Many mammals are social. The most basic social behaviour is when the actions of one conspecific are directed toward another, what we call the dyadic interaction. Both intrinsic and extrinsic factors may affect an individuals propensity to interact with other members of a population. I used a social cervid, elk (Cervus elaphus), as a model species to test the importance of intrinsic and extrinsic factors of sociality on dyadic interactions. Dyadic interactions not only form the basis for social structure and information transfer within a population, but are also routes of pathogen transmission. My objective in this thesis was thus twofold: to improve our understanding of sociobiology, but also to gain insight into how sociality may underlie the transmission of communicable wildlife disease. I used a hierarchical, autecological approach from DNA, through individual, dyad, group, subpopulation, and ultimately population to explore the effects of intrinsic factors (e.g., sex and pairwise genetic relatedness) and extrinsic factors (e.g., season, conspecific density, habitat, and elk group size) on sociality. Elk in Riding Mountain National Park (RMNP), Manitoba, Canada, are exposed to the causal agent of bovine tuberculosis (Mycobacterium bovis; TB); however, spatial variation in apparent disease prevalence suggests that TB can only persist in one subpopulation within the Park. Using the natural RMNP system and a captive elk herd that I manipulated, I explored factors that influence interaction rates and durations (as a proxy for pathogen transmission) among elk. Sexual segregation in elk results in seasonal and sex-based differences in interaction rate and duration; with interactions peaking in autumn-winter for both sexes. Female-female dyads interact more frequently than male-male dyads. However, male-male dyads interact for longer durations than do female-female dyads. Interaction rate and duration did not covary with pairwise relatedness. Conspecific density also had sex-specific results for interaction rate and duration. Whereas male-male dyadic interaction rates increase with density, female-female dyads increase until they reach a threshold and subsequently reduce their interaction rates at high density. I observed density dependence in interaction rates in experimental trials and from field data. Furthermore, social networks revealed that social familiarity (i.e., heterogeneity of interactions) can be both frequency- and- density dependent depending on the strength of the relationship (i.e., number of repeat interactions). Density also affected the likelihood that an interaction would occur; however, this was modified by vegetation association used by elk. My results reveal several ecological and evolutionary implications for information transfer and pathogen transmission. In particular, I show that seasonal inter-sex routes of transfer may exist and that transfer is likely to be density-dependent. Finally, I conclude that such transfer is modified by available resources.

Many mammals are social. The most basic social behaviour is when the actions of one conspecific are directed toward another, what we call the dyadic interaction. Both intrinsic and extrinsic factors may affect an individuals propensity to interact with other members of a population. I used a social cervid, elk (Cervus elaphus), as a model species to test the importance of intrinsic and extrinsic factors of sociality on dyadic interactions. Dyadic interactions not only form the basis for social structure and information transfer within a population, but are also routes of pathogen transmission. My objective in this thesis was thus twofold: to improve our understanding of sociobiology, but also to gain insight into how sociality may underlie the transmission of communicable wildlife disease. I used a hierarchical, autecological approach from DNA, through individual, dyad, group, subpopulation, and ultimately population to explore the effects of intrinsic factors (e.g., sex and pairwise genetic relatedness) and extrinsic factors (e.g., season, conspecific density, habitat, and elk group size) on sociality. Elk in Riding Mountain National Park (RMNP), Manitoba, Canada, are exposed to the causal agent of bovine tuberculosis (Mycobacterium bovis; TB); however, spatial variation in apparent disease prevalence suggests that TB can only persist in one subpopulation within the Park. Using the natural RMNP system and a captive elk herd that I manipulated, I explored factors that influence interaction rates and durations (as a proxy for pathogen transmission) among elk. Sexual segregation in elk results in seasonal and sex-based differences in interaction rate and duration; with interactions peaking in autumn-winter for both sexes. Female-female dyads interact more frequently than male-male dyads. However, male-male dyads interact for longer durations than do female-female dyads. Interaction rate and duration did not covary with pairwise relatedness. Conspecific density also had sex-specific results for interaction rate and duration. Whereas male-male dyadic interaction rates increase with density, female-female dyads increase until they reach a threshold and subsequently reduce their interaction rates at high density. I observed density dependence in interaction rates in experimental trials and from field data. Furthermore, social networks revealed that social familiarity (i.e., heterogeneity of interactions) can be both frequency- and- density dependent depending on the strength of the relationship (i.e., number of repeat interactions). Density also affected the likelihood that an interaction would occur; however, this was modified by vegetation association used by elk. My results reveal several ecological and evolutionary implications for information transfer and pathogen transmission. In particular, I show that seasonal inter-sex routes of transfer may exist and that transfer is likely to be density-dependent. Finally, I conclude that such transfer is modified by available resources.

Vom Frühjahr 2010 bis Frühsommer 2011 wurden Verhalten, intra- und interspezifische Interaktionen sowie Raumnutzung von Wisenten (Bison bonasus), Exmoor-Ponys (Equus ferus) und Dybowski-Hirschen (Cervus nippon hortulorum) auf einer 14 ha großen Gemeinschaftsanlage im Tierpark Sababurg untersucht. 15 Individuen wurden dazu insgesamt für 605 Stunden mittels focal-animal und scan sampling beobachtet. Die Mittelwerte der Hauptverhaltensweisen über den gesamten Beobachtungszeitraum ergaben bei den Wisenten Tagesanteile von 47,73 % für Nahrungsaufnahme, 43,05 % für Ruheverhalten, 6,79 % für Fortbewegung und 1,68 % für Komfortverhalten. Die Exmoor-Ponys fraßen 58,62 % des Tages, ruhten 34,54 %, nutzten 3,87 % für die Fortbewegung und verbrachten 1,37 % mit Komfortverhalten. 47,22 % der Zeit wurden von den Dybowski-Hirschen zur Nahrungsaufnahme verwendet, 43,64 % geruht, 4,55 % zur Fortbewegung genutzt und 2,44 % mit Komfortverhalten verbracht. All diese Verhaltensweisen waren Bestandteil eines artspezifischen Tagesrhythmus aus alternierenden Nahrungsaufnahme- und Ruhephasen. Die prozentualen Anteile und somit auch die Ausprägung dieses Tagesrhythmus variierten dabei allerdings teilweise stark zwischen den einzelnen Jahreszeiten und den damit verbundenen klimatischen und umweltbedingten Veränderungen. Die Individuen aller drei Arten verhielten sich dabei völlig normal im Rahmen der nachgewiesenen Bandbreite des natürlichen Verhaltens ihrer wildlebenden Artgenossen, unbeeinflusst von den Gehegebedingungen und Eingriffen des Zuchtmanagements. Interspezifische Interaktionen traten regelmäßig auf, positive Kontakte blieben dabei allerdings auf die Exmoor-Ponys und die Dybowski-Hirsche beschränkt. Agonistische Interaktionen waren meist lediglich als einfache Drohgebärden oder als indirekte Interaktionen ohne körperlichen Kontakt feststellbar. Eine feste Artenhierarchie ist etabliert und wird von allen Tieren akzeptiert, wobei die Wisente die dominante Rolle einnehmen, gefolgt von den Exmoor-Ponys und den Dybowski-Hirschen. Die Habitatnutzungsanalyse ergab für alle drei Arten bevorzugte Aufenthaltsbereiche und genutzte Gehegeobjekte. Dabei war wie das Verhalten auch die Habitatnutzung stark von den jahreszeitlich veränderlichen Umweltbedingungen beeinflusst. Während die offeneren Areale insbesondere im Frühjahr und Herbst genutzt wurden, bevorzugten die Tiere im Sommer und Winter eher die geschützteren Anlagenbereiche, im Sommer hauptsächlich als Schattenbereich, im Winter als Wetterschutz. Zusammenfassend betrachtet funktioniert das Zusammenleben der Wisente, Exmoor-Ponys und Dybowski-Hirsche im Tierpark Sababurg ohne Probleme und kann als Beispiel-Projekt für die zukünftige Forschung über die Möglichkeiten für Beweidungsprojekte mit heimischen Wildtieren im Naturschutz dienen.

No description available.

It is generally accepted that the Marginal Value Theorem (MVT) describes optimal foraging strategies. Some research findings, however, indicate that in natural conditions foragers not always behave according to the MVT. To address this inconsistency, in a series of computer simulations, I examined the behaviour of four types of foragers having specific foraging efficiencies and using the MVT and alternative strategies in 16 simulated landscapes in an ideal environment (no intra- and inter-species interactions). I used data on elk (Cervus elaphus) to construct the virtual forager. Contrary to the widely accepted understanding of the MVT, I found that in environments with the same average patch quality and varying average travel times between patches, patch residence times of some foragers were not affected by travel times. I propose a mechanism responsible for this observation and formulate the perfect forager theorem (PFT). I also introduce the concepts of a foraging coefficient (F) and foragers hub (), and formulate a model to describe the relationship between the perfect forager and other forager types. I identify situations where a forager aiming to choose an optimal foraging strategy and maximize its cumulative consumption should not follow the MVT. I describe these situations in a form of a mathematical model. I also demonstrate that the lack of biological realism and environmental noise are not required to explain the deviations from the MVT observed in field research, and explain the importance of scale in optimal foraging behaviour. I also demonstrate that smart foraging, which is a set of rules based on key ecological concepts: the functional response curve (FRC), satisficing, the MVT, and incorporates time limitations, should allow for fitness maximization. Thus, it should be an optimal behavior in the context of natural selection. I also demonstrate the importance of the FRC as a driver for foraging behaviors and argue that animals should focus more on increasing the slope of their FRC than on choosing a specific foraging strategy. Natural selection should, therefore, favor foragers with steep FRC. My findings introduce new concepts in behavioural ecology, have implications for animal ecology and inform wildlife management.

The experiments described in this thesis investigated nutritional and physiological aspects of antler growth in red deer stags. The initial experiment (Section 3) examined the effects of winter nutrition on subsequent antler casting date and velvet antler weight. Mature stags on two different farm types (hill country, Farm H and irrigated lowland, Farm L) were offered three levels of winter nutrition, two levels of a concentrate supplement (ad libitum pellets and 1/2 ad libitum pellets) and a basal hay ration. On both properties liveweight gains occurred in supplemented groups and liveweight losses in unsupplemented groups. At antler casting there were significant differences in liveweight of approximately 10 kg between fully supplemented and unsupplemented groups. Realimentation of winter liveweight losses subsequently occurred so that by the following rut the effects of winter undernutrition had been eliminated. On Farm H poor winter nutrition (hay only) resulted in a significant delay in casting date (13 days) and lower velvet antler yields (0.24 kg), than in stags offered the ad libitum concentrate ration. Stags on Farm H were 13 kg lighter at commencement of the trial than at Farm L and the differences in treatment effects obtained between farms may have been due to differences in body condition at commencement of the trial. An association was demonstrated between liveweight and date of antler casting, with heavier stags casting earlier than lighter stags. There was no effect of age of stag on casting date. Of the liveweights recorded, liveweight prior to the rut showed the best relationship with casting date, possibly because the seasonal nature of liveweight change meant that a weight recorded at this time gave the best indicator of the true frame size of a stag. Both age and liveweight significantly affected velvet antler weight, with increases of velvet antler weight of 0.26 kg between 3 and 4-year-old stags and of 0.30 kg between 4 and 5-year-old stags at the same liveweight. Within an age group velvet weight increased by 0.12 kg for each 10 kg increase in pre-rut liveweight. The experiments described in Section 4 comprised studies on antler growth and composition. In order to obtain data on antler growth and composition individual antlers were removed sequentially from mature red deer stags between 28 and 112 days after casting of hard antlers. Contralateral antlers were removed after stripping of velvet. Wide variation occurred in antler casting date (53 days) compared to date of velvet stripping (24 days). The duration of the period of antler growth may therefore be governed more by date of casting than by date of velvet stripping. Mean duration of the antler growth period was 164 days. Growth in length of the antler appeared to follow a sigmoid curve. However, between 28 and 112 days after casting, rates of elongation were close to linear. Mean length of hard stripped antlers was 0.71 m and between 28 to 112 days after casting mean rate of antler elongation was 0.62 cm/day. Over this period indivdual antlers increased in fresh weight at a rate of 13.7 g/d, with heaviest weight recorded 112 days after antler casting, at approximately 130% of final hard antler weight. Between 28 and 91 days of growth, volume of blood in the antler increased linearly at a rate of 194 ml/kg. Three phases of mineralization were demonstrated in developing antlers. Tips of growing antlers were cartilaginous and poorly mineralized. A zone of mineralization occurred 5.0 to 7.5 cm behind the antler tip which corresponded histologically to the transition from mineralized cartilage to trabecular bone. The second phase of mineralization occurred through continued accretion of trabecular bone in the antler shaft. The third phase, described as "terminal mineralization" in this study, appeared to be associated with a rapid increase in density of cortical bone in the periphery of the antler shaft. Terminal mineralization (between 91 and 112 days after casting of hard antlers) coincided with the slowing of growth in length, a decrease in relative blood volume in the antler and an increase in levels of plasma testosterone. These events occurred close to the summer solstice. At velvet stripping individual antlers had a mean weight of 1.12 kg and contained 81.1% dry matter (DM). Fat free organic matter (FFOM) and ash concentration in DM were 36.6 and 60.0%, respectively. Peak daily rates of FFOM and ash deposition occurred between 91 days and 112 days after casting, at rates of 1.4% of hard antler FFOM and 1.6% of hard antler ash. For a stag producing 2.24 kg of hard antler mean rates of FFOM and ash deposition over this period were 9.3 and 18.3 g/d, respectively. On a whole antler basis calcium concentration in antler ash remained constant, at around 35%. Therefore peak rate of antler calcium deposition would be 6.4 g/d. In the final experiment (Section 5) mature stags were offered a maintenance ration of greenfeed oats during the period of peak calcium requirement for antler growth and the kinetics of calcium metabolism were examined using a radio-isotope (⁴⁵Ca). Rates of faecal endogenous loss were low and at approximately 6.4 mg/kg BW per were half the estimated requirements of ARC (1980) for sheep and cattle. Availability of calcium from greenfeed oats was low (mean, 37%) and less than 30% of total calcium requirements were derived from the diet. Poorly mineralized skeletal bones indicated that the shortfall in antler calcium was derived from the skeleton. In spite of a severely negative calcium balance stags were capable of maintaining high and apparently normal rates of antler calcium deposition (mean, 44 mg/kg BW per day). Antlers appear to be acting as a sink with calcium being irreversibly deposited in the antler and lost to the animal's body. On the assumption therefore that antler calcium behaves like calcium lost during lactation a kinetic model of calcium metabolism in the stag was developed.

No description available.

The eco-epidemiology of bovine tuberculosis (Tb) in wild deer (mainly red deer Cervus elaphus) in New Zealand was investigated. Bovine Tb is caused by Mycobacterium bovis. Specific aims were to clarify the likely routes of infection in deer, and to determine the status of deer as hosts of Tb, the likely rates and routes of inter- and intra-species transmission between deer and other wildlife hosts, the role of deer in spreading Tb, and the likely utility of deer as sentinels of Tb presence in wildlife. As the possum (Trichosurus vulpecula) is the main wildlife host of Tb, the research also included some investigation of transmission routes in possums. Patterns of infection were measured in 994 deer killed between 1993 and 2003. Tb prevalence varied between areas (range 8–36%). Few deer had generalised infection, with 21–68% of infected deer having no visible lesions, depending on the area. The retropharyngeal lymph nodes and oropharyngeal tonsils were commonly infected. No dependent fawns less than 0.75 years old were infected, indicating intra-species transmission is rare in wild deer. Where possums were not controlled, the net (cumulative) force of infection in young (1–4 y) deer was 0.10–0.24 per year in males and 0.09–0.12 per year in females, but much lower in older deer (less than 0.05 per year). Possum control reduced the net force of infection quickly, and eventually to zero. However, Tb persisted in possum-controlled areas through immigration of infected deer and, for almost a decade, through the survival of resident deer infected before possum control. Tb was lost from infected deer at an exponential rate of 0.13 per year, mostly as a result of deer recovering from infection rather than dying from it. Wild deer do die of Tb, but there was no discernible effect on age structure. The occurrence of infection in deer was not linked to the local deer or possum density at their kill sites (i.e. in their home range), but the area-wide prevalence of Tb in deer was closely correlated with Tb levels in possums, which were in turn correlated with area-wide measures of possum density. For wild deer in New Zealand, Tb is a persistent but usually inconsequential disease of the lymphatic system. It is acquired mainly by young independent deer, usually orally via the tonsils, and probably as a result of licking infected possums. Many species fed on deer carrion, including possums. Most possums encountering carrion did not feed on it, but a few fed for long periods. Other scavengers such ferrets (Mustela furo), hawks (Circus approximans), and weka (a hen-sized flightless native bird; Gallirallus australis) fed in a way that probably increased the infectivity of carrion to possums. Commercial deer hunting may have facilitated the historical establishment of Tb in possums. Scavenging (including cannibalism) and interactions with dead and dying possums are identified for the first time as potentially important routes for transmission of Tb to possums, and I develop new hypotheses involving peri- and post-mortem transmission in possums that explain many of the epidemiological patterns that are characteristic of the disease in possum. In continuous native forest, deer home range size averaged 250 hectares for six young females, and over twice that for two males. Over 90% of infected deer are likely to die within 2 km (females) or 6 km (males) of where they acquired Tb, but deer could occasionally carry Tb up to 30 km. Deer will be useful as sentinels, but only where other sentinels are rare, because the force of infection for a deer with a single infected possum in its home range is only 0.004 per year, compared to greater than 0.2 per year for deliberately released pigs. Deer are occasionally capable of initiating new cycles of infection in wildlife, but deer control is not essential to eradicate Tb from wildlife.