The Aderidae (Coleoptera: Tenebrionoidea), commonly known as the puppet beetles or ant-like leaf beetles, is a moderately diverse beetle family with approximately 1,000 described species. Most species are small in size (1–4mm) and little is known regarding their natural history though adults are generally active on the surfaces of foliage, often collected using beating and sweeping methods. The family is widely distributed biogeographically and many species exhibit secondary sexually dimorphic characters of their antennae and hind legs. The characterization and identification of tribes, subtribes, and genera of Aderidae from all biogeographic regions is incredibly difficult and previous authors have expressed the need for a worldwide revision of the higher classification for the group – the puppet beetles are one of the taxonomically and systematically poorest known families in all of Coleoptera. The primary goal of this study was to reconstruct the evolutionary history of the beetle family Aderidae worldwide and examine the evolution of sexually dimorphic characters using current phylogenetic methods while incorporating both morphological and molecular data.

First, the Aderidae of New Zealand were revised in order to understand intraspecific morphological character variation and examine the structural diversity of sexually dimorphic features within the genus Zenascus. Three new genera and six new species were described and keys to the genera and species of the region were presented. The phylogenetic relationships of species in the genus Zenascus were reconstructed using parsimony and a correlation between gland pore abundance of the hind legs and the degree of antennal modification of males was found. Intraspecific variation of morphological characters that were previously used in higher classification of the Aderidae indicated that these character systems may be evolutionarily labile and not very useful for higher-level aderid classification.

These intraspecific findings were then used to inform the following two studies, which examined the Aderidae at the generic level using molecular and morphological data respectively. The molecular study represented the first comprehensive phylogenetic analysis of the puppet beetles that used over 150 exemplars representing all four tribes and eight aderid subtribes. The resulting phylogenies were used to characterize and establish major clades within the family and provide a robust phylogenetic classification for the group. Nomenclature changes include parsing the family into two subfamilies, the Aderinae and Cnopinae, and further dividing the Aderinae into five highly supported clades, which are designated as tribes (Aderini, Gompeliini, Euglenini, Phytobaenini, and Syzetonini). Ancestral state reconstructions for morphological characters that have been previously used to establish taxonomic relationships were evaluated within this phylogenetic context. The character of elytral pubescence was found to be phylogenetically informative for several clades whereas the degree of posterior cephalic constriction did not support major groupings. Metafemoral modifications provided phylogenetic signal at the generic level. Additionally, character mapping of sexually dimorphic characters was performed to investigate the evolutionary pathways of these character systems and determine whether dimorphism has arisen through character state gain in males, character state loss in females, or both. Sexual dimorphism of the hind legs appears to have a single evolutionary origin within the Aderidae, with an ancestral state of sexual dimorphism for the Syzetonini clade. Secondary loss of this dimorphism has occurred twice and resulted from both loss of the trait in males leading to monomorphism without ornamentation, and gain of the trait in females leading to monomorphism with ornamentation in both sexes.

Finally, the world genera of Aderidae were reviewed and redescribed and an identification key provided for all extant, adult genera. This study summarized the currently known taxonomic information, clearly defined the morphological limits of each aderid genus, and reassigned previously described species into established genera to enable future taxonomic work. Scanning electron micrographs for all main body regions were provided for the majority of genera in order to enable future character and state recognition and specimen identification. Three genera were newly described while twelve genus-level synonymies were established for the family. Additionally, a total of 362 generic reassignments were established based on type material examined.

The current natural history knowledge for aderid species is extremely limited, so in order to disentangle the ecological, behavioral, and biogeographical processes that underlie the macroevolutionary patterns presented here, additional life history and alpha-level taxonomic studies are required. The phylogenetic framework provided within these works will serve as a backbone for these future studies and will hopefully ignite additional research on the puppet beetles.

Interest in the California Floristic Province as a study region for scientists interested in biodiversity, evolution, systematics, and phylogeography has been increasing over the last several years. The amaurobiid spider genus Callobius (Chamberlin) occurs throughout the Northern Hemisphere, but is particularly common in western North America and particularly diverse in the California Floristic Province. An understanding of the evolutionary history of Callobius would contribute a great deal to this growing body of work, but the genus has received little attention since being revised in 1972. Since that time, a great deal has changed in the way biodiversity is studied. The Hennigian revolution changed the fundamental framework of systematics, and molecular techniques based on DNA sequences have brought enormous inferential power to bear on questions of systematics and population genetics, as well as almost every other discipline in organismal biology.

In my doctoral work I have approached Callobius in the context of phylogenetic systematics and the biogeography of the California Floristic Province. First, I have taken a broad approach to the morphology of Callobius, particularly the morphology of the copulatory organs, as it might apply to cladistic investigations both within Callobius and more broadly among spiders. Second, I have undertaken a phylogenetic analysis of the subfamily Amaurobiinae, to illuminate evolutionary pattern within Callobius as well as between Callobius and other amaurobiine genera. And thirdly, I have used geometric morphometrics and population genetics to look for divergence within the widespread species Callobius severus.

I have produced an atlas to the genitalic morphology of Callobius, and propose a morphological diagnosis that is more consistent with principles of homology than those previously in use. I have also found and described the male of Callobius pauculus, previously known only from females. Although my phylogenetic analysis did not resolve the relationships between the amaurobiine genera, it does offer some support for the monophyly of Callobius and identifies a clade of California Floristic Province neoendemic species. My study of Callobius severus supports infra-specific structure, and suggests that the geography of the California Floristic Province is influencing the evolution of Callobius in patterns similar to its influence on other taxa.

Abstract Comparative Biogeography of Dune-Restricted Insects in the Desert Southwest by Matthew H Van Dam Doctor of Philosophy in Environmental Science, Policy and Management University of California, Berkeley Professor Kipling W. Will, Chair

The focus of my thesis is on the biogeography of dune-restricted insects, there are three main inquiries I addressed in my thesis: (1) Testing the effect distance may have on dispersal in the sand dunes fauna of the desert Southwest of North America. (2) A statistical evaluation of phylogenetic signal contained in different character systems in Trigonoscuta, using novel reproductive characters to delimit species. (3) Incorporating more precise natural history data into niche modeling and an evaluation of its effect on adaptation to climate change in the giant flower-loving flies Rhaphiomidas.

(1) I am specifically testing the hypothesis that Pleistocene river corridors mediated dispersal among sand dune-restricted taxa and aquatic taxa found in desert springs. From the seven different genera of dune restricted taxa and four aquatic desert taxa I examined, it appears that some taxa have used river corridors of sandy habitat to disperse. Although the patterns are largely concordant they differ markedly in dispersal times. Some having long distance dispersal occurring in the Pleistocene while others only disperse at the local level having disjunct distributions the result of much older vicariance events. Using both the phylogenetic pattern and date estimates allows for my conclusions not to be drawn from pattern alone. In addition I examined the effect distance has on jump dispersal and vicariance. I incorporated GIS estimates of dispersal distance in a connectivity matrix to model the connectivity of river corridors. Model selection was then used to see if the constraints and distance effect improved the fit of the model to the data, indicating if there was an actual effect of the Pleistocene river corridors used as dispersal pathways. This application of patterns, distance and dates allows for more statistical testing of biogeographic hypothesis and has implications for biogeography as a whole to explicitly test the effect of distance on biogeographic reconstructions.

(2) The genus Trigonoscuta consists of 65 described species with 98 subspecies. I investigated the validity of these species using a combined approach of morphology and molecular data to be able to properly delimit and reliably identify the species. For delimiting the species I examined the morphology of the male endophallus. The endophallus is the internal structure of the male genitalia, which is everted during mating. I derived a novel technique to evert the endophallus consistently. The characters of the endophallus alone repeated the relationships seen in the molecular data and providedcharacters to delimit and describe species using morphological criteria. Using the consistency index (CI), I evaluated the contribution of each morphological character across all the equally most parsimonious trees. I then used this data to see if there was statistical support for which character system provided more phylogenetic resolution. This supported the idea that using the endophallus outperformed more traditional characters and should be utilized in phylogenetic reconstruction more often.

(3) I explored how partitioning climate data by day influences niche model predictions, as estimated by the MaxEnt machine-learning algorithm, for taxa with constrained phenologies. I utilized the giant flower-loving flies Rhaphiomidas because of their discrete phenologies as my focal taxon. I compared the results of partitioned by day data with WorldClim data that is partitioned by moth to examine what effect using more precise data has on species distribution models. I also examined how phylogenetic signal in both life history traits and climate tolerances can be used to identify how they adapt to different climates.