Developing Behavioral Indices of Population Viability: A Case Study of California Sea Lions in the Gulf of California, Mexico

abstract: Despite years of effort, the field of conservation biology still struggles to incorporate theories of animal behavior. I introduce in Chapter I the issues surrounding the disconnect between behavioral ecology and conservation biology, and propose the use of behavioral knowledge in population viability analysis. In Chapter II, I develop a framework that uses three strategies for incorporating behavior into demographic models, outline the costs of each strategy through decision analysis, and build on previous work in behavioral ecology and demography. First, relevant behavioral mechanisms should be included in demographic models used for conservation decision-making. Second, I propose rapid behavioral assessment as a useful tool to approximate demographic rates through regression of demographic phenomena on observations of related behaviors. This technique provides behaviorally estimated parameters that may be applied to population viability analysis for use in management. Finally, behavioral indices can be used as warning signs of population decline. The proposed framework combines each strategy through decision analysis to provide quantitative rules that determine when incorporating aspects of conservation behavior may be beneficial to management. Chapter III applies this technique to estimate birthrate in a colony of California sea lions in the Gulf of California, Mexico. This study includes a cost analysis of the behavioral and traditional parameter estimation techniques. I then provide in Chapter IV practical recommendations for applying this framework to management programs along with general guidelines for the development of rapid behavioral assessment.Dissertation/ThesisM.S. Biology 201

Frequency-dependent two-sex models : a new approach to sex ratio evolution with multiple maternal conditions

© The Author(s), 2016. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Ecology and Evolution 6 (2016): 6855–6879, doi:10.1002/ece3.2202.Mothers that experience different individual or environmental conditions may produce different proportions of male to female offspring. The Trivers-Willard hypothesis, for instance, suggests that mothers with different qualities (size, health, etc.) will use different sex ratios if maternal quality differentially affects sex-specific reproductive success. Condition-dependent, or facultative, sex ratio strategies like these allow multiple sex ratios to coexist within a population. They also create complex population structure due to the presence of multiple maternal conditions. As a result, modeling facultative sex ratio evolution requires not only sex ratio strategies with multiple components, but also two-sex population models with explicit stage structure. To this end, we combine nonlinear, frequency-dependent matrix models and multidimensional adaptive dynamics to create a new framework for studying sex ratio evolution. We illustrate the applications of this framework with two case studies where the sex ratios depend one of two possible maternal conditions (age or quality). In these cases, we identify evolutionarily singular sex ratio strategies, find instances where one maternal condition produces exclusively male or female offspring, and show that sex ratio biases depend on the relative reproductive value ratios for each sex.National Science Foundation Graduate Research Fellowship Grant Number: 1122374; National Science Foundation Grant Numbers: DEB1145017, DEB1257545; European Research Council Grant Number: 322989; Woods Hole Oceanographic Institution Academic Programs Offic

The Ecology and Evolution of Human Reproductive Behavior

The complexity of human reproductive behavior has necessitated its examination through a variety of scientific disciplines, each focusing on specific elements of our biology, behavior, and society. However, this complexity also necessitates that we reintegrate the information learned from each discipline into a single framework, one rooted in the evolutionary principles that have shaped the development of all life on earth. In this dissertation, I use this framework to explore human reproductive behavior, with a particular focus on sexual coercion and fertility-mediated sexual behavior. In Chapter 1, I introduce the approach taken in this document, identify several key limitations, and outline the general structure. In Chapter 2, I conduct a comprehensive and interdisciplinary review that includes the fundamentals of sexual conflict and reproductive strategies; the evolution of human reproductive characteristics in response to socio-cognitive demands; the aspects of human sociality expected to influence reproductive behavior; the identified trends in human mating behavior; the proposed pressures behind concealed ovulation in primates; the essentials of the menstrual cycle; and the existing evidence for behavioral fertility in humans. In Chapter 3, I use a game-theory model to investigate the emergence of sexually coercive behavior across a variety of species, including humans, in which male coercion is a non-developmentally-determined reproductive strategy to identify several ecological and behavioral characteristics that predict the emergence of coercive behavior generally consistent with observed trends. In Chapter 4, I use face-trait research to investigate the degree to which women recognize and discriminate between images of men with personality traits associated with different male reproductive strategies as well as how these preferences might be mediated by her relationship and fertility status. In Chapter 5, explore the intersection of fertility, fertility belief, and sexuality, specifically testing the hypothesis that a woman’s sexual interest shifts in response to her fertility while taking into consideration her beliefs regarding her fertility. Finally, in Chapter 6, I review the primary take-home messages of this work and recommend that future research take these into consideration as they move forward. By taking an interdisciplinary approach rooted in evolutionary biology, this work reveals the need for an understanding of human reproductive behavior that incorporates a wider view of reproductive ecology. In doing so, we can gain a more accurate, comprehensive, and nuanced understanding of human reproductive behavior

ALL IN DUE TIME: MULTI-TRAIT ASSESSMENT OF ELK ACCLIMATION TO TRANSLOCATION

Wildlife translocation – the intentional movement of animals – is a crucial conservation tool for restoring species and halting global biodiversity decline. However, this practice is challenging for wildlife, and animals must adjust to their release landscapes for restoration to be successful. The period following release is a vulnerable time for translocated wildlife and determining when and how animals eventually acclimate following releases allows researchers to efficiently tailor post-release management to each species’ needs, thus maximizing the success of translocations while minimizing costs of an already expensive conservation practice. In this dissertation, I investigate changes in the physiological, behavioral, and social dynamics of 106 elk (Cervus canadensis) during the 6-8 years following their release to Missouri, U.S.A. in 2011-2013. I define the acclimation period throughout this work as the duration of time prior to stabilization in each investigated response relative to time from release. In Chapter 1, I analyzed changes in glucocorticoid metabolites (fGCMs) as an indicator of physiological acclimation. Fecal GCM levels declined following translocation and subsequently stabilized relative to days from release at approximately 42 days. The fast physiological acclimation by Missouri elk relative to other species suggests relatively low sensitivity by elk to translocation and effective use of temporary post-release management efforts. In Chapter 2, I investigated changes in elk spatial behavior (movements and resource selection patterns) using location data from GPS-collars deployed on all translocated elk. Changes in resource selection and monthly individual range sizes and overlap relative to time from release stabilized within the first year of translocation. Sexes varied in their post-release movement dynamics, with females showing faster and stronger evidence of acclimation following translocations that occurred during the parturition season. Significant temporal dynamics in selection for multiple resource covariates indicated that elk did not demonstrate a simple forage-refuge tradeoff while acclimating to their release landscape. In Chapter 3, I investigated dynamics in elk mating structure using paternity analysis on DNA extracted from tissue samples of all translocated elk and subsequently captured adults and calves. Following a translocation tactic favoring releases of young-aged males, initial polygyny in the restored Missouri population was low; however, polygyny levels increased and stabilized to expected values within four years of the last translocation event. Importantly, initial dampened polygyny may facilitate retention of genetic variation by maximizing the genetic contribution of more founding individuals. In Chapter 4, I investigated retention of genetic diversity over initial generations following release and projected future losses over a management-relevant time period. The Missouri elk population retained relatively high levels of genetic diversity as evidenced by minimal losses in allelic richness and expected heterozygosity (He), and we projected similarly stable He levels for the next 130 years (loss \u3c 10%). Together, these results suggest translocated wildlife acclimate to their release landscapes in a continuum of response, with behavior lagging physiological responses, and larger-scaled population processes, such as mating structure, sitting at the ultimate end of this spectrum. Investigating the manifold changes of translocated animals as they acclimate to their release landscape represents an opportunity to improve post-release monitoring and assessment while directly informing dynamic management needs of restored populations

Population dynamics of Ctenosaura bakeri

The Ctenosaura bakeri is an iguana species endemic to the island of Utila, a small island off the eastern coast of Honduras. It is currently one of the species of the genus Ctenosaura most threatened with extinction, having its conservation status labelled as "Critically Endangered" by the IUCN Red List. The goals of this paper are to give some insights on the intrinsic trend of the whole population and to analyse the influence of the greater threats to the survival of the species (such as sex dependent hunting and habitat destruction). We will use a transition matrix approach to investigate the intrinsic trend of the population and we will provide arguments for the estimation of the different parameters. For the influence of the threats we will take a deterministic approach using systems of ODEs and DDEs, investigating the stationary points and their stability and giving prediction through simulations for the evolution of the population. We will also introduce a model for the occurence of hybridization with another iguana species of the island. The achieved results are summarized and still open questions stated at the end

The Effects of Sex, Energy, and Environmental Conditions on the Movement Ecology of Migratory Bats

Lack of knowledge about the behaviour of migratory species during the migratory period is a major barrier to conservation efforts. In this thesis I focus primarily on differences between the sexes of the bat Lasionycteris noctivagans, during spring migration. Females are pregnant during spring migration and this overlap between migration and reproduction may affect the time and energy management of females as compared to males. In Chapter 2 I examine spring migration phenology of bats at a stopover site. Females arrived earlier than males, likely to give their pups a long growing season. Fat stores appeared to reflect a strategy to provision for upcoming lactation demands. In Chapter 3 I explore stopover behaviour and I show that despite the use of torpor to minimize roosting energy expenditure, cold weather extends stopover duration. There was no sex difference in the length of time spent at stopover. By regressing the time of night bats were captured against their fat and lean mass I demonstrate that bats have greater fat and lean masses closer to dawn, and therefore are likely using stopover periods to refuel. In Chapter 4 I compare sex and seasonal differences in daytime torpor use at stopover. I found that in spring bats used torpor for fewer hours than in autumn, even after accounting for the effect of ambient temperature. Further, females used torpor for fewer hours than males. I propose that these seasonal differences are due to lower prey abundance and predictability in spring; sex differences may be attributable to a higher foraging intensity by females compared to males. Finally, in Chapter 5 I use a radio-telemetry array in southwestern Ontario, Canada to track the spring and autumn long distance migratory movements of L. noctivagans, Lasiurus borealis, and Lasiurus cinereus. No migration corridors were identified in either season. Estimated migration speeds indicate that multiday stopovers are also used in the autumn. These studies combined show that sex affects the spring migration time and energy management of bats. Bat migration research is still in its formative stages and my studies provide new information on bat migration in North America

Evolutionary demography of structured two-sex populations and sex ratios

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution September 2015Males and females may differ in stage-specific survival, maturation, fertility, or mating availability. These demographic differences, in turn, affect population growth rates, equilibrium structure, and evolutionary trajectories. Models considering only a single sex cannot capture these effects, motivating the use of demographic two-sex models for sexually reproducing populations. I developed a new two-sex modeling framework that incorporates population structure and multiple life cycle processes through transition rate matrices. These models can be applied to a variety of life histories to address both ecological and evolutionary questions. Here, I apply the model to the effects of sex-biased harvest on populations with various mating systems. Demographic considerations also affect evolutionary projections. I derived matrix calculus expressions for key evolutionary quantities in my two-sex models, including the invasion fitness, selection gradient, and second derivatives of growth rates (which have many applications, including the classification of evolutionary singular strategies). I used these quantities to analyze the evolution of the primary sex ratio, under various sex- and stage-specific offspring costs and maternal conditions. Demographic two-sex models lend insight into complex, and sometimes counterintuitive, results that are not captured by models lacking population structure. These findings highlight the importance of demographic structure in ecology and evolution.This work was supported by a National Science Foundation Graduate Research Fellowship under Grant 1122374, NSF Grants DEB1145017 and DEB1257545 (to H. Caswell), Advanced Grant 322989 from the European Research Council (to H. Caswell), and theWoods Hole Oceanographic Institution Academic Programs Office

Studies on the biology of the brown hare (Lepus europaeus) with particular reference to behaviour

Behaviour of the brown hare (Lepus europaeus) was studied on the Somerset levels, U.K., between 1977 and 1987. The hare is exclusively nocturnal when nights are longest and part-diurnal when they are shortest but the transition is not smooth. There is a linear dominance hierarchy between bucks. The nearer a doe is to oestrus, the higher the rank of her escort. Some pre-partum does monitor the prospective birthsite. They are only occasionally escorted by bucks. In contrast, a non-monitoring pregnant doe was regularly escorted. Does lie up within 250m of their litters but take a more circuitous route when visiting them and 'detour when foxes are encountered. Four Autumn litters were suckled for between six and nine weeks. Sucking leverets usually sniff-noses when meeting up, followed by a short period of locomotor play after which they go quietly to the nursing point. Some weaned leverets continued to meet up. The principal component of locomotor play is 'streaking' - running top speed back and forth down a familiar route. Leverets disperse after nursing to a daily increasing extent. Leveret distress screams were audible from a distance of 550m. As a visible signal, the buck's white tail flag seems to serve no purpose. The doe uses her tail flag to lead her litter. Only does shake their tails which they do exclusively in the near presence of bucks; the bucks then sniff the ground underneath. Hares were not observed to signal to each other by body postures or to use their ears for purposes other than acoustic. Does convey a threat to bucks by flattening their ears and lifting their muzzles. Adults, particularly bucks; sniff a partner's nose for olfactory information as an alternative when the ano-genital region is inaccessible, but are frequently threatened when so doing. All chin-marking was by bucks of which 80% were solitary. Hares approached in the open by foxes stand bipedal when, on average, the fox is about 30m distant. The relevance of the behaviour patterns are discussed

Education and Living: Volume 2

In Education and Living, a two-volume work, Borsodi elaborated the model of the School of Living. Most of volume one consists of a critique of “mis-education.” Most of that critique focuses on the problems of centralization; centralization of industry, the economy, politics and education. The second volume of Education and Living explains Borsodi’s vision of achieving decentralization in detail. The second volume is in two parts: Right-Education and Re-Education. It explains how to educate for the “Normal” human being and for achieving the “Normal” way of living. This is not the “normal” of the bell curve, the average of a population. This is normal in terms of what a human being is innately capable of. It is not the “normal” of “the noble savage” but the norm of a person whose capacity for life, whose capacity for action, has been nurtured by education and training for life; who is the product of a School of Living. Volume 2 provides an insightful description of what and how a person of high individual and social potential can be developed. -- Bill Shar

Foraging, roosting and survival of Natterer’s bats, Myotis Nattereri, in a commercial coniferous plantation

I studied Natterer's bats in Tentsmuir Forest, a 9143-hectare commercial coniferous plantation on the NE coast of Fife, Scotland that has been planted predominantly with Scots and Corsican pine. Two maternity colonies of approximately 111 & 68 adult females regularly use bat boxes within the forest as summer and maternity roosts. The foraging ecology, roost dynamics, population structure and survival rates of the Natterer's bats present were determined in this study by radio tracking and banding. The two colonies occupied home ranges of 4.43 & 6.46 km2 with densities of approximately 25 & 10 adult females per km2 . Individual bats had a mean foraging home range of 0.47 km2 that contained 1-3 core foraging areas, which did not overlap with the core areas of other individual bats. Adult female Natterer's bats preferentially foraged in stands of mature Corsican pines and around water bodies; grazing meadows and arable land adjacent to the forest were avoided. They also foraged extensively in mature Scots pine, the predominant habitat. However, it was underused relative to its availability. Out of a set of five different bat box types, the bats preferentially chose old square wooden and round woodcrete boxes to use as summer and maternity roosts. I found that the bats also roosted in a specific type of natural cavity in double-Ieadered Corsican pines that has not been recorded before. Bats changed roost sites every 2.5 days on average. Natural tree roosts were used significantly more than bat boxes; however, there were no significant difference in the number of day's bats occupied the different roosts. Once a female was older than 1 year old the annual survival rate was between 0.79-0.87. Juveniles had a first year survival rate of 0.38-0.58. It was not possible to calculate male survival rates. These findings shed new light on how commercial plantations are a valuable foraging and roosting habitat for Natterer's bats and given the large amount of commercial coniferous forest now planted in the U.K. this has important implications for planning and implementing conservation management of these species. Chapter 7 contains an action plan with recommendations on how to improve existing and new coniferous forests to make them more "bat friendly" and to promote bat conservation