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.     

Capital energy drives production of multiple clutches whereas income energy fuels growth in female collared lizards Crotaphytus collaris

Animals invest energy in reproduction that is obtained at two distinct times relative to the reproductive cycle. Energy obtained during egg production is referred to as income energy whereas stored energy acquired prior to reproduction is capital energy. Similar to most ectotherms, squamate reptiles are generally hypothesized to be capital breeders. Nearly all squamates in which income/capital energy investment has been examined thus far produce only one clutch per reproductive season. Although it is likely that squamates producing multiple seasonal clutches fuel first clutches with capital energy, either capital or income energy may be used to produce later clutches. We first monitored female eastern collared lizards over 14 reproductive seasons to confirm that the number of clutches females produce seasonally is a plastic response to variable environmental parameters, and to examine the effects of female body condition at the beginning of the reproductive season on clutch production. Clutch production varied annually and both the size and number of clutches were positively correlated with body condition. We then tested the competing predictions of the income and capital hypotheses experimentally by supplementing the diets of female collared lizards in situ for one season. Diet‐supplementation had no effect on the number of clutches produced but increased growth rates of gravid females. We further tested the competing predictions of these two hypotheses by examining variation in maternal energy investment per clutch using preserved specimens collected near our primary field site. Clutch size was highly correlated with female body size. Together, our results suggest that variation in reproductive output by female collared lizards is linked to stored capital energy rather than income energy, similar to most ectotherms.     

King eiders use an income strategy for egg production: a case study for incorporating individual dietary variation into nutrient allocation research

The use of stored nutrients for reproduction represents an important component of life-history variation. Recent studies from several species have used stable isotopes to estimate the reliance on stored body reserves in reproduction. Such approaches rely on population-level dietary endpoints to characterize stored reserves (“capital”) and current diet (“income”). Individual variation in diet choice has so far not been incorporated in such approaches, but is crucial for assessing variation in nutrient allocation strategies. We investigated nutrient allocation to egg production in a large-bodied sea duck in northern Alaska, the king eider (Somateria spectabilis). We first used Bayesian isotopic mixing models to quantify at the population level the amount of endogenous carbon and nitrogen invested into egg proteins based on carbon and nitrogen isotope ratios. We then defined the isotopic signature of the current diet of every nesting female based on isotope ratios of eggshell membranes, because diets varied isotopically among individual king eiders on breeding grounds. We used these individual-based dietary isotope signals to characterize nutrient allocation for each female in the study population. At the population level, the Bayesian and the individual-based approaches yielded identical results, and showed that king eiders used an income strategy for the synthesis of egg proteins. The majority of the carbon and nitrogen in albumen (C: 86 ± 18%, N: 99 ± 1%) and the nitrogen in lipid-free yolk (90 ± 15%) were derived from food consumed on breeding grounds. Carbon in lipid-free yolk derived evenly from endogenous sources and current diet (exogenous C: 54 ± 24%), but source contribution was highly variable among individual females. These results suggest that even large-bodied birds traditionally viewed as capital breeders use exogenous nutrients for reproduction. We recommend that investigations of nutrient allocation should incorporate individual variation into mixing models to reveal intraspecific variation in reproductive strategies.     

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.     

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.     

Short-term versus long-term effects of food intake on reproductive output in a viviparous snake, Vipera aspis

Feeding rates influence reproductive output in many kinds of animals, but we need to understand the timescale of this influence before we can compare reproductive energy allocation to energy intake. A central issue is the extent to which reproduction is fuelled by long-term energy stores ("capital" breeding) versus recently-acquired resources ("income" breeding). Our data on free-living aspic vipers show that there is no simple answer to this question: reproductive frequency is determined by long-term energy stores, offspring size is influenced by maternal food intake immediately before ovulation, and litter size is influenced by both long-term stores and short-term energy acquisition. Thus, offspring size in free-living vipers reflects the mother's energy balance over the preceding year (via a trade-off between litter size and offspring size) as well as her energy balance in the current breeding season. Hence, different components of a given reproductive output (litter) are not only functionally linked, but also respond to different temporal scales of prey availability. A female's body size has little effect on her reproductive output. Attempts to quantify reproductive energy allocation must take into account the fact that different reproductive traits (such as offspring size versus number) may respond to energy availability over different timespans. Thus, although the aspic viper is a typical "capital breeder" in terms of its reliance on stored reserves for maternal "decisions" concerning reproductive frequency, it is to some degree a facultative "income breeder" with respect to the determination of offspring size and litter size.     

Relative contribution of lipid sources to eggs of lesser scaup

Studies of how birds mobilize nutrients to eggs have traditionally considered a continuum of possible allocation strategies ranging from income breeding (rely on food sources found on the breeding grounds) to capital breeding (rely on body reserves stored prior to the breeding season). For capital breeding, stored body reserves can be acquired either on or away from the breeding grounds, but it has been difficult to quantify the relative contribution of each, precluding identification of key habitats for acquiring nutrients for clutch formation. During 2006–2009, we explored the importance of spring‐staging habitats versus breeding‐area habitats for egg‐lipid formation in female lesser scaup Aythya affinis using stable carbon (δ¹³C) isotope analyses. Although δ¹³C values for abdominal lipid reserves brought to the breeding grounds overlapped those of local amphipods, we were able to quantify the importance of local plant carbohydrates (seeds of emergent wetland plants) to the production of eggs. We compared the importance of local wetland seeds (overall δ¹³C: ?29.1 ± 0.9‰ SD) to combined lipid stores and lipids from local amphipods (overall δ¹³C: ?23.8 ± 2.2‰). Local seeds and stored body lipids contributed equally to egg lipid formation across years but we found evidence of annual variation in their relative importance. Wetland seeds contributed 39% (SE = 10%) to egg lipid production, and the importance of this source varied by year (90% CI = 47–75% in 2006, 13–42% in 2007, 29–65% in 2008, and 7–30% in 2009). In contrast to earlier studies that suggest lesser scaup predominantly employ a capital breeding strategy, our results suggest that in some years females may attain half of their energy for clutch formation from foods on the breeding grounds.     

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.     

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.     

Utilization of pre-existing energy stores of female Aedes aegypti mosquitoes during the first gonotrophic cycle

Pre-existing energy reserves may play an important role in regulating the utilization of blood meal proteins in female anautogenous mosquitoes. Determining the fate of reserves derived from the sugar meal and larval food during the first gonotrophic cycle would help to elucidate the relative contributions of larval and adult nutrition to survival and reproduction. We measured the allocation of pre-blood-meal reserves to egg production or energy production during the first gonotrophic cycle by using [14C]-labeled female Aedes aegypti mosquitoes. Feeding adults [3,4-14C]-glucose labeled the glycogen and sugar stores (approximately 50%), lipid stores (approximately 25%), and protein and amino acid stores (approximately 25%). During the first gonotrophic cycle, about 60% of the glycogen and sugar stores were metabolized and all were used for energy production. About 33% of the labeled protein and 72% of the labeled amino acid stores were metabolized, with about 9% being transferred to the eggs and the rest oxidized. About 30% of the lipid was metabolized, with about 65% being transferred to the eggs and the rest oxidized. Feeding [1-14C]-oleic acid to larvae effectively labeled adult lipid stores with about 75% of the label in lipid stores and 16% in proteins and 6% in glycogen. During the first gonotrophic cycle, about 35% of the labeled lipid stores were metabolized, with equal amounts being oxidized and transferred to the eggs. None of the other maternal stores labeled by fatty acid were metabolized during the first gonotrophic cycle. These results show that carbohydrate reserves are a critical source of energy during the first gonotrophic cycle, while lipid reserves are used equally for energy production and provisioning the eggs.     

Sexual patterns of prebreeding energy reserves in the common frog Rana temporaria along a latitudinal gradient

The ability to store energy is an important life history trait for organisms facing long periods without energy income, and in particular for capital breeders such as temperate zone amphibians, which rely on stored energy during reproduction. However, large scale comparative studies of energy stores in populations with different environmental constraints on energy allocation are scarce. We investigated energy storage patterns in spring (after hibernation and before reproduction) in eight common frog Rana temporaria populations exposed to different environmental conditions along a 1600  km latitudinal gradient across Scandinavia (range of annual activity period is 3–7  months). Analyses of lean body weight (eviscerated body mass), weight of fat bodies, liver weight, and liver fat content, showed that 1) post-hibernation/pre-breeding energy stores increased with increasing latitude in both sexes, 2) males generally had larger energy reserves than females and 3) the difference in energy stores between sexes decreased towards the north. Larger energy reserves towards the north can serve as a buffer against less predictable and/or less benign weather conditions during the short activity period, and may also represent a risk-averse tactic connected with a more pronounced iteroparous life history. In females, the continuous and overlapping vitellogenic activity in the north may also demand more reserves in early spring. The general sexual difference could be a consequence of the fact that, at the time of our sampling, females had already invested their energy into reproduction in the given year (i.e. their eggs were already ovulated), while the males' main reproductive activities (e.g. calling, mate searching, sexual competition) occurred later in the season.     

Exploring the origin of egg protein in an oviparous water snake (Natrix maura)

Reptiles are typical capital breeders that fuel reproduction by the use of lipids stored in fat bodies. However, little is known about the origin (exogenous or endogenous) of egg protein. We have examined the origin of egg protein by means of the analysis of protein content of liver and carcass (skeletal muscle) of the oviparous snake Natrix maura throughout an annual cycle. We have also measured monthly variation of the digestive-content mass and the ovarian mass. Results showed that protein in liver peaked during vitellogenesis according to the role of the liver in the synthesis of vitellogenin. Partial correlations showed the path of protein from the prey (digestive-content) to the liver, and finally to the ovaries, as well as an inverse relation between carcass protein and ovarian mass. Carcass muscle is the only body part that may act as a potential reserve for endogenous protein, although we did not find significant variation in carcass protein during female reproduction. As females with large follicles did not stop foraging activity, we assumed that egg protein was derived from the diet as partial correlations indicated. Our results suggest that N. maura is a capital breeder for lipids and tend to be income breeder for protein. This conclusion contrasts with that observed in capital breeders for which egg protein was derived from muscle. We discuss the idea that flexibility in the origin of egg protein could affect the body condition in post-reproductive females.     

Capital breeding and allocation to life-history demands are highly plastic in lizards

The use of stored resources to fuel reproduction, growth, and self-maintenance in the face of uncertain nutrient availability is a tactic common to many organisms. The degree to which organisms rely on stored resources in response to varied nutrients, however, is not well quantified. In this study, we used stable isotope methods to quantify the use of stored versus incoming nutrients to fuel growth and egg and fat body development in lizards under differing nutrient regimes. We found that the degree of capital breeding is a function of an individual's body condition. Furthermore, given sufficient income, lizards in poor condition can allocate simultaneously to storage, growth, and reproduction and "catch up" in body size and reproductive allocation to better-conditioned animals. Using natural variation in the δ(13)C of environmental nutrient pulses, we also found a high degree of variation in capital breeding in a lizard community. These findings demonstrate that capital breeding in lizards is not simply a one-way flow of endogenous stores to eggs but is a function of the condition state of individuals and seasonal nutrient availability. We use our findings to comment on capital breeding in lizards and the utility of the capital-income concept in general.     

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.     

Detecting life history trade-offs: measuring energy stores in “capital” breeders reveals costs of reproduction

Life history trade-offs should be detectable as negative correlations between the relevant traits (e.g. reproductive output versus energy storage), but may be masked by variation in resource levels among individuals. One way to detect underlying trade-offs, at least in organisms that rely on stored energy for reproduction (“capital breeders”), may be to monitor an individual's energy stores before and after reproduction. We analysed energy stores and reproductive output in Eulamprus tympanum, a viviparous scincid lizard that stores energy for reproduction in its tail. One predicted trade-off (that between the size and number of offspring in a litter) is consistently observed, and is detectable with minimal information. Another predicted trade-off (that between offspring size and subsequent energy reserves) is not apparent in our data, perhaps because of constraints imposed by correlations among other traits. Finally, trade-offs between reproductive output and subsequent energy stores are evident in this species, but are only detectable with information on the extent of pre-reproductive as well as post-reproductive energy stores. For “capital breeders”, non-destructive measurement of pre- and post-reproductive energy stores may greatly enhance our ability to detect significant life history trade-offs. Received: 10 July 1996 / Accepted: 16 January 1997     

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.     

Maternal condition influences phenotypic selection on offspring

1. Environmentally induced maternal effects are known to affect offspring phenotype, and as a result, the dynamics and evolution of populations across a wide range of taxa. 2. In a field experiment, we manipulated maternal condition by altering food availability, a key factor influencing maternal energy allocation to offspring. We then examined how maternal condition at the time of gametogenesis affects the relationships among early life-history traits and survivorship during early development of the coral reef fish Pomacentrus amboinensis. 3. Maternal condition did not affect the number of embryos that hatched or the number of hatchlings surviving to a set time. 4. We found no significant difference in egg size in relation to the maternal physiological state. However, eggs spawned by supplemented mothers were provisioned with greater energy reserves (yolk-sac and oil globule size) than nonsupplemented counterparts, suggesting that provision of energy reserves rather than egg size more closely reflected the maternal environment. 5. Among offspring originating from supplemented mothers, those with larger yolk-sacs were more likely to successfully hatch and survive for longer periods after hatching. However, among offspring from nonsupplemented mothers, yolk-sac size was either inconsequential to survival or offspring with smaller yolk-sac sizes were favoured. Mothers appear to influence the physiological capacity of their progeny and in turn the efficiency of individual offspring to utilize endogenous reserves. 6. In summary, our results show that the maternal environment influences the relationship between offspring characteristics and survival and suggest that energy-driven selective mechanisms may operate to determine progeny viability.     

Metabolic costs of capital energy storage in a small‐bodied ectotherm

Reproduction is energetically financed using strategies that fall along a continuum from animals that rely on stored energy acquired prior to reproduction (i.e., capital breeders) to those that rely on energy acquired during reproduction (i.e., income breeders). Energy storage incurs a metabolic cost. However, previous studies suggest that this cost may be minimal for small‐bodied ectotherms. Here I test this assumption. I use a laboratory feeding experiment with the European green crab Carcinus maenas to establish individuals with different amounts of energy storage. I then demonstrate that differences in energy storage account for 26% of the variation in basal metabolic costs. The magnitudes of these costs for any individual crab vary through time depending on the amount of energy it has stored, as well as on temperature‐dependent metabolism. I use previously established relationships between temperature‐ and mass‐dependent metabolic rates, combined with a feasible annual pattern of energy storage in the Gulf of Maine and annual sea surface temperature patterns in this region, to estimate potential annual metabolic costs expected for mature female green crabs. Results indicate that energy storage should incur an ~8% increase in metabolic costs for female crabs, relative to a hypothetical crab that did not store any energy. Translated into feeding, for a medium‐sized mature female (45 mm carapace width), this requires the consumption of an additional ~156 mussels annually to support the metabolic cost of energy storage. These results indicate, contrary to previous assumptions, that the cost of energy storage for small‐bodied ectotherms may represent a considerable portion of their basic operating energy budget. An inability to meet these additional costs of energy storage may help explain the recent decline of green crabs in the Gulf of Maine where reduced prey availability and increased consumer competition have combined to hamper green crab foraging success in recent years.     

Maternal effects in Daphnia: what mothers are telling their offspring and do they listen?

Maternal effects can significantly impact offspring performance. Provisioning of offspring with energy stores can quantitatively alter their growth rates, survivorship, and future fecundity, and influence population regulatory mechanisms. In this paper, we show that maternal effects can also qualitatively affect offspring reproduction (i.e. their mode of reproduction). The freshwater herbivore Daphnia pulex can change the amount of energy allocated between asexual and ephippial egg production. Our experiments on individuals, experiencing “step‐up” or “step‐down” food manipulations, reveal that offspring qualitatively shift their energy allocation away from asexual reproduction to ephippial egg production when there is a simple mismatch between maternal and offspring food environments. We show that the response is asymmetric with respect to changes in food level, ephippial egg production is higher with a greater mismatch between environments, and that the effect can be observed in dynamic experimental populations. These results point to a “generational memory” that could challenge our interpretation of field patterns and mechanisms influencing population dynamics in Daphnia–algal systems.     

Alternative reproductive tactics in snail shell‐brooding cichlids diverge in energy reserve allocation

Life history theory predicts that the amount of resources allocated to reproduction should maximize an individual's lifetime reproductive success. So far, resource allocation in reproduction has been studied mainly in females. Intraspecific variation of endogenous energy storage and utilization patterns of males has received little attention, although these patterns may vary greatly between individuals pursuing alternative reproductive tactics (ARTs). ARTs are characterized by systematic variation of behavioral, physiological, and often morphological traits among same‐sex conspecifics. Some individuals may rely on previously accumulated reserves, because of limited foraging opportunities during reproduction. Others may be able to continue foraging during reproduction, thus relying on reserves to a lesser extent. We therefore predicted that, if male tactics involve such divergent limitations and trade‐offs within a species, ARTs should correspondingly differ in energy reserve allocation and utilization. To test this prediction, we studied short‐term and long‐term reserve storage patterns of males in the shell‐brooding cichlid Lamprologus callipterus. In this species, bourgeois males investing in territory defense, courtship, and guarding of broods coexist with two distinct parasitic male tactics: (1) opportunistic sneaker males attempting to fertilize eggs by releasing sperm into the shell opening when a female is spawning; and (2) specialized dwarf males attempting to enter the shell past the spawning female to fertilize eggs from inside the shell. Sneaker males differed from other male types by showing the highest amount of accumulated short‐term and long‐term fat stores, apparently anticipating their upcoming adoption of the nest male status. In contrast, nest males depleted previously accumulated energy reserves with increasing nest holding period, as they invest heavily into costly reproductive behaviors while not taking up any food. This conforms to a capital breeder strategy. Dwarf males did not accumulate long‐term fat stores at all, which they can afford due to their small behavioral effort during reproduction and their continued feeding activity, conforming to an income breeder strategy. Our data confirm that the resource storage patterns of males pursuing ARTs can diverge substantially, which adds to our understanding of the coexistence and maintenance of alternative reproductive patterns within species.