A central goal of life history theory is to understand the selective factors that generate the diversity of reproductive patterns observed in nature. Within lifetime changes in reproductive investment will determine an organism's fitness; however, this area of life history theory has received less attention than comparisons among population that characterize life history traits as a single population mean. Reproductive allocation can be affected by multiple cues; the integration of these cues across an organism's lifetime generates the diversity in life history strategies observed in nature. Life history studies should examine the interacting effects of multiple cues on life history strategies to generate better predictions and generalizations of age-related changes in reproductive investment. An individual's life history strategy is inherently multivariate consisting of a coordinated suite of life history traits that, when combined across the organism's lifetime, determines its fitness. Life history strategies can therefore be described as a trajectory through multivariate space defined by life history traits. Here I describe life history trajectory analysis, a multivariate analytical approach for quantifying and comparing phenotypic change in life history strategies; this methodology is adapted from an analytical framework originally described for studies of morphological evolution. Life history trajectories have attributes (magnitude, direction, and shape) that can be quantified and statistically compared among taxa to determine if life history patterns are predictable. Using the life history trajectory analysis, I demonstrate the effect of prior experience on reproductive allocation in the burying beetle Nicrophorus orbicollis. The effect of prior experience resulted in a terminal investment or accentuated response to age-based cues, or resulted in a conservative investment strategy or reproductive restraint. In the livebearing fish Gambusia affinis, females adjust the level of reproductive investment to current reproduction based on age- or environment-based cues. Age-0 females decreased the level of reproductive investment to current reproduction in late summer prior to the onset of fall and winter months. Old females, on the other hand, increased the level of reproductive investment as the summer progressed. The reproductive restraint and terminal investment patterns exhibited by age-0 and age-1 females, respectively, were consistent with the predictions from the cost of reproduction hypothesis. These studies demonstrate how the life history trajectory analysis provides an analytical tool to test predictions of life history theory. Additionally, I provide evidence that organisms use multiple cues to determine the level of reproductive investment and that the strength of the effect of each cue will depend on the age of an individual.

Tesi realitzada al Instituto Pirenaico de Ecología (CSIC) / Widespread species have traditionally received much less attention than rare and endemic ones. However, they are crucial in macroecological patterns and in ecosystem structure and functioning. Thus, understanding the characteristics that allow widespread organisms to extend over large areas has a high interest from both theoretical and applied perspectives. One of the most frequent hypotheses to explain the success of widespread plants is that they show much wider ecological niches, and thus a high life-history and demographic variability. However, studies are often very specific and carried out over small spatio-temporal scales, which hinders a general understanding of intraspecific variation in widespread taxa. In this thesis, we span a large spatio-temporal scale and a large environmental gradient to analyze the magnitude and the possible causes of natural variation in the in the range centre and the northern periphery of the widespread herb Plantago coronopus. More precisely, we analyze variability in population dynamics, life-history traits, and genetic diversity in up to 22 populations in Europe and North Africa. We aim to explore the relation of such variability with the position of populations within the species’ range, since peripheral populations are traditionally expected to show a lower and more variable performance with respect to central populations. Additionally, we aim to analyze the effects of the most relevant environmental factors in population and individual performance at different spatial scales. In the first chapter, we found higher values in central populations in some vital rates, such as fecundity and growth, but recruitment and density were higher in northern peripheral populations, and there were no clear differences between regions in temporal variability of vital rates. Differences in population performance across the species’ range seemed to be correlated with local precipitation and intraspecific competition. In the second chapter, differences in mean values and variability of vital rates between central and peripheral areas led to no differences in stochastic population growth rates. In addition, recruitment was the most influential vital rate for population growth rates at different spatial scales, and we found the same pattern of differentiation in population dynamics in response to environmental conditions within central and peripheral regions. In the third chapter, we reported high variation among populations in seed traits along a steep environmental stress gradient. Moreover, patterns in seed production were opposite at the fruit and the individual scale, as a strategy of populations to maximize fitness in each set of local conditions. Finally, in the fourth chapter, we found no relationship within populations between phenotypic variability and genetic diversity. Phenotypic variation was mainly shaped by precipitation variability, suggesting adaptive variation, whereas genetic diversity was correlated with the central vs. peripheral position, probably in close relation with some random demographic processes experienced by populations in the past. Despite genetic diversity was higher in central populations, our results contradicted classical hypotheses predicting a lower demographic performance towards species’ range edges. In fact, environmental conditions seemed to have a higher influence on plant performance than the position of populations within the species’ range, which calls for the necessity of distinguishing between geographical periphery and ecological marginality in demographic studies. Overall, our study highlights the versatility of P. coronopus at different spatial scales in response to varying environmental conditions, complementing similar findings of previous research on the same taxon at smaller spatial scales. Such life-history variability seems to be a key factor for widespread plants to extend over large and heterogeneous ranges. / Las especies de amplia distribución han recibido tradicionalmente poca atención, a pesar de su importancia para la estructura y el funcionamiento de los ecosistemas. En esta tesis, se analiza la variabilidad demográfica, de historia vital y genética en una planta de amplia distribución en Europa y el norte de África (Plantago coronopus), en un total de 22 poblaciones a lo largo de gran parte del rango latitudinal de la especie (centro y periferia norte). Se pretende analizar la magnitud y las causas de esta variabilidad intraespecífica en relación con la posición de las poblaciones dentro del rango y con los principales factores medioambientales. Las poblaciones periféricas mostraron una menor diversidad genética, pero no mostraron en general un peor o más variable comportamiento demográfico en cuanto a densidad o tasa de crecimiento poblacional, contradiciendo así las hipótesis clásicas centro-periferia. Se encontró un mismo patrón de diferenciación demográfica dentro de las regiones tanto central como periférica, en relación con la variación en el régimen de precipitaciones. La tasa de reclutamiento de nuevos individuos fue el proceso del ciclo vital con mayor importancia para el funcionamiento de las poblaciones. Se encontró también una gran variación entre poblaciones en las características de las semillas (número, tamaño, mucílago y proporción de dos tipos de semilla dimórficos) en relación con el gradiente de estrés ambiental. Finalmente, la variación fenotípica dentro de las poblaciones se relacionó con la variabilidad ambiental, mientras que la diversidad genética se correlacionó con la posición central vs. periférica de las poblaciones y posiblemente con la historia demográfica de la especie. Globalmente, este estudio muestra la importancia de distinguir entre periferia geográfica y marginalidad ecológica, y sugiere que el éxito de las plantas de amplia distribución reside en una gran variabilidad demográfica y de historia vital a diferentes escalas espaciales.

Life history traits affect the timing and pattern of maturation, reproduction, and survival during an organism’s lifecycle and are the major components influencing Darwinian fitness. Co-evolved patterns of these traits are known as life history strategies (LHS) and variation occurs between individuals, populations, and species. The polymorphisms underlying LHS are important targets of natural selection, yet the underlying genes and physiological mechanisms remain largely unknown. Mapping the genetic basis of a LHS and subsequently unraveling the associated physiological mechanisms is a challenging task, as complex phenotypes are often polygenic. However, in several systems discrete LHS are maintained within the population and are inherited as a single locus with pleiotropic effects. These systems provide a promising starting point for investigation into LHS mechanisms and this thesis focuses on one such strategy - the Alba polymorphism in Colias butterflies. Alba is inherited as a single autosomal locus, expressed only in females, and simultaneously affects development rate, reproductive potential, and wing color. Alba females are white, while the alternative morph is yellow/orange. About 28 of 90 species exhibit polymorphic females, though whether the Alba mechanism and associated tradeoffs are conserved across the genus remains to be determined. In this thesis I primarily focus on the species Colias croceus and integrate results from lipidomics, transcriptomics, microscopy, and genomics to gain insights to the proximate mechanisms underlying Alba and Alba’s evolution within the genus. Lipidomics confirm that, consistent with findings in New World species, C. croceus Alba females have larger abdominal lipid stores than orange, an advantage which is temperature dependent and arises primarily due to mobilized lipids. Gene expression data suggests differences in resource allocation, with Alba females investing in reproduction rather than wing color, consistent with previous findings in other Colias species. Additionally, I identify a morphological basis for Alba’s white wing color. Alba females from C. croceus, an Old World species, and Colias eurytheme, a New World species both exhibit a significant reduction in pigment granules, the structures within the wing scale that contain pigment. This is a trait that seems to be unique to Colias as other white Pierid butterflies have an abundance of pigment granules, similar to orange females. I also map the genetic basis of Alba to a single genomic region containing an Alba specific, Jockey-like transposable element insertion. Interestingly this transposable element​ is located downstream of BarH-1, a gene known to affect pigment granule formation in Drosophila. Finally, I construct a phylogeny using a global distribution of 20 Colias species to facilitate investigations of Alba’s evolution within the genus. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 4: Manuscript.</p>

I investigated streambed stability in spawning microhabitats of warmwater fishes in the upper Roanoke River, Virginia. Spawning microhabitats used by four reproductive guilds (egg-clusterers, mound-builders, egg-buriers, and pit-builders) were identified and ranges of 15 microhabitat variables used by each guild were compared to available microhabitat conditions to investigate habitat selection. Habitat usage by egg-clusterers was most characterized by selection for spawning rocks in the cobble size range, substrate roughness elements in the cobble to boulder size range, high roughness Reynolds number and moderate water column velocity. The mound-building bluehead chub (Nocomis leptocephalus) was distinct in its selection of substrate in the small gravel range, low water velocity and non-turbulent flow. Egg-buriers were the least distinct of the four guilds, exhibiting much variation in habitat use among the component species. However, all species used areas with small substrate (sand to gravel range), high velocity, and high turbulence. The pit-building central stoneroller (Campostoma anomalum) was distinct in its usage of areas with high velocity and turbulence, gravel sized substrate, and low embeddedness. Stability of each guild's spawning microhabitats was empirically evaluated through analysis of tracer particle movement and repeated surveying of bed elevation along stream transects. Logistic regression equations developed from tracer particle data predicted that microhabitats selected by egg-clusterers are among the most stable of all available habitats during high flows. Microhabitats utilized by mound-builders, egg-buriers, and pit-builders are predicted to be less stable. Repeat transect surveying corroborates model predictions in that egg-burier habitats experienced changes in bed elevation in high flows, while egg-clusterer habitats did not. / Master of Science

Einer der ausschlaggebenden Faktoren für das Übertragungspotenzial von Malaria ist die Häufigkeit ihres Überträgers, der Anopheles-Mücke. Da diese Insekten besonders empfindlich auf ihre Umgebung reagieren, wurde in vielen Laborexperimenten das Verhältnis zwischen Umweltfaktoren und Malaria untersucht. In natürlichen Umgebungen ist über diesen Zusammenhang wenig bekannt. Untersuchungsgegenstand dieser Arbeit mit multidisziplinärem Ansatz ist die Beziehung zwischen verschiedenen Umwelteinflüssen und der Häufigkeit von Anopheles-Mücken in natürlicher Umgebung sowie die möglichen diesem Prozess zugrundeliegenden Mechanismen. Zunächst lässt sich anhand von Feldstudien und empirischen dynamischen Modellen eine verzögerte Auswirkung der Umwelt auf die Mückenhäufigkeit feststellen, was wiederum einen Einfluss auf die Entwicklung der Larvenstadien bedeuten könnte. Im Anschluss werden mögliche Hintergründe untersucht mit Hilfe eines kontrollierten Versuchsaufbaus, in dem Anopheles-coluzzii-Larven und Erwachsenentiere auf Umweltbedingungen reagieren. Dabei lässt sich feststellen, dass Temperatur und Besatzdichte vielschichtige Auswirkungen auf Larvenentwicklung und -sterblichkeit haben. Darüber hinaus gibt es Übertragungseffekte auf die Alterung und Fruchtbarkeit der Erwachsenentiere. Abschließend zeigt sich durch die Entwicklung eines vollständigen Modells für den Lebenszyklus von Anopheles-Mücken, dass die ermittelten Umwelteinflüsse auf Larven und erwachsene Mücken einen komplexen Einfluss auf die Größe der Mückenpopulation haben. Anhand mehrere mathematischer Methoden werden in dieser Arbeit Informationen aus Feldsammlungen und Versuchsdaten erfolgreich kombiniert. Dies unterstreicht die Bedeutung multidisziplinärer Ansätze für das Verständnis der komplexen Dynamik zwischen Mücken und der Umwelt. So entsteht ein detailliertes Bild der Lebensmerkmale der Mücken, die von der Umwelt beeinflusst werden, und deren Auswirkungen auf die Populationsdynamik. / One of the main determinants of malaria transmission is the abundance of its vector, the Anopheles mosquito. As these insects are susceptible to the environment in which they live, many laboratory experiments have explored the connection between environmental factors and the abundance of mosquitoes. However, little is known about this relationship in natural settings. In this work, I used a multidisciplinary approach to determine the relationship between environmental drivers and Anopheles abundance and the possible mechanisms that characterise these relationships. First, using field study data and an empirical dynamic modelling approach, I identified a delayed effect of the environment on mosquito abundance, suggesting an impact on the development of larval stages. Second, I explored the possible mechanisms behind this process using a controlled experimental setup in which Anopheles coluzzii larvae and adults responded to the environmental conditions. I identified a complex response to temperature and rearing density of larval development and mortality, as well as carry-over effects on adult ageing and fecundity. Finally, by developing a full model for the life cycle of Anopheles mosquitoes, I determined that the identified environmental impacts on larvae and adult mosquitoes have a complex impact on the mosquito population size. Taken together, this work combined the information from field collections and experimental data successfully using multiple mathematical methods, highlighting the importance of multidisciplinary approaches in understanding the complex dynamics between mosquitoes and the environment. With this approach, we obtained a detailed picture of the mosquito life traits that are impacted by the environment and its effects on mosquito population dynamics.

Master of Science / Department of Biology / David C. Hartnett / The effects of bison activities on forb diversity and cover have been well-established, but less is known about how forb life history is altered by bison-mediated habitat changes. This study had three main objectives: 1) to evaluate the hypothesis that release from aboveground competition with grasses may contribute to the increased cover and diversity of forbs in prairie grazed by bison relative to ungrazed prairie, 2) to determine whether differences in forb reproductive effort between grazed and ungrazed habitats were size-dependent, and 3) to look for evidence of a trade-off between allocation to vegetative and sexual reproduction. The growth, biomass allocation, and sexual reproduction of six common unpalatable perennial species were measured and compared between bison-grazed and ungrazed tallgrass prairie burned at 2-year intervals: Ambrosia psilostachya, Artemisia ludoviciana, Baptisia australis, Psoralidium tenuiflorum, Solidago canadensis, and Vernonia baldwinii. Vegetative reproduction was also measured for B. australis¸ S. canadensis, and V. baldwinii. Light availability, canopy density and height, and percent cover of neighboring plants were measured in each studied individual’s immediate neighborhood and compared between habitats to establish the possibility of differing aboveground competition. Aboveground competition may be lower in bison-grazed habitats, as evidenced by differences in habitat characteristics and plant performance found in this study. In bison-present habitats, sexual reproduction was elevated for all six species and average plant size was greater for all species except A. psilostachya. Vegetative reproduction was not clearly different between habitats for all three species examined. Sexual reproduction increased with size for all species, and the relationship differed significantly between habitats for all species except A. psilostachya. Allocation to vegetative reproduction was not generally related to aboveground biomass, nor was there a clear trade-off between allocation to vegetative and sexual reproduction. The results of this study provide evidence that release from aboveground competition with grasses promotes the growth and sexual reproduction of the studied species of forb, and that differences in sexual reproduction are not entirely size-dependent. Patterns in allocation to vegetative reproduction were less clear and were not clearly tied to sexual reproductive allocation.

Published Article / In this article we use the concepts identity, agency and social justice education as a lens to explore the role of life history research in the study of the interconnection between emerging teacher identities, critical agency and social justice education. By exploring the life history of a white woman pre-service teacher, this study foregrounds the use of life history research to help teacher educators to understand the contexts through which student teachers' identities are constructed, and how these identities feed into agency and a stance to bring about social change.

The thrust of this work is to study Israelite pre-exilic writing of religious literature. The beginning of literacy is considered from an archaeological perspective; especially, in the pre-exilic Israelite community. The study of scribes and their services assist in the quest for understanding pre-exilic religious writing in Israel. The Bible attests to pre-exilic religious writing despite the often inferred ‘anachronism.’ The issue of post-exilic composition of all Old Testament books is a matter of debate as opposed to pre-exilic writing of some religious sources which is a matter that can be historically verified. / Biblical & Ancient Studies / M.A. (Biblical Archeology)

The aim of this project is to examine the relationships between two approaches to the measurement of life history strategies. The traditional method, termed here the biodemographic approach, measures developmental characteristics like birthweight, gestation length, inter-birth intervals, pubertal timing, and sexual debut. The alternative method under exploration, termed here the psychological approach, measures a suite of cognitive and behavioral traits such as altruism, sociosexual orientation, personality, mutualism, familial relationships, and religiosity. Although both approaches are supported by a large body of literature, they remain relatively segregated. This study draws inspiration from both views, integrating measures that assess developmental milestones, including birthweight, prematurity, pubertal timing, and onset of sexual behavior, as well as psychological life history measures such as the Mini-K and a personality inventory. Drawing on previous theoretical work on the fundamental dimensions of environmental risk, these measures are tested in conjunction with several scales assessing the stability of early environmental conditions, including both "event-based" measures that are defined with an external referent, and measures of internal schemata, or the predicted psychological sequelae of early events. The data are tested in a three-part sequence, beginning with the measurement models under investigation, proceeding to an exploratory analysis of the causal network, and finishing with a cross-validation of the structural model on a new sample. The findings point to exciting new directions for future researchers who seek to integrate the two perspectives.

The following thesis presents the results of a mixed-design study (quasi-experimental and true experimental) testing an integrated model of human risk-taking behavior, defined statistically as a preference for variance in outcomes. The research presented examines the relationships among early-life environmental conditions (i.e., harshness and unpredictability), life-history strategy, and risk-taking behavior for primary resources under various "resource-budget" conditions consisting of deprivation and non-deprivation in two areas: (1) Social-inclusion and (2) caloric "Energy-budget." Two hundred and forty seven (N=247) university students participated in the research. In session one, participants completed multiple questionnaires assessing levels of environmental harshness and unpredictability experienced during development and individual life-history strategy. In session two, participants were pseudo-randomly assigned to experience laboratory induced deprivation or non-deprivation in one of two possible areas: Social-inclusion or caloric "Energy-budget." Following the experimental manipulations, participants played two different behavioral risk-taking tasks: (1) The Wheel Spin Risk Task which required participants to select between a low variance "safe" wheel and a high variance "risky" wheel in an attempt to earn either points or food rewards (depending upon study condition). (2) The Operant Risk Taking Task which required participants to select between a low variance "safe" keyboard key which produced constant rewards and a high variance "risky" keyboard key which produced variable rewards (points or food, depending upon study condition). The results of the multivariate analyses supported main effects only (no moderation) between the characteristics of the early-life environment, life-history strategy, and the experimental manipulations on risk-taking behavior. Specifically, early-life harshness was significantly associated with a faster life-history strategy in participants. Participants with a faster life-history strategy were significantly more likely to select the risky spin wheel on the Wheel Spin Risk Task than were slower life-history strategy participants who were more likely to select the safe spin wheel. Furthermore, participants who experienced the deprivation experimental manipulations behaved more risky on the Operant Risk-Taking Task (for reward amount) than did participants exposed to the non-deprivation manipulations in the study. Interestingly, this effect was domain-general in that deprivation in either Social-inclusion or Energy-budget was associated with risk taking for both social points and for food rewards. The results of this study suggest that life-history strategy is predictive of instrumental risk-taking behavior for reward amount and that deprivation in adaptive areas like Social-inclusion and Energy-budget enhances risk-taking behavior for primary rewards in a domain-general manner as opposed to a domain-specific manner.