Early is better: seasonal egg fitness and timing of reproduction in a zooplankton life-history model

Timing of reproduction influences future prospects of offspring and therefore the reproductive value of parents. Early offspring are often more valuable than later ones when food availability and predation risk fluctuate seasonally. Marine zooplankton have evolved a diversity of life history strategies in response to seasonality. We present a state-dependent life history model for the annual and herbivorous high-latitude copepod Calanoides acutus . Individuals are characterised by four states; developmental stage, structural size, energy reserves and vertical location. There are two habitats, a surface habitat with seasonal predation risk and food availability, and a safer deep habitat with no food and low metabolism (diapause). Optimal life histories (diapause and energy allocation strategies) are found by dynamic programming. Seasonal egg fitness (reproductive value) emerges from the model and peak values are typically before the feeding season. Disentangling the fitness components, we conclude that seasonality in egg fitness is caused both by environmental seasonality in food and predation risk and by time-constraints on development and diapause preparation. Realised egg production, as predicted from population simulations, does not match the seasonal peak in offspring fitness but is delayed relative to peak egg fitness. We term this an 'internal life history mismatch' as constraints and tradeoffs cause sub-optimal birth dates for most eggs whereas mothers maximise their reproductive value by high number of eggs rather than few and optimally timed eggs. The earliest eggs have a disproportionately high contribution to population recruitment, emphasising the importance of early eggs and the need to understand seasonal patterns in offspring fitness.     

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.     

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.     

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.     

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.     

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.     

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.     

Best squirrels trade a long life for an early reproduction

Age at primiparity plays a crucial role in population dynamics and life-history evolution. Long-term data on female North American red squirrels were analysed to study the fitness consequences of delaying first reproduction. Early breeders were born earlier, had a higher breeding success and achieved a higher lifetime reproductive success than females who delayed their first reproduction, which suggests a higher quality of early breeders. However, early breeders had similar mass when tagged, and similar number of food caches available at one year of age as late breeders. Nevertheless, we found evidence of survival costs of early primiparity. Early breeders had a lower survival between one and two years of age than late breeders and a lower lifespan. Our study points out that two reproductive tactics co-occurred in this population: a tactic based on early maturity at the cost of a lower survival versus a tactic based on delayed maturity and long lifespan. High quality individuals express the most profitable tactic by breeding early whereas low quality individuals do the best of a bad job by delaying their first reproduction.     

Costs of reproduction and terminal investment by females in a semelparous marsupial

Evolutionary explanations for life history diversity are based on the idea of costs of reproduction, particularly on the concept of a trade-off between age-specific reproduction and parental survival, and between expenditure on current and future offspring. Such trade-offs are often difficult to detect in population studies of wild mammals. Terminal investment theory predicts that reproductive effort by older parents should increase, because individual offspring become more valuable to parents as the conflict between current versus potential future offspring declines with age. In order to demonstrate this phenomenon in females, there must be an increase in maternal expenditure on offspring with age, imposing a fitness cost on the mother. Clear evidence of both the expenditure and fitness cost components has rarely been found. In this study, we quantify costs of reproduction throughout the lifespan of female antechinuses. Antechinuses are nocturnal, insectivorous, forest-dwelling small (20-40 g) marsupials, which nest in tree hollows. They have a single synchronized mating season of around three weeks, which occurs on predictable dates each year in a population. Females produce only one litter per year. Unlike almost all other mammals, all males, and in the smaller species, most females are semelparous. We show that increased allocation to current reproduction reduces maternal survival, and that offspring growth and survival in the first breeding season is traded-off with performance of the second litter in iteroparous females. In iteroparous females, increased allocation to second litters is associated with severe weight loss in late lactation and post-lactation death of mothers, but increased offspring growth in late lactation and survival to weaning. These findings are consistent with terminal investment. Iteroparity did not increase lifetime reproductive success, indicating that terminal investment in the first breeding season at the expense of maternal survival (i.e. semelparity) is likely to be advantageous for females.     

Egg maturation and oviposition strategy of a capital breeder,Cleorodes lichenaria,feeding on lichens at the larval stage

Abstract 1. Capital breeders rely solely on resources acquired before breeding, whereas income breeders may obtain considerable amounts of resources following the commencement of reproduction. In income breeders oviposition occurs over a longer time period with a small number of eggs ready to be laid at the start of adult life, whereas capital breeders reproduce more rapidly and contain higher numbers of mature eggs at eclosion relative to potential fecundity. 2. This study explored egg maturation and oviposition strategy of a nocturnal geometrid moth, Cleorodes lichenaria. It was predicted, based on the long larval period of C. lichenaria and the known biology of other geometrids, that this lichen‐feeding geometrid moth concentrates egg production at the beginning of its breeding period and that external nutrients have a limited role on its fecundity. 3. Approximately 46% of eggs were ready to be laid in newly hatched females and approximately 40% of their potential fecundity was realised during the first night, after which the number and the weight of eggs decreased steadily. Feeding or drinking did not increase the fecundity of females. Pupal mass correlated positively with realised fecundity and ovigeny index. 4. Reproduction of C. lichenaria is based solely on larval‐derived resources. It is suggested that females are able to resorb thoracic tissues to increase fecundity and that this ability is probably linked to relatively long lifespan and low ovigeny index of this species, characteristics rarely observed in other geometrid moths.     

Early reproductive investment,senescence and lifetime reproductive success in female Asian elephants

The evolutionary theory of senescence posits that as the probability of extrinsic mortality increases with age, selection should favour early‐life over late‐life reproduction. Studies on natural vertebrate populations show early reproduction may impair later‐life performance, but the consequences for lifetime fitness have rarely been determined, and little is known of whether similar patterns apply to mammals which typically live for several decades. We used a longitudinal dataset on Asian elephants (Elephas maximus) to investigate associations between early‐life reproduction and female age‐specific survival, fecundity and offspring survival to independence, as well as lifetime breeding success (lifetime number of calves produced). Females showed low fecundity following sexual maturity, followed by a rapid increase to a peak at age 19 and a subsequent decline. High early life reproductive output (before the peak of performance) was positively associated with subsequent age‐specific fecundity and offspring survival, but significantly impaired a female's own later‐life survival. Despite the negative effects of early reproduction on late‐life survival, early reproduction is under positive selection through a positive association with lifetime breeding success. Our results suggest a trade‐off between early reproduction and later survival which is maintained by strong selection for high early fecundity, and thus support the prediction from life history theory that high investment in reproductive success in early life is favoured by selection through lifetime fitness despite costs to later‐life survival. That maternal survival in elephants depends on previous reproductive investment also has implications for the success of (semi‐)captive breeding programmes of this endangered species.     

Heterogeneity in individual quality overrides costs of reproduction in female reindeer

Reproductive allocation at one age is predicted to reduce the probability of surviving to the next year or to lead to a decrease in future reproduction. This prediction assumes that reproduction involves fitness costs. However, few empirical studies have assessed whether such costs may vary with the age at primiparity or might be overridden by heterogeneities in individual quality. We used data from 35 years’ monitoring of individually marked semi-domestic reindeer females to investigate fitness costs of reproduction. Using multi-state statistical models, we compared age-specific survival and reproduction among four reproductive states (never reproduced, experienced non-breeders, reproduced but did not wean offspring, and reproduced and weaned offspring) and among contrasted age at primiparity. We assessed whether reproductive costs occurred, resulting in a trade-off between current reproduction and future reproduction or survival, and whether early maturation was costly or rather reflected differences in individual quality of survival and reproduction capabilities. We did not find any evidence for fitness costs of reproduction in female reindeer. We found no cost of gestation and lactation in terms of future reproduction and survival. Conversely, successful breeders had higher survival and subsequent reproductive success than experienced non-breeders and unsuccessful breeders, independently of the age at primiparity. Moreover, it was beneficial to mature earlier, especially for females that successfully weaned their first offspring. Successful females at early primiparity remained successful throughout their life, clearly supporting the existence of marked among-female differences in quality. The weaning success peaked for multiparous females and was lower for first-time breeders, indicating a positive effect of experience on reproductive performance. Our findings emphasize an overwhelming importance of individual quality and experience to account for observed variation in survival and reproductive patterns of female reindeer that override trade-offs between current reproduction and future performance, at least in the absence of harsh winters.     

Natural history of small mammals of subtropical montane areas in central Taiwan

Based on samples collected in 1989–90, I report on several aspects of the natural history of species of small mammals from seasonal montane areas in subtropical central Taiwan, including their reproduction, demography, sex ratio and body size. The climatic seasonality in the areas is striking. The wet season lasts from May through September and the dry season from October through April. The phenology of plant life is related to this seasonality: vegetative growth occurs primarily in the wet season, but fruit and seed production is concentrated in the dry season. Variation in the reproduction and demography are analysed between the wet and the dry season for six species, Apodemus semotus, Niviventer culturatus. Eothenomys melanogaster, Microtus Kikuchii, Anourosorex syuamipes and Soriculus fumidus . Two murid rodents. A semotus and N. culturatus. are capable of breeding year-round, whereas the microtine rodents, E. melanogaster and M. Kikuchii , have discrete breeding seasons. The shrews, A. squamipes and S. fumidus are both seasonal breeders. but they differ in the timing of their breeding cycles. While S. fumidus breeds primarily in the late dry season, A. syuamipes reproduces in the wet season. Finally, seasonal variation in reproduction is discussed in relation to phylogenetic and physio-ecological characteristics of these six species.     

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.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

SEASONAL DECLINES IN OFFSPRING FITNESS AND SELECTION FOR EARLY REPRODUCTION IN NYMPH-OVERWINTERING GRASSHOPPERS

In this study, I examine the effects of natural and experimentally induced variation in life cycle timing on offspring fitness in Arphia sulphurea and Chortophaga viridifasciata, to understand the selective pressures shaping phenology in these two species of nymph-overwintering grasshoppers. Because these species lack embryonic diapause, hatching varies over a two month range under natural conditions. I used a cold treatment to delay hatching of some egg pods and extend the natural range of hatching dates. Due to the shorter time for growth and poorer growing conditions late in the fall, late-hatching nymphs of both species grew to a smaller size before winter and suffered higher overwinter mortality, compared to early nymphs. In addition, late nymphs that did survive the winter became reproductive later in the following year's breeding season. Size- dependent mortality of offspring during the winter is a strong selective pressure favoring early reproduction in these species. Female adult life history traits appear responsive to the seasonal declines in offspring fitness, in that late-maturing females began reproducing sooner after adult maturation and reproduced at a more rapid rate, even at the expense of having shorter adult longevity and producing fewer total egg pods. Experimental manipulations were crucial in understanding the fitness consequences of intrapopulation variation in the timing of specific life-cycle events for these species.     

Does time of the season influence filial cannibalism in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>?

According to life-history theory, filial cannibalism by fish that breed over one season only should be more beneficial early than late in the season if they eat eggs to invest energy into later clutches. Also, filial cannibalism may be more costly late in the season if finding ripe females for replacing eaten eggs is harder then. On the other hand, offspring hatching early may have a competitive advantage over fry hatching late and hence provide higher fitness to the parent. Using data collected over three successive years, I tested if sand goby males are more prone to eat of their eggs early than late in the reproductive season. I found no difference in the amount of eggs eaten or in the frequency of males eating the whole clutch between early and late in the season. Furthermore, there was no difference in the frequency of males who ate parts of their clutches, early compared to late. This might reflect a trade-off between quality (early hatching offspring) and quantity (producing as many offspring as possible over a long reproductive season). If so, the lack of seasonal pattern of filial cannibalism found in sand gobies might be the result of opposing selection pressures.     

Adult feeding and host type modulate the life history traits of the capital breeder Zabrotes subfasciatus

Income breeding animals support reproduction by the intake of energy and molecular building blocks during adulthood. Capital breeders, in contrast, fuel reproduction by the intake of materials during the larval stage. There exists, however, some controversy as to whether adult feeding in capital breeders actually increases fitness. We tested whether individuals feed as adults in the weevil Zabrotes subfasciatus Boheman, whether they equally accept carbohydrate- or nitrogen-rich food (sugar or yeast), whether food ingestion varies with the available host type and whether feeding as adults alters life history traits. The hosts tested were three varieties of Phaseolus vulgaris, Carioca, Jalo, and Preto; Vigna unguiculata Frade; and Vicia faba. We also tested whether sugar ingestion affects life history traits differently in the presence of different host types. We found that Z. subfasciatus feed as adults and equally accept sugar and yeast, and food ingestion depends on host type. Sugar and yeast significantly promote oviposition of larger eggs on Carioca seeds and enhance adult emergence, and sugar ingestion increases longevity. Moreover, the insects avoid feeding when on V. faba seeds, on which they did not oviposit. Mean fitness values showed the following hierarchy: Carioca = Jalo > Frade > Preto > V. faba. These results show that the capital breeder Z. subfasciatus feed as adults, which increases life history traits and suggests the existence of a hierarchy of performance on different hosts, with the P. vulgaris varieties Carioca and Jalo offering the best conditions.     

Pre-breeding energy requirements: thyroid hormone, metabolism and the timing of reproduction in house sparrows Passer domesticus

We measured thyroid hormone (T3) levels and energy expenditure of pre-breeding house sparrows Passer domesticus in relation to the timing of breeding and reproductive success. The onset of reproduction was synchronised in two waves, separated by a three-week interval. On an annual basis, early breeders (birds that bred for the first time during the first wave) made significantly more breeding attempts, laid significantly more egg and raised 2.3 times more chicks to fledging than late breeders (birds that bred for the first time during the second wave). By the end of March, about one month before the first egg was laid in the population, plasma titres of testosterone in males and estradiol in females were still low and did not differ between early and late breeders. However, birds that subsequently bred early had higher titres of plasma triiodothyronine (T3) than birds that started to breed late. We show for the first time in a free-living bird population that Basal Metabolic Rate (BMR) is positively correlated with the plasma titre of T3. Differences in plasma T3 accounted for 48% of the inter-individual variation in BMR. Elevated T3 levels indicate that energy requirements increase prior to breeding. Although early breeding appears to be advantageous in terms of the number of offspring raised on an annual basis, the increased energy requirements prior to breeding are thought to delay the onset of reproduction in those birds that cannot afford the additional energy expenditure early in the season.     

Female investment in offspring size and number shifts seasonally in a lizard with single-egg clutches

The timing of reproduction strongly influences reproductive success in many organisms. For species with extended reproductive seasons, the quality of the environment may change throughout the season in ways that impact offspring survival, and, accordingly, aspects of reproductive strategies may shift to maximize fitness. Life-history theory predicts that if offspring environments deteriorate through the season, females should shift from producing more, smaller offspring early in the season to fewer, higher quality offspring later in the season. We leverage multiple iterations of anole breeding colonies, which control for temperature, moisture, and food availability, to identify seasonal changes in reproduction. These breeding colonies varied only by the capture date of the adult animals from the field. We show that seasonal cohorts exhibit variation in key reproductive traits such as inter-clutch interval, egg size and hatchling size consistent with seasonal shifts in reproductive effort. Overall, reproductive effort was highest early in the season due to a relatively high rate of egg production. Later season cohorts produced fewer, but larger offspring. We infer that these results indicate a strategy for differential allocation of resources through the season. Females maximize offspring quantity when environments are favorable, and maximize offspring quality when environments are poor for those offspring. Our study also highlights that subtle differences in methodology (such as capture date of study animals) may influence the interpretation of results. Researchers interested in reproduction must be conscious of how their organism’s reproductive patterns may shift through the season when designing experiments or comparing results across studies.