Behavioral sciences

AbstractDifferences and similarities in exploration and risk-taking behaviours of two Myotis bat species. Laura Michele Scott Behaviours that are repeatable across circumstances and time determine an individual's personality. Personality and behavioural variation are subject to selective pressures, including risks related to the use of different habitat types. I explored the ecological and evolutionary consequences of habitat selection by comparing the behaviour of two sympatric bat species, Myotis leibii and M. lucifugus. These species display overlap in roosting preferences, however, M. leibii tend to roost in crevices on the ground, while M. lucifugus tend to roost in crevices or cavities that are raised off the ground. I hypothesized that the habitat selection patterns of these two species create behavioural reaction norms at the species level. I predicted that ground roosting behaviour favours bolder personality and more exploratory and active traits when compared with bats that do not ground roost. I examined inter- and intra-specific variation in behaviour using a modified, three-dimensional open-field test and quantified the frequency and duration of behaviours such as flying, landing, and crawling. Bats were continuously video-recorded over 1-hour nocturnal and diurnal trials. I used a priori mixed models with combinations of individual characteristics and life-history traits to select the models that best describe each species. We found that M. leibii (n = 15) displayed more exploratory and bolder behaviours than M. lucifugus while on the ground (n = 21) and higher overall activity during the trial. I also found that M. leibii displayed crawling behaviours and movements consistent with foraging while on the ground which is a rare behaviour in bats and only observed in a few species (Desmodus rotundus and Mystacina tuberculate to my knowledge). Future research should explore biomechanical adaptations associated with ground-foraging in M. leibii.

Author Keywords: Bats, Behaviour, Exploration, Myotis leibii, Myotis lucifugus, Roosting

The Pearl River Estuary (PRE) and eastern Taiwan Strait (ETS) populations of Indo-Pacific humpback dolphins (Sousa chinensis) contain ~600 and 100 catalogued individuals, respectively. Population demography is important for conservation actions but few animals have been sexed by conventional methods. Longitudinal analysis of tooth-rake scars on dorsal fins showed scars disappear within 7 months; sexing adults by scarring is likely not impacted by juvenile scarring. Using dorsal fin photographs, sex assigned for 87% of catalogued PRE adults (n=300) and for 93% of ETS adults (n=60), using scars hypothesized from male-male competition, was in concordance with sex assigned by DNA, calf association, and ventral photographs. Scarring was higher in presumed males than females and in PRE females than ETS females. Female:male sex ratios were 3:2 (PRE) and 2:1 (ETS), though this likely results from biases in photo-identification methods. Social analysis with presumed sexes showed strong female-female associations in both populations but stronger female-male and male-male associations in PRE. These results support sex differentiation by scarring, which was a non-invasive approach, and sex assignment for many PRE and ETS individuals.

Author Keywords: Indo-Pacific humpback dolphins, photo-identification, scarring, sex differentiation, sexes, social structure

Memories which typically require the hippocampus (HPC) can become represented in structures outside of the HPC, and therefore resistant to HPC damage, but, the properties of these memories are poorly understood. Some research has suggested that the HPC continually contributes to memories that are resistant to hippocampal damage, and without this support, they are weaker and more susceptible to loss. However, this hypothesis has yet to be tested experimentally. We examined this possibility in rats by assessing decay and extinction of a context fear memory that had become independent of the HPC via repeated learning episodes. We found that HPC-independent context fear memories decay and extinguish faster without continued HPC support, suggesting that the HPC plays a continued role in long-term memory. We also provide new evidence of a persistent interaction between the HPC and other memory systems, which strengthens non-HPC representations so that they withstand HPC damage at longer intervals.

Author Keywords: consolidation, context fear, hippocampus, memory, retrograde amnesia

Caribou experience direct and indirect negative effects of harassment from biting flies, influencing behavior and activity on several spatial and temporal scales. I used systematic insect collection surveys during the summers of 2011 and 2012 to examine the spatial and temporal distributions of black flies (Simuliidae), mosquitoes (Culicidae), and deer flies and horse flies (Tabanidae) in a managed boreal forest in northern Ontario. Mosquitoes had a positive association with densely treed habitats, whereas black flies more often occurred in open areas, and tabanids had a strong presence in all habitat types. Habitats in proximity to large bodies of water had fewer biting flies than inland areas. Young stands supported higher abundances of tabanids despite vegetation community type. Next, I tested for seasonal effects of biting fly abundance on caribou activity by modelling the seasonal trend in abundance for each fly family for each year and compared this to an index of daily activity for 20 radio-collared female caribou in 2011 and 10 females in 2012. I modeled this index of caribou activity for each animal in each year and extracted the set of partial correlation coefficients from multiple regressions to test for effects of biting fly abundances on caribou activity. Caribou reduced their daily activity when tabanids were more numerous, and increased activity when mosquitoes were numerous. This divergent response may reflect a difference in the efficacy of moving to reduce harassment, owing to the stronger flight capabilities of tabanids.

Author Keywords: Activity, Anthropogenic Disturbance, Behaviour, Insect harassment, Temporal distribution, Woodland Caribou

Inducible defenses are plastic responses by an organism to the perception of predation risk. This dissertation focuses on three experiments designed to test the hypothesis that plastic ability is limited by energetic constraints. Chapter 1 provides a general introduction to phenotypic plasticity research and the theoretical costs and limitations affecting the expression of plastic traits. In Chapter 2, I tested the hypothesis that costs of early plasticity may be manifested by a reduced response to risk in later life stages. I found that amphibian embryos are able to detect and respond to larval predators, but that the energetic cost of those plastic responses are not equivalent among behavioural, growth, and morphological traits, and their expression differs between closely-related species. Chapter 3 explicitly examines the relationship between food resource availability and plasticity in response to perceived predation risk during larval development. Food-restricted tadpoles showed limited responses to predation risk; larvae at food saturation altered behaviour, development, and growth in response to predation risk. Responses to risk varied through time, suggesting ontogeny may affect the deployment of particular defensive traits. Chapter 4 examines the influence of maternal investment into propagule size on the magnitude of the plastic responses to predation risk in resulting offspring. I found that females in better body condition laid larger eggs and that these eggs, in turn, hatched into larvae that showed greater morphological plasticity in response to predation risk. Maternal investment can therefore affect the ability of offspring to mount morphological defenses to predation risk. Last, Chapter 5 provides a synthesis of my research findings, identifying specific factors constraining the plastic responses of prey to perceived predation risk. Overall, I found constraints on plastic responses imposed by the current environment experienced by the organism (resource availability), the prior experience of the organism (predator cues in the embryonic environment), and even the condition of the previous generation (maternal body condition and reproductive investment). Together, these findings both provide new knowledge and create novel research questions regarding constraints limiting phenotypic variation in natural populations.

Author Keywords: behaviour, inducible defense, Lithobates pipiens, morphometrics, phenotypic plasticity, predation risk

ABSTRACT

Calving site selection and fidelity in a restored elk (Cervus elaphus) herd in Bancroft

Ontario, Canada.

Michael R. Allan

Parturition site selection by ungulates is believed to be influenced by forage abundance and concealment from predators. In 2011 and 2012, I used vaginal implant transmitters and movements to identify calving sites for 23 GPS collared elk (Cervus elaphus) from a restored herd. I tested the hypothesis that maternal elk used sites with higher forage and denser concealment compared to pre-calving sites at micro and macrohabitat levels. I detected no significant microhabitat differences from direct measurements of vegetation. At the macrohabitat scale, based on proximity of landcover classes, mean distances to hardwood forests was significantly less for calving (153 m) than pre-calving sites (198 m).

Site fidelity is hypothesized to offer security in terms of familiarity to an area. I tested the hypothesis that females demonstrated fidelity to their previous year's location during pre-partum, parturition, post partum, breeding and winter periods. Elk were more philopatric during parturition and post partum than during breeding. Compared to winter elk were more philopatric during pre-partum, parturition and post-partum periods. Expressed as distance between consecutive-year calving locations, site fidelity varied with 27% of females exhibiting high (<1 km), 18% moderate and 55% (>2.9 km) low fidelity.

I measured nearest-neighbour distances at calving time, exploring the hypothesis that females distance themselves from conspecifics. Elk increased the average distances

to collared conspecifics during parturition; however, sample sizes were small. This strategy might influence calving site selection.

Rapid movement prior to parturition, low site fidelity and spacing-out of females during parturition appear to be strategies to minimize predator risk and detection. Little evidence of selection for vegetation structure suggests this may not be limiting to these elk.

Author Keywords: calving, elk, fidelity, movement, parturition, selection

Stressors are often an inescapable part of an organism's life. While the effects of many stressors have been well studied individually, potential interactions between stressors exist that may result in greater than additive negative effects. Stressors may be linked by conflicting demands on energy budgets, interfering with important physiological pathways, or necessitating incompatible adaptive responses. Using Ranavirus (FV3) and larval dragonfly predators (Anax spp.) in a 2x2 factorial experiment on green frog (Lithobates clamitans) tadpoles, I investigate the interactions in behaviour, morphology, and metabolism when both stressors were applied in concert. I demonstrate that activity and feeding are reduced additively by both stressors, and tadpoles increase distance between conspecifics in FV3-exposed tanks, but only in the absence of predators. I also note decreases in mass, and a non-significant marginal increase in metabolic rate of tadpoles exposed to FV3. Interestingly, I provide evidence that FV3 can compromise morphometric responses through antagonistic interactions with perceived predation risk exposure, which may result in significantly elevated mortality even when either stressor is present in sub-lethal quantities. Thus, I conclude that sub-lethal exposure to stressors can nonetheless have substantial impacts on organisms and a more integrative approach to examining the impacts of stressors on individual physiology and fitness is necessary.

Author Keywords: Behaviour, Interaction, Morphology, Predation Risk, Ranavirus, Tadpoles

Polar bears are generally described as solitary, but features of their life cycles and

habitats regularly necessitate interaction. Effective conspecific assessment, including accurate

recognition and discrimination, likely confers benefits, especially to females accompanied by

dependent young. Individuals in the Southern (SH) and Western (WH) Hudson Bay

subpopulations are ideal for studying polar bear social behaviours because of the prolonged

high densities of the ice-free season. First, I looked outside family groups to model their fine

scale sociospatial organization on land. Capture locations were more likely to correspond to

family groups when there were fewer neighbouring bears, when a greater proportion of

neighbours were female, and when the focal individual and neighbours were significantly

related. Second, I looked within the family group to assess offspring recognition. Of 288

offspring in 207 family groups captured in the SH subpopulation from 1999 through 2013, only

one case of adoption (of a singleton) was observed.

Author Keywords: Adoption, Kin Recognition, Logistic Regression, Maternity Analysis, Social Discrimination, Sociospatial

Stopover ecology is a topic that surges in relevancy as choices made by migrants during stationary periods (stopover sites) may not only have important individuals' fitness consequences but also can affect population dynamics. I used MOTUS automated telemetry array to study fall stopover duration of Blackpoll Warbler (BLPW) and departure decisions of BLPW and Canada Warbler (CAWA) in relation to various predictors. I affixed radio-transmitters on 55 BLPWs and 32 CAWAs at two banding stations in Ontario in September-October 2014-2015. Radio-tagged individuals were tracked through the MOTUS network across southeastern Canada. I developed models relating age class, fat score, Julian date and stopover movement types to Blackpolls' stopover duration. I also examined whether there were species-related differences of wind selectivity when resuming migration. No explanatory variable significantly influenced BLPW's stopover duration. Both species tended to depart under increased tailwind assistance, but with no difference in the effect of wind conditions between the two species. This study provides further evidence supporting the relevance of local wind conditions as a key factor affecting the departure likelihood, especially when migrating birds face an ecological barrier.

Author Keywords: Cardellina canadensis, departure decisions, minimum stopover length, MOTUS, overland fall migration, Setophaga striata

Most grassland bird populations are in decline, so it is becoming increasingly important to understand how they use agricultural field types and form their communities. I performed point counts in cultural meadow, intensive agriculture, and non-intensive agriculture areas in 2011 and 2012. Generalized linear models were used to determine the habitat relationships of six focal species. I found that non-intensive agriculture was used most often and intensive agriculture was often avoided, but there were exceptions which indicate habitat use can be species-specific. I determined in which habitats competition was likely occurring and which species pairs were competing in 2011. In 2012, I experimentally tested these relationships by introducing artificial competitors onto sites. By comparing presence-absence data from 2011 to 2012, I found evidence of habitat-mediated interspecific and conspecific attraction involving Bobolink and Grasshopper Sparrow. This research contributes to the current understanding of grassland bird community ecology and conservation.

Author Keywords: agriculture, BACI, community ecology, habitat use, species at risk, species interactions