A central question in life history theory is, what combination of traits and behaviors will lead to the highest reproductive success throughout a lifetime. The trade-off between current and future reproduction is central to the lifetime reproductive success of an organism. If there is a cost to reproduction, then allocation of energy to current reproduction will come at a cost to future reproduction. We expect young individuals to favor future reproduction over current reproduction and that balance shifts to current reproduction as they age (i.e. terminal investment hypothesis). However, how this transition from an emphasis on future reproduction to emphasis on current reproduction changes throughout a lifetime should depend on environmental factors like mortality and resource availability. We test for the generality of terminal investment across three species of poecilliid fishes in a range of environments. We found evidence of terminal investment in all three species in both high and low mortality environments and high and low resource availability environments. In general, high mortality or high resource availability tended to result in a decreased slope of the relationship between reproductive allocation and body size. Terminal investment appears to be general, even though there was an effect of high mortality and resource availability, it was not sufficient to completely preclude terminal investment.

The cost of reproduction hypothesis suggests that allocation into current reproduction constrains future reproduction. How organisms accrue reproductive costs may differ between species and with varying levels of resource quality. Burying beetles are model organisms for testing for the cost of reproduction because of their unique natural history; beetles utilize small vertebrate carcasses for reproduction and they and their offspring feed exclusively on these discrete resources. Burying beetles also can utilize a large range of carcass sizes for reproduction. We tested for the cost of reproduction in two species of burying beetles, Nicrophorus marginatus and Nicrophorus guttula found in Central Utah by breeding beetles on a range of carcass sizes (5g, 10g, 20g, 30g, 40g, and 50g carcasses). We also used a manipulation experiment to force beetles into over-allocating energy into reproduction to assess reproductive costs. For both species, reproduction was costly, with beetles suffering reduced lifespan and reduced lifetime fecundity with increased resource quality. Both species also showed clear signs of senescence, having reduced brood size and lower efficiency as individuals aged. Females did not show indications of terminal investment in terms of female mass change, unlike the previously studied Nicrophorus orbicollis, which gained less mass after each reproductive attempt as it aged. Nicrophorus marginatus consistently outperformed N. guttula in terms of total number of offspring produced for all carcass sizes. Nicrophorus guttula populations may continue to persist with N. marginatus by exploiting a less desirable but more abundant resource.

In order to maximize fitness, individuals must partition their limited resources among competing physiological processes, creating negative statistical associations between processes known as “life-history trade-offs”. Evidence indicates that individuals tend to decrease their reproductive investment when confronted with a significant immunological challenge in order to increase investment in immune defense. This trade-off is often accompanied by a significant decrease in the sexual signal, which provides an honest signal of the male’s infection status to potential mates. However, if individual residual reproductive value is low, they may instead increase their reproductive investment to maximize reproductive success before the end of their life (a.k.a. terminal investment). Here, we investigate the potential for terminal investment in the ground cricket Allonemobius socius by inoculating males with varying dosages of an immune challenge. We predicted that both high dose and advanced male age would induce terminal investment. Furthermore, we predicted that terminally investing males would produce a dishonest signal by increasing their signaling effort. We found that upon infection We found that upon infection, young males and old males differentially alter their reproductive strategy. Young males exhibited the classic deceleration of reproductive effort. However, old males increased their calling song energetics and decreased their parental investment (nuptial gift size), suggesting that old males are dishonestly signaling their condition to the female.

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.

Age is often a neglected factor in ecological studies. However, age-related changes in reproduction and survival of organisms may strongly influence population dynamics. The Gotlandic population of collared flycatchers is a perfect system for studying age-related changes in the wild, as the exact age and reproductive history of most individuals is known. Collared flycatchers (Ficedula albicollis) on Gotland show the typical pattern of age-related changes in survival and reproductive success; both factors show an increase early in life and a decrease late in life. This thesis presents a broad study not only of age-related patterns of reproduction and immunity, but also proposes the mechanisms driving these patterns. My results show that in addition to survival probability and reproductive performance, reproductive costs and life-history trade-offs also change with progressing age. There is a significant increase in reproductive performance at the population level during first years of life which result from selection against low quality phenotypes. On the individual level this pattern is best explained by an optimization of reproductive effort. However, high quality individuals have higher reproductive success as early as their first breeding event and are long-lived. Thus, they seem to adopt a different strategy than lower quality, short-lived individuals. Differences in individual quality seem to be shaped by the developmental conditions experienced as nestlings. Fledglings with longer tarsi, but lower body mass become long-lived, high quality adults. Young individuals breeding for first time pay higher costs of reproduction. They also express a limited ability to reduce these costs by breeding in high quality territories when compared to middle-aged individuals. Young individuals seem to invest more into self-maintenance, whereas old individuals reduce the level of self-maintenance (measured as immune response) and redistribute their investment towards reproduction. Thus, old individuals are limited in their ability to reduce reproductive costs under favorable conditions, especially as they also senesce, which pattern is also shaped by individual quality. Variation in individual quality appears to have an strong effect on age-related survival probability, reproductive performance, reproductive costs, and even life-history decisions. Therefore, taking this factor into account in studies of life-history patterns is necessary to obtain reliable results.