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.     

Snakes allocate amino acids acquired during vitellogenesis to offspring: are capital and income breeding consequences of variable foraging success?

Reproductive allocation strategies have been historically described as lying on a continuum between capital and income breeding. Capital breeders have been defined as species that allocate stored reserves to reproduction, whereas income breeders have been defined as species that allocate relatively recently‐ingested food resources to reproduction. Snakes are considered capital breeders because they efficiently store large amounts of nutrients and energy, potentially enough to support an entire reproductive bout without feeding. We examined the abilities of five viviparous snake species to allocate income to follicles during vitellogenesis. We fed ¹⁵N‐labelled L‐leucine to experimental females of each species during vitellogenesis, whereas control females were fed unlabelled meals. After ovulation, we measured yolk ¹⁵N p.p.m. using mass spectrometry. Maternal scale samples taken before labelling were used to estimate endogenous ¹⁵N concentrations, which should represent ‘capital’. Scale samples taken at ovulation were used to determine whether snakes assimilated ¹⁵N‐labelled‐leucine from labelled diets. Yolks and post‐ovulatory scales of labelled females were significantly more enriched in ¹⁵N than those of unlabelled females in all species, indicating significant assimilation and allocation of income‐derived amino acids to the yolk during vitellogenesis. The lack of among‐species differences suggests that all species allocated income amino acids to vitellogenesis. The results obtained in the present study suggest that proportional utilization of income or capital depends on the frequency and timing of foraging success during reproductive events. Therefore, capital and income breeding may be consequences of both life‐history and environmental constraints on foraging success, rather than strategies of reproductive allocation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 390–404.     

Testing the reproductive and somatic trade‐off in female Columbian ground squirrels

Energetic trade‐offs in resource allocation form the basis of life‐history theory, which predicts that reproductive allocation in a given season should negatively affect future reproduction or individual survival. We examined how allocation of resources differed between successful and unsuccessful breeding female Columbian ground squirrels to discern any effects of resource allocation on reproductive and somatic efforts. We compared the survival rates, subsequent reprodction, and mass gain of successful breeders (females that successfully weaned young) and unsuccessful breeders (females that failed to give birth or wean young) and investigated “carryover” effects to the next year. Starting capital was an important factor influencing whether successful reproduction was initiated or not, as females with the lowest spring emergence masses did not give birth to a litter in that year. Females that were successful and unsuccessful at breeding in one year, however, were equally likely to be successful breeders in the next year and at very similar litter sizes. Although successful and unsuccessful breeding females showed no difference in over winter survival, females that failed to wean a litter gained additional mass during the season when they failed. The next year, those females had increased energy “capital” in the spring, leading to larger litter sizes. Columbian ground squirrels appear to act as income breeders that also rely on stored capital to increase their propensity for future reproduction. Failed breeders in one year “prepare” for future reproduction by accumulating additional mass, which is “carried over” to the subsequent reproductive season.     

Allocation from capital and income sources to reproduction shift from first to second clutch in the flesh fly, Sarcophaga crassipalpis

Understanding how resources are allocated between survival and reproduction is fundamental to the study of the evolution of life histories. Reproductive resources can come from two intrinsic resource pools, stored reserves (capital) acquired before reproduction or income acquired during reproduction. The variety of reproductive strategies in insects is remarkable and reproductive allocation encompasses the complete range of allocation strategies from pure capital breeders to pure income breeders. However, most organisms probably use a blend of capital and income and this blend is likely dynamic, changing between reproductive bouts in response to internal and external conditions. We used stable isotopes to quantify the allocation of capital and income resources to reproduction in the flesh fly, Sarcopha crassipalpis and assessed how allocation patterns change over multiple bouts of reproduction. Sarcophaga crassipalpis shifts from a slight investment of capital in the first clutch to an almost pure income breeder in the second clutch. We discuss the relationship between activity and allocation, and the potential for this system to understand how allocation patterns change in response to environmental stress.     

Resource use for reproduction depends on spring arrival time and wintering area in an arctic breeding shorebird

Resources for egg production may come from body reserves stored before breeding (“capital breeders”) or from food acquired at the breeding site (“income breeders”). Arctic migrants were long thought to be capital breeders, because they often arrive at a time when local food availability is still limited. However, later evidence suggested that arctic breeding shorebirds are primarily income breeders, or that they use a mixed strategy depending on laying date. We explored the relationship between laying date and resource use for reproduction in the pectoral sandpiper Calidris melanotos breeding in the Alaskan arctic by contrasting carbon isotope (δ¹³C) values of the local diet and of maternal plasma, cellular blood, feather and claw with those of the eggs produced. Our results revealed that early breeding females utilize resources for egg production that were acquired recently at staging areas, whereas later breeding females mostly relied on nutrients derived from local food sources. These findings suggest that the resource allocation strategy used for reproduction differs among females, and varies depending on the timing of arrival and the start of reproduction. The arrival date at the breeding ground and laying date may critically depend on non‐breeding season events such as winter habitat choice, staging areas or migration routes. By comparing maternal feather δ¹³C, claw δ¹³C and feather δD, we examined whether non‐breeding season events influenced the use of resources for egg production through variation in capture date or clutch initiation date. Female pectoral sandpipers originating from moulting areas characterized by higher (more positive) δD signatures were caught earlier and started laying earlier, and they used stored resources for reproduction. Using regional maps of δD values for precipitation in the wintering sites in South America, we compared the spatial variation in the observed feather δD signatures. This analysis indicated that female pectoral sandpipers with higher δD signatures, presumably coming from more north‐easterly wintering sites in southern America, started laying earlier and used mostly stored resources for egg production, compared to females that wintered (or at least moulted) further south. Our results thus show that winter moulting habitat is linked to breeding resource allocation strategy in this high‐arctic breeding shorebird.     

Fitness consequences of alternative life histories in water striders,Aquarius remigis (Heteroptera: Gerridae)

Using field and laboratory observations and experiments over 3 years, I investigated whether reproductive trade-offs shape individual life histories in two natural populations of the water strider, Aquarius remigis, in which univoltine and bivoltine life cycles coexist. Both later eclosion dates and food shortages, even after adult eclosion, induced diapause in females, thus deferring reproduction to the following spring. Adult body size was positively affected by food availability during juvenile development. Higher food levels also increased the reproductive output of females, but not their longevity or oviposition period. When compared to spring breeders (univoltine life cycle), direct (summer) breeders (bivoltine life cycle) experienced reduced lifetime egg numbers and longevity, as well as reduced survivorship of their second-summer-generation offspring; these reproductive costs offset, at least in part, the advantage in non-decreasing populations of having two generations per year. Fecundity was correlated with body size, and among summer-generation females direct breeders were larger than non-breeders. The time remaining before the onset of winter and/or the time since adult eclosion augmented cumulative energy uptake, and consequently the lipid reserves and winter survival probability of non-breeding (diapausing) summer adults approaching hibernation. Overwintered spring reproductives died at faster rates than non-reproductive summer individuals despite greater food availability in spring, indicating a mortality cost of reproduction. Body length correlated with absolute and not with proportional lipid content but showed no consistent relationship with survivorship in the field. These results are in agreement with current theory on the evolution of insect voltinism patterns, and further indicate high degrees of life history flexibility (phenotypic plasticity) in the study populations in response to variable environmental factors (notably photoperiod and food availability). This may be related to their location in a geographic transition zone from uni- to bivoltine life cycles.     

Long-term shifts in the cyclicity of outbreaks of a forest-defoliating insect

Optimisation of reproductive investment is crucial for Darwinian fitness, and detailed long-term studies are especially suited to unravel reproductive allocation strategies. Allocation strategies depend on the timing of resource acquisition, the timing of resource allocation, and trade-offs between different life-history traits. A distinction can be made between capital breeders that fuel reproduction with stored resources and income breeders that use recently acquired resources. In capital breeders, but not in income breeders, energy allocation may be decoupled from energy acquisition. Here, we tested the influence of extrinsic (weather conditions) and intrinsic (female characteristics) factors during energy storage, vitellogenesis and early gestation on reproductive investment, including litter mass, litter size, offspring mass and the litter size and offspring mass trade-off. We used data from a long-term study of the viviparous lizard, Lacerta (Zootoca) vivipara. In terms of extrinsic factors, rainfall during vitellogenesis was positively correlated with litter size and mass, but temperature did not affect reproductive investment. With respect to intrinsic factors, litter size and mass were positively correlated with current body size and postpartum body condition of the previous year, but negatively with parturition date of the previous year. Offspring mass was negatively correlated with litter size, and the strength of this trade-off decreased with the degree of individual variation in resource acquisition, which confirms theoretical predictions. The combined effects of past intrinsic factors and current weather conditions suggest that common lizards combine both recently acquired and stored resources to fuel reproduction. The effect of past energy store points out a trade-off between current and future reproduction.     

The timing of resource availability does not affect reproductive allotment or the rate of oocyte development in the flesh fly,Sarcophaga crassipalpis

1. The flesh fly, Sarcophaga crassipalpis, is anautogenous and largely relies on adult‐acquired income resources for reproduction, but allocates some larvally derived capital into the first clutch. Therefore, the timing of adult resource acquisition may be important for both reproductive timing and magnitude of capital vs. income resources allocated to reproduction. Specifically, we predict that flesh flies that wait longer to acquire adult income resources will allocate greater quantities of larvally derived capital to the first clutch. 2. To test how reproductive allocation in flesh flies responds to the timing of adult protein availability, we provided pulses of protein only on day 3, 6, 9, or 12 after eclosion, a series of times equivalent to the onset of oogensis and early, middle and late oogenic development in individuals fed ad libitum. Protein pulses contained isotopically distinct carbon (¹³C), allowing us to distinguish between larval capital and adult‐income resources allocated towards reproduction. 3. Neither the timing of oocyte development nor reproductive allotment (egg number by egg size) was altered by the timing of protein availability. 4. There was no effect of adult protein acquisition timing on the quantity of larvally derived somatic capital vs. adult‐acquired income carbon allocated to reproduction. While flesh flies have remarkable pre‐feeding plasticity in reproductive timing, they appear to have little post‐feeding plasticity in allocation of stored reserves towards reproduction.     

What is the appropriate timescale for measuring costs of reproduction in a `capital breeder' such as the aspic viper?

Before we can quantify the degree to which reproductive activities constitute a cost (i.e., depress an organism's probable future reproductive output), we need to determine the timescale over which such costs are paid. This is straightforward for species that acquire and expend resources simultaneously (income breeders), but more problematical for organisms that gather resources over a long period and then expend them in a brief reproductive phase (capital breeders). Most snakes are capital breeders; for example, female aspic vipers (Vipera aspis) in central western France exhibit a 2- to 3-year reproductive cycle, with females amassing energy reserves for one or more years prior to the year in which they become pregnant. We use long-term mark-recapture data on free-living vipers to quantify the appropriate timescale for studies of reproductive costs. Annual survival rates of female vipers varied significantly during their cycle, such that estimates of survival costs based only on years when the females were ‘reproductive’ (i.e., produced offspring) substantially underestimated the true costs of reproduction. High mortality in the year after reproducing was apparently linked to reproductive output; low energy reserves (poor body condition) after parturition were associated with low survival rates in the following year. Thus, measures of cost need to consider the timescale over which resources are gathered as well as that over which they are expended in reproductive activities. Also, the timescale of measurement needs to continue long enough into the post-reproductive period to detect delayed effects of reproductive ‘decisions’. This revised version was published online in July 2006 with corrections to the Cover Date.     

Do purely capital layers exist among flying birds? Evidence of exogenous contribution to arctic-nesting common eider eggs

The strategy of relying extensively on stored resources for reproduction has been termed capital breeding and is in contrast to income breeding, where needs of reproduction are satisfied by exogenous (dietary) resources. Most species likely fall somewhere between these two extremes, and the position of an organism along this gradient can influence several key life-history traits. Common eiders (Somateria mollissima) are the only flying birds that are still typically considered pure capital breeders, suggesting that they depend exclusively on endogenous reserves to form their eggs and incubate. We investigated the annual and seasonal variation in contributions of endogenous and exogenous resources to egg formation in eiders breeding at the East Bay colony in the Canadian Arctic. We collected prey items along with females and their eggs during various stages of breeding and used two complementary analytical approaches: body reserve dynamics and stable isotope [δ(13)C, δ(15)N] mixing models. Indices of protein reserves remained stable from pre-laying to post-laying stages, while lipid reserves declined significantly during laying. Similarly, stable isotope analyses indicated that (1) exogenous nutrients derived from marine invertebrates strongly contributed to the formation of lipid-free egg constituents, and (2) yolk lipids were constituted mostly from endogenous lipids. We also found evidence of seasonal variation in the use of body reserves, with early breeders using proportionally more exogenous proteins to form each egg than late breeders. Based on these results, we reject the hypothesis that eiders are pure capital layers. In these flying birds, the fitness costs of a strict capital breeding strategy, such as temporary loss of flight capability and limitation of clutch and egg size, may outweigh benefits such as a reduction in egg predation rate.     

Effects of reproductive strategies on pollutant concentrations in pinnipeds: a meta‐analysis

In marine mammals such as pinnipeds (seals, sea lions and walruses), reproductive strategies reveal how species acquire, store and allocate energy to offspring. During lactation, females can allocate energy acquired from concurrent resources (income breeding); or utilize energy stored prior to reproduction (capital breeding). Mothers transfer a large proportion of energy to their pups via lipid rich milk, meaning that pollutants such as polychlorinated biphenyls (PCBs) and mercury (Hg) will also transfer, raising concern for negative health effects. To quantify the effect of reproductive strategy on maternal pollutant transfer in pinnipeds, we developed a proxy for income and capital breeding by focusing on the lactational component of reproduction, and arranged species along this gradient. We found a strong positive relationship between lipid content in milk and degree of capital breeding. We tested this gradient against maternal pollutant transfer expressed as a concentration ratio by meta‐analysis. In mother‐pup pairs, the concentration ratio of PCBs was one order of magnitude higher than for mercury. PCB concentrations in pups and juveniles were similar to adult females, but mercury was always lower in young offspring than in females. We found no effect of reproductive strategy between studies investigating mother‐pup pairs and non‐related females and juveniles (< 1 year old), however data were strongly biased towards capital breeders. Our results suggest that either: 1) reproductive strategy does not affect pollutant bioaccumulation; or 2) a lack of income breeder data prevents us from testing the overall effect of reproductive strategy on maternal pollutant transfer. The finding that PCB concentrations in juveniles are similar to females is of concern due to early life stage exposure. We recommend data collection from income breeding species such as the sea lions to elucidate whether reproductive strategy, and potentially other life history traits, has an overall effect on maternal pollutant transfer.     

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.     

Reproductive phenology of a food-hoarding mast-seed consumer: resource- and density-dependent benefits of early breeding in red squirrels

The production of offspring by vertebrates is often timed to coincide with the annual peak in resource availability. However, capital breeders can extend the energetic benefits of a resource pulse by storing food or fat, thus relaxing the need for synchrony between energy supply and demand. Food-hoarding red squirrels (Tamiasciurus hudsonicus) breeding in the boreal forest are reliant on cones from a masting conifer for their nutrition, yet lactation is typically completed before the annual crop of cones is available for consumption such that peaks in energy supply and demand are not synchronized. We investigated the phenological response of red squirrels to annual variation in environmental conditions over a 20-year span and examined how intra- and inter-annual variation in the timing of reproduction affected offspring recruitment. Reproductive phenology was strongly affected by past resource availability with offspring born earlier in years following large cone crops, presumably because this affected the amount of capital available for reproduction. Early breeders had higher offspring survival and were more likely to renest following early litter loss when population density was high, perhaps because late-born offspring are less competitive in obtaining a territory when vacancies are limited. Early breeders were also more likely to renest after successfully weaning their first litter, but renesting predominantly occurred during mast years. Because of their increased propensity to renest and the higher survival rates of their offspring, early breeders contribute more recruits to the population but the advantage of early breeding depends on population density and resource availability.     

Capital and income breeding and the evolution of colony founding strategies in ants

Ability to store resources that will be used for reproduction represents a potential life history adaptation because storage permits feeding and reproduction to be decoupled spatially and/or temporally. The two ends of a continuum involve acquiring all resources prior to reproduction (capital breeding) or acquiring all resources during the reproductive period (income breeding). Traditional life history theory examines tradeoffs between costs and benefits of such strategies, but this theory has not been integrated into life history studies of ants, even though founding queens have the analogous strategies of fully claustral (capital breeding) and semi-claustral (income breeding). This study demonstrates that facultatively semi-claustral queens of the seed-harvester ant Pogonomyrmex desertorum exhibit phenotypic plasticity during colony founding because unfed queens produced few, small minims, whereas ad libitum fed queens produced larger, heavier minims and additional brood. Fed queens also lost less mass than unfed queens despite their producing more brood. Overall, foraging provides queens with a suite of benefits that likely offset potential negative effects of foraging risk. Life history studies across a diverse array of taxa show that capital breeding is consistently associated with low availability and/or unpredictability of food, i.e., environmental conditions that favor prepackaging of reproductive resources. Such a broad and consistent pattern suggests that similar factors favored the evolution of fully claustral (capital breeding) colony founding in ants. Overall, these data suggest that ant researchers should revise their conventional view that fully claustral colony founding evolved because it eliminated the need for queens to leave the nest to forage. Instead, colony founding strategies should be examined from the perspective of environmental variation, i.e., availability and predictability of food. I also provide a functional scenario that could explain the evolution of colony founding strategies in ants. Received 16 November 2005; revised 1 March 2006; accepted 29 March 2006.     

Condition-dependent reproductive success in bighorn ewes

Individual bighorn ewes ( Ovis canadensis ) at 4–14 years of age were 1.5% heavier preceding years when they weaned a lamb than preceding years when their lamb died before weaning. Intra-individual differences in mass between years of successful and unsuccessful reproduction appeared independent of a ewe's multiyear average mass. Relative mass loss both in the winter before and in the winter after a given repro-ductive episode increased with reproductive success. Long-term monitoring of individual mass and reproductive success is a promising technique to study life histories in capital breeders, because it allows to partially account for differences in reproductive potential.     

Are Kirindy sifaka capital or income breeders? It depends

The capital and income breeding framework has only recently been used to explain variation in female reproductive strategies in primates. The application of this framework to primates and other mammals with long reproductive cycles has not been consistent. We evaluated data on Verreaux's sifaka (Propithecus verreauxi verreauxi) in the Kirindy Forest of western Madagascar to determine whether they are capital or income breeders. We found that Verreaux's sifaka can be classified as either capital or income breeders, depending on how these concepts are operationalized. These conflicting findings highlight why the capital/income framework is currently problematic and must be standardized if it is to be a useful framework for primatologists.     

Timing of diapause initiation, metabolic changes and overwintering survival of the spruce budworm, Choristoneura fumiferana

1. Four groups of spruce budworm larvae, Choristoneura fumiferana , of the same physiological stage (at the beginning of diapause) were exposed to natural temperatures, starting in July, August, September and October. Post-diapause emergence and certain metabolites were monitored throughout the overwintering period.
2. Larval exposure to high temperatures for long periods before winter had a significant effect on temperature requirements for subsequent diapause development and apparently caused a certain amount of mortality, especially for those larvae that entered diapause earlier in the season.
3. Substantial loss of lipid and glycogen reserves was related to intensity of high temperature exposure during the pre-winter period, which in turn was associated with the timing of diapause initiation.
4. Patterns of glycerol accumulation during the winter were also influenced by temperature conditions in the early phase of diapause development.
5. Among several possible mortality factors, an extended cool period in spring was suggested as a potential cause of overwintering mortality, which was closely associated with a deficiency in energy reserves caused by high temperature exposure before winter.
6. These results stress the significant impact of pre-winter conditions on the overwintering process and highlight the importance of an ecophysiological approach to insect overwintering biology.     

Reproductive Seasonality in Female Capuchins (<Emphasis Type="Italic">Cebus capucinus</Emphasis>) in Santa Rosa (Area de Conservación Guanacaste), Costa Rica

The income-capital breeding model was developed to explain birth seasonality and reproductive strategies in female animals in relation to the abundance of food energy in the environment. An income breeder uses currently available energy and acts so as to maximize either maternal survival or weanling survival, depending on the relationship between timing of births and abundance of food energy. A capital breeder stores energy reserves for future reproductive use. Here we examined energetic influences on reproductive seasonality in a population of female white-faced capuchins (Cebus capucinus) living in a seasonal dry forest in Costa Rica. Our objectives were to determine: 1) the degree of fruiting seasonality in capuchin food trees and 2) the temporal relationship between capuchin births/conceptions and fruit abundance. Our sample included 25 yr of birth data (N = 100 births), 4 yr of capuchin fruit tree phenology data, and 18 mo of ovarian hormone data, which we used to calculate gestation lengths and estimate conception dates. Using circular statistics, we found that the mean peak in fruit abundance occurs in June, and that this population of capuchins reproduces seasonally, with 44% of births occurring within a 3-mo period (May to July, mean month = May). We propose that white-faced capuchins can be generally classified as income breeders that maximize maternal survival instead of weanling survival and that they time infant births such that the most energy expensive period of reproduction, mid-to-late lactation, occurs during the mean peak in fruit abundance.     

Egg investment in response to helper presence in cooperatively breeding Tibetan ground tits

Life‐history theory predicts a trade‐off between current and future reproduction to maximize lifetime fitness. In cooperatively breeding species, where offspring care is shared between breeders and helpers, helper presence may influence the female breeders’ egg investment, and consequently, survival and future reproductive success. For example, female breeders may reduce egg investment in response to helper presence if this reduction is compensated by helpers during provisioning. Alternatively, female breeders may increase egg investment in response to helper presence if helpers allow the breeders to raise more or higher quality offspring successfully. In the facultatively cooperative‐breeding Tibetan ground tit Pseudopodoces humilis, previous studies found that helpers improve total nestling provisioning rates and fledgling recruitment, but have no apparent effects on the number and body mass of fledglings produced, while breeders with helpers show reduced provisioning rates and higher survival. Here, we investigated whether some of these effects may be explained by female breeders reducing their investment in eggs in response to helper presence. In addition, we investigated whether egg investment is associated with the female breeder's future fitness. Our results showed that helper presence had no effect on the female breeders’ egg investment, and that egg investment was not associated with breeder survival and reproductive success. Our findings suggest that the responses of breeders to helping should be investigated throughout the breeding cycle, because the conclusions regarding the breeders’ adjustment of reproductive investment in response to being helped may depend on which stage of the breeding cycle is considered.     

Capital or income breeding? A theoretical model of female reproductive strategies

Energy storage is an important component of life-history variation.A distinction is recognized between species that provision offspringusing energy gained concurrently (income breeders) and thosethat provision offspring using energy stores accumulated atan earlier time (capital breeders). Although this distinctionhas been recognized for some time, surprisingly little attentionhas been paid to the general adaptive value of the 2 strategies.Here, we present a simple, general framework for modeling femalereproductive strategies. We show that our framework can be applicableeither to annual breeders that aim to maximize the energy deliveredto their offspring before independence, or to species with shorterreproductive cycles that aim to maximize reproductive rate,given that their offspring must build up a given level of reservesbefore independence. For both scenarios, we show that the costsof accumulating capital can lead to pure income breeding, purecapital breeding, or a mixture of the 2 strategies. Our modelallows the effects of a variety of parameters to be assessed.Length of gestation, offspring metabolism, efficiency of energytransfer from mother to offspring, and the relative rates ofenergy gain by females with and without offspring are all importantfactors. The cost associated with accumulated capital is a particularlycritical determinant of the strategy adopted. More detailedapproaches to specific systems may provide a greater understandingof the factors promoting different maternal strategies for offspringprovisioning.