Sexual conflict arises in biparental species because of the fitness payoffs of shifting the costs of care onto their partner. The negotiation model asserts that parents actively monitor their partner’s level of investment and adjust their own in response generally resulting in partial compensation when a member of the pair reduces their level of investment. The willingness of one parent to compensate for the other’s change is found to be widely variable. Habitat or resource quality available to pairs may explain such variation. It is predicted that the level of compensation by one partner will increase with decreasing resource quality. I tested this prediction with the biparental burying beetle, <i>Nicrophorus orbicollis</i>. Under natural conditions, burying beetles apply costly social immune molecules to carcasses of small vertebrates to preserve nutritional value for young. The goals of this research were (1) to determine if males immunologically compensate when females are immunologically handicapped; and (2) determine the impact of resource quality on compensation. Changes in lysozyme-like-activity (LLA) and phenoloxidase (PO) production in oral and anal secretions were used to quantify social immune investment. Pairs were provided a mouse carcass of one of three qualities: freshly thawed, aged for 3 days, or aged for 7 days. As expected, female LLA decreased once injected with lipopolysaccharide (LPS) irrespective of carcass quality. Injections caused significant down regulation of oral PO in females. Male LLA increased as the quality of carcasses decreased when paired with handicapped females. Males showed no changes in PO across treatments or carcass types. My results demonstrate that males compensate for a change in maternal investment and the level of compensation increases as resource quality decreases.

In dense mating aggregations, such as insect and anuran choruses, signals produced at the same time can overlap and interfere with one another, reducing the ability of receivers to discriminate between individual signals. Thus, evolution by sexual selection is expected to result in mating signal timing strategies that avoid overlap. Patterns of signal alternation between competing males are commonly observed in leks and choruses across taxa. In some species, however, signalers instead deliberately overlap, or ‘synchronize’, their mating signals with neighboring conspecifics. Given the assumed high cost of reduced mate attraction when signals overlap, mating signal synchronization has remained an evolutionary puzzle. Synchronization may be beneficial, however, if overlapping signals reduce the attraction of nontarget receivers (predator avoidance hypothesis). Synchronized signals could also constructively interfere, increasing female attraction to the mating aggregation (the beacon effect hypothesis). I investigate these functions of synchronized signaling in two species of tree frogs that synchronize their mating calls: the pug-nosed tree frog (<i>Smilisca sila</i>) and the Ryukyu Kajika frog (<i>Buergeria japonica</i>). To examine the trade-offs imposed by call synchronization in each species, I conduct a series of field and laboratory playback experiments on target (female frogs) and nontarget (eavesdropping predators) receivers of frog calls. Results from these experiments support both hypotheses, suggesting that synchronized frog calls can reduce the attraction of predators and attract mates to the chorus. In addition, I found reduced preferences for fine-scale call timings in female <i>S. sila</i> and <i>B. japonica</i>, deviating from the expected preferences observed in many other anuran and non-anuran species. Thus, while males may enjoy multiple benefits from synchronized mating signals, relaxed sexual selection for non-synchronous signals may be key to the evolution and maintenance of mating signal synchrony.

<p dir="ltr">Bat population numbers are declining in the Midwestern United States. Reasons for decline are multifaceted (habitat degradation, fatalities at wind turbines, White Nose Syndrome, and declining insect populations), and many species are listed as endangered (Myotis sodalis, Myotis septentrionalis, Perimyotis subflavus). Other species in the Midwestern United States have no conservation status (Eptesicus fuscus), or are only listed as a species of concern (Lasiurus cinereus, Lasiurus borealis, Lasionycteris noctivagans). Bats play a crucial role in our ecosystems, providing both ecological and economic benefit as pollinators and insect population regulators. Thus, conserving these species is vital. To gain better insight into the ecology of Midwestern bat species I studied five species in two respects. First, I investigated the availability of roosts for a colony of endangered Myotis sodalis near Indianapolis. This colony has withstood high levels of urbanization and habitat degradation. Therefore, understanding what aspects of the roosting area has allowed for continued use by the colony is crucial for future conservation efforts. I used an Akaike’s Information Criteria approach to rank models that best differentiate between the current roosting area and surrounding landscape. I identified that the roosting area contained a greater number of large standing dead trees (Snags >42.6 cm) that are able to serve as primary roosts for the colony. These results demonstrate that a colony of Indiana bats may be able to withstand urbanization if they have enough large DBH snags available in the area. Future conservation efforts in a heavily urbanized environment should focus on the maintaining large snags as primary roosts. Second, I studied the morphological variation of four species (Lasiurus cinereus, Lasiurus borealis, Lasionycteris noctivagans, and Eptesicus fuscus) to parse out differences that may lead to niche specialization. These four species share habitat and foraging range and therefore may directly compete for resources. However, despite declining insect populations these four species have moderately stable populations. I collected approximately 30 craniums and mandibles for each species and compared the linear size differences between landmarks and the overall shape variation from these landmarks. Due to different phylogenies and body sizes, the four species were different from one another in all 24 linear measurements. In regards to shape variation, the Eptesicini bat craniums had characteristics of more gracile species. In addition, the mandibles of Eptesicini were highly distinct. Lasionycteris noctivagans was the least durable and Eptesicus fuscus had 10 specialization for hard bodied prey consumption. These results suggest potential niche specialization due variation in morphology.</p>

<p dir="ltr">Rapid ʻōhiʻa death (ROD) is a disease complex caused by two <i>Ceratocystis fungi</i>, <i>C. lukuohia</i> and <i>C. huliohia</i>, that is devastating the keystone tree of the Hawaiian Islands, ʻōhiʻa lehua (<i>Metrosideros polymoropha</i>). The causal agents of ROD were identified in 2015 and I began researching entomological aspects of the complex in 2016. Much like other <i>Ceratocystis</i> diseases, my colleagues and I suspected that beetles and frass might be involved in the system. Together, we identified four species of invasive ambrosia beetles (Coleoptera: Curculionidae) that contribute to the spread of ROD: <i>Xyleborinus saxesenii, Xyleborus affinis, Xyleborus ferrugineus</i>, and <i>Xyleborus perforans</i>. Both ROD-<i>Ceratocystis</i> fungi and the ambrosia beetles inhabit the xylem of ʻōhiʻa. When these beetles create their home galleries, they produce frass particles infested with resting chlamydospores that can be transported in the environment through the soil, wind, and water. Secondly, the beetles are capable of vectoring the fungi directly to stressed trees via viable propagules attached to their exoskeleton. The natural progression of this research was to investigate the chemical ecology of the system, therefore building the foundations for management strategies to reduce the spread of ROD. In addition, I satisfied my curiosity to explore the fungal mutualisms of these beetles through the use of phylogenetics.</p><p dir="ltr">In Chapter 1, I review the literature describing ROD and the four ROD-associated ambrosia beetle species. I report all of the research to date regarding ROD, including current monitoring and management strategies. Then, I introduce ambrosia beetles and the Xyleborini tribe, focusing on the life history of the ROD-associated beetles and current literature describing the use of semiochemicals to control them.</p><p dir="ltr">In Chapter 2, I determine the volatile organic compounds associated with the ROD <i>Ceratocystis</i> – ʻōhiʻa pathosystem and the response of the associated beetles to those compounds. I investigated the volatiles produced by <i>C. lukuohia</i> and <i>C. hulihia</i> in culture in addition to when inoculated into ʻōhiʻa seedlings. Then, I describe olfactometer assays to determine if the ROD-associated beetles are attracted to the volatiles emitted from ROD-<i>Ceratocysti</i>s in culture.</p><p dir="ltr">In Chapter 3, I investigate semiochemicals for attracting and repelling ambrosia beetles in ʻōhiʻa forests. I describe separate trapping experiments, first, testing the attraction of beetles to 100% ethanol and 1:1 methanol ethanol. Second, we investigate the use of two beetle repellent products, one with verbenone and the other with verbenone + methyl salicylate active ingredients.</p><p dir="ltr">In Chapter 4, I describe the testing of the repellent, verbenone, in the SPLAT^® Verb formulation, to deter ambrosia beetle attack from both healthy ʻōhiʻa trees and trees infested with ROD-<i>Ceratocystis</i>. Over two field seasons, we monitored ambrosia beetle attacks on trees treated with verbenone and measured the abundance of verbenone released from the repellents over time during the first season.</p><p dir="ltr">In Chapter 5, I investigate the ambrosia fungi of the ROD-associated beetles and native Hawaiian ambrosia beetles on the Island of Hawaiʻi. We isolated a dozen fungal symbionts from the mycetangia of ambrosia beetles, most of which are first reports in Hawaiʻi, and use phylogenetics to investigate putative new species of <i>Raffaelea</i> and <i>Ambrosiozyma.</i></p><p dir="ltr">Finally, in Chapter 6, I synthesize the results and future directions of the aforementioned chapters. Together, these dissertation chapters provide insights into ambrosia beetle monitoring and management strategies in Hawaiʻi and beyond. I describe the groundwork for understanding the pathosystem from a chemical ecology perspective and touch on the understudied world of Hawaiʻi fungi and potential pathogens.</p>

Full thesis document accompanied by all data files and R scripts to reproduce analyses in both data chapters (2 & 3). / Animals should favour breeding locations that maximize their lifetime reproductive output. Parents ought to rear young in sites that positively affect offspring but sometimes these same sites inflict costs on parents. How parents balance their own needs against those of their offspring when selecting a site for reproduction remains unclear, particularly in animals that provide extended parental care at one location. Further, few studies have linked variation in life history traits to reproductive site choices within a single species. In this thesis, I addressed both knowledge gaps by studying the plainfin midshipman (Porichthys notatus), a marine toadfish where fathers provide sole parental care to broods of young in intertidal and shallow subtidal nests for up to two months. In Chapter 2, I measured the costs incurred by caring males and the benefits conferred to offspring in nest sites along a tidal gradient. Males suffered similar rates of body condition deterioration in all nests across the intertidal gradient. Young developed more quickly in intertidal nests compared to subtidal nests, but broods in the highest intertidal nests suffered the highest mortality rates, despite receiving more parental care from the males at these nest sites. We found the most competitive males in lower intertidal nests, a trend that agrees with life history theory—in species with relatively slow offspring development, parents should accrue greater reproductive benefits from nest sites where offspring benefits are highest. In Chapter 3, I describe a laboratory experiment designed to examine how warm water and air exposure (two abiotic conditions that vary dramatically within the intertidal) affect development and survival of plainfin midshipman young, and how these effects vary with maternal traits. Exposure to warm water enhanced embryonic and larval development rates; this effect was attenuated by air exposure. Juveniles raised in warm water also exhibited superior swimming performance, while air-exposed young suffered higher mortality rates. Although larger juveniles emerged from larger eggs, development rates were similar across egg sizes. Offspring survival increased with maternal body condition in cold water but decreased with maternal body condition in warm water. Juvenile body sizes increased with maternal condition in cold water without emersion, and in warm water with emersion—the two ecologically relevant rearing environments. Thus, low condition mothers might accrue greater benefits by depositing their eggs in nests at higher tidal elevations—where development is more rapid—further supporting the idea that among-individual variation in the expression of life history traits might influence nest site preferences in these fish. In sum, my research (Chapters 2 & 3) elucidates the link between life history traits and the spatial component of animal reproductive strategies. / Thesis / Master of Science (MSc)

La communication chimique, c'est-à-dire le transfert d'information d'un émetteur à un receveur via signaux moléculaires, est présente dans tous les embranchements animaux. Longtemps négligée, l'étude récente de ces processus chez les oiseaux suggère que les signaux chimiques (ou ‘chémosignaux') ont une fonction beaucoup plus importante que longtemps anticipée. Les pétrels hypogés (ordre: procellariiformes) fournissent un modele approprié pour examiner ces questions. Ces oiseaux marins possèdent en effet une neuro-anatomie olfactive développée, une odeur musquée caractéristique et des traits d'histoire de vie favorisant l'évolution d'une composante olfactive aux comportements sociaux. En utilisant une combinaison de méthodes, existantes et spécifiquement développées, d'écologie comportementale, de chimie analytique et de statistique multivariées, nous avons examiné le rôle des chémosignaux dans l'écologie du pétrel bleu (Halobaena caerulea), un pétrel hypogée de l'océan Subantarctique. Nous avons ainsi démontré que la sécrétion uropygiale des pétrels bleus, leur source principale de substances chimiques exogènes, contient des informations sociales telles que l'espèce, le sexe et l'identité (i.e. une signature chimique). De plus, cette information est encore présente, de manière quasi-identique, sur le plumage des oiseaux et participe donc vraisemblablement à l'odeur des individus. En termes de perception des signaux chimiques, nous avons établi que les pétrels bleus sont capables de percevoir et distinguer entre les odeurs de différentes espèces de pétrels, ainsi qu'entre les odeurs de différents conspécifiques. Cependant, aucune capacité de discrimination olfactive intersexuelle n'a été observée. Longtemps restreinte aux comportements de recherche alimentaire et d'orientation, l'étude de l'olfaction aviaire est en pleine expansion pour désormais intégrer des aspects sociaux. Nos résultats fournissent en ce sens une première étude multidisciplinaire du sujet. La clarification de l'origine, de la nature et de la fonction de la communication chimique chez les oiseaux devrait avoir des implications éco-évolutives majeures pour la compréhension de leur biologie. / Chemical communication, the transfer of information from an emitter to a receiver via molecular signals, occurs in all animal phyla. Although such processes have been largely overlooked in birds, recent results suggest that chemical signals may play a more significant role than previously assumed in the social lives of birds. Procellariiform seabirds, and burrow-nesting petrels in particular, are appropriate models to investigate these questions. They indeed possess a well-developed olfactory anatomy, a noticeable musky scent and a life-history which favours the evolution of olfactory-mediated social behaviours (Chapter1).We have explored the role of chemical signals in the ecology of the blue petrel (Halobaena caerulea), a burrow-nesting seabird from the Subantarctic Ocean, using existing and innovative methods from field ornithology, analytical chemistry and multivariate statistics (Chapter 2). We first demonstrate that the uropygial secretions of these birds, their main source of endogenous chemical substances, contain social information including species, sex and individual identity (i.e. a chemical signature). We also show evidence that these signals are still present, in a virtually identical form, on the plumage of the birds and are thus a likely contributor to the animals' scent (Chapter 3). Furthermore, we show that blue petrels, as receivers of the sociochemical information, are able to discriminate between the odours from different species, and between the odours of different conspecifics. There is however no evidence of olfactory capabilities of sex discrimination in this species ( Chapter 4).The study of avian olfactory behaviours, historically limited to foraging and orientation (Chapter 5), is rapidly expanding to incorporate social functions. In this regard, our results provide the first multidisciplinary case-study of avian chemical communication. The elucidation of the origin, nature and function of chemical communication in birds has major eco-evolutionary implications for our understanding of avian ecology (Chapter 6).

In societies where all individuals are reproductively totipotent and yet, at a given time only one of them reproduces, it is interesting to examine the factor(s) that may influence and predict who will be the reproductive. I am investigating various behavioural, morphological and physiological parameters in the primitively eusocial wasp Ropalidia marginata, and their role in determining the current reproductive and her future successors. In several group-living species, especially in primitively eusocial ones, a strong link between behavioural dominance and reproductive dominance is observed. Hence, I am also investigating the possible determinants of behavioural dominance in R. marginata. I have carried out my study on artificially constituted pairs of wasps as well as in natural colonies in laboratory cages, which represent the founding phase and the established phase in the colony cycle, respectively. Chapter 1: Behavioural and Reproductive Dominance in Pairs of R. marginata Age and body size had no effect on behavioural dominance in pairs of R. marginata, whereas prior experience of behavioural dominance affected future dominance status, indicating presence of winner- and loser-effects. Dominance ranks are relatively stable. This is different from what has been found in colonies, where dominance ranks sometimes change on a daily basis. Body size had no effect, whereas age and behavioural dominance had a significant effect on reproductive dominance in pairs, with older individuals and more dominant individuals having a higher probability of becoming the reproductive. Since no relationship was found between age and behavioural dominance, we predict that the underlying mechanisms by which age and behavioural dominance affect reproductive dominance and independent of each other. This study gives a clear indication that age and behavioural dominance are important variables that determine the reproductive individual during the founding phase of the colony. Chapter 2: Comparison of Dominance Indices and Recommendations for their Use When several individuals interact with each other as in colonies, in a differential and sometimes in a preferential manner, it is difficult to attribute dominance ranks to individuals. Dominance indices are employed to simplify these interactions and rank individuals in dominance hierarchies. Since the rationale behind using a particular dominance index is seldom given in behavioural literature, a comparison of three dominance indices was carried out in second part of the thesis. Each index was gauged on how similar are its ranks as compared to other two indices. Indices were also compared based on the number of untied or unique ranks they attributed. The index that gave least number of ties in ranks was assumed to be better than others. In addition to data from R. marginata colonies, I used data from R. cyathiformis colonies (a congeneric species which behaves more like a typical primitively eusocial species), and artificial data sets, to increase variability in the interaction patterns. We found that each of the indices had their own advantages and disadvantages. In species like R. marginata and R. cyathiformis, where only a few pairs show interactions, and among those who do, very few show reversals, Frequency-based Dominance Index (FDI) is the recommended index of choice. Studies like these will help in understanding how dominance indices operate under certain situations before applying them to construct hierarchies. Chapter 3: Behavioural and Reproductive Dominance in Colonies of R. marginata Age does not affect behavioural dominance, whereas winner and loser effects exist in colonies of R. marginata, just as in pairs. When analysed in detail, I found that colonies of R. marginata showed fewer proportion of pairs interacting, and lower frequency/hour/pair of dominance-subordinate interactions as compared to experimentally paired individuals (from 1st chapter). However, the dominance displays and behaviours were much more intense and severe in colonies. After dominance hierarchies are already established in colonies, frequent need to show dominance behaviour may not arise, due to familiarity between interacting individuals. However, since individuals are possibly aware of each others’ strengths due to past interactions, dominance behaviours are much more severe when contests do happen. My results show that there might be some similarities in terms of determinants of behavioural dominance between pairs and colonies, but the expression of behavioural dominance is quite different. From earlier work it was already known that if the queen/reproductive of the colony disappears or is experimentally removed, one of the individuals shows extreme levels of aggression. This individual, referred to as the potential queen (PQ), will go on to become the next queen of the colony. Her behavioural profile, from the emergence till she establishes herself as the next queen have been well studied earlier. What was not known were the factor(s) that determine the identity of the PQ. It was also unclear what happens when the queen as well as the PQ are both removed, simultaneously or in quick succession. To test whether there is a longer reproductive hierarchy in R. marginata, the queen and the first potential queen of a nest were removed. I found that successive potential queens emerged as readily as the first potential queen, and with dominance profiles comparable to the first PQ, indicating that a reproductive hierarchy indeed exists, at least up to five PQ’s. It was also found that these potential queens were acceptable to all other individuals, as there was not a single act of behavioural dominance directed toward any potential queen. It was also observed that all PQs went on to become queens if the previous queen or PQ was not returned. When tested for various morphological, physiological, behavioural and life history traits (factors possibly influencing the position of an individual in the reproductive hierarchy), we found that age is the only variable that emerges as an important predictor of reproductive succession, with older animals having a higher chance to succeed as next queens of the colony, although even age is not an absolute predictor. Unlike in the pairs, in colonies of R. marginata behavioural dominance is not a good predictor of an individual’s ability to be the queen or the potential queens. The four most important findings of my study are: (i) the first demonstration of winner and loser effects in social insects; (ii) the demonstration that behavioural dominance influences reproductive dominance in pairs but not in colonies; (iii) demonstration of a long reproductive queue among individuals of a colony; and (iv) discovering that age is an important predictor of the identity of the queen and the future queens of the colony. I believe these findings will add significantly to our growing knowledge of the social biology of R. marginata. Finally, my work shows that pairs of R. marginata, representing the founding phase of the colony, behave more like a typical primitively eusocial species, whereas colonies which represent the established phase of the colony cycle behave more like highly eusocial species. Finding the characters of two different forms of sociality in the same species in different phases of the colony cycle makes R. marginata an excellent model system to study evolution of eusociality.

<p>In this thesis, I experimentally investigate feeding selectivity in fallow deer (Dama dama), with respect to plant secondary compounds, especially tannins, which can decrease the quality of foods. I found that fallow deer avoided foods with higher amounts of tannic acid and Quebracho tannin, even though the deer ate some high-tannin food. The food choice was strongly dependent on the context in which the food was presented, so that the food choice in relation to tannin content was relative rather than absolute. When high-tannin food occurred at low frequency, the deer ate proportionally less from this type of food, at least when the difference in tannin content between the two foods was large. A basic implication is that an unpalatable plant type could benefit from its unpalatability, especially when occurring at low frequency. In experiments with two patches, the finding of a stronger within- than between-patch selectivity was mirrored in associational effects. First, low-tannin, palatable food was more eaten when occurring in a high-tannin patch, which corresponds to neighbour contrast susceptibility. Second, high-tannin, unpalatable food in a less defended patch was less eaten, which corresponds to neighbour contrast defence. A proximate cause of the associational effects can be the presence of a simultaneous negative contrast, which was experimentally demonstrated in an additional study. Individual differences in selectivity were present early in life and were consistent over five years, and selectivity was correlated with foraging exploratory behaviour. The results from this thesis suggest that fallow deer are selective in their food choice with respect to tannins from the beginning, and that the frequency of occurrence of different foods, but also the distance between foods and the complexity of presentation, influence the food choice. It is also suggested that a foraging behavioural syndrome is present in mammalian herbivores.</p>

<p>The evolutionary goal of individuals is reproduction and sexual selection favours traits improving reproductive success. When males invest less than females in offspring, males have potentially a higher reproductive rate than females. This typically results in sex-specific reproductive strategies of male-male competition and female choice of mating partner. Under polyandry, sexual selection can continue after copulation as sperm competition and cryptic female choice. This thesis focuses on male and female pre- and post-copulatory reproductive strategies in the promiscuous red junglefowl, <i>Gallus</i> <i>gallus ssp.</i>, and its domestic subspecies the domestic fowl, <i>Gallus gallus</i> <i>domesticus</i>. Males impose high re-mating rates on females, which triggers female resistance in copulations. In addition, when sexual harassment increases, females re-mate at times of day when male mating propensity is lower, to avoid intense sexual harassment. Males allocate sperm supplies differentially according to (i) variation in female polyandry and own competitive ability, (ii) earlier sperm investment in a female, and (iii) female reproductive quality, signalled by female comb size. Males also perform ‘aspermic’ copulations (i.e. copulations with no semen transfer), which inhibit polyandry and in turn reduce sperm competition. In mating opportunities with relatives, males do not avoid inbreeding. However, females avoid inbreeding before copulation through kin recognition and after copulation by selecting against related males’ sperm. These results show that selection on males to re-mate at higher rates than females and copulate indiscriminately according to partner relatedness, trigger counteracting female responses, creating the potential for sexual conflict over fertilisation. Teasing apart pre- and post-copulatory strategies and the contribution of each sex therefore becomes crucial in order to understand the evolution of reproductive strategies and the mechanisms affecting paternity.</p>

<p>When breeding wild species in captivity, the animals may gradually become more adapted to captivity and therefore less suited for reintroduction which is the ultimate goal for some species. This study measured the activity budget of European bison (<em>Bison bonasus</em>) in six enclosures in Sweden with the aim to find out how the characteristics of the enclosures – with and without pasture - influenced the activity budget. The results show that there were significant differences in the activity budget, i.e. the activity was higher in the enclosures with pasture than in enclosures with barren ground. However, since barren enclosures were smaller than naturalistic, it was not possible to exclude the effect of size. Judged from observations of bison in the wild, there seems to be a direct correlation between food availability and ranging, indicating that enclosure characteristics affect activity more than size. The bison foraged differently in the two enclosure categories, but the total amount of time spent on feeding did not differ. A feeding enrichment experiment showed to have more positive effects in the barren enclosures than in the naturalistic ones, as the amount of time of inactivity decreased in the former. Since the genetic characteristics of all Swedish bison are very similar, the differences between the two enclosure groups indicate that the animals still have the ability to respond appropriately to improved environmental quality. Hence the next step in the assessment of the suitability of these bison for release would be to study them during an acclimatisation program.</p>