
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A peer-reviewed open-access journal ZooKeysTh e39: North 161–181 American (2010) species of Charadra Walker, with a revision of the Charadra pata (Druce)...161 doi: 10.3897/zookeys.39.432 RESEARCH ARTICLE www.pensoftonline.net/zookeys Launched to accelerate biodiversity research The North American species of Charadra Walker, with a revision of the Charadra pata (Druce) group (Noctuidae, Pantheinae) B. Christian Schmidt1,†, Gary G. Anweiler2,‡ 1 Canadian Food Inspection Agency, Canadian National Collection of Insects, Arachnids and Nematodes, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6 2 E.H. Strickland Entomological Museum, 218 Earth Sciences Building, Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada, T6G 2E9 † urn:lsid:zoobank.org:author:C3C5392A-EBF8-41B9-99BE-364A8C2FBB7F ‡ urn:lsid:zoobank.org:author:87B63195-32B4-4FAF-8732-54242BF1FAA9 Corresponding authors: B. Christian Schmidt ([email protected]), Gary G. Anweiler (ggan- [email protected]) Academic editor: J.D. Lafontaine | Received 29 January 2010 | Accepted 5 February 2010 | Published 18 March 2010 urn:lsid:zoobank.org:pub:F4D24A8D-3EBF-4688-910D-413E328F92BF Citation: Schmidt BC, Anweiler GG (2010) Th e North American species of Charadra Walker, with a revision of the Cha- radra pata (Druce) group (Noctuidae, Pantheinae) In: Schmidt BC, Lafontaine JD (Eds) Contributions to the systematics of New World macro-moths II. ZooKeys 39: 161–181. doi: 10.3897/zookeys.39.432 Abstract Th e North American species of the genus Charadra Walker are reviewed, and the species of the yellow- hindwing (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. Keywords Charadra, new species, taxonomy, Arizona, New Mexico, Mexico, Quercus Copyright B.C. Schmidt, G.G. Anweiler. 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. 162 B. Christian Schmidt & Gary G. Anweiler / ZooKeys 39: 161–181 (2010) Introduction Th e genus Charadra Walker as currently understood is comprised of three yellow-hind- winged species (C. pata (Druce), C. oligarchia Dyar, and C. patafex Dyar) and fi ve grey- hindwinged species (C. deridens (Guenée), C. ingenua J.B. Smith, C. dispulsa Morri- son, 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 spe- cies 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 dou- ble 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 recog- nition 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 spe- cies 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 taxonC. 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 slide- mounted 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 Th e North American species of Charadra Walker, with a revision of the Charadra pata (Druce)...163 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 neighbour- joining 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://bar- codinglife.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. 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 Histo- ry]), London, United Kingdom. CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ot- tawa, 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. Systematics Charadra Walker Charadra Walker, 1865: 445. Type species: Charadra contigua Walker, 1865, a junior subjective synonym of Charadra deridens (Gueneé). Th e generic placement of Charadra in relation to other pantheine genera, especial- ly 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; dor- sal 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 164 B. Christian Schmidt & Gary G. Anweiler / ZooKeys 39: 161–181 (2010) 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. Charadra pata (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 de- scription, which consists of: “Trisuloides patens [sic] Druce, is a Charadra which I had named basifl avabefore 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 avais 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).