Parasites often have profound effects on the survival and evolution of their hosts, and hence on the structure and health of entire ecosystems. Yet basic questions, such as the degree of virulence of a given parasite on its host, and factors influencing which hosts in a population are at the greatest risk of infection, are vexingly difficult to resolve. The western blacklegged tick-western fence lizard (Ixodes pacificus-Sceloporus occidentalis) system is important, primarily because I. pacificus, a vector of the Lyme disease spirochete Borrelia burgdorferi, is dependent on S. occidentalis for blood meals in its subadult stages, and this lizard possesses an innate immune response that removes the Lyme disease pathogen from attached ticks. My study focused on two aspects of the I. pacificus-S. occidentalis interaction. In Chapter 1, I investigated factors correlating with the intensity of I. pacificus infestations on S. occidentalis. Infection intensity (parasites per host) is often highly variable within a host population, though certain individuals, such as males, tend to be more heavily infected. Previous work in the I. pacificus-S. occidentalis system suggests that differences in behavior, such as the frequency of territorial patrols, may contribute to variation in tick intensity among lizards. I therefore hypothesized that lizard traits that correlate with dominance would also correlate with infestation intensity. Specifically, I predicted that larger and more colorful males would have higher infestation intensities than less impressive animals. In this chapter, I also focused on site selection by ticks infesting S. occidentalis. Skin folds on the necks of these lizards (nuchal pockets) may function to divert ectoparasites away from eyes, ears, and other potentially vulnerable structures. I therefore also looked for factors correlating with tick attachment in these pockets. I sampled ticks on adult male S. occidentalis in the spring and summer, which is the seasonal peak for both S. occidentalis territorial behavior and subadult I. pacificus abundance. After determining the site of infestation and intensity of ticks on these lizards, I re-infested lizards with laboratory-reared I. pacificus larvae, and again quantified tick intensity and attachment location. Contrary to expectation, no host traits correlated with tick intensity among ticks naturally infesting lizards, and lab-reared larval intensity was negatively correlated with lizard body size. As expected, ticks acquired by lizards naturally concentrated inside nuchal pockets, and I also observed this site preference among ticks in lab-based experimental infestations. Although the general pattern, lab-reared ticks were more varied in the sites on which they fed. There was a negative correlation between infestation intensity and the proportion of ticks attached in nuchal pockets. Unsurprisingly, the most reliable predictor of tick intensity and site selection was the season. In Chapter 2, I explored how tick attachment affects male S. occidentalis contest behavior. I. pacificus infestation has been shown to have negative physiological impacts on S. occidentalis, but mechanisms linking physiological changes to ultimate fitness consequences have been largely underexplored. I hypothesized that tick infestation reduces male S. occidentalis fighting ability by reducing O2 carrying ­­capacity­, or by obstructing or damaging vulnerable structures on their hosts. I held fifty half-hour trials between pairs of size- and ventral badge-matched male S. occidentalis, with one male in each pair infested with lab-reared I. pacificus larvae. I found that tick infestation negatively correlated with aggressive behavior in these staged contests. In support of reduced O2 capacity as the mechanism of reduced aggression, my ecologically relevant infestation intensities seemed to cause significant declines in hematocrit among experimentally infested lizards relative to controls. However, the site at which ticks attached did not significantly correlate with the aggressiveness of their lizard hosts. This is one of only a handful of studies to address the direct effect of I. pacificus on S. occidentalis. My study demonstrates that tick infestation can be detrimental to the fitness of their lizard hosts even without the transmission of pathogens. This insight may prove informative in future work on the ecology of I. pacificus-borne diseases in the western United States. This study is also one of only a few to use parasite infection to induce an asymmetry in fighting ability in intrasexual contests.

Investigating ecological questions at the scale of individual organisms is necessary to understand and predict the biological consequences of environmental conditions. For small organisms this can be challenging because we need tools with the appropriate accuracy and resolution to record and quantify their ecological interactions. Unfortunately, many of our existing tools are only appropriate for medium to large organisms or those that are wide ranging, inhibiting our ability to investigate the ecology of small organisms at fine scales. In Chapter 1, I tested a novel workflow for recording animal movements at very fine spatial and temporal scales. The workflow combined direct observation and the mapping of locations onto high-resolution uncrewed aerial vehicle (UAV) imagery loaded on hand-held digital devices. Observers identified landscape features they recognized in the imagery and estimated positions relative to those features. I found this approach was approximately twice as accurate as consumer-grade GPS devices with a mean and median error of 0.75 m and 0.30 m, respectively. I also found that performance varied across landscape features, with accuracy highest in areas that had more visual landmarks for observers to use as reference points. In addition to sub-meter accuracy, this method was cost-effective and practical, requiring no bulky equipment and allowing observers to easily record locations away from their own location. While this workflow could be used to record locations in a variety of situations, it will be most cost-effective when also using high-resolution environmental data from a UAV. In Chapter 2, I used the workflow described in Chapter 1 to investigate blunt-nosed leopard lizard (Gambelia sila) thermoregulation at fine-scales. Recent research has suggested that the spatial distribution of temperatures is important to consider for ectotherm thermoregulation, but this work has been limited to simple artificial environments. My goal was to investigate this idea in a complex natural system for the first time. I tracked lizard movement and body temperatures at a desert site from May to July 2021. I used machine learning to combine high-resolution environmental data from a UAV with microclimate temperature data to create a model of the spatial distribution of environmental temperatures over time. I found that including information about the spatial distribution of temperatures improved the models of lizard thermoregulatory accuracy and movement rate. Because these response variables are important aspects of ectotherm energetics, this suggests that the spatial distribution of temperatures may be an important, but often overlooked, component of habitat quality. Going forward, identifying better methods to quantify the spatial distribution of temperatures would provide insights into the specific responses of ectotherms to different spatial distributions. In this work I used recent technological advances in UAVs to investigate ecological questions at the scale of a small organism. The methods developed here provide insights into the importance of the spatial distribution of temperatures for a small ectotherm. Further efforts to develop, test and utilize tools for fine-scale ecological research will advance our ability to understand species’ interactions with current conditions and predict their responses to future changes.

No description available.

Mate competition is a prominent component of sexual selection theory. Typically, males attempt to mate with the most females possible and females attempt to mate with the highest quality males possible. In the polygynous female-defense mating system of Anolis carolinensis, males compete directly for females through territorial behavior. Inter-male competition is intense due to an average polygyny ratio of 1 male to 3 females despite a 1:1 adult sex ratio. Through high levels of territorial behavior (e.g., 100 displays/h, 27 m patrol distances/h, 70% of day in defense-related activities), males attempt to exclude other males from resident females who, in turn, both store sperm and ovulate a single-egg clutch at weekly intervals over a 4-month breeding season. Paternity of hatchlings in 16 naturally occurring breeding groups was analyzed to determine the extent to which the territorial resident male was able to prevent other males from fathering offspring of his resident females. Lizards residing in or neighboring a resident male's territory were collected and RAPD-PCR was used to determine the paternity of hatchlings. Of the 48 hatchlings from 26 females, resident territorial males fathered 52% of hatchlings; 15% were fathered by a male whose territory bordered that of the resident male and 21% were fathered by a smaller male living covertly within the resident male's territory. Paternity for the remaining 12% of hatchlings belonged to an unsampled male. Given that females mated with multiple males, laboratory-based controlled matings were conducted where females were sequentially paired with two males and RAPD-PCR was used to analyze which of the two males fathered the subsequent hatchlings to determine the mechanism of sperm precedence. Regardless of mating order, only one male of the pair fertilized the eggs. Male A. carolinensis have reproductive strategies present in addition to defending resident females and female A. carolinensis have options in addition to simply mating with the resident male. While sperm precedence is present in this species, it is not based on mating order, but may involve both the number of sperm deposited in the female's tract as well as the quality of those sperm. / Ph. D.

Adult green anoles, Anolis carolinensis, exhibit numerous sex differences resulting from divergent strategies for maximizing reproductive success. I focused on the ontogeny of sex differences in behavior in juveniles, in relation to adult sex differences, by documenting the behavior of free-ranging juveniles, examining the structure and use of headbobbing displays, and determining the role of the androgen testosterone (T) in producing behavioral sex differences. Field observations indicated that juvenile males eat and forage actively more often than juvenile females. This divergent feeding behavior may result from sexual selection, given that body size is a major factor in determining the reproductive success of males. Analyses of headbobbing displays, used by adults in aggressive and sexual interactions, revealed that juvenile males and females each give the same three A, B, and C display types described for adults. However, there may be a maturational component to display structure, as juvenile displays differ from those of adults in within-display temporal structure, and are not as stereotyped. Concerning display use, social context affects neither the types of display interactions observed nor the rates of displays and related behaviors. However, size affects nearly every aspect of display behavior. Both juvenile males and females show increased display rates and probabilities of expressing display-related behaviors with increasing body size, although in the largest juveniles, male display rates become higher than those of females. These results, like those from analyses of display structure, suggest a maturational component to display use, perhaps mediated by changes in the underlying motivational states of juveniles. Consistent with the divergence in display rates in large juveniles, males of approximately 30 d of age and older have higher plasma T concentrations than females. Furthermore, juvenile males and females that have been given T implants each respond with increased behavior levels, approaching those of breeding adult males. These analyses indicate that sexual dimorphisms in behavior in adults likely arise through underlying physiological differences between males and females that mediate the expression of behavior, rather than through fundamental sex differences in the ability to perform these behaviors. / Ph. D.

Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: In the present study I examined the phylogeography of the rupicolous Cape girdled lizard, Cordylus cordylus. Samples were collected across the species distribution range from 63 localities in the Eastern and Western Cape and Free State provinces of South Africa, yielding a total sample size of 207 specimens. Four DNA loci, two nuclear (PRLR, PTPN12) and two mitochondrial (16S rRNA, ND2), were sequenced. Bayesian inference, maximum likelihood and maximum parsimony methods were employed to test evolutionary relationships among populations, followed by population structure analyses, divergence time estimations and niche modelling. My results confirm the species monophyly and revealed the presence of two distinct clades. Clade 1 comprised specimens from the western and southern portions of the Western Cape coast, while clade 2 comprised specimens from the southern and eastern Cape coast and adjacent interior of the Eastern and Western Cape and Free State provinces. An area of sympatry between the two clades was observed in the Breede river valley. The divergence time estimates revealed an Early Pliocene (4.31 Ma), Late Miocene (6.01 Ma) divergence for each of the two clades retrieved. Phylogeographic data suggest that clade 1 is younger (lower haplotypic and nucleotide diversity), in comparison to clade 2. Furthermore, the niche modelling shows that C. cordylus occupies a wide range of unfavourable habitats. The absence of marked phylogeographic patterning within clades is very uncharacteristic for a rupicolous vertebrate species. The ecological pliability and generalist nature of C. cordylus presumably contributed to the observed phylogeographic pattern and have facilitated the absence of within clade differentiation. Moreover, I suggest that microclimatic variables, rather than geographic barriers influence the genetic structuring of C. cordylus.

Loess is a silty material produced via glacial grinding and its deposition is usually associated with colder periods when there is a considerable increase in atmospheric dust transport. The magnetic susceptibility of loess-palaeosol sequences has been the subject of extensive research within palaeoclimatology as it has proven to be reliable climate proxy. Most research on magnetic susceptibility has been focused on thick deposits found in Europe and Asia, in particular the Chinese Loess Plateau where it is possible to discern more than 2.5 million years of climate variation in deposits several hundred meters thick. The climate development throughout the Quaternary has been characterized by glacials where there has been a significant ice growth, and interglacials where ice sheets have retreated. The Last Glacial Maximum was the most recent time of ice sheets reaching their maximum extent, occurring at some point between 26.5 and 18 ka BP. Aeolian reworking of material produced in conjunction with the Irish Sea Ice Stream (ISIS) occurred approximately 5000 years after the LGM. The ISIS limit were located just north of the Lizard Peninsula, and impinged on the Isles of Scilly. Loess deposits in SW England has largely been ignored in previous research in favour of the previously mentioned Chinese loess, but may be important climate archives in understanding the climate development of the North Atlantic. As such, the scope of this thesis is investigating the magnetic susceptibility of loess deposits in Cornwall and the Isles of Scilly in the far SW England. This is done by analyzing 36 samples from Lowland Point and 24 samples from Chynhalls Point, both located at the Lizard Peninsula in Cornwall, as well as 20 samples from Porthcressa and 14 samples from Gimble Porth, both located at the Isles of Scilly. The instruments used for analysis are the Bartington MS2B and the Agico MFK-1 FA Kappabridge, where all 96 samples were analyzed on the former while the 36 Lowland Points samples and the 20 Porthcressa samples were also analyzed on the latter. The Bartington MS2B operates at two frequencies, 465 Hz and 4650 Hz, while the Agico MFK-1 FA Kappabridge operates at three frequencies, 976 hz, 3904 Hz, and 15616 Hz. The results from this study suggest that the Lizard Peninsula sample sites exhibit signs of magnetic enhancement through pedogenic processes while the sample sites at the Isles of Scilly consist of mainly unaltered material, likely due to an overlying sollifluction unit inhibiting magnetic enhancement. As a consequence of the different frequencies, the results were lognormalized to permit a direct comparison between the instruments. While the variability in magnetic susceptibility was strikingly similar, the results of the Kappabridge were shifted to ~20% higher values than those of the Bartington which may be attributed to the Kappabridges higher precision when analyzing weakly magnetic material. A similar shift is not evident when measuring the calibration samples of each instrument. In determining the mean annual precipitation after deposition, the Lizard Peninsula samples sites suggest an exposure to an annual precipitation between 300-400 mm/yr, compared to modern rainfall between 900-1000 mm/yr. The data from the Isles of Scilly were insufficient for calculating the mean annual precipitation due to the lack of evidence for magnetic enhancement. / Loess är benämningen för material i siltstorlek som produceras då glacial aktivitet maler ned den underliggande berggrunden. Detta material är tillräckligt litet för att fångas upp av vindar och därefter transporteras i atmosfären, för att därefter avsättas då vindhastigheten minskar. Under kalla perioder i jordens historia ökar denna transport då vindhastigheten ofta varit högre än under varma perioder. Under de varmare perioderna och med tiden så utsätts loess för jordmånsbildande processer och bildar palaeosoler. Den magnetiska susceptibiliteten hos loess-palaeosol-sekvenser har under flera årtionden varit ämnet för utbredd forskning då detta har visat sig vara en tillförlitlig klimatproxy för att förstå klimatutvecklingen under jordens gångna historia. I synnerhet har loess-palaeosol-sekvenser i Kina varit av intresse då det går att utröna mer än 2,5 miljoner år av klimatutveckling i avlagringar som på sina platser är flera hundra meter tjocka. Under den senaste istidens maximala utbredning mellan 26500 och 18000 år sedan passerade en isström över den Irländska sjön (Irish Sea Ice Stream, sammankopplad med inlandsisen), förbi Cornwalls norra kust och stötte samman med Scillyöarna. Det tog sedan ungefär 5000 år tills vindtransport av avsatt material påbörjades. Loessavlagringar i sydvästra England har känts till under årtionden men har till stor del ignorerats till fördel för de tidigare nämnda avlagringarna i Kina, men de kan agera som viktiga klimatarkiv för tiden efter den senaste istiden och därmed klimatutvecklingen i Nordatlanten. Denna rapport undersöker således den magnetiska susceptibiliteten i loess från två platser på Lizard-halvön i Cornwall, och två platser på Scillyöarna. Totalt analyserades 96 prover, 36 av dem från Lowland Point, 24 av dem från Chynhalls Point, båda belägna på Lizard-halvön, samt 20 prover från Porthcressa och 14 prover från Gimble Porth, båda belägna på Scillyöarna. Resultaten från denna studie påvisar att provplatserna på Lizard-halvön visar tecken på magnetisk förstärkning på grund av jordmånsbildande processer, medan provplatserna på Scillyöarna huvudsakligen består av oförändrat material, troligen på grund av att dessa täckts av annat material under isens tillbakadragning, vilket förhindrat jordmånsbildning. Resultaten från Lizard-halvön möjliggör beräkning av årsmedelnederbörden sedan materialet först avsattes, vilket beräknades till mellan 300-400 mm/år jämfört med modern årsmedelnederbörd mellan 900-1000 mm/år. Dessvärre möjliggjorde inte resultaten från provplatserna på Scillyöarna en liknande beräkning.

Context. In recent years, Software Quality Assurance (SQA) has become a crucial part of software development processes. Therefore, modern software development processes led to an increase in demand for manual and automated code quality assurance. Manual code quality reviews can be a time-consuming and expensive process with varying results. Thus, automated code reviews turn out to be a preferred alternative for mitigating this process. However, commercial and open-source static code analyzer tools often offer deep analysis with long lead time. Objectives. In this thesis work, the main aim is to introduce an early code quality assurance tool, which features a combination of software metrics. The tool should be able to examine code quality and complexity of a telecommunication grade software product such as source code of specific Ericsson product by Ericsson. This tool should encapsulate complexity and quality of a software product with regards to its efficiency, scope, flexibility, and execution time. Methods. For this purpose, the background section of the thesis is dedicated to in-depth research on software metrics included in well-known static code analyzers. Then, development environment, under investigation source code of Ericsson product, and collected software metric for evaluation were presented. Next, according to each software metric’s characteristics, point of interest, and requirement, a set of steps based on a Susman’s action research cycle were defined. Moreover, SWAT, a suitable software analytics toolkit, employed to extract conducted experiment data of each software metric from a static analyzer code named Lizard in order to detect most efficient software metrics. Outcome of conducted experiment demonstrates relationship of selected software metrics with one another. Results. The chosen software metrics were evaluated based on a variety of vital factors especially actual data from number of defects of specific Ericsson product. Highly effective software metrics from investigations in this thesis work were implemented as a new model named hybrid model to be utilized as an early quality assurance. Conclusions. The proposed model, which consist of well-performing software metrics, demonstrate an impressive performance as an early code quality indicator. Consequently, the utilized model in this master thesis could be studied in a future research to further investigate the effectiveness and robustness of this tool an early quality assurance.

We studied the Flat-tailed Horned Lizard, Phrynosoma mcallii, and the Sonoran Horned Lizard, P. goodei, in an area of narrow sympatry near Yuma, Arizona, and found they overlapped broadly in use of available food resources, body size, and growth rates. We compared diet (Chapter 2), growth and reproduction (Chapter 3), and survivorship (Chapter 4) of P. mcallii and P. goodei during two years of drought followed by a year of higher-than-average rainfall. We predicted that P. mcallii would be more tolerant of drought conditions than its congener, since P. mcalliiis found only in an extremely arid region while P. goodei is part of a more northerly-clade and that P. goodei is excluded from the sandy habitat of P. mcallii due largely to the paucity of rainfall and the lack of moisture-holding ability of the sand. During the extended period of drought food became limited, horned lizards lost mass, and there was less growth and reproduction. While both species showed strong differences between dry and wet years, the within-year differences between species were generally small. When resources were abundant we witnessed rapid growth for both P. mcallii and P. goodei and a much higher rate of reproduction. While both species showed similar growth patterns, P. mcallii had a smaller adult body size, larger hatchling size, and seasonally-delayed reproduction compared to P. goodei; differences we propose are adaptations for frequent periods of drought, but which may be disadvantageous in years of abundant resources. We used mark-recapture analysis to derive estimates of detection probability and survival rates. Contrary to prediction, survival rate estimates were higher for P. goodei than for P. mcallii, and higher in dry years than in wet years. However, decreased probability of detection and increased emigration in the wet year confounded survival rate estimates. We also reviewed the conservation and management of P. mcallii, a species of conservation concern (Chapter 5). We proposed using fine-scale scat surveys rather than mark-recapture surveys for the long-term monitoring of P. mcallii.

Survival is a prerequisite for successful reproduction, and is thus intertwined with fitness. Some physiological systems can improve survival, like the immune system, but compete with other processes for resources. Because animals evolved with resource limitation, it is important to understand how these resource-allocation decisions are made. To meet this end, I performed four investigations addressing how life-history characteristics shift in side-blotched lizards (Uta stansburiana) using laboratory studies and multi-year field sampling. First, I measured metabolic rates in response to different immune challenges and different energy states in male lizards. I found that, surprisingly, cutaneous biopsies were associated with a downregulation of metabolic rate, lipopolysaccharide injection did not affect metabolic rate, and the fastest-healing individuals had the largest decrease in metabolic rate, regardless of feeding treatment. In my second experiment, female lizards were challenged with a cutaneous biopsy and follicle-stimulating hormone to force a trade-off between the reproductive and immune systems. I learned that follicle-stimulating hormone increased metabolic rates and cutaneous biopsies decreased them, and that a combination of both treatments reduced metabolic rate the most. In my third experiment, I sampled wild lizards from northern populations in eastern Oregon and southern populations in southern Utah. In the first year, longer-lived northern lizards had higher circulating corticosterone concentration, decreased reproductive investment, and increased microbiocidal ability, but the subsequent year did not hold to these trends. A subsequent common-garden experiment revealed that southern lizards were faster at healing cutaneous wounds, but lost much more body mass than the northern individuals, which healed more slowly but maintained body mass. Finally, I have conducted a 5-year field investigation addressing the life-history trade-offs associated with urbanization, which reveals interesting effects of precipitation on survival and reproductive effort at urban and rural sites.