Sperm structure in five species of cicadine cicadas (Albanycada albigera, Azanicada zuluensis, Platypleura capensis, P. hirtipennis and Pycna semiclara) and five species of cicadettine cicadas (Melampsalta leucoptera, Quintilia walkeri, Stagira simplex, Xosopsaltria thunbergi and Monomatapa matoposa) was investigated by light and electron microscopy. In addition, spermiogenesis in cicadas was described; the information was derived from two cicadettines (Diceroprocta biconica and M. matoposa) and three cicadines (Kongota punctigera, P. capensis and P. semiclara). Mature spermatozoa of all species investigated are elongate and filiform, consisting of three distinct regions: the head (acrosome and nucleus), mid-piece and tail. All species produce more than one discrete length of nucleated, motile sperm, a form of sperm polymorphism termed polymegaly. Polymegaly is expressed in three ways; sperm have uni-, bi- or trimodal nucleus and tail lengths. Besides the differences in length, there are also notable differences in the size of nuclei. The anterior parts of sperm heads are embedded in an elongate homogenous matrix forming spermatodesmata. The conical acrosome is deeply invaginated posteriorly, and sits on top of the nucleus. The acrosomal contents are differentiated internally with a tubular substructure and a subacrosomal space. The anterior of the nucleus intrudes into the posterior section of the subacrosomal space. Anteriorly the acrosome is laterally flattened; posteriorly it extends as two tubular processes on either side of the nucleus that gradually decrease in diameter. The homogenously electron-dense nucleus is pointed anteriorly and is generally cylindrical, although posteriorly there is a lateral invagination that extends part-way along the nucleus. This invagination houses fine granular material of the putative centriolar adjunct which does not form in close proximity to the centriole and hence may not be a true centriolar adjunct. The lamellate disposition of the centriolar adjunct material within the sperm-midpiece of cicadettine cicadas is distinct, and separates these cicadas from their cicadine counterparts in which the centriolar adjunct material is non-lamellate. Vesicle-like elements that are associated with both the posterior nucleus and the centriolar adjunct are also found within the invagination. Immediately posterior of and adjoining the centriolar adjunct is a pair of mitochondrial derivatives that are elongated and extend for almost the entire length of the tail. Except for size the architecture of short and long spermatozoa is generally similar in all species. The absence of accessory bodies in cicada sperm suggests that within the Cicadomorpha, the families Cicadidae and Cercopidae are closely related. Only long nuclei were observed in the fertilized eggs of A. zuluensis indicating that sperm with long nuclei might be favoured for fertilization. Spermiogenesis involves: (a) development of the acrosome from a proacrosomal granule; (b) development of the nucleus, characterized by elongation and streamlining with a simultaneous condensation of chromatin; (c) development of the axoneme from the centriole; (d) amalgamation of individual small mitochondria to form elongated mitochondrial derivatives in which cristae are arranged into regularly spaced lamellae; and (f) elimination of cytoplasm. The presence of a manchette, a transient microtubular organelle, which surrounds the acrosome, nucleus and mitochondrial derivatives, is a characteristic feature of spermiogenesis. The gross morphology of the reproductive tract in both male and female cicadas exhibits an organization similar to that in most oviparous insects. The non-functional spermatheca is the only exceptional feature in the female reproductive tract. Its role has been taken over by the common oviduct which, subsequently, has become modified into a swollen, differentiated structure with a dual role of receiving oocytes from the paired ovaries and storage of spermatozoa. Testis mass varies between cicada species; this variation might be linked to the intensity of sperm competition which has been found to be positively correlated with relative investment in spermatogenesis. Based on the preliminary findings of this study, K. punctigera, with its larger testis relative to body size, would be the ideal candidate to show the greatest levels of sperm competition. Accessory glands in both male and female A. zuluensis, D. biconica, P. hirtipennis and O. quadraticollis are very long; this character might be of phylogenetic significance. Despite being notoriously refractory spermiocladistics is potentially valuable in systematic and phylogenic studies of cicadas, especially at the subfamily level.

With our contemporary biota under increasing threat of extinction, it is of interest to understand where, why and how biological diversity is generated. If focussed on appropriate taxa, phylogeographic and phylogenetic studies can assist in the identification of both places and processes central to the origin and maintenance of biological diversity. It is explained why southern Africa presents a perfect test-bed for exploring such mechanisms of diversification and why cicadas (Hemiptera: Cicadidae) have proved very suitable tools for studies of historical biogeography. This study then exemplifies these points by providing the first large-scale investigation of the historical biogeography of the tribe Platypleurini Schmidt, 1918, with emphasis on the genus Platypleura Amyot & Seville, 1843 in southern Africa. Standard methods of DNA sequencing provided data from portions of the mitochondrial small subunit ribosomal 16S RNA (16S) and cytochrome oxidase subunits I (COI) and II (COII); and the nuclear elongation factor 1 alpha (EF-1α) from 400 ethanol-preserved specimens. These data were analysed using standard phylogenetic methods and a time scale of diversification was estimated using a Bayesian framework and both fossil data and DNA substitution rates. The results showed that the tribe is too recent to be of Gondwanan origin. The lack of monophyly of the genera represented in both Asia and Africa showed that the tribe needs formal taxonomic revision. Diversification of the African platypleurine genera coincides with aridification in the early Oligocene. Dispersal of Asian platypleurine taxa coincides with the meeting of Africa and Eurasia in the mid-Oligocene. Two radiations within African Platypleura are hypothesised; one distributed over most of sub-Saharan Africa and the second restricted to southern Africa, with clades restricted within regional biomes. Within each of the three focal biomes, cryptic taxonomic diversity was confirmed, suggesting that, even in relatively well understood groups such as the southern African platypleurine cicadas, molecular data can identify further diversity. Although each focal taxon was restricted to non-overlapping biomes, comparison of the three biomes highlighted interactions between palaeoclimates and fixed landscape features (coastal topography, river catchments and escarpments) as causative agents of vicariance, dispersal, extinction and diversification of these volant insects. The results of using co-distributed species for comparative study cautions against making inferences based on single-taxon datasets and highlights the need to use many, evolutionarily independent taxa when identifying mechanisms of diversification. The dating analyses imply that within-species lineage diversification occurred overwhelmingly within the Pleistocene, a trend that is being increasingly recognised in print for other biota. Some caveats about using phylogenetic approaches to estimate ancestral areas are illustrated. Several recommendations are made regarding additional taxa and data sources for understanding the origin and maintenance of biological diversity.

In 2021, 17-year periodical cicadas (Magicicada spp.) in Brood X emerged throughout much of the eastern United States. Due to their abundance and lack of predator avoidance during their short six-week period on the above-ground landscape, many species of animals consume cicadas and in turn have short-term habitat use and population responses. The response of bats to periodical cicadas is largely unknown. Using a before-after-control-impact study design, we deployed ultrasonic acoustic detectors during the summers of 2020–2022 within and just outside the range of the cicada emergence along the Potomac River corridor of Maryland and Virginia. For total bat activity, there was significantly more acoustic activity within the range of the cicada emergence the year during and the year following emergence than where emergence did not occur. Additionally, for all individual species and phonic groups, bat activity increased the year during and year after the emergence within the range of periodical cicadas compared to the year prior to emergence. Our study demonstrates that periodical cicadas may serve as a resource pulse that can cause an increase in bat activity and suggests at least a short-term local to regional population increase from immigration the year during and from recruitment the year following. Bats of the mid-Atlantic have been negatively affected by numerous stressors including white-nose syndrome, wind turbine development, and land use change. One species suffering widespread population decline over the last decade in the mid-Atlantic has been the little brown bat (Myotis lucifugus). Understanding the space use and resource selection of remaining populations is critical for management and species recovery. Nevertheless, there are difficulties with studying the movement of insectivorous bats due to their small size and high vagility. Traditional foraging studies of Myotis spp. have used fixed station telemetry and associated multi-azimuthal observations to estimate the location of individuals. However, single azimuth observations are often unable to be included in these studies leading to more potential location estimates missing from movement models. During the summer of 2022, I radio-tagged little brown bats from a maternity colony along the Potomac River in Maryland and collected bearings on nine individuals using fixed station telemetry from sunset until bats returned to the maternity roost. Location estimates were obtained from both single and multi-azimuthal observations. I fit movement models for each individual and used auto-correlated kernel density estimation (AKDE) to estimate space use. I also assessed second and third order habitat selection of individual bats. The average 95% AKDE for males was 889 ha ± 424.6 and 699.3 ha ± 129 for adult females. I found bats had higher predicted use of habitat that was closer in distance to water and wetlands and further from open habitat at the second order and closer to water and open habitat at the third order of resource selection. Habitat associations were similar to previous little brown bat resource selection studies. However, estimates of space use were larger than those using other movement models. This study was novel in that it incorporated single azimuth observations and periodicity into models of space use, thereby increasing sample size and offering a new framework for future telemetry studies. These estimates of space use can be used by land managers to protect the habitat of this imperiled species. / Master of Science / In 2021, 17-year periodical cicadas (Magicicada spp.) in Brood X emerged throughout much of the eastern United States in high abundance. As an abundant food resource, many animals consume periodical cicadas while they are above ground during the summers. This has allowed for short-term population increases and changes in reproduction in some species. However, the response of bats to periodical cicadas is largely unknown. To study this, I deployed ultrasonic acoustic detectors that recorded six species/species groups of bats during the summers of 2020–2022 within and just outside the range of the cicada emergence along the Potomac River Corridor, Maryland, and Virginia. I observed significantly more total bat acoustic activity within the range of the cicada during the emergence year and the year following than where emergence did not occur. Also, for all individual species and species groups, bat activity increased in the year during and after the emergence within the range of periodical cicadas compared to the year before emergence. My study demonstrates that periodical cicadas cause an increase in bat activity and may suggest a population increase in the areas where the emergence has occurred. Bats of the mid-Atlantic have experienced population declines due to numerous stressors including white-nose syndrome caused by the fungus Pseudogymnoascus destructans, wind turbine development, and habitat fragmentation. One species that has been heavily affected by white-nose syndrome is the little brown bat (Myotis lucifugus). To help conserve the species, understanding their home range and their resource needs is critical. However, little brown bats are small and agile fliers which poses difficulties for tracking them. Traditional foraging studies of species in this genus have used telemetry to obtain triangulations and location estimates of individuals but often leave out data collected from a single bearing as it is difficult to calculate the true location of the animal. During the summer of 2022, I radio-tagged little brown bats from a maternity colony along the Potomac River in Maryland and collected bearings on nine individuals using fixed station telemetry from sunset until bats returned to the maternity roost. We estimated locations from both triangulations and single bearings. I calculated movement models for each individual and fit them to an auto-correlated kernel density estimator (AKDE) of space use. The average 95% auto-correlated kernel density estimates of space use (AKDE) for males was 889 ha ± 424.6 and 699.3 ha ± 129 for adult females. I also assessed the habitat selection of individuals within their home range and across the nearby landscape. I found that bats preferred to use areas closer to water and wetlands and preferred to use areas closer to open habitat within their home range but preferred to use areas further from open habitat across the landscape. Habitat associations were similar to previous little brown bat resource selection studies. However, estimates of space use were larger than those using other movement models. My study was novel in that it incorporated single azimuth observations and periodicity into models of space use, thereby increasing sample size and offering a new framework for future telemetry studies. These estimates of space use can be used by land managers to protect the habitat of this imperiled species.

Models that are used for predicting species' potential distributions are important tools that have found applications in a number of areas of applied ecology. The majority of these models can be classified as correlative, as they rely on strong, often indirect, links between species distribution records and environmental predictor variables to make predictions. Correlative models are an alternative to more complex mechanistic models that attempt to simulate the mechanisms considered to underlie the observed correlations with environmental attributes. This study explores the influence of the type and quality of the data used to calibrate correlative models. In terms of data type, the most popular techniques in use are group discrimination techniques, those that use both presence and absence locality data to make predictions. However, for many organisms absence data are either not available or are considered to be unreliable. As the available range of profile techniques (those using presence only data) appeared to be limited, new profile techniques were investigated and evaluated. A new profile modelling technique based on fuzzy classification (the Fuzzy Envelope Model) was developed and implemented. A second profile technique based on Principal Components Analysis was implemented and evaluated. Based on quantitative model evaluation tests, both of these techniques performed well and show considerable promise. In terms of data quality, the effects on model performance of false absence records, the number of locality records (sample size) and the proportion of localities representing species presence (prevalence) in samples were investigated for logistic regression distribution models. Sample size and prevalence both had a significant effect on model performance. False absence records had a significant influence on model performance, which was affected by sample size. A quantitative comparison of the performance of selected profile models and group discrimination modelling techniques suggests that different techniques may be more successful for predicting distributions for particular species or types of organism than others. The results also suggest that several different model design! sample size combinations are capable of making predictions that will on average not differ significantly in performance for a particular species. A further quantitative comparison among modelling techniques suggests that correlative techniques can perform as well as simple mechanistic techniques for predicting potential distributions.