abstract: Understanding the diversity, evolutionary relationships, and geographic distribution of species is foundational knowledge in biology. However, this knowledge is lacking for many diverse lineages of the tree of life. This is the case for the desert stink beetles in the tribe Amphidorini LeConte, 1862 (Coleoptera: Tenebrionidae) – a lineage of arid-adapted flightless beetles found throughout western North America. Four interconnected studies that jointly increase our knowledge of this group are presented. First, the darkling beetle fauna of the Algodones sand dunes in southern California is examined as a case study to explore the scientific practice of checklist creation. An updated list of the species known from this region is presented, with a critical focus on material now made available through digitization and global aggregation. This part concludes with recommendations for future biodiversity checklist authors. Second, the psammophilic genus Trogloderus LeConte, 1879 is revised. Six new species are described, and the first, multi-gene phylogeny for the genus is inferred. In addition, historical biogeographic reconstructions along with novel hypotheses of speciation patterns within the Intermountain Region are given. In particular, the Kaibab Plateau and Kaiparowitz Formation are found to have promoted speciation on the Colorado Plateau. The Owens Valley and prehistoric Bouse Embayment are similarly hypothesized to drive species diversification in southern California. Third, a novel phylogenomic analysis for the tribe Amphidorini is presented, based on 29 de novo partial transcriptomes. Three putative ortholog sets were discovered and analyzed to infer the relationships between species groups and genera. The existing classification of the tribe is found to be highly inadequate, though the earliest-diverging relationships within the tribe are still in question. Finally, the new phylogenetic framework is used to provide a genus-level revision for the Amphidorini, which previously contained six valid genera and 253 valid species. This updated classification includes more than 100 taxonomic changes and results in the revised tribe consisting of 16 genera, with three being described as new to science. / Dissertation/Thesis / Doctoral Dissertation Evolutionary Biology 2018

Insects fulfill important roles within all ecosystems, including deserts, and interact directly and indirectly with the endemic and endangered species at the Ash Meadows National Wildlife Refuge (AMNWR). Here I investigate the variability of species richness, diversity, abundance and community composition of four insect groups between stabilized sand dune habitats, unstabilized sand dune habitats and non-sand dune habitats. The insects examined in this thesis include antlions (Myrmeleontidae), bees (Apiformes), darkling beetles (Tenebrionidae) and velvet ants (Mutillidae). As the impact and interactions of two of the insect groups, antlions and velvet ants, are largely unknown within any ecosystem, this thesis also includes a faunal study of velvet ants at AMNWR and their spatial and temporal variation at the refuge. Generalized linear mixed models were used to identify significant differences in richness, diversity and abundance for each insect group between the stabilized sand dune, unstabilized sand dune and non-sand dune habitats. Species richness differed between habitats for antlions, beetles and velvet ants over the study period. Diversity differed between habitats for antlions and beetles, and abundance differed between habitats for all groups over the study period. Nearly every habitat supported some unique species. The faunal survey of velvet ants revealed minor variation in flight times between species and a unique pattern of temporal niche partitioning in one species. Habitat preferences were observed for seven of 42 velvet ant species at AMNWR. In a comparison of the diversity of velvet ants between AMNWR and the Nevada Test Site (NTS), six velvet ant species that occurred at the NTS were not found at AMNWR. Diagnoses and a key are provided for the velvet ants of AMNWR. This thesis increases the number of known terrestrial invertebrates at the refuge and provides a comparison of terrestrial insect distribution and habitat use at AMNWR. These investigations contribute to the goals and objectives of the U. S. Fish and Wildlife Service to obtain basic inventories and understand the terrestrial habitat use of invertebrates at AMNWR.

<p dir="ltr">Sepidiini is a large and morphologically diverse tribe of darkling beetles (Tenebrionidae) in the subfamily Pimeliinae, containing ~1,000 species and subspecies. Despite the presence of many large and charismatic species and the cultural significance of some of its members, the toktokkie beetles, this tribe has been lacking revision at all taxonomic levels. To develop a framework for taxonomic revisionary work, in Chapter 1, I reconstructed a phylogeny of the tribe using targeted enrichment sequencing data. I also scored a comprehensive suite of diagnostic characters for the tribe to test in a phylogenetic context. Based on the results of the study, I proposed revising the subtribe Oxurina Koch, 1955, <b>sens. nov. </b>(now containing the genera <i>Oxura </i>Kirby, 1918, and <i>Miripronotum </i>Louw, 1979) and moving the genera <i>Decoriplus </i>Louw, 1979, <i>Pterostichula </i>Koch, 1952, <i>Stenethmus </i>Gebien, 1937b, and <i>Synhimba </i>Koch, 1952 to a new subtribe Stenethina <b>subtr. nov. </b>The tree topology also supports revising or synonymizing the genera <i>Dichtha </i>Haag Rutenberg, 1871, and <i>Amiantus </i>Fåhraeus, 1870,<i> </i>and revising the genus <i>Somaticus </i>Hope, 1840.</p><p dir="ltr">In Chapter 2, I performed a partial revision of the subtribe Hypomelina Koch 1955. A new genus <i>Bufoniopsis </i><b>gen. nov. </b>is erected containing one newly described species <i>Bufoniopsis hypnosis</i> <b>sp. nov. </b>The genus <i>Hypomelus </i>Solier, 1843, and all of its species are redescribed, and three new species are described: <i>Hypomelus johnprinei</i> <b>sp. nov</b>., <i>Hypomelus lorettalynnae</i> <b>sp. nov.</b>, and <i>Hypomelus tomhalli</i> <b>sp. nov</b>. One species was moved from <i>Hypomelus </i>to <i>Triangulipenna</i><i> </i>Louw 1979<i> </i>and redescribed, <i>Triangulipenna vulipnus </i>(Haag Rutenberg, 1873)<i> </i><b>comb. nov. </b><i>Triangulipenna </i>was redescribed and three new species were described, <i>Triangulipenna tylerchildersi </i><b>sp. nov.</b>, <i>Triangulipenna dollypartonae</i> <b>sp. nov</b>., and <i>Triangulipenna ralphstanleyi</i> <b>sp. nov</b>. A revised key to the genera of Hypomelina is provided as well as keys to the species of <i>Hypomelus </i>and <i>Triangulipenna</i>.</p><p dir="ltr">Toktokkie beetles—members of the subtribe Molurina Solier, 1843—are known for their substrate tapping behavior which is a form of sexual communication in which beetles tap their abdomens on the ground to locate mates. Despite the fact that this behavior is well-known among people in southern Africa, very little research has been done on the topic. In Chapter 3, I document variation in tapping patterns across the subtribe Molurina. Three characters of tap trains were found to vary: length of tap trains, tapping rate, and tapping pattern. Ancestral state reconstruction was performed for each of the characters. Characters were found to be only partially linked to ancestry. Additionally, in any given locality sampled, no two species had the same “song”. This suggests that signal partitioning could be influencing species “songs”.</p>

<p>The tribe Edrotini is the largest component group of the largest tenebrionid subfamily, Pimeliinae, in the Americas, with 427 described species/subspecies in 55 genera. However, the group is taxonomically impeded, with the last comprehensive revision published nearly 115 years ago. This is particularly regrettable since members of this tribe are ubiquitous in arid regions throughout the Americas and are exceptionally diverse in their morphology and behaviors. To provide phylogenetic context and a foundation for taxonomic work, in Chapter 1 I sample a majority of genera and reconstruct the first phylogeny of the Edrotini, using targeted enrichment sequencing. My results indicate major changes are required to both edrotine tribal composition and generic concepts. In combination with a suite of eight morphological characters I use this phylogeny to reconstruct ancestral states and test for characters correlated with stridulation the tribe. I find stridulation is strongly correlated with two morphological characters and propose a defensive function for these structures</p> <p>In Chapter 2, I use the molecular phylogeny in combination with 100 morphological characters to evaluate all Edrotini genera and members of five related tribes with constrained parsimony analyses. Based on the results thirteen genera are transferred from the Edrotini and the tribal classification is revised, with 35 genera recognized and description of a further five recommended. One neotype and seven lectotypes are designated for type species. A dichotomous key to genera is provided. Thirty-one current genera are redescribed; two species described; and four genera described, including four species. One subgenus is elevated to genus and three genera are placed as subgenera pending a species-level revision of the clade. Six genera, four subgenera, and one species are synonymized. Eleven species are transferred to the correct genus and one replacement name proposed. </p> <p>And in Chapter 3, I revise the genus <em>Edrotes</em> LeConte to include eight species distributed across arid southwestern North America. All species are redescribed, of which three are brought out of synonymy. A neotype for <em>E. rotundus</em> (Say) is designated. The synonymic position of five species is amended. An illustrated key to <em>Edrotes</em> species is included. A molecular phylogeny of all species is generated and used to infer divergences dates and historical biogeography for the genus. The most recent common ancestor of <em>Edrotes</em> is dated to the late Miocene or early Pliocene and inferred to have inhabited the Sonoran Desert. </p>