Zootaxa 2909: 27 37 (2011) www.mapress.com/zootaxa/ Copyright 2011 Magnolia Press Article ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) A new species of Stenomorpha Solier (Coleoptera: Tenebrionidae: Pimeliinae: Asidini) from Cuatrociénegas, Mexico with a key to the furcata species group AARON D. SMITH 1,3, KELLY B. MILLER 2 & QUENTIN D. WHEELER 1 1 College of Liberal Arts and Sciences, Arizona State University, PO Box 876505, Tempe, AZ 85287-6505, U.S.A. E-mail: pimeliinae@asu.edu, Quentin.Wheeler@asu.edu 2 Department of Biology and Museum of Southwestern Biology, University of New Mexico, MSC03 2020, Albuquerque, NM 87131 0001 USA. E-mail: kbmiller@unm.edu 3 Corresponding author Abstract A new species, Stenomorpha roosevelti Smith, Miller, and Wheeler, n. sp., is described from the Cuatrociénegas Protected Area in Coahuila, Mexico. Three related species are transferred into Stenomorpha, producing the following new combinations: Stenomorpha furcata (Champion), Stenomorpha wickhami (Horn), and Stenomorpha granicollis (Blaisdell). A new informal group, the furcata species group, comprised of these four species is recognized within Stenomorpha. Diagnoses of the group and its species are provided, along with distribution data and a key to the species. Key words: Darkling beetles, Asidini, Stenomorpha, Mexico, taxonomy Resumen Una nueva especie, Stenomorpha roosevelti Smith, Wheeler, y Miller, n. sp., se describe del Área de Protección de Flora y Fauna Cuatrociénegas en Coahuila, México. Tres especies relacionadas se transfieren a Stenomorpha, produciendo las nuevas combinaciones: Stenomorpha furcata (Champion), Stenomorpha wickhami (Horn), y Stenomorpha granicollis (Blaisdell). Un nuevo grupo informal, la especie-grupo del furcata, abarcado de estas cuatro especies dentro de Stenomorpha. Las diagnosis del grupo y de sus especies se proporcionan, junto con datos de la distribución y una llave a las especies. Introduction Asidine darkling beetles (Tenebrionidae: Pimeliinae: Asidini) are a large tribe of flightless mid- to large-sized (4.5 35mm) tenebrionids found in arid and semi-arid regions around the world. The North American (N.A.) component of the tribe occurs from southern Mexico, throughout the western United States, and into southwestern Canada. It currently contains nearly three hundred species in 27 genera (Papp 1961), making it one of the most species-rich tenebrionid groups in North America. Stenomorpha Solier, with approximately 71 current species, is the largest New World genus in the tribe. Stenomorpha was originally erected by Solier (1836) to hold three new central Mexican species (Stenomorpha blapsoides Solier, S. costata Solier, and Trichiasida subpilosa (Solier)). The replacement name Euschides was provided by LeConte (1851) due to the similarity between Stenomorpha and Stenomorphus Dejean, 1831. Horn (1870) concluded that no characters reliably separated Euschides from the Mediterranean genus Asida Latreille (1802) and synonymized it, along with Philolithus Lacordaire (in LeConte 1858) and Pelecyphorus Solier (1836), under Asida. In the Biologia Centrali-Americana, Champion (1884, 1892) followed Horn and described new Mexican asidine species in Asida. However, Casey (1912) removed Asida from the New World by resurrecting both Stenomorpha and its replacement name Euschides and moving all of the previously described North American species, along with the numerous species he described, into these genera and 14 new genera he erected. Wilke (1921) synony- Accepted by A. Cline: 10 May 2011; published: 8 Jun. 2011 27
mized Euschides under Stenomorpha. Since then, little work has been done on the genus beyond the description of a few new species (Blaisdell 1933, 1936; Pierce 1954). Generic boundaries within and between Stenomorpha and many of Casey s asidine genera remain poorly defined (Brown 1971) and need to be tested within a phylogenetic framework. A revision of the N.A. asidine genera utilizing both morphological and molecular data (Smith, in prep.) clearly indicates that a number of Casey s (1912) genera will need to be synonymized with Stenomorpha. As such, we have chosen to transfer three closely related species currently placed in Asidina Casey and Stethasida Casey into Stenomorpha, where analysis of their morphology indicates that they belong (see below). An informal species group is described to contain Stenomorpha furcata (Champion) n. comb., Stenomorpha wickhami (Horn) n. comb., Stenomorpha granicollis (Blaisdell) n. comb., and an interesting new species: Stenomorpha roosevelti from the Cuatrociénegas Protected Area in Coahuila, Mexico. Material and methods Specimens. Approximately 126 specimens from the furcata species group were examined during this study, including the type material of S. furcata, S. granicollis, and S. wickhami. Specimens and/or distributional data were graciously provided by the following private collectors and institutions; collection codens follow Evenhuis (2011), when available: ADSC AMNH ASUT BFLC BMNH BYUC CASC CIDA CSCA EMEC FMNH KWBC MCZC NMNH OSUC RLAC TAMU TTUZ UAIC UCDC UNMC WFBM Aaron D. Smith Collection, Phoenix, AZ, USA. American Museum of Natural History, New York, NY, USA. (Lee Herman) Frank F. Hasbrouck Insect Collection, Arizona State University, Tempe, AZ, USA. Brackenridge Field Laboratory Insect Collection, University of Texas, Austin, TX, USA. (John Abbott) The Natural History Museum, London, United Kingdom. (Max Barclay) Monte L. Bean Life Science Museum, Brigham Young University, Provo, UT, USA. (Shawn Clark) California Academy of Sciences, San Francisco, CA, USA. (Dave Kavanaugh) Orma J. Smith Museum of Natural History, College of Idaho, Caldwell, ID, USA. (William Clark) California State Collection of Arthropods, Sacramento, CA, USA. (Charles Bellamy) Essig Museum of Entomology, University of California, Berkeley, CA, USA. (Cheryl Barr) Field Museum of Natural History, Chicago, IL, USA. (James Boone) Kirby W. Brown Collection, Paradise, CA, USA. Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA. (Philip Perkins) National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. (Warren Steiner and David Furth) C. A. Triplehorn Insect Collection, Ohio State University, Columbus, OH, USA. (Charles Triplehorn and Norm Johnson) Rolf L. Aalbu Collection, Sacramento CA, USA. Texas A&M University Insect Collection, College Station, TX, USA. (Edward Riley) Invertebrate Zoology Collection, Texas Tech University, Lubbock, TX, USA. (James Cokendolpher) University of Arizona Insect Collection, Tucson, AZ, USA. (Wendy Moore) University of California, Davis, Bohart Museum of Entomology, CA, USA. (Fran Keller and Steve Heydon) Museum of Southwestern Biology, University of New Mexico, NM, USA. (Dave Lightfoot and Sandra Brantley) W. F. Barr Entomological Collection, University of Idaho, Moscow, ID, USA. (James Johnson) Morphological parameters. Measurements were taken using digital calipers or an optical micrometer attached to a Nikon SMZ 1500 stereomicroscope. Length (L) was measured along the midline from the anterior margin of the pronotum to the apex of the elytra. Width was measured across the widest point of the elytra (EW) and pronotum (PW). Color was determined under fiber optic illumination and magnification. Images of specimens 28 Zootaxa 2909 2011 Magnolia Press SMITH ET AL.
or characters were taken using a BK Plus Imaging system (R. Larimer, www.visionarydigital.com). Montaged images were assembled using Zerene Stacker (zerenesystems.com/stacker/) and backgrounds were cleaned up in Adobe Photoshop CS5. Setae are acuminate and defined as either simple (circular in cross section and hair-like) or spiniform (thickened, circular or occasionally flattened in cross section, and rigid). Punctation closely follows the conventions used in Paulsen (2005) and is defined in terms of density: either confluent (separated by less than a puncture diameter), dense (separated by between 1 and 2 puncture diameters), moderate (separated by 2 4 puncture diameters), sparse (separated by more than 4 puncture diameters), or impunctate; and size: either coarse (rounded pits, diameter greater than 0.10mm), large (diameter.06 0.10mm), moderate (diameter.03 0.059mm), or fine (diameter less than.03mm). Species recognition. The phylogenetic species concept of Wheeler and Platnick (2000) is employed to define a species as the smallest aggregation of (sexual) populations or (asexual) lineages diagnosable by a unique combination of character states. This species concept is appropriate due to its emphasis on character transformations between species and the lack of data, beyond adult morphology, available for many of the asidine species. Species were maintained or erected based on the presence of autapomorphic morphological characters and/or a unique combination of homoplastic characters shared by all of the specimens assigned to a species. Recognized species should be considered as scientific hypotheses based on the available data and, as such, their validity can be tested as more information is gathered. Key to the Stenomorpha furcata species group 1 Pronotal disc impunctate and tuberculate, tubercles glossy, each bearing a long decumbent hair laterally; posterolateral pronotal apices acute and slightly prolonged; Sonora, Mexico.................................... S. granicollis (Blaisdell) - Pronotal disc finely to moderately punctate and lacking tubercles; posterolateral pronotal apices obtuse, not prolonged.... 2 2(1 ) Pronotal disc and lateral margin moderately to densely clothed in erect or recurved setae; prosternum lacking tubercles..... 3 - Pronotal disc and lateral margin sparsely clothed in minute decumbent setae; prosternum tuberculate.. S. furcata (Champion) 3(2) Dorsum clothed in short erect light brown setae; tarsi with plantar surface bearing light brown spiniform setae along lateral margins, medially glabrous or nearly glabrous; Arizona and California............................ S. wickhami (Horn) - Dorsum clothed in long blackish-brown recurved and erect setae (fig. 13); tarsi with plantar surface densely clothed in long golden-orange setae (fig. 15), lacking a glabrous medial region; Coahuila, Mexico....................S. roosevelti, n. sp. furcata species group Diagnosis. The furcata species group is placed within Stenomorpha based on the following shared characters: Ligula large and tumid, width one-fifth or more mentum width, length approximately 3X width, strongly curved and raised above anterior margin of mentum (Fig.14). Scutellum gradually raised above scutum; mesonotum with two deep rounded paramedial depressions. Protibial spurs subequal. Mesotrochantin large and exposed. Apical segment of maxillary palps enlarged, sexually dimorphic, securiform in females, larger and isotriangular or scalene in males (Fig.14). Members of the furcata species group are distinguished from other Stenomorpha species by a unique combination of the following characters: Mentum transverse, sub-trapezoidal, nearly filling buccal cavity. Submentum distinctly projecting into buccal cavity; width approximately four-fifths that of buccal cavity, laterally exposing a quadrate space before postgena. Pronotum gibbose (viewed laterally); prosternal process declivous behind procoxae. Elytron with marginal costa, bifurcate or not, and one non-bifurcated discal costa. Protibia with outer apical margin unarmed or forming a small indistinct spine with a ring of socketed spiniform setae surrounding the entire margin (Fig. 9). The furcata group can also be separated from the remainder of the species currently in Asidina Casey by the gibbose pronotum (viewed laterally) and lack of prominent protibial spines. The sub-trapezoidal mentum and wide submentum (exposing only small quadrate pockets between the submentum and postgenal processes) exhibited in the furcata group are also present in Asidina confluens (LeConte) and A. semilaevis (Horn), as opposed to the more oval mentum and narrower submentum (exposing large rounded spaces between the submentum and postgenal processes) present in Asidina parallela (LeConte) and A. rugicollis Triplehorn and Brown. Stethasida Casey species A NEW SPECIES OF STENOMORPHA FROM CUATROCIÉNEGAS Zootaxa 2909 2011 Magnolia Press 29
are separated from the furcata group based on their raised prosternal process (declivous in the furcata group species) and Trichiasida Casey species are separated by their large protibial spines. Species description Stenomorpha roosevelti Smith, Miller, and Wheeler, NEW SPECIES (Figs. 4, 8, 11, 13 15) Type material. HOLOTYPE (sex undetermined, probably male) labeled: a) Mexico: Coahuila, Cuatro- / Cienegas Prot. Area / 23. VII. 2010 / 26.90886 N / 102.1185327 W / K. Miller, K. Wetherill et al ; b) Site C4 / ~9.7 km SW Cuatrocienegas / bajada slope, travertine slope / black light and gen. night / collecting ; c) on red paper, HOLOTYPE / Stenomorpha / roosevelti / Smith, Miller & Wheeler, 2011. (UNMC) Description (Holotype). Length: 12.2mm, EW: 6.4mm, PW: 5.2. Integument black, reflective; densely clothed in simple recurved blackish-brown setae interspersed with longer erect setae dorsally; greatest elytral width slightly wider than pronotum, 2.5X longer. Head: Densely punctate, both moderate and fine punctures present, moderately sized punctures each with a simple blackish-brown seta; frontoclypeal suture neither impressed or indicated, epistomal suture faintly visible, not indented at margin; medial depression on frons absent; labral margin broadly emarginate; clypeal membrane concealed. Apical segment of maxillary palps expanded, isotriangular. Mentum transverse, sub-trapezoidal, nearly filling buccal cavity. Submentum width approximately four-fifths that of buccal cavity, laterally exposing a quadrate space before postgena. Postgenal processes rounded, extending to nearly midline of mentum. Antenna densely clothed in dark brown erect simple setae; antennal segment 10 subquadrate, with tomentose sensilla separated, curved around apex. Eye weakly reniform. Prothorax: Pronotum gibbose; pronotal lateral margins arcuate, widest at middle; marginal costa strongly reflexed, crenate; anterior apices produced, acute; posterior apices obtuse; posterior margin weakly bisinuate, medially weakly depressed. Pronotal disc moderately punctate, each puncture of moderate size and nearly filled with a long recurved or erect simple seta. Prosternal process longitudinally impressed, declivous behind procoxae. Prosternum moderately punctate, punctures fine, with decumbent brown simple setae. Pterothorax: Scutellum exposed, gradually posteriorly raised above scutum; mesonotum with two deep rounded paramedial depressions. Elytra elongate oval, widest posterior to middle, strongly declivous near apex. Elytral humera weakly constricted before base, apices acute, not reflexed. Marginal costa tuberculate, weakly raised, bifurcated in basal third; inner branch weakly indicated, not reaching outer branch at base or apical elytron declivity. Discal costa of elytron tuberculate, weakly raised from behind humera to apical third of elytron, parallel to suture. Elytral disc moderately punctate, punctures large; densely clothed in long recurved blackish-brown setae interspersed with longer erect setae along costae and margins. Legs: Femora and tibia densely clothed with decumbent erect dark brown simple setae, pro-, meso-tibia also with sparse socketed black spiniform setae. Femora transversely rugose; tibia weakly rugose; tarsi with plantar surface densely pubescent throughout, setae simple, golden-orange, becoming sparser, darker on distal tarsomere; protibia with outer apical margin forming a small indistinct spine with a ring of socketed spiniform setae running entire margin; inner apical margin with two subequal spurs; combined protarsal segments approximately one-half length of protibia. Abdomen: Moderately punctate with decumbent simple setae. Diagnosis. Stenomorpha roosevelti is easily separated from other members of the furcata group by its dorsal vestiture (composed of both recurved and erect simple blackish-brown setae), its weakly indicated elytral carinae, and by the dense golden-orange pubescence on the ventral portion of the tarsi. Etymology. This species is named in honor of the 26th President of the United States, Theodore Roosevelt, for his contributions to conservation. Remarks. Known only from the type specimen, collected in the Cuatrociénegas Protected Area in Coahuila, Mexico (see discussion for details). Stenomorpha furcata (Champion), new combination (Figs. 1, 5, 11) Asida furcata Champion 1892:499 Asidina furcata (Champion) 1892:499. Casey 1912:172 30 Zootaxa 2909 2011 Magnolia Press SMITH ET AL.
FIGURES 1 4. Dorsal habitus of the Stenomorpha furcata species-group: 1) Stenomorpha furcata (male); 2) Stenomorpha granicollis (male); 3) Stenomorpha wickhami (male); 4) Stenomorpha roosevelti (holotype). A NEW SPECIES OF STENOMORPHA FROM CUATROCIÉNEGAS Zootaxa 2909 2011 Magnolia Press 31
FIGURES 5 8. Lateral view of the Stenomorpha furcata species-group: 5) Stenomorpha furcata (male); 6) Stenomorpha granicollis (male); 7) Stenomorpha wickhami (male); 8) Stenomorpha roosevelti (holotype). Type material. HOLOTYPE, by monotypy, (sex undetermined) labeled: a) Orange bordered white circle reading Type ; b) Villa Lerdo, / Durango. / Hoge. ; c) Asida / furcata, Ch. ; d) Sp. figured; e) B.C.ACol.IV.1. / Asida / furcata. (BMNH) Additional material examined. (33 specimens). MEXICO: Durango and Coahuila. USA: Texas (Brewster, Presidio, Terrell, Upton, Val Verde Counties). 32 Zootaxa 2909 2011 Magnolia Press SMITH ET AL.
Diagnosis. S. furcata can be distinguished from other members of the furcata group by its sparsely clothed and finely punctate pronotal disc, which is more strongly gibbose in lateral view than are those of the other species, and its tuberculate prosternum. Remarks. Triplehorn and Brown (1971) presented the first records of S. furcata within the United States and discussed variation within and between S. furcata specimens from differing localities. They examined differences in elytral sculpturing and marginal carina bifurcation between populations and concluded that, while there are slight geographical variations, all of the observed variation fell within the range of S. furcata. FIGURES 9 10. Protibia, outer apical margin: 9) Stenomorpha wickhami; 10) Asidina parallela. FIGURE 11. Distribution of furcata group species. Stars ( ) = Stenomorpha granicollis, circle ( ) = Stenomorpha roosevelti, squares ( ) = Stenomorpha wickhami, and triangles ( ) = Stenomorpha furcata.. Stenomorpha granicollis (Blaisdell), new combination (Figs. 2, 6, 11) Asida (Stethasida) granicollis Blaisdell 1923:256 Stethasida granicollis (Blaisdell). Papp 1961: 29. A NEW SPECIES OF STENOMORPHA FROM CUATROCIÉNEGAS Zootaxa 2909 2011 Magnolia Press 33
Type material. HOLOTYPE, by original designation, (sex undetermined) locality label: a) Guaymas Mex., April 14 1921, E P Van Duzee Collector (CASC). California Academy of Sciences type No. 1165. Additional material examined. (10 specimens). MEXICO: Sonora. Diagnosis. Stenomorpha granicollis is easily distinguished from other members of the furcata group by its impunctate and tuberculate pronotum and its acute and somewhat prolonged posterolateral pronotal apices. Remarks. Blaisdell (1923) originally described S. granicollis in the genus Asida and used Casey s genus Stethasida as a subgenus. The species is not mentioned again until Papp (1961) lists it as species of Stethasida. All known specimens have been collected within a small area in western Sonora around Guayamas. FIGURE 12. Habitat at Stenomorpha roosevelti collection site (type locality). Stenomorpha wickhami (Horn), new combination (Figs. 3, 7, 9, 11) Asida wickhami Horn 1894:420 Asidina wickhami (Horn). Casey 1912:171 = Asidina liberta Casey 1912:171. [Synonymy by Triplehorn and Brown 1971] Type material. LECTOTYPE (sex undetermined), designated here, labeled: a) Riverside / Ariz. ; b) 64 ; c) HoloTYPE / 3932 ; d) A. / Wickhami / Horn; e) MCZ / Type / 33979 (MCZ). Additional material examined. (79 specimens). USA: Arizona (Gila, Coconino, Pinal, Yavapai, Maricopa, Pima, Greenlee, and Mohave Counties), California (San Bernardino County). Diagnosis. Stenomorpha wickhami can be distinguished from other members of the furcata group by the dense coating of short erect light brown setae on the dorsum, venter, and legs and the weakly crenulated elytral base. It is 34 Zootaxa 2909 2011 Magnolia Press SMITH ET AL.
superficially similar to Asidina parallela (LeConte) and the two species co-occur in at least part of their ranges. Stenomorpha wickhami differs from A. parallela as per the species group characters, including the larger, well defined protibial spine (Fig. 10), smaller more laterally rounded mentum, and the larger rounded spaces separating the submentum from the postgena. Also, on each elytron S. wickhami has a discal costa, and a marginal costa that is nearly always bifurcated. However, the carina on each elytron of A. parallela appears to be the inner branch of a bifurcated marginal costa, though it is not connected to the raised carinate marginal costa at either end. The discal costa is entirely absent. Remarks. Variation in the bifurcation of the elytral marginal costa in S. wickhami is discussed in Triplehorn and Brown (1971). Discussion The type specimen of S. roosevelti was collected as part of a larger survey for undescribed and endemic arthropods within the Cuatrociénegas Protected Area conducted by the Museum of Southwestern Biology s Division of Arthropods. The Cuatrociénegas region is in a desert basin bordered by mountains, making it relatively isolated from the more typical surrounding Chihuahuan desert habitat. While the basin receives relatively little rainfall, it does have a variety of aquatic habitats (including marshes, playa lakes, and streams) due to an abundance of natural springs. Indeed, Cuatrociénegas is probably best known for its diversity of endemic fish and aquatic invertebrates (Minckley 1984, Dinger et al. 2005). Major features of the terrestrial habitat at Cuatrociénegas include the presence of gypsum dunes and salt flats with halophytic grasses and shrubs. The single specimen of S. roosevelti was collected on a bajada slope dominated by Larrea tridentata (creosote), Prosopis gladulosa (honey mesquite), Grusonia bradtiana (cactus), and Jatropha dioica (leatherstem) several kilometers away from the nearest unvegetated dunes (Karen Wetherill, pers. comm.). However; two characters, dense pubescence on the tarsi plantar surfaces and a thick clothing of long setae over nearly the entire body, indicate that S. roosevelti may be associated with dunes. Both characters are uncommon within the Asidini, though present in a number of other dune- dwelling tenebrionids (Edrotes, Asbolus, etc.). In addition, the only two known North American asidine species exhibiting both characters (Trichasida hirsuta (LeConte) and Stethasida muricatula (LeConte) are either found on or near dunes, or in desert arroyos with sandy or very fine clay soil. None of the other members of the furcata group have been associated with dune habitats, and none exhibit these characters. It is worth noting that S. furcata has been collected just outside of the Cuatrociénegas basin, 44 km from the collection site of S. roosevelti, and may potentially occur in the basin. Other than this one instance, the known ranges for the furcata group species are widely separated. The closest collecting sites for S. granicollis and S. wickhami to S. roosevelti are nearly 900 km and 1100 km away respectively. Both S. roosevelti and S. granicollis appear to have fairly limited distributions, while S. furcata has one of the widest ranges of any known asidine, with the most distant collecting points being nearly 800 km apart. It is likely that additional species within the furcata group will be discovered as more intense sampling is done on other unique and isolated habitats such as Cuatrociénegas. Acknowledgments For his valuable advice and comments on the manuscript we thank R.L. Aalbu. For assistance with imaging we wish to thank R. Dornburg and A. Ramirez. For providing habitat information and images for Cuatrociénegas we thank Karen Wetherill. For assistance in project organization, permitting and logistical support we would like to thank D.C. Lightfoot (research project coordinator, co-pi and MSBA collection manager), D. Bustos (research project coordinator and Resource Program Manager, White Sands National Monument), I.G. Gutierrez (Cuatrociénegas coordinator, Director, Área de Protección de Flora y Fauna Cuatrociénegas), and V.A. Rodriguez (research colleague and permit coordinator, Universidad Juarez del Estado de Durango, Coahuila, Mexico). We also wish to thank the private collectors and museum curators who graciously loaned us the specimens used in this study. This publication was supported by a grant from the US National Park Service Sister Parks Program (#NSP- 1262-RICO- 673). A NEW SPECIES OF STENOMORPHA FROM CUATROCIÉNEGAS Zootaxa 2909 2011 Magnolia Press 35
FIGURES 13 15. Stenomorpha roosevelti: 13) Recurved setae on elytron; 14) Mentum and apical segment of maxillary palp; 15) Protarsi, venter. 36 Zootaxa 2909 2011 Magnolia Press SMITH ET AL.
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