White, Bradley N

Line transect sampling and photographic-identification (photo-ID) are common survey techniques for estimating the abundance and distribution of cetaceans. Combining these approaches in the field ('combined LTPI' surveys) and using data from both components has the potential for generating comprehensive ecological knowledge that can be far more valuable than when these techniques and their data are used independently. In this thesis, I evaluated the results and conclusions from these two methods, used singly and in tandem, by investigating the population dynamics of two humpback dolphin (Sousa chinensis spp.) populations: the large and widely distributed Chinese white dolphin (S. c. chinensis) of the Pearl River estuary (PRE), and the small and geographically isolated subspecies of Taiwanese white dolphin (S. c. taiwanensis) in the eastern Taiwan Strait. Data from combined LTPI surveys in Hong Kong waters, at the eastern edge of the PRE, revealed a shift in space use with individuals spending less time in these waters than at the start of surveys. Data from combined LTPI surveys in Taiwan provided further support for a subspecies restricted to the central western waters, and identified a commonly used area at the northern part of their limited range. These two case studies demonstrated an overall efficacy of combined LTPI surveys in ecological studies of cetaceans. However, a multi-criteria analysis revealed that combined LTPI surveys with a line transect focus (e.g., Hong Kong) performed better than a LTPI survey with a photo-ID focus (e.g., Taiwan) when considering ecological aspects of the study populations, labour and data requirements, and ecological output. Even so, the photo-ID focus of Taiwan's monitoring program led to better assessments of individual space use patterns, likely helped by the Taiwanese white dolphin population's smaller size and intensive photographic effort. In both cases, the ecological output of combined LTPI surveys could be improved by expanding the study area or extending the field season or frequency of surveys. Overall, I showed that by following a set of general guidelines, different iterations of the combined LTPI approach (i.e., photo-ID focus or LT focus) can serve as powerful tools for uncovering multi-dimensional ecological information on cetaceans.

Author Keywords: abundance, cetacean, distribution, line transect sampling, multi-criteria analysis, Photo-ID

Interactions between hosts and pathogens play a crucial role in their adaptation, evolution and persistence. These interactions have been extensively studied in model organisms, yet it is unclear how well they represent mechanisms of disease response in primary vectors in natural settings. The objective of my thesis was to investigate host-pathogen interactions in natural host populations exposed to raccoon rabies virus (RRV). RRV is endemic to North America, that causes acute encephalopathies in mammals and is commonly regarded as 100% lethal if untreated; however variable immune responses have been noted in natural reservoirs. In order to further understand variable immune responses to RRV, my thesis examined (i) potential immunogenetic associations to RRV using genes intimately associated with an immune response, (ii) the nature of immune responses triggered in the host after infection, and (iii) viral expression and genetic variation, to provide insight into factors that may influence RRV virulence.

Immunogenetic variation of RRV vectors was assessed using major histocompatibility complex (MHC) DRB alleles. Associations were found between specific MHC alleles, RRV status, and viral lineages. Further, similarities at functionally relevant polymorphic sites in divergent RRV vector species, raccoons and skunks, suggested that both species recognize and bind a similar suite of peptides, highlighting the adaptive significance of MHC and contemporary selective pressures.

To understand mechanisms of disease spread and pathogenesis, I screened for variation and expression of genes indicative of innate immune response and patterns of viral gene expression. RRV activated components of the innate immune system, with transcript levels correlated with the presence of RRV. These data indicate that timing of the immune response is crucial in pathogenesis. Expression patterns of viral genes suggest they are tightly controlled until reaching the central nervous system (CNS), where replication increases significantly. These results suggest previous molecular mechanisms for rabies host response derived from mouse models do not strictly apply to natural vector populations. Overall my research provides a better understanding of the immunological factors that contribute to the pathogenesis of RRV in a natural system.

Author Keywords: immune response, major histocompatibility complex, rabies, raccoons, skunks, virus

The endangered North Atlantic right whale (Eubalaena glacialis) has been internationally protected from whaling since 1935 but recovery has been slow compared to the southern right whale (Eubalaena australis) due to anthropogenic mortalities and poor reproduction. Prey availability, genetic variability, and alleles of genes associated with reproductive dysfunction have been hypothesized to contribute to low calf production. The North Atlantic Right Whale DNA Bank and Database contains 1168 samples from 603 individuals. I added 115 new genetic profiles to the database which now contains profiles for 81% of individuals alive since 1980. Paternity assignments using these profiles resulted in 62% of sampled calves being assigned a father and only 38% of candidate males being assigned a paternity. This may suggest false exclusion due to genotyping errors or the existence of an unknown group of males. The use of the DNA database allowed for the identification of 10 deceased individuals which has implications for identifying cause of death and reducing mortalities. However, genetic identification is dependent on the time of post-mortem sample collection which influences DNA quantity and quality. An assessment for variations in methylenetetrahydrofolate reductase, a candidate gene associated with reproductive dysfunction, revealed six females heterozygous for a synonymous A/T variant in exon four which may influence reproductive success through changes in enzyme production, conformation or activity.

Author Keywords: Eubalaena glacialis, Forensic Identification, Genetic Profiling, North Atlantic Right Whale, Paternity, Reproductive Dysfunction

Most delphinids produce narrowband frequency-modulated whistles with a high level of plasticity to communicate with conspecifics. It is important to understand geographic variation in whistles as signal variation in other taxa has provided insight into the dispersal capabilities, genetic divergence and isolation among groups, and adaptation to ecological conditions. I investigated whistle variation of Indo-Pacific humpback dolphins (Sousa chinensis chinensis), Taiwanese humpback dolphins (S. c. taiwanensis) and Australian humpback dolphins (S. sahulensis) to test whether differences in whistles support the hypotheses of population structure, regional and species differences in the genus Sousa, which were based on morphological and genetic data. I also investigated important factors that may contribute to local distinctiveness in whistles including behavioural state, group size, and the influence of vessel noise. Multivariate analyses of seven acoustic variables supported the hypotheses of population structure, regional and species differences. Acoustic diversification between groups is likely influenced by behaviour and social contexts of whistles, and environmental noise. The use of sound to identify discrete groups of humpback dolphins may be important in future studies where genetic and morphological studies may not reveal recent differentiation or are difficult to conduct.

Author Keywords: Bioacoustics, Cetacean, Geographic variation, Population biology, Sousa, Whistle characteristics

The purpose of this thesis is to quantify the habitat characteristics, density patterns and environmental niches of two groups of Indo-Pacific humpback dolphins: Chinese white dolphins (CWD) of the Pearl River estuary (PRE), and Taiwanese white dolphins (TWD, =Taiwanese humpback dolphin, THD) found in the eastern Taiwan Strait (ETS). Much work has already been done on the habitat use of CWDs in parts of the PRE, so the purpose of my first two chapters was to advance knowledge of the TWD to a comparable level. Chapter 2 contains the first published description of the relatively shallow, inshore, estuarine habitat of the TWD. General environmental characteristics and observed group sizes were consistent with other populations of humpback dolphins, and group sizes were not correlated with the environmental variables measured during surveys. Chapter 3 investigated density patterns of TWDs, finding spatiotemporal heterogeneity across the study area. Humpback dolphin densities fluctuated from year to year, but some parts of the study area were consistently used more than others. Environmental characteristics again did not influence dolphin densities, though more dolphins than expected were sighted in waters adjacent to major land reclamations, which may be related to the location of these areas close to major rivers. In Chapter 4, niches of the TWD and CWDs found in the PRE were compared using species distribution models, which indicated significant niche overlap. This may be due to niche conservatism maintaining similar fundamental niches between the two groups since their historical split >10,000 years ago, or a result of the intrinsic biotic factors that influence occurrence data affecting the hypervolume dimensions of each realized niche in similar ways. Geographic predictions indicate that most of the TWD's range has likely been surveyed, and that there may be connectivity between PRE humpback dolphins and at least one neighbouring putative population due to continuous predicted suitable habitat in waters that remain poorly surveyed. Overall, my thesis demonstrates that density patterns may vary over time, but on a broad temporal scale, these two allopatric groups of Indo-Pacific humpback dolphins have similar habitat requirements in geographically isolated, but environmentally similar locations.

Author Keywords: density, habitat, Indo-Pacific humpback dolphin, niche overlap, Sousa chinensis, species distribution model