Size delays female senescence in a medium sized marsupial: The effects of maternal traits on annual fecundity in the northern brown bandicoot (Isoodon macrourus)

The degree to which females allocate resources between current reproduction, future fecundity and survival is a central theme in life history theory. We investigated two hypotheses proposed to explain patterns of reproductive investment, terminal investment and senescence, by examining the effects of maternal traits (age and maternal mass) on annual fecundity in female northern brown bandicoots, Isoodon macrourus (Marsupialia: Peramelidae). We found that annual fecundity in females declined in their final year of reproduction, indicating reproductive senescence. Maternal mass significantly influenced the rate of senescence and, in turn, a female's lifetime reproductive output. Mass had little effect on fecundity in 1st and 2nd year females, but a positive relationship with fecundity in 3rd year females. This meant that heavy, 3rd year females did not suffer the decline in fecundity shown in light 3rd year females. For 1st year females, mass and leg length increased between their first and second reproductive seasons, indicating a temporary shift, from the allocation of resources to reproduction, to increasing condition or structural size post their first breeding event. There were no net changes to body mass in subsequent years. We suggest that this year of post‐reproductive growth has important consequences for senescent effects on reproduction. Overall, results provided support for the effects of senescence on annual fecundity. Our findings were not consistent with the terminal investment hypothesis; reproductive output did not increase in females' final reproductive season despite a rapid decline in survival. However, this notion cannot be entirely dismissed; other measures of reproductive performance not examined here (e.g. offspring mass) may have provided an indication that females did increase their effort at the end of their lifespan. This study highlights the difficulty of measuring reproductive costs and the importance of understanding the combined effects of specific characteristics of an individual when interpreting reproductive strategies in iteroparous organisms.     

The adaptive value of energy storage and capital breeding in seasonal environments

Timing of reproduction in a seasonal cycle is a life history trait with important fitness consequences. Capital breeders produce offspring from stored resources and, by decoupling feeding and reproduction, may bend the constraints caused by seasonality in food or predation. Income breeders, on the other hand, produce offspring from concurrent food intake, with the disadvantage of less flexibility, but with high efficiency and no inventory costs of carrying stores. Here, we assess relative profitability of capital and income breeding in herbivorous zooplankton inhabiting seasonal, high-latitude environments. We apply a state-dependent life history model where reproductive values are used to optimise energy allocation and diapause strategies over the year. Three environmental scenarios were modelled: an early, an intermediate, and a late feeding season. We found that capital breeding was most important in the early season. Capital breeding ranged from 7–9% of the eggs produced but, because of the high reproductive value of early eggs, capital breeding ranged from 9–30% when measured in terms of reproductive value. The main benefit of capital breeding was reproduction prior to the feeding season – when the reproductive value of an egg peaked. In addition, capital breeding was also used to increase egg production rates at times of income breeding. For individuals born late in the season the model predicted a two-year cycle instead of the typical annual life cycle. These individuals could then reap the benefits of early reproduction and capital breeding in their second year instead of income breeding late in the first year. We emphasize the importance of evaluating reproductive strategies such as capital and income breeding from a complete life cycle perspective. In particular, knowing the seasonality in offspring fitness is essential to appreciate evolutionary and population-level consequences of capital breeding.     

When to parasitize? A dynamic optimization model of reproductive strategies in a cooperative breeder

We consider a cooperatively breeding group and find the optimal pattern of reproductive parasitism by a subordinate helper as a function of its body size, and hence the share of reproduction obtained by the subordinate. We develop the model for the social system of the cooperatively breeding cichlid fish Neolamprologus pulcher but the general framework is also applicable to other cooperative systems. In addition to behaving cooperatively by sharing tasks, sexually mature male cichlid helpers may directly parasitize the reproduction of dominant breeders in the group. We investigate the relative influence of life history and behavioural variables including growth, parasitism capacity, future reproductive fitness benefits and costs, relatedness and expulsion risk on the optimal reproductive strategy of subordinates. In a detailed analysis of the parameter space we show that a male helper should base its decision to parasitize primarily on an increase in expulsion risk resulting from reproductive parasitism (punishment), intra-group relatedness and the parasitism capacity. If expulsion risk is high then helpers should not parasitize reproduction at medium body size but should parasitize either when small or large.     

Heuristic optimization of the general life history problem: A novel approach

Summary The general life history problem concerns the optimal allocation of resources to growth, survival and reproduction. We analysed this problem for a perennial model organism that decides once each year to switch from growth to reproduction. As a fitness measure we used the Malthusian parameterr, which we calculated from the Euler-Lotka equation. Trade-offs were incorporated by assuming that fecundity is size dependent, so that increased fecundity could only be gained by devoting more time to growth and less time to reproduction. To calculate numerically the optimalr for different growth dynamics and mortality regimes, we used a simplified version of the simulated annealing method. The major differences among optimal life histories resulted from different accumulation patterns of intrinsic mortalities resulting from reproductive costs. If these mortalities were accumulated throughout life, i.e. if they were senescent, a bangbang strategy was optimal, in which there was a single switch from growth to reproduction: after the age at maturity all resources were allocated to reproduction. If reproductive costs did not carry over from year to year, i.e. if they were not senescent, the optimal resource allocation resulted in a graded switch strategy and growth became indeterminate. Our numerical approach brings two major advantages for solving optimization problems in life history theory. First, its implementation is very simple, even for complex models that are analytically intractable. Such intractability emerged in our model when we introduced reproductive costs representing an intrinsic mortality. Second, it is not a backward algorithm. This means that lifespan does not have to be fixed at the begining of the computation. Instead, lifespan itself is a trait that can evolve. We suggest that heuristic algorithms are good tools for solving complex optimality problems in life history theory, in particular questions concerning the evolution of lifespan and senescence.     

Evaluation of reproductive costs for Weddell seals in Erebus Bay, Antarctica

1. Organisms balance current reproduction against future survival and reproduction, which results in life-history trade-offs. These trade-offs are also known as reproductive costs and may represent significant factors shaping life-history strategy for many species. 2. Using multistate mark-resight models and 26 years of mark-resight data (1979-2004), we estimated the costs of reproduction to survival and reproductive probabilities for Weddell seals in Erebus Bay, Antarctica and evaluated whether this species either conformed to the 'prudent parent' reproductive strategy predicted by life-history theory for long-lived mammals or alternatively, incurred costs to survival in order to reproduce in a variable environment (flexible-strategy hypothesis). 3. Results strongly supported the presence of reproductive costs to survival (mean annual survival probability was 0.91 for breeders vs. 0.94 for nonbreeders), a notable difference for a long-lived mammal, demonstrating that investment in reproduction does result in a cost to survival for Weddell seals, contrary to the prudent parent hypothesis. 4. Reproductive costs to subsequent reproductive probabilities were also present for first-time breeders (mean probability of breeding the next year was 31.3% lower for first-time breeders than for experienced breeders), thus supporting our prediction of the influence of breeding experience. 5. We detected substantial annual variation in survival and breeding probabilities. Breeding probabilities were negatively influenced by summer sea-ice extent, whereas weak evidence suggested that survival probabilities were affected more by winter sea-ice extent, and the direction of this effect was negative. However, a model with annual variation unrelated to any of our climate or sea-ice covariates performed best, indicating that further study will be needed to determine the appropriate mechanism or combination of mechanisms underlying this annual variation.     

Seasonal changes in condition, reproduction and fecundity in the wild European rabbit (Oryctolagus cuniculus)

A total of 1248 rabbits ( Oryctolagus amiculus ) were shot on farmland in Cambridgeshire, UK over a period of three years. This provided pooled estimates of changes in reproductive and body condition and of fecundity during the annual cycle. Both males and females showed a significant annual cycle in reproductive condition as indicated by changes in the size of the gonads and accessory glands of reproduction. No females were pregnant during October, November and December, but males with active spermatogenesis were found during every month of the year. The distribution of pregnancies through the year showed that the reproductive season varied greatly between individuals. Peak fecundity occurred during April, May and June. On average, each female conceived 23.9 and suckled 17.2 young per breeding season. Proportionately more young died during the early stages of the reproductive season. The body weight of adult males, but not of females, varied seasonally. Body condition (kidney fat index) in adults was maximal at the start of the breeding season and minimal at the end.     

Social and demographic effects of anthropogenic mortality: a test of the compensatory mortality hypothesis in the red wolf

Whether anthropogenic mortality is additive or compensatory to natural mortality in animal populations has long been a question of theoretical and practical importance. Theoretically, under density-dependent conditions populations compensate for anthropogenic mortality through decreases in natural mortality and/or increases in productivity, but recent studies of large carnivores suggest that anthropogenic mortality can be fully additive to natural mortality and thereby constrain annual survival and population growth rate. Nevertheless, mechanisms underlying either compensatory or additive effects continue to be poorly understood. Using long-term data on a reintroduced population of the red wolf, we tested for evidence of additive vs. compensatory effects of anthropogenic mortality on annual survival and population growth rates, and the preservation and reproductive success of breeding pairs. We found that anthropogenic mortality had a strong additive effect on annual survival and population growth rate at low population density, though there was evidence for compensation in population growth at high density. When involving the death of a breeder, anthropogenic mortality was also additive to natural rates of breeding pair dissolution, resulting in a net decrease in the annual preservation of existing breeding pairs. However, though the disbanding of a pack following death of a breeder resulted in fewer recruits per litter relative to stable packs, there was no relationship between natural rates of pair dissolution and population growth rate at either high or low density. Thus we propose that short-term additive effects of anthropogenic mortality on population growth in the red wolf population at low density were primarily a result of direct mortality of adults rather than indirect socially-mediated effects resulting in reduced recruitment. Finally, we also demonstrate that per capita recruitment and the proportion of adults that became reproductive declined steeply with increasing population density, suggesting that there is potential for density-dependent compensation of anthropogenically-mediated population regulation.     

Herbivores and the evolution of the semelparous perennial life-history of plants

The relationship between a plant and its potential enemies changes drastically after reproduction has started. Using a dynamic modelling approach we study the effects of herbivores and pathogens, that are attracted by reproducing plants, on optimal allocation of resources, and life-history strategies. We assume that the level of attack increases with the investment in reproduction, which may lead to a reduction of current years reproductive success, a reduction of storage efficiency or an increase of plant mortality. If herbivores or pathogens attracted by flowering plants mainly reduce current years reproductive success, the optimal life-history is annual or iteroparous perennial if the attack is an all or nothing event. If the level of consumption increases with the number of herbivores attracted, the optimal life-history is most likely iteroparity with or without mast years. Only under very restricted conditions this may lead to semelparity. If herbivores mainly reduce the efficiency of the resources stored for next year, the optimal life-history is iteroparity. If herbivores mainly reduce survival, the optimal solution is likely to be mast years or semelparity. For parameter values that are realistic for Cynoglossum officinale, a semelparous perennial from calcereous soils, the model predicts that reproduction should start in the third year and that 99% of the available resources at the end of season should be invested in reproduction and only 1% saved for growth next year. With such an investment only c. 1% of the plants would survive after reproduction, so the optimal life-history is close to semelparity. For the small fraction of plants that reproduce more than once, years of vegetative growth only and years with reproduction should alternate. Multiple reproduction is rare in C. officinale. However, such a life history is very common for plants known as semelparous perennial. Although the available empirical evidence is, as yet, circumstantial rather than conclus ive we propose that reproduction related mortality mediated through herbivores or pathogens may play a role in the evolution of the semelparous perennial life-history.     

Environmental and social cues can be used in combination to develop sustainable breeding techniques for goat reproduction in the subtropics

Goat breeds from subtropical latitudes show different annual reproductive cycles. Some of them display large seasonal variations in their annual breeding season, while others display a moderate seasonality or sexual activity all year round. This reproductive seasonality causes seasonality of milk, cheese and meat productions and, as a consequence, induces wide variation in producer incomes. To solve this problem and provide methods allowing producers to breed animals during the anestrous period and stabilize production all year round, it is necessary to have a deep knowledge of their annual sexual activity and to identify the environmental factors controlling the timing of the annual reproductive cycle. Then, it is possible to build on these knowledge sustainable breeding techniques adapted to the environmental, economic and social characteristics of the local breeding system. In this review, I will illustrate this strategy through the example of our experiments in subtropical goats. First, we determined the characteristics of the annual breeding season in both male and female goats. Second, we identified the photoperiod as the major environmental factor controlling the timing of this annual breeding season. Third, we used the photoperiod to stimulate indirectly the sexual behavior of does. Indeed, we used photoperiodic treatments to stimulate the sexual activity of bucks during the non-breeding season. These sexually active male goats were then used to induce and synchronize the estrous behavior and ovulatory activity of anestrous females in confined or grazing conditions by using the 'male effect'. Under subtropical conditions, these results constitute an original manner to control the reproductive activity of local goats using the photoperiod combined with the 'male effect.'     

To breed or not to breed: past reproductive status and environmental cues drive current breeding decisions in a long-lived amphibian

Iteroparity is an adaptive response to uncertainty in reproductive success. However, spreading reproductive success over multiple reproduction events during a lifetime is constrained by adult mortality and the stochasticity associated with interactions between external factors and physiological states. The acquisition of information about environmental conditions during the growth of progeny and sufficient resources during the non-reproductive period are key factors for breeding success. Consequently, we hypothesized that long-lived animals may skip a breeding opportunity when information about unfavourable environmental conditions is available. In addition, nutritional constraints could prevent an animal from replenishing its reserves sufficiently to invest in the current breeding period. We investigated these questions using capture–recapture data from a 5-year study on a large population of yellow-bellied toads in a forest in north-eastern France. We took advantage of various advances in multi-state capture–recapture models (e.g. unobservable states and mixture models) to test our hypotheses. Our results show that the combined effects of rainfall deficit and the breeding/non-breeding state of individuals during the past breeding season affect breeding probability during the following breeding opportunity. We also found that females breed less frequently than males, suggesting that the overall energy cost of reproduction differs between genders. Finally, the results indicate that toad survival appears to be negatively influenced by rainfall deficits. We discuss the yellow-bellied toad’s reproductive behaviour in term of bet-hedging strategy and life history trait evolution.     

Breeding frequency in Grey-headed Albatrosses Thalassarche chrysostoma

Although Grey-headed Albatrosses Thalassarche chrysostoma are usually regarded as biennial breeders, taking a year off following a successful breeding attempt, a small proportion of successful birds attempt to breed annually. This proportion was higher at Marion Island (5.4%) than at Bird Island, South Georgia (1.0%), suggesting that conditions are more favourable at Marion Island. This hypothesis is supported by higher average breeding success and shorter lags following both successful and failed breeding attempts at Marion Island. Factors favouring reproduction at Marion Island may include reduced intraspecific competition (given the much smaller breeding population) and/or more predictable food supply (owing to production of meso-scale eddies associated with the Indian Ocean Ridge). Although annual breeding appeared to increase the risk of adult mortality, with several birds that attempted to breed annually found dead the following year, at least some birds greatly enhanced their reproductive output, with one male raising five chicks in five successive years. Contrary to life-history theory, there was no evidence that older birds were more likely to attempt annual breeding because of declining reproductive value.     

To breed or not to breed: a model of partial migration

Migration is used by a number of species as a strategy for dealing with a seasonally variable environment. In many migratory species, only some individuals migrate within a given season (migrants) while the rest remain in the same location (residents), a phenomenon called ‘partial migration’. Most examples of partial migration considered in the literature (both empirically and theoretically) fall into one of two categories: either species where residents and migrants share a breeding ground and winter apart, or species where residents and migrants share an overwintering ground and breed apart. However, a third form of partial migration can occur when non‐migrating individuals actually forgo reproduction, essentially a special form of low‐frequency reproduction. While this type of partial migration is well documented in many taxa, it is not often included in the partial migration literature, and has not been considered theoretically to date. In this paper we present a model for this partial migration scenario and determine under what conditions an individual should skip a breeding opportunity (resulting in partial migration), and under what conditions individuals should breed every chance they get (resulting in complete migration). In a constant environment, we find that partial migration is expected to occur when the mortality cost of migration is high, and when individuals can greatly increase their fecundity by skipping a year before breeding. In a stochastic environment, we find that an individual should skip migration more frequently with increased risk of a bad year (higher probability and severity), with higher mortality cost of migration, and with lower mortality cost of skipping. We discuss these results in the context of empirical data and existing life history theory.     

Trade-offs between storage and survival affect diapause timing in capital breeders

Many organisms spend the unfavourable part of the year, such as the winter season, in diapause or dormancy and reproduce in spring shortly after emergence. Reserves are acquired prior to diapause to cover metabolic costs and in some species also reproduction (capital breeding) directly after diapause. Storage is then a component of future reproduction, and capital breeders consequently pay a pre-breeding cost of reproduction as they risk dying while obtaining and carrying the reserves. How large should the reserves be, and to what extent should optimal storage, and thereby timing of diapause, depend on predation risk and reproductive strategy? We present a general and simplistic life history model of an arthropod (e.g. crustaceans or insects) that is exposed to background mortality risk when it accumulates reserves before diapause. The model optimizes diapause timing and resultant reserves for income, mixed and capital breeders, and predicts how mortality risk affects the degree of capital breeding. For income breeders, timing of diapause is insensitive to the risk while obtaining reserves as they, regardless of risk, acquire the minimum amount needed to survive the winter. For capital breeders, the higher the risk the earlier the diapause and less is consequently stored. Mixed breeders diapause late and store as much as pure capital breeders when exposed to low risk, but behave as income breeders and diapause early when mortality is high. Our model shows that the degree of capital breeding impacts phenology of diapause in a risk-dependent manner. This prediction should impact how diapause timing is thought of across a wide range of taxa, including the much studied marine copepods. Timing of diapause, including triggers and cues, can only be understood when the diversity of reproductive strategies and the adaptive value of storage is taken into account.     

Individual breeding decisions and long-term reproductive strategy in the King Penguin Aptenodytes patagonicus

Although studied for 35 years, knowledge of the reproductive biology of the King Penguin Aptenodytes patagonicus remains incomplete. The chick requires more than 12 months of care, which extends the breeding cycle, including moult, to more than one year, i.e. the King Penguin is neither annual nor biennial. In an attempt to resolve ambiguities in the literature and to elucidate the long-term breeding strategy of the species, we studied breeciing frequency at the individual level, considering the decision to breed in relation to breeding history over the previous few years. Although adult birds attempted to breed annually (0.83 breeding attempts per year), successful rearing occurred, at best, every two years only (maximum of 0.41 fledged chick per pair). Comparing successive years, the number of breeding birds in the colony was stable but the number of fledged chicks varied from 29 to 278 over eight years. These results suggest that King Penguins adopt (as individuals) an opportunistic reproductive strategy, in that they usually lay an egg every year, even when failure is certain. Nevertheless, the decision to breed was not entirely blind, and we identified groups of birds that invested differentially in breeding attempts. The decision to breed was related to the previous breeding frequency, i.e. 81% of the birds that had bred continuously in the past started a new breeding attempt, but only 67% of birds that had missed a year did so. In intermittent breeders, birds that had bred frequently, more often started a new breeding attempt than birds that had largely missed years (71% versus 57%, respectively). Classes of breeders could correspond to age classes, to birds of different breeding quality or to alternative breeding strategies coexisting in the species. Testing the hypothesis that reproductive effort increases with age should be possible in future.     

The evolution of intermittent breeding

A central issue in life history theory is how organisms trade off current and future reproduction. A variety of organisms exhibit intermittent breeding, meaning sexually mature adults will skip breeding opportunities between reproduction attempts. It’s thought that intermittent breeding occurs when reproduction incurs an extra cost in terms of survival, energy, or recovery time. We have developed a matrix population model for intermittent breeding, and use adaptive dynamics to determine under what conditions individuals should breed at every opportunity, and under what conditions they should skip some breeding opportunities (and if so, how many). We also examine the effect of environmental stochasticity on breeding behavior. We find that the evolutionarily stable strategy (ESS) for breeding behavior depends on an individual’s expected growth and mortality, and that the conditions for skipped breeding depend on the type of reproductive cost incurred (survival, energy, recovery time). In constant environments there is always a pure ESS, however environmental stochasticity and deterministic population fluctuations can both select for a mixed ESS. Finally, we compare our model results to patterns of intermittent breeding in species from a range of taxonomic groups.     

Not putting all their eggs in one basket: bet‐hedging despite extraordinary annual reproductive output of desert tortoises

Bet‐hedging theory makes the counter‐intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short‐term reproductive output to minimize the risk of reproductive failure in the long‐term. We investigated the long‐term reproductive output of an Agassiz's desert tortoise (Gopherus agassizii) population and conformance to a bet‐hedging strategy of reproduction in an unpredictable but comparatively productive environment. Most females reproduced every year, even during periods of low precipitation and poor germination of food plants, and the mean percentage of reproducing females did not differ significantly on an annual basis. Although mean annual egg production (clutch size × clutch frequency) differed significantly among years, mean clutch size and mean clutch frequency remained relatively constant. During an El Niño year, mean annual egg production and mean annual clutch frequency were the highest ever reported for this species. Annual egg production was positively influenced by maternal body size but clutch size and clutch frequency were not. Our long‐term results confirm earlier conclusions based on short‐term research that desert tortoises have a bet‐hedging strategy of producing small clutches almost every year. The risk of long‐term reproductive failure is minimized in unpredictable environments, both through time by annually producing multiple small clutches over a long reproductive lifespan, even in years of low resource availability, and through space by depositing multiple annual clutches in different locations. The extraordinary annual reproductive output of this population appears to be the result of a typically high but unpredictable biomass of annual food plants at the site relative to tortoise habitat in dryer regions. Under the comparatively productive but unpredictable conditions, tortoises conform to predictions of a bet‐hedging strategy of reproduction with relatively small but consistent clutch sizes. Published 2015. This article is a U.S. Government work and is in the public domain in the USA, Biological Journal of the Linnean Society, 2015, 115 , 399–410.     

Variation in probability of first reproduction of Weddell seals

1. For many species, when to begin reproduction is an important life-history decision that varies by individual and can have substantial implications for lifetime reproductive success and fitness. 2. We estimated age-specific probabilities of first-time breeding and modelled variation in these rates to determine age at first reproduction and understand why it varies in a population of Weddell seals in Erebus Bay, Antarctica. We used multistate mark-recapture modelling methods and encounter histories of 4965 known-age female seals to test predictions about age-related variation in probability of first reproduction and the effects of annual variation, cohort and population density. 3. Mean age at first reproduction in this southerly located study population (7.62 years of age, SD=1.71) was greater than age at first reproduction for a Weddell seal population at a more northerly and typical latitude for breeding Weddell seals (mean=4-5 years of age). This difference suggests that age at first reproduction may be influenced by whether a population inhabits the core or periphery of its range. 4. Age at first reproduction varied from 4 to 14 years, but there was no age by which all seals recruited to the breeding population, suggesting that individual heterogeneity exists among females in this population. 5. In the best model, the probability of breeding for the first time varied by age and year, and the amount of annual variation varied with age (average variance ratio for age-specific rates=4.3%). 6. Our results affirmed the predictions of life-history theory that age at first reproduction in long-lived mammals will be sensitive to environmental variation. In terms of life-history evolution, this variability suggests that Weddell seals display flexibility in age at first reproduction in order to maximize reproductive output under varying environmental conditions. Future analyses will attempt to test predictions regarding relationships between environmental covariates and annual variation in age at first reproduction and evaluate the relationship between age at first reproduction and lifetime reproductive success.     

Population response to environmental productivity throughout the annual cycle in a migratory songbird

Environmental factors affect migratory animal populations in every phase of their annual cycle and have significant impacts on breeding success and survival. The Breeding Bird Survey provides a long-term database for examining population trends in North American birds, allowing us to examine large-scale environmental factors that influence population abundance. We examined plant productivity as measured by normalized difference vegetation index (NDVI) over a 24-year period from 1983–2006 in bird conservation regions (BCRs) that overlapped Bullock's oriole (Icterus bullockii) breeding, moult, and wintering ranges to ask whether plant productivity in 1 year influences population abundance in the subsequent breeding season. Bullock's orioles have a moult-migration strategy, with a stopover moult in the Mexican monsoon region, which necessitates examining each stationary phase of the bird's annual cycle to understand the impacts of environmental factors on population abundance. Our results show increased breeding abundance in three (Great Basin, Coastal California and Shortgrass Prairies) of the six BCRs in which the species breeds following years with high NDVI values. We did not detect a response of breeding abundance to high NDVI values in the previous year in either the moulting region or in their primary over-wintering area in central Mexico. Our results demonstrate that large-scale annual variation in primary productivity on the breeding grounds can have an impact on breeding abundance in the following season, but further studies on migratory connectivity and on ecological mechanisms during the non-breeding seasons are needed to understand why we did not detect an influence of productivity during these periods.     

Diversity of age‐specific reproductive rates may result from ageing and optimal resource allocation

This paper reports the results of a dynamic programming model which optimizes resource allocation to growth, reproduction and repair of somatic damage, based on the disposable soma theory of ageing. Here it is shown that different age‐dependent patterns of reproductive rates are products of optimal lifetime strategies of resource partitioning. The array of different reproductive patterns generated by the model includes those in which reproduction begins at the maximum rate at maturity and then declines to the end of life, or increases up to a certain age and then drops. The observed patterns reflect optimal resource allocation shaped by the level of extrinsic mortality. A continuous decline in the reproductive rate from the start of reproduction is associated with high extrinsic mortality, and an early increase in the reproductive rate occurs under low extrinsic mortality. A long‐lived organism shows a low reproductive rate early in life, and short‐lived organisms start reproduction at the maximum rate.     

Natural selection of life history attributes: An analytical approach

The life history attributes which maximize fitness can be established analytically through Fisher's equation for reproductive value. Maximizing the reproductive value at age zero is equivalent to maximizing the ultimate rate of increase. As an example of the usefulness of this equality it is shown that when survivorship is uniformly reduced, the corresponding optimal maternal frequency is unaltered, even though the ultimate rate of increase is lowered by a known amount. A general life history model is proposed which links these demographic determinants of rate of increase with the energy utilization alternatives (as among maintenance, growth, and reproduction) characterizing an individual organism's development. Since the energy partitioning alternatives at any age may depend on previous allocations, an organism state variable is introduced to describe the domain over which the maximization of reproductive value may take place. Further, if the reproductive value is to be a maximum at age zero, it must be maximized at every age. An optimal life history, then, is characterized by the energy allocations which maximize sequential reproductive values. Further examples of the utility of the model focus on growth vs reproduction decisions under biomass specific life history attributes. It is shown that if births per unit energy is a linear or convex function, then an organism will not simultaneously grow and reproduce. Determinant growth, biomass at first reproduction, and explicit calculation of the maximum ultimate rate of increase are also illustrated.