The life history consequences of admixture between intentionally and unintentionally introduced populations of Trissolcus japonicus in Europe

<p>Identifying sources of variation in individual reproductive success is crucial to our understanding of population dynamics and evolutionary ecology. In many systems, the determinants of success are not well known. Where species have parental care, for example, determinants of success can be particularly challenging to partition between parents and offspring. In this thesis I investigate drivers and consequences of variable life histories, for a small reef fish that exhibits male parental care (the common triplefin Forsterygion lapillum). I examined the influence of individual life history, phenotype and behaviour on (1) the performance of recently settled juveniles, and (2) the reproductive success adult males. I made field-based observations of adult males during the breeding season, measured their phenotypic traits (body size and condition) and used their otoliths to reconstruct life history characteristics (hatch dates and mean growth rates). My life history trait reconstructions suggested two alternate pathways to ’success’ for adult males. Successful males hatched earlier and therefore had a developmental ’head start’ over less successful males (i.e., males with eggs > male territory holders without eggs > floaters). Alternatively, males can apparently achieve success by growing faster: for males born in the same month, those with eggs grew faster than those with territories and no eggs, and both groups grew faster than floaters. These results suggest that accelerated growth rate may mediate the effects of a later hatch date, and that both hatch dates and growth rates influence the success of adult males, likely through proximate effects on individual phenotypes. Identifying sources of variation in individual reproductive success is crucial to our understanding of population dynamics and evolutionary ecology. In many systems, the determinants of success are not well known. Where species have parental care, for example, determinants of success can be particularly challenging to partition between parents and offspring. Male parental care is common among fishes, where resources such as high quality territories and mates often may be limiting. In such systems, individual success of offspring may result from distinct life history pathways that are influenced by both parental effects (e.g., timing of reproduction) and by the offspring themselves (e.g., ’personalities’). These pathways, in turn, can induce phenotypic variation and affect success later in life. The drivers and consequences of variable life histories are not well understood in the context of reproductive success. In this thesis I investigate drivers and consequences of variable life histories, for a small reef fish that exhibits male parental care (the common triplefin Forsterygion lapillum). I examined the influence of individual life history, phenotype and behaviour on (1) the performance of recently settled juveniles, and (2) the reproductive success adult males. I made field-based observations of adult males during the breeding season, measured their phenotypic traits (body size and condition) and used their otoliths to reconstruct life history characteristics (hatch dates and mean growth rates). Some males showed no evidence of territorial defence and were defined as ’floaters’; others defended territories, and a subset of these also had nests with eggs present. Adult male body size was significantly higher for males that defended breeding territories, and body condition was significantly higher for the males that had eggs (i.e., had successfully courted females). My otolith-based reconstructions of life history traits suggested two alternate pathways to ’success’ for adult males. Successful males hatched earlier and therefore had a developmental ’head start’ over less successful males (i.e., males with eggs > male territory holders without eggs > floaters). Alternatively, males can apparently achieve success by growing faster: for males born in the same month, those with eggs grew faster than those with territories and no eggs, and both groups grew faster than floaters. These results suggest that accelerated growth rate may mediate the effects of a later hatch date, and that both hatch dates and growth rates influence the success of adult males, likely through proximate effects on individual phenotypes. I evaluated the effects of variable life history in a complimentary lab-based study. Specifically, I manipulated the developmental environments (feeding regime and temperature) for young fish and evaluated the direct effects on life history traits and phenotypes. Then, I conducted an assay to quantify the indirect effects of developmental environment, life history traits, and phenotypes on aggression and performance of young fish. These developmental environments did not have a clear, overall effect on juvenile phenotype or performance (i.e. behavioural aggression and the ability to dominate a resource). Instead, individuals (irrespective of developmental environment) that grew faster and/or longer pelagic larval durations had increased odds of dominating a limited resource. I attributed the non-significant direct effect of developmental environment to within-treatment mortality and variation among individuals in terms of their realised access to food (i.e., dominance hierarchies were apparent in rearing chambers, suggesting a non-uniform access to food). Fish that were more likely to dominate a resource were also more aggressive (i.e., more likely to engage in chasing behaviours). Fish that were larger and more aggressive established territories that were deemed to be of higher ’quality’ (inferred from percent cover of cobble resources). Overall, this study suggests a complex interplay between social systems, phenotype and life history. Developmental environments may influence phenotypes, although behavioural differences among individuals may moderate that effect, contributing to additional variation in phenotypes and life history traits which, in turn, shape the success of individuals. Collectively, my thesis emphasises the consequences of life history variability on success at multiple life stages. These results may be relevant to other species that exhibit male parental care or undergo intense competition for space during early life stages. In addition, my results highlight interactions between life history, phenotype and behaviour that can have important implications for population dynamics and evolutionary ecology.</p>

Dental microstructure records life history events: A histological study of mandrills (Mandrillus sphinx) from Gabon

Life history events, from mating and voltinism to migration and emergence, are governed by external and historically predictable environmental factors. The ways humans have altered natural environments during the Anthropocene have created myriad and compounding changes to these historically predictable environmental cues. Over the past few decades, there has been an increased interest in the control temperature exerts on life history events as concern over climate change has increased. However, temperature is not the only life history cue that humans have altered. In stream ecosystems, flow and light serve as important life history cues in addition to temperature. The timing and magnitude of peak flows can trigger migrations, decreases in stream temperature may cause a stream insect to enter diapause, and photoperiod appears to prompt spawning in some species of fish. Two or more of these cues may interact with one another in complex and sometimes unpredictable ways. Large dams and increasing impervious cover in urban ecosystems have modified flows and altered the timing of spawning and migration in fish. Precipitation draining hot impervious surfaces increases stream temperature and adds variability to the general pattern of stream warming from climate change. The addition of artificial light in urban and suburban areas is bright enough to eliminate or dampen the photoperiod signal and has resulted in caddisfly emergence becoming acyclical. The resulting changes in the timing of life history events also have the potential to influence the evolutionary trajectory of an organism and its interactions with other species. This paper offers a review and conceptual framework for future research into how flow, temperature, and light interact to drive life history events of stream organisms and how humans have changed these cues. We then present some of the potential evolutionary and ecological consequences of altered life history events, and conclude by highlighting what we perceive to be the most pressing research needs.

When faced with behavioural options differing in energetic gain and mortality risk due to predation, an individual's best compromise to the conflicting demands of growth and survival will depend upon both its current energetic state and the future opportunity for growth. Such state- and time-dependent tradeoffs are often investigated using dynamic programming. By specifying the relationship between fitness and the state variable of interest at the time of some relevant life history event. fitness-maximizing solutions for all state and time combinations can be found. To date, however, no dynamic programming model has considered the possibility that animals may be capable of delaying life history events beyond the time period modelled. In such cases, in addition to being influenced by future life history events. short term behavioural responses to foraging-predation risk tradeoffs may also indirectly affect the timing of those events. I use dynamic programming (1) to investigate the effects of body size and time of year on patterns of risk-taking behaviour in animals capable of postponing life history events, and (2) to explore the outcome of such individual decisions on the subsequent timing of life history events and the states of individuals undergoing those events. In doing so. I relax the basic dynamic programming assumption of a finite time horizon and allow individuals to postpone initiating the life history event until some future favourable period of time. Such delays are frequently observed in anadromous fishes, including coho salmon, Oncorhynchus kisutch; hence, I use the relevant features of their biology to develop the model and illustrate the general problem of interest.

Many wild populations are suffering from the loss of genetic diversity caused by habitat fragmentation, while the degree of diversity loss differs among species and populations based on their life history characteristics. Trillium camschatcense , an understory perennial plant, has undergone intensive habitat fragmentation in the Tokachi region, Hokkaido, Japan. Although demographic deteriorations, such as reduced seed production, were already reported, genetic consequences of fragmentation have not been studied with reference to its life history. Here, we examined how life history events (e.g., growth and reproduction) and the stochasticity therein influence genetic diversity in two (each large and small) fragmented T. camschatcense populations. Genetic diversity was evaluated using genome‐wide 2,008 single nucleotide polymorphisms (SNPs). In the small population, genetic diversity of newly germinated seedlings was significantly lower than that of matured life history stages, and effective number of breeders ( N b ) was smaller than that of the large population. Simulations using a matrix population model showed that the diversity loss at seedlings is caused by genetic drift during reproduction, which was intensified by smaller N b . Besides, simulations using randomly perturbed transition matrices suggested that stasis at juvenile stages, which is a common characteristics of T. camschatcense , maintains genetic diversity by buffering stochastic decrease, possibly contributing to population viability. While previous studies showed the importance to facilitate reproduction and recruitment for demographic recovery, this study highlighted the crucial roles of juvenile survival in terms of genetic diversity for the conservation of fragmented T. camschatcense populations in the Tokachi region.

Knowledge of the timing of major life history events in aquatic species is important for informing conservation and resource management planning. Accordingly, surveys of environmental DNA (eDNA) have been performed to determine the efficacy of eDNA for providing information on life history events, primarily focusing on the timing of events associated with spawning, and these studies have proved successful. However, spawning represents only one part of the life history, and therefore, information on eDNA shedding during other life history stages is needed to fill gaps in knowledge. Here, we explored eDNA shedding during early life history (from fertilized eggs until near yolk sac absorption) in Chinook Salmon (Oncorhynchus tshawytscha) at three biomasses in a laboratory environment. We found that fertilized eggs shed little eDNA prior to hatching. Hatching coincided with a spike in eDNA, and we observed a significant and positive relationship between eDNA concentration and the number of hatched eggs. The concentration of eDNA shed by larvae after hatching was not consistent across post‐hatch sampling days, suggesting developmental and behavioral changes associated with larval ontogeny may affect eDNA shedding rate. These results indicate that eDNA data may be used to identify hatch timing and verify successful reproduction in oviparous aquatic fishes. The application of eDNA to early life history broadens the capacity of eDNA‐based methods for assessing population status and trends.

Age at first intercourse in twins reared apart: Genetic influence and life history events

The life history pattern of recent humans is uniquely derived in many of its aspects including an extended post-reproductive lifespan combined with short interbirth intervals. A number of theories have been proposed to explain the evolution of this unusual pattern. However most have been difficult to test due to the fragmentary nature of the hominin fossil record and the lack of methods capable of inferring such later life history events. In search of a method we tested the hypothesis that the physiologically impactful events of parturition and menopause are recorded in dental cementum microstructure. We performed histomorphological analyses of 47 teeth from 15 individuals with known life history events and were able to detect reproductive events and menopause in all females. Furthermore, we found that other stressful events such as systemic illnesses and incarceration are also detectable. Finally, through the development of a novel analytical method we were able to time all such events with high accuracy (R-squared = 0.92).

Most research on the demography of wild animal populations has focused on characterizing the variation in the mortality of organisms as a function of intrinsic and environmental characteristics. However, such variation in mortality is difficult to relate to functional life history components (e.g. reproduction, dispersal, hibernation) due to the difficulty in monitoring biological processes at a sufficiently fine timescale. In this study, we used a 10-year individual-based data set with an infra-annual timescale to investigate both intra- and inter-annual survival patterns according to intrinsic and environmental covariates in an introduced population of a small hibernating rodent, the Siberian chipmunk. We compared three distinct periods related to particular life history events: spring reproduction, summer reproduction and hibernation. Our results revealed strong interactions between intrinsic and temporal effects. First, survival of male chipmunks strongly decreases during the reproduction periods, while survival is high and equal between sexes during hibernation. Second, the season of birth affects the survival of juveniles during their first hibernation, which does not have long-lasting consequences at the adult stage. Third, at an inter-annual scale, we found that high food resource availability before hibernation and low chipmunk densities specifically favour subsequent winter survival. Overall, our results confirm that the well-known patterns of yearly and inter-individual variation of mortality observed in animals are themselves strongly variable within a given year, suggesting that they are associated with various functional components of the animals' life history.

Weaning represents a pivotal ontogenetic process for mammals, marking the transition from parental provisioning to independent foraging. In monophyodont species, distinct growth layer groups that are deposited in their teeth as the animals age represent a permanent chronological archive of physiological events across the animals' lifetimes. Thus, biochemical analysis of annual dentine increments provides a means to explore animal physiological history. We examined the age-specific pattern, individual-level variations, and sex-related differences in dentine nitrogen isotopic values in the Indo-Pacific humpback dolphin (Sousa chinensis) using tooth samples collected from 38 carcasses that stranded ashore in the Pearl River Delta region, southeast China, between 2007 and 2018. The longitudinal isotopic records archived in dolphin teeth offered insights into their foraging ecology and individual ontogenetic dietary life history. The overall pattern of δ15N isotopic values in the incremental layers, analyzed under a hierarchical Bayesian framework, indicates that humpback dolphins typically undergo an ontogenetic dietary shift (i.e., wean) before reaching the age of 3 yr (mean: 2.394 ± 0.143 yr), albeit there is considerable individual heterogeneity (range: 1.548-4.180 yr), with males consistently weaning ∼3.5 mo earlier than females. Our study underscores the importance of quantifying ontogenetic parameters at the individual level, as overlooking individual variations in life history events (such as the age of weaning) may introduce biases in the broader population-level life history metrics. The application of hierarchical Bayesian modeling proved effective in quantifying individual heterogeneity and factoring it into the estimates of ontogenetic dietary shifts-an important component in analyses of broader population-level processes.

Climatic warming is associated with organisms breeding earlier in the season than is typical for their species. In some species, however, response to warming is more complex than a simple advance in the timing of all life history events preceding reproduction. Disparities in the extent to which different components of the reproductive phenology of organisms vary with climatic warming indicate that not all life history events are equally responsive to environmental variation. Here, we propose that our understanding of phenological response to climate change can be improved by considering entire sequences of events comprising the aggregate life histories of organisms preceding reproduction. We present results of a two-year warming experiment conducted on 33 individuals of three plant species inhabiting a low-arctic site. Analysis of phenological sequences of three key events for each species revealed how the aggregate life histories preceding reproduction responded to warming, and which individual events exerted the greatest influence on aggregate life history variation. For alpine chickweed (Cerastium alpinum), warming elicited a shortening of the duration of the emergence stage by 2.5 days on average, but the aggregate life history did not differ between warmed and ambient plots. For gray willow (Salix glauca), however, all phenological events monitored occurred earlier on warmed than on ambient plots, and warming reduced the aggregate life history of this species by 22 days on average. Similarly, in dwarf birch (Betula nana), warming advanced flower bud set, blooming, and fruit set and reduced the aggregate life history by 27 days on average. Our approach provides important insight into life history responses of many organisms to climate change and other forms of environmental variation. Such insight may be compromised by considering changes in individual phenological events in isolation.

A temporal separation of energetically costly life history events like reproduction and maintenance of the integumentary system is thought to be promoted by selection to avoid trade-offs and maximize fitness. It has therefore remained somewhat of a paradox that certain vertebrate species can undergo both events simultaneously. Identifying potential costs of overlapping two demanding life history stages will further our understanding of the selection pressures that shape the temporal regulation of life history events in vertebrates. We studied free-living tropical Slaty brush-finches (Atlapetes schistaceus), in which individuals spontaneously overlap reproduction and moult or undergo both events in separation. To assess possible costs of such an overlap we quantified feather quality and flight performance of individuals in different states. We determined individual’s life history state by measuring gonad size and scoring moult stage, and collected a newly grown 7th primary wing feather for later analysis of feather quality. Finally, we quantified flight performance for each individual in the wild. Overlapping individuals produced lighter and shorter wing feathers than individuals just moulting, with females decreasing feather quality more strongly during the overlap than males. Moreover, overlapping individuals had a reduced flight speed during escape flights, while their foraging flight speed was unaffected. Despite overlappers being larger and having a smaller wing area, their lower body mass resulted in a similar wing load as in breeders or moulters. Individuals measured repeatedly in different states also showed significant decreases in feather quality and escape flight speed during the overlap. Reduced escape flight speed may represent a major consequence of the overlap by increasing predation risk. Our data document costs to undergoing two life history stages simultaneously, which likely arise from energetic trade-offs. Impairments in individual quality and performance may represent important factors that select for temporal separation of life history stages in other species.

Mammalian brain development and our grandmothering life history

All life histories are a complex of trade-offs that depend on individual conditions—including size, age, and sex—and adaptively mold suites of traits through the biased intergenerational transmission of successful physiological strategies. I explore intrapopulation variation in how female rhesus macaques on Cayo Santiago have negotiated a central life history trade-off: when to start reproducing. I emphasize how evolutionary quantitative genetic models require explicit links between hypothesized or measured patterns of selection and the genetic substrates that influence phenotypes and change intergenerationally. Perhaps counterintuitively, I also show how genetic models offer valuable insights on how environments, including those provided by mothers and other kin, affect offspring development and later female life histories. The emerging picture of female macaque maturation is one of the great flexibility and environmental responsiveness coupled with an important genetic component that is significantly entangled with later life history events.

Fasting ameliorates oxidative stress: A review of physiological strategies across life history events in wild vertebrates