Aaron D. Smith

Finding Our Way through Phenotypes

Imagine if we could compute across phenotype data as easily as genomic data; this article calls for efforts to realize this vision and discusses the potential benefits.

A REVISION OF THE ELEODES (SUBGENUS CAVERNELEODES) WITH NEW SPECIES AND NOTES ON CAVE BREEDING ELEODES (TENEBRIONIDAE: AMPHIDORINI)

Abstract. Subgenus Caverneleodes of the genus Eleodes is diagnosed and revised. Six new species from the United States: California (E. microps); Utah and Northern Arizona (E. wynnei), Central Arizona (E. wheeleri), Southern New Mexico (E. guadalupensis), and Mexico (E. thomasi and E. grutus) are described. The biogeography of the subgenus is discussed. Diagnoses and a key are provided to known species of Caverneleodes. Relationships with other Eleodes are discussed. Cave associated Amphidorini are surveyed.

The Tenebrionidae of California: A Time Sensitive Snapshot Assessment

Abstract Due to a diversity of habitats and its geologic history, the US state of California hosts a spectacular assemblage of darkling beetle species (Coleoptera: Tenebrionidae). In addition to being part of the California Floristic Province, one of 34 global biodiversity hotspots identified by Conservation International, California also has additional areas which are parts of the Great Basin, Mojave, and Sonoran deserts. California is divided into nine floristic regions. Each region is assessed in terms of faunal composition and endemism. A “snapshot” of our present knowledge of the Tenebrionidae indicates that 447 currently recognized species, representing 108 genera, occur in California of which one hundred and ninety are endemic. California is compared to other nearby regions in diversity and endemism. An analysis of currently valid species vs a more realistic species account based on unpublished records of likely synonyms and known species yet to be described in the scientific literature is presented. The California Floristic Region, rather than other more arid parts of California, has the highest number of total and endemic species. Because of their high diversity and endemism, tenebrionids could potentially provide a valuable tool for monitoring the environment for conservation purposes.

A Revision of Eleodes Subgenus Eleodes Eschscholtz (Coleoptera: Tenebrionidae)

ABSTRACT Keys, descriptions and figures are provided for the identification of 30 species assignable to the nominate subgenus of the tenebrionid genus Eleodes. Three species groups can be recognized: the grandicollis group, containing only E. grandicollis Mannerheim; the obscurus group, containing E. obscurus (Say), E. acutus (Say) and E. suturalis (Say); and the dentipes group, containing E. dentipes Eschscholtz, E. acuticaudus LeConte, E. adumbratus Blaisdell, E. armatus (LeConte), E. curvidens Triplehorn & Cifuentes, E. discinctus Blaisdell, E. eschscholtzi Solier, E. femoratus LeConte, E. gracilis LeConte, E. hispilabris (Say), E. loretensis Blaisdell, E. mexicanus Blaisdell, E. mirabilis Triplehorn, E. moestus Blaisdell, E. muricatulus Triplehorn, E. rossi Blaisdell, E. rugosus Perbosc, E. samalayucae Triplehorn, E. sanmartinensus Blaisdell, E. scyropterus Triplehorn, E. spinipes Champion, E. sponsus LeConte, E. subcylindricus Casey, E. tenuipes Casey, E. vanduzeei Blaisdell, and a new species described herein, Eleodes fiski Triplehorn. Eleodes amadeensis Blaisdell and Eleodes striatipennis Blaisdell are synonymized under Eleodes armatus (LeConte). Eleodes subpinguis Blaisdell is synonymized under Eleodes hispilabris (Say). Eleodes wickhami Horn is synonymized under E. eschscholtzi Solier. The status of Eleodes femoratus LeConte 1851 under E. militaris Horn 1870, is reversed based on priority.

Pythiopina, an enigmatic subtribe of darkling beetles (Coleoptera: Tenebrionidae: Pedinini): taxonomic revision, microtomography, ecological niche models and phylogenetic position

Morphological, anatomical, and distributional data concerning the South African endemic beetle subtribe Pythiopina (Tenebrionidae: Pedinini) are revised. Five species, representing two genera, are recognized. Included in this total is one new species (Meglyphus mariae Kamiński sp.n.). The following species are placed in synonymy: Meglyphus ciliatipes [=Meglyphus calitzensis syn.n.]; Meglyphus laenoides [=Meglyphus andreaei syn.n.; =Meglyphus namaqua syn.n.]. Microtomographic models for all valid Pythiopina species, including the holotype of the newly described species, are presented and analysed. Endoskeleton morphology (specifically characters of the tentorium and metendosternite) proved to be informative at the specific and generic levels. An identification key is provided to all known species of the subtribe. Environmental niche models are presented for the majority of species. A molecular phylogeny of Pedinini based on six genetic loci (28S: D1–D3 region; 28S: D4–D5 region, COII, ArgK, CAD2, wg) was also produced to explore the phylogenetic position of Pythiopina. This analysis is the first to include representatives of all seven subtribes of Pedinini, and supports a sister relationship between Pythiopina and the Palaearctic subtribe Dendarina. Results also suggest the existence of a second pair of sister taxa within Pedinini (in addition to Melambiina) with an amphitropical African distribution.

Catalogue of Tenebrionidae (Coleoptera) of North America

Abstract This catalogue includes all valid family-group (8 subfamilies, 52 tribes, 14 subtribes), genus-group (349 genera, 86 subgenera), and species-group names (2825 species, 215 subspecies) of darkling beetles (Coleoptera: Tenebrionidae) known to occur in North America1 and their available synonyms. Data on extant, subfossil and fossil taxa are given. For each name the author and year and page number of the description are provided, with additional information (e.g., type species for genus-group names, author of synonymies for invalid taxa) depending on the taxon rank. Several new nomenclatural acts are included. One new genus, Lepidocnemeplatia Bousquet and Bouchard, is described. Spelaebiosis Bousquet and Bouchard [for Ardoinia Özdikmen, 2004], Blapstinus marcuzzii Aalbu [for Blapstinus kulzeri Marcuzzi, 1977], and Hymenorus campbelli Bouchard [for Hymenorus oculatus Doyen and Poinar, 1994] are proposed as new replacement names. Supporting evidence is provided for the conservation of usage of Tarpela micans (Fabricius, 1798) nomen protectum over Tarpela vittata (Olivier, 1793) nomen oblitum. The generic names Psilomera Motschulsky, 1870 [= Stenomorpha Solier, 1836], Steneleodes Blaisdell, 1909 [= Xysta Eschscholtz, 1829], Ooconibius Casey, 1895 and Euconibius Casey, 1895 [= Conibius LeConte, 1851] are new synonyms (valid names in square brackets). The following 127 new synonymies of species-group names, listed in their original combination, are proposed (valid names, in their current combination, placed in square brackets): Bothrasida mucorea Wilke, 1922 [= Pelecyphorus guanajuatensis (Champion, 1884)]; Parasida zacualpanicola Wilke, 1922 [= Pelecyphorus asidoides Solier, 1836]; Stenosides kulzeri Pallister, 1954, Stenosides bisinuatus Pallister, 1954, and Parasida trisinuata Pallister, 1954 [= Pelecyphorus dispar (Champion, 1892)]; Asida favosa Champion, 1884 and Asida similata Champion, 1884 [= Pelecyphorus fallax (Champion, 1884)]; Ologlyptus bicarinatus Champion, 1884 [= Pelecyphorus indutus (Champion, 1884)]; Parasida laciniata Casey, 1912 and Parasida cristata Pallister, 1954 [= Pelecyphorus liratus (LeConte, 1854)]; Parasida esperanzae Wilke, 1922 and Parasida mixtecae Wilke, 1922 [= Pelecyphorus longipennis (Champion, 1884)]; Parasida tolucana Casey, 1912 [= Pelecyphorus scutellaris (Champion, 1884)]; Parasida purpusi Wilke, 1922 [= Pelecyphorus tristis (Champion, 1884)]; Astrotus nosodermoides Champion, 1892 [= Pelecyphorus erosus (Champion, 1892)]; Astrotus seticornis var. humeralis Champion, 1884 [= Pelecyphorus seticornis (Champion, 1884)]; Pactostoma breviuscula Casey, 1912, Pactostoma exoleta Casey, 1912, Pactostoma luteotecta Casey, 1912, Pactostoma monticola Casey, 1912, Pactostoma obtecta Casey, 1912, and Pactostoma sigillata Casey, 1912 [=Pelecyphorus anastomosis (Say, 1824)]; Ologlyptus canus Champion, 1884 and Ologlyptus sinuaticollis Champion, 1884 [= Pelecyphorus graciliformis (Solier, 1836)]; Gonasida elata reducta Casey, 1912, Gonasida elata prolixa Casey, 1912, and Gonasida aucta Casey, 1912 [= Philolithus elatus compar (Casey, 1912)]; Gonasida alaticollis Casey, 1912 [= Philolithus elatus difformis (LeConte, 1854)]; Gonasida gravida Casey, 1912 [= Philolithus elatus elatus (LeConte, 1853)]; Pelecyphorus aegrotus limbatus Casey, 1912 [= Philolithus aegrotus aegrotus (LeConte, 1861)]; Pelecyphorus corporalis Casey, 1912, Pelecyphorus reptans Casey, 1912, Pelecyphorus socer Casey, 1912, Pelecyphorus abscissus Casey, 1912, Pelecyphorus fumosus Casey, 1912, Pelecyphorus parvus Casey, 1912, Pelecyphorus morbillosus pacatus Casey, 1912, Pelecyphorus morbillosus sobrius Casey, 1912, Pelecyphorus piceus Casey, 1912, Pelecyphorus piceus crudelis Casey, 1912, Pelecyphorus snowi Casey, 1912, and Pelecyphorus subtenuis Casey, 1912 [= Philolithus morbillosus (LeConte, 1858)]; Bothrasida sanctae-agnae Wilke, 1922 [= Stenomorpha funesta (Champion, 1884)]; Asida flaccida Horn, 1896 [= Stenomorpha embaphionides (Horn, 1894)]; Asida angustula Casey, 1890, Stethasida stricta Casey, 1912, Stethasida muricatula languida Casey, 1912, Stethasida pertinax Casey, 1912, Stethasida socors Casey, 1912, Stethasida angustula inepta Casey, 1912, Stethasida tenax Casey, 1912, and Stethasida vegrandis Casey, 1912 [= Stenomorpha muricatula (LeConte, 1851)]; Stethasida obsoleta expansa Casey, 1912, Stethasida obsoleta opacella Casey, 1912, Stethasida brevipes Casey, 1912, Stethasida torpida Casey, 1912, Stethasida convergens Casey, 1912, Stethasida discreta Casey, 1912, Stethasida longula Casey, 1912, Stethasida adumbrata Casey, 1912, Stethasida occulta Casey, 1912, Stethasida tarsalis Casey, 1912, Stethasida unica Casey, 1912, and Pelecyphorus laevigatus Papp, 1961 [= Stenomorpha obsoleta (LeConte, 1851)]; Trichiasida eremica Wilke, 1922 [= Stenomorpha difficilis (Champion, 1884)]; Trichiasida lineatopilosa Casey, 1912 [= Stenomorpha hirsuta (LeConte, 1851)]; Trichiasida tenella Casey, 1912 [= Stenomorpha hispidula (LeConte, 1851)]; Trichiasida duplex Casey, 1912 [= Stenomorpha villosa (Champion, 1884)]; Alaudes squamosa Blaisdell, 1919, Alaudes testacea Blaisdell, 1919, and Alaudes fallax Fall, 1928 [= Alaudes singularis Horn, 1870]; Edrotes barrowsi Dajoz, 1999 [=Edrotes ventricosus LeConte, 1851]; Nyctoporis tetrica Casey, 1907 and Nyctoporis maura Casey, 1907 [= Nyctoporis aequicollis Eschscholtz, 1831]; Nyctoporis pullata Casey, 1907 [= Nyctoporis sponsa Casey, 1907]; Eleodes tibialis forma oblonga Blaisdell, 1909 [= Eleodes tibialis Blaisdell, 1909]; Eleodes (manni var.) variolosa Blaisdell, 1917 [= Eleodes constrictus LeConte, 1858]; Eleodes cordata forma sublaevis Blaisdell, 1909, Eleodes cordata forma intermedia Blaisdell, 1909, Eleodes cordata forma oblonga Blaisdell, 1909, Eleodes cordata forma elongata Blaisdell, 1909, and Eleodes (cordata var.) adulterina Blaisdell, 1917 [= Eleodes cordata Eschscholtz, 1829]; Eleodes hornii var. monticula Blaisdell, 1918 and Eleodes manni sierra Blaisdell, 1925 [= Eleodes fuchsii Blaisdell, 1909]; Eleodes parvicollis var. squalida Blaisdell, 1918 [= Eleodes parvicollis Eschscholtz, 1829]; Eleodes reflexicollis Mannerheim, 1843 and Eleodes parvicollis forma farallonica Blaisdell, 1909 [= Eleodes planata Eschscholtz, 1829]; Eleodes indentata Blaisdell, 1935 [= Eleodes rotundipennis LeConte, 1857]; Eleodes intricata Mannerheim, 1843 [= Eleodes scabrosa Eschscholtz, 1829]; Eleodes horni fenyesi Blaisdell, 1925 [= Eleodes tenebrosa Horn, 1870]; Eleodes cordata var. horrida Blaisdell, 1918 [= Eleodes tuberculata Eschscholtz, 1829]; Eleodes oblonga Blaisdell, 1933 [= Eleodes versatilis Blaisdell, 1921]; Eleodes dentipes marinae Blaisdell, 1921 [= Eleodes dentipes Eschscholtz, 1829]; Eleodes carbonaria forma glabra Blaisdell, 1909 [= Eleodes carbonaria carbonaria (Say, 1824)]; Eleodes granosa forma fortis Blaisdell, 1909 [= Eleodes granosa LeConte, 1866]; Eleodes pilosa forma ordinata Blaisdell, 1909 [= Eleodes pilosa Horn, 1870]; Trogloderus costatus pappi Kulzer, 1960 [= Trogloderus tuberculatus Blaisdell, 1909]; Trogloderus costatus mayhewi Papp, 1961 [= Trogloderus vandykei La Rivers, 1946]; Bolitophagus cristatus Gosse, 1840 [= Bolitotherus cornutus (Fabricius, 1801)]; Eleates explanatus Casey, 1890 [= Eleates depressus (Randall, 1838)]; Blapstinus sonorae Casey, 1890 [= Blapstinus brevicollis LeConte, 1851]; Blapstinus falli Blaisdell, 1929 [= Blapstinus castaneus Casey, 1890]; Blapstinus brunneus Casey, 1890 and Blapstinus coronadensis Blaisdell, 1892 [=Blapstinus histricus Casey, 1890]; Blapstinus hesperius Casey, 1890 [=Blapstinus intermixtus Casey, 1890]; Blapstinus cinerascens Fall, 1929 [= Blapstinus lecontei Mulsant and Rey, 1859]; Blapstinus niger Casey, 1890 and Blapstinus cribricollis Casey, 1890 [= Blapstinus pimalis Casey, 1885]; Blapstinus arenarius Casey, 1890 [= Blapstinus pratensis LeConte, 1859]; Blapstinus gregalis Casey, 1890 [= Blapstinus substriatus Champion, 1885]; Blapstinus hydropicus Casey, 1890 [= Blapstinus sulcatus LeConte, 1851]; Blapstinus hospes Casey, 1890 [= Blapstinus vestitus LeConte, 1859]; Notibius reflexus Horn, 1894 [= Conibius opacus (LeConte, 1866)]; Notibius affinis Champion, 1885 [=Conibius rugipes (Champion, 1885)]; Conibius parallelus LeConte, 1851 [= Conibius seriatus LeConte, 1851]; Nocibiotes rubripes Casey, 1895 [=Nocibiotes caudatus Casey, 1895]; Nocibiotes gracilis Casey, 1895 and Nocibiotes acutus Casey, 1895 [=Nocibiotes granulatus (LeConte, 1851)]; Conibius alternatus Casey, 1890 [= Tonibius sulcatus (LeConte, 1851)]; Pedinus suturalis Say, 1824 [= Alaetrinus minimus (Palisot de Beauvois, 1817)]; Menedrio longipennis Motschulsky, 1872 [= Tenebrio obscurus Fabricius, 1792]; Hymenophorus megops Hatch, 1965 and Telesicles magnus Hatch, 1965 [= Hymenorus sinuatus Fall, 1931]; Andrimus concolor Casey, 1891 and Andrimus convergens Casey, 1891 [= Andrimus murrayi (LeConte, 1866)]; Mycetochara marshalli Campbell, 1978 [= Mycetochara perplexata Marshall, 1970]; Phaleria globosa LeConte, 1857 [= Phaleria picta Mannerheim, 1843]. The following subspecies of Trogloderus costatus LeConte, 1879 are given species rank: Trogloderus nevadus La Rivers, 1943, Trogloderus tuberculatus Blaisdell, 1909, and Trogloderus vandykei La Rivers, 1946. The following taxa, previously thought to be junior synonyms, are considered valid: Amphidora Eschscholtz, 1829; Xysta Eschscholtz, 1829; Helops confluens (Casey, 1924). Two new combinations are proposed: Stenomorpha spinimana (Champion, 1892) and Stenomorpha tenebrosa (Champion, 1892) [from the genus Parasida Casey, 1912]. The type species [placed in square brackets] of the following 12 genus-group taxa are designated for the first time: Lagriola Kirsch, 1874 [Lagriola operosa Kirsch, 1874]; Locrodes Casey, 1907 [Emmenastus piceus Casey, 1890]; Falacer Laporte, 1840 [Acanthopus cupreus Laporte, 1840 (= Helops contractus Palisot de Beauvois, 1812)]; Blapylis Horn, 1870 [Eleodes cordata Eschscholtz

Available data support protection of the Southwestern Willow Flycatcher under the Endangered Species Act

ABSTRACT Zink (2015) argued there was no evidence for genetic, morphological, or ecological differentiation between the federally endangered Southwestern Willow Flycatcher (Empidonax traillii extimus) and other Willow Flycatcher subspecies. Using the same data, we show there is a step-cline in both the frequency of a mtDNA haplotype and in plumage variation roughly concordant with the currently recognized boundary between E. t. extimus and E. t adastus, the subspecies with which it shares the longest common boundary. The geographical pattern of plumage variation is also concordant with previous song analyses differentiating those 2 subspecies and identified birds in one low-latitude, high-elevation site in Arizona as the northern subspecies. We also demonstrate that the ecological niche modeling approach used by Zink yields the same result whether applied to the 2 flycatcher subspecies or to 2 unrelated species, E. t. extimus and Yellow Warbler (Setophaga petechia). As a result, any interpretation of those results as evidence for lack of ecological niche differentiation among Willow Flycatcher subspecies would also indicate no differentiation among recognized species and would therefore be an inappropriate standard for delineating subspecies. We agree that many analytical techniques now available to examine genetic, morphological, and ecological differentiation would improve our understanding of the distinctness (or lack thereof) of Willow Flycatcher subspecies, but we argue that currently available evidence supports protection of the Southwestern Willow Flycatcher under the Endangered Species Act.

Review of the genus Trichiotes Casey (Coleoptera: Tenebrionidae: Pimeliinae: Edrotini), with the description of a new species and a preliminary checklist of the Tenebrionidae from Cuatrociénegas, Mexico

The rarely collected, previously monotypic, darkling beetle genus Trichiotes is reviewed, including new state and country records for T. seriatus Casey, 1907 and a new species, Trichiotes lightfooti Wirth & Smith, n. sp., is described from the Cuatrociénegas Protected Area in Coahuila, Mexico. Differences in stridulatory structures between the species are discussed and the occurrence of a waxy, extracuticular pruinescence is reported for Trichiotes. A species key to the genus is provided and a checklist of the known Tenebrionidae from the Cuatrociénegas region is presented.

Caribanosis gen. nov. from Hispaniola (Pimeliinae: Stenosiini) with taxonomic notes on the tribes belopini and stenosini (Coleoptera: Tenebrionidae)

Abstract. Rhypasma Pascoe, 1862 is transferred from the tribe Stenosini (Pimeliinae) to Belopini (Lagriinae). Caribanosis gen. nov. is described and placed in Stenosini (Tenebrionidae: Pimeliinae) to hold Rhypasma quisqueyanus Garrido et Varela, 2011. The following new combination is established: Caribanosis quisqueyanus (Garrido et Varela, 2011), comb. nov. Caribanosis is similar to the South American genus Grammicus Waterhouse, 1845 but differs in having a single central pronotal keel, not two lateral keels as in Grammicus. Both are members of the subtribe Stenosina.

A New Genus and Species of Stridulating Edrotini (Coleoptera: Tenebrionidae: Pimeliinae) from West Texas, with Notes on Stridulation Within the Tribe

Abstract. Eremocantor marioni, gen. nov. et sp. nov. is described for an unusual species of stridulating darkling beetle discovered in West Texas, USA. Stridulation in other members of the tribe Edrotini (Tenebrionidae: Pimeliinae) is discussed, including Edrotes LeConte and the previously unrecognized sound producing genera Pimeliopsis Champion, Pescennius Champion, Trichiotes Casey, and Oxygonodera Casey. A key to the known stridulating edrotine genera is also provided.