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Organisms must overcome environmental limitations to optimize their investment in life history stages to maximize fitness. Human-induced climate change is generating increasingly variable environmental conditions, impacting the demography of prey items and, therefore, the ability of consumers to successfully access resources to fuel reproduction. While climate change effects are especially pronounced in the Arctic, it is unknown whether organisms can adjust foraging decisions to match such changes. We used a 9-year blood plasma δ13C and δ15N data set from over 700 pre-breeding Arctic common eiders (Somateria mollissima) to assess breeding-stage and inter-annual variation in isotopic niche, and whether inferred trophic flexibility was related to colony-level breeding parameters and environmental variation. Eider blood isotope values varied both across years and breeding stages, and combined with only weak relationships between isotopic metrics and environmental conditions suggests that pre-breeding eiders can make flexible foraging decisions to overcome constraints imposed by local abiotic conditions. From an investment perspective, an inshore, smaller isotopic niche predicted a greater probability to invest in reproduction, but was not related to laying phenology. Proximately, our results provide evidence that eiders breeding in the Arctic can alter their diet at the onset of reproductive investment to overcome increases in the energetic demand of egg production. Ultimately, Arctic pre-breeding common eiders may have the stage- and year-related foraging flexibility to respond to abiotic variation to reproduce successfully. [ABSTRACT FROM AUTHOR]

Breeding habitat choice and investment decisions are key contributors to fitness in animals. Density of individuals is a well-known cue of habitat quality used for future breeding decisions, but accuracy of density cues decreases as individuals disperse from breeding sites. Used nests remain an available information source also after breeding season, but whether such information is used for breeding decisions is less well known. We experimentally investigated whether migratory, cavity-nesting pied flycatchers (Ficedula hypoleuca) prospect potential breeding sites after breeding season and use old nests as a cue for future breeding decisions. In late summer 2013, forest sites were assigned to four treatments: (1) sites including nest boxes with old nests of heterospecifics (tits), (2) sites including suitable but empty nest boxes, (3) sites with unsuitable nest boxes, or (4) sites without any nest boxes. In the following year, we investigated pied flycatcher habitat choice and reproductive investment according to these "past" cues while also controlling for additional information sources present during settlement. Flycatchers preferred sites where tits had been perceived to breed in the previous year, but only if great tits were also currently breeding in the site and had a relatively high number of eggs. Old flycatchers avoided sites previously treated with suitable but empty cavities, whereas young flycatchers preferred sites where tits had apparently bred in the previous year. Also egg mass, but not clutch size or clutch mass, was affected by the combination of past treatment information and current tit abundance. [ABSTRACT FROM AUTHOR]

Many animals build new nests every breeding season instead of saving time by reusing old ones. One hypothesis is that nest reuse leads to increased predation risk if predators memorize nest locations and revisit these sites. Here we examine patterns in the prevalence of facultative nest reuse. Further, we relate nest reuse and timing of breeding to nest predation risk, clutch size and nestling survival. We analyse 1570 breeding attempts of the Eurasian sparrowhawk (Accipiter nisus) from Denmark (1977–1997) and from two sites in Norway (1985–2017). The probability of reuse varied between study areas, increased in replacement clutches, and was lower in adults compared to 1-year-old breeders. Pairs reusing nests laid their first egg on average 2.6 ± 1.0 SE days later than those building new nests, suggesting they are compensating for an already late breeding schedule. Indeed, reuse increased nest predation risk, but we discovered no other productive effects of reuse. In non-predated nests, late breeders had both smaller clutches and lower nestling survival. We propose that nest predation is a contributing driver to the behaviour of building a new nest each year, whereas nest reuse is a strategy to compensate for delayed onset of breeding, mainly used by inexperienced males. [ABSTRACT FROM AUTHOR]

Live-bearing reproduction (viviparity) has evolved from egg-laying (oviparity) independently many times and most abundantly in squamate reptiles. Studying life-history trade-offs between the two reproductive modes is an inherently difficult task, as most transitions to viviparity are evolutionarily old and/or are confounded by environmental effects. The common lizard (Zootoca vivipara) is one of very few known reproductively bimodal species, in which some populations are oviparous and others viviparous. Oviparous and viviparous populations can occur in sympatry in the same environment, making this a unique system for investigating alternative life-history trade-offs between oviparous and viviparous reproduction. We find that viviparous females exhibit larger body size, smaller clutch sizes, a larger reproductive investment, and a higher hatching success rate than oviparous females. We find that offspring size and weight from viviparous females was lower compared to offspring from oviparous females, which may reflect space constraints during pregnancy. We suggest that viviparity in common lizards is associated with increased reproductive burden for viviparous females and speculate that this promoted the evolution of larger body size to create more physical space for developing embryos. In the context of life-history trade-offs in the evolution of viviparity, we suggest that the extent of correlation between reproductive traits, or differences between reproductive modes, may also depend on the time since the transition occurred. [ABSTRACT FROM AUTHOR]

The fitness consequences of joining a group are highly dependent on ecological context, especially for non-kin. To assess the relationships between cooperation and environment, we examined variation in colony reproductive success for a harvester ant species that nests either solitarily or with multiple, unrelated queens, a social strategy known as primary polygyny. We measured the reproductive investment of colonies of solitary versus social nesting types at two sites, one with primarily single-queen colonies, and the other with a majority of polygynous nests. Our results were consistent with the hypothesis that cooperative nesting by unrelated ant queens is likely a selection response to difficult environments, rather than a strategy to maximize reproduction under favorable conditions. Fewer colonies at the primarily polygynous site reproduced than at the site with primarily single queen nests, and those that did had lower reproductive investment, as measured by number and total mass of reproductives. Assessment of ecological conditions also support the harsh environment hypothesis. Colony density in the multi-queen population was higher, and nearest neighbor distances were lower for non-reproducing than reproducing colonies. To more directly test the hypothesis that colony reproduction was ecologically constrained, we experimentally supplemented food resources for a subset of colonies at the primary polygyny site. Supplemented colonies increased reproductive investment levels to equal that of colonies at the single-queen population, further indicating that environmental pressures are severe where primary polygyny is dominant, and may drive the evolution of non-kin cooperation in this context. [ABSTRACT FROM AUTHOR]

Intraspecific variation in behavior and diet can have important consequences for population and ecosystem dynamics. Here, we examine how differences in reproductive investment and spatial ecology influence individual diet specialization in male and female southern sea otters ( Enhydra lutris nereis). We hypothesize that greater reproductive constraints and smaller home ranges of females lead to more pronounced intraspecific competition and increased specialization. We integrate stable carbon (δC) and nitrogen (δN) isotope analysis of sea otter vibrissae with long-term observational studies of five subpopulations in California. We define individual diet specialization as low ratios of within-individual variation (WIC) to total population niche width (TNW). We compare isotopic and observational based metrics of WIC/TNW for males and females to data on population densities, and movement patterns using both general linear and linear mixed-effects models. Consistent with our hypothesis, increasing population density is associated with increased individual diet specialization by females but not by males. Additionally, we find the amount of coastline in a sea otter's home range positively related with individual dietary variability, with increased range span resulting in weaker specialization for both males and females. We attribute our results to sex-based differences in movement, with females needing to specialize in their small ranges to maximize energy gain, and posit that the paradigm of individual prey specialization in sea otters with increased intraspecific competition may be a pattern driven largely by females. Our work highlights a potentially broader role of sex in the mechanistic pressures promoting and maintaining diet specialization. [ABSTRACT FROM AUTHOR]

In mutualistic interactions, partners obtain a net benefit, but there may also be costs associated with the provision of benefits for a partner. The question of whether aphids suffer such costs when attended by ants has been raised in previous work. Transgenerational effects, where offspring phenotypes are adjusted based on maternal influences, could be important in the mutualistic interaction between aphids and ants, in particular because aphids have telescoping generations where two offspring generations can be present in a mature aphid. We investigated the immediate and transgenerational influence of ant tending on aphid life history and reproduction by observing the interaction between the facultative myrmecophile Aphis fabae and the ant Lasius niger over 13 aphid generations in the laboratory. We found that the effect of ant tending changes dynamically over successive aphid generations after the start of tending. Initially, total aphid colony weight, aphid adult weight and aphid embryo size decreased compared with untended aphids, consistent with a cost of ant association, but these differences disappeared within four generations of interaction. We conclude that transgenerational effects are important in the aphid-ant interactions and that the costs for aphids of being tended by ants can vary over generations. [ABSTRACT FROM AUTHOR]

A new method is presented to estimate individuals' (1) age at maturation, (2) energy acquisition rate, (3) energy expenditure for body maintenance, and (4) reproductive investment, and the multivariate distribution of these traits in a population. The method relies on adjusting a conceptual energy allocation model to individual growth curves using nonlinear mixed-effects modelling. The method's performance was tested using simulated growth curves for a range of life-history types. Individual age at maturation, energy acquisition rate and the sum of maintenance and reproductive investment rates, and their multivariate distribution, were accurately estimated. For the estimation of maintenance and reproductive investment rates separately, biases were observed for life-histories with a large imbalance between these traits. For low reproductive investment rates and high maintenance rates, reproductive investment rate estimates were strongly biased whereas maintenance rate estimates were not, the reverse holding in the opposite situation. The method was applied to individual growth curves back-calculated from otoliths of North Sea plaice ( Pleuronectes platessa) and from scales of Norwegian spring spawning herring ( Clupea harengus). For plaice, maturity ogives derived from our individual estimates of age at maturation were almost identical to the maturity ogives based on gonad observation in catch samples. For herring, we observed 51.5 % of agreement between our individual estimates and those directly obtained from scale reading, with a difference lower than 1 year in 97 % of cases. We conclude that the method is a powerful tool to estimate the distribution of correlated life-history traits for any species for which individual growth curves are available. [ABSTRACT FROM AUTHOR]

When females and males differ in their timing of maximum reproductive effort, this can result in sex-specific seasonal cycles in body mass. Such cycles are undoubtedly under strong selection, particularly in bats, where they affect flying ability. Flying foxes (Old World fruit bats, Pteropus spp.) are the largest mammals that can sustain powered flight and therefore face critical trade-offs in managing body reserves for reproduction, yet little is known about body mass dynamics in this group. I investigated body mass changes in relation to reproductive behaviour in a large colony of grey-headed flying foxes ( Pteropus poliocephalus). In this polygynous mammal, females were predicted to maximise reproductive effort during lactation and males during the breeding season. As predicted, female body condition declined during the nursing period, but did not vary in relation to sexual activity. By contrast, males accumulated body reserves prior to the breeding season, but subsequently lost over 20% of their body mass on territory defence and courtship, and lost foraging opportunities as they also defended their day roost territories at night. Males in better condition had larger testes, particularly during territory establishment, prior to maximum sexual activity. Thus, the seasonality of female mass reflected the high metabolic load that lactation imposes on mothers. However, male mass followed a pattern akin to the 'fatted male phenomenon', which is commonly observed in large polygynous mammals with seasonal reproduction, but not in bats. This shows the importance of body reserves for reproduction in flying foxes, despite their severe constraints on body mass. [ABSTRACT FROM AUTHOR]

Non-direct effects of predation can be an important component of the total effect of predation, modulating animal population and community dynamics. The isolated effects of predation risk on the spatial organisation of the breeding bird community, however, remains poorly studied. We investigated whether an experimentally increased predation risk prior to reproduction affected breeding territory selection and subsequent reproductive strategies in three Mediterranean cavity-nesting birds, i.e., the little owl Athene noctua, European roller Coracias garrulus and scops owl Otus scops. We found that territories used the previous year were more likely to be re-occupied when they belonged to the safe treatment rather than to the risky treatment. The first choice of breeders of all three species was for safe territories over risky ones. When all breeding attempts in the season (i.e., final occupation) were considered, breeders also preferred safe to risky sites. In addition, little owls laid larger eggs in risky territories than in safe territories. Our study provides experimental evidence of a rapid preventive response of the three most abundant species in a cavity-nesting bird community to a short-term manipulation of predation risk. This response highlights the key role of the non-direct effects of predation in modulating avian community organisation. [ABSTRACT FROM AUTHOR]

Selection on offspring size and timing of birth or hatching could have important consequences for maternal investment strategies. Here we show consistent viability selection on hatchling body length across 2 consecutive years in a lizard that lays several clutches per season. There was no effect of hatching date on survival to maturity. However, both early hatching and large hatchling size increased adult size, which has a positive effect on total reproductive output. Earlier hatching also led to an earlier onset of reproduction. Overall, increased survival probability for large hatchlings and a positive effect of clutch size on recruitment suggest consistent directional selection on both egg size and clutch size within and across years. Because offspring size and timing of hatching are strongly affected by environmental and maternal effects, there should be potential for strong transgenerational effects on reproductive output in this species. We briefly discuss the implications of these results for the evolutionary ecology of maternal investment and population fluctuations in short-lived lizards. [ABSTRACT FROM AUTHOR]

Maternal hormones can have substantial phenotypic effects in the progeny of many vertebrates. It has been proposed that mothers adaptively adjust hormone levels experienced by particular young to optimize their reproductive output. In birds, systematic variation in egg hormone levels has been related to different female reproductive strategies. Because in many bird species prospects of the offspring change seasonally and with brood number, strategic adjustment of yolk androgen levels would be expected. To test this idea, we induced pied flycatcher ( Ficedula hypoleuca) females to nest twice during the same season by removing their first clutches shortly after clutch completion. We collected eggs of first and replacement clutches to measure yolk concentrations of androstenedione (A4) and testosterone (T) and captured the females that laid these clutches for phenotypic measurements. Although average egg androgen levels were remarkably consistent within females, hormone patterns differed considerably between first and replacement clutches. Eggs of replacement clutches were heavier with larger yolks compared to first clutches, but they contained on average lower levels of androgens. Within clutches, androgen concentration increased over the laying sequence in the first clutch, but decreased or remained more constant over the laying sequence in the replacement clutch. Mean yolk T, but not A4 levels, were negatively associated with laying date for both breeding attempts. Moreover, females in good body condition produced eggs containing lower levels of androgens than females in poor condition. Our results are consistent with the idea that differences in yolk androgen levels may be one mechanism underlying seasonal variation in reproductive success and it is possible that changes in egg androgen patterns may reflect a change in female reproductive strategy. High within-female consistency also highlights the possibility that there may be some underlying genetic variation in yolk androgen levels. [ABSTRACT FROM AUTHOR]

Although arthropod egg size is an evolutionarily and ecologically significant trait, there is still a poor understanding of the specific factors determining it. For butterflies there is evidence from an interspecific comparison that egg size is related to adult size, suggesting a morphological constraint. Using laboratory populations of the tropical butterfly Bicyclus anynana, we show that larger eggs produce larger hatchlings, and that there is considerable intra-populational variation in egg size. However, the correlation between egg size and female size is weak and explains only 1% of the variation found within a population; even development time is a slightly better predictor for egg size than female size. We conclude that there is no evidence that body size imposes a constraint on the evolution of egg size within butterfly populations. However, populations that have diverged in body size under artificial selection show correlated responses in egg size. Thus, correlations between body size and egg size may represent an emergent property, visible only when a large range of differences in body size is considered. [ABSTRACT FROM AUTHOR]

Exploiters of short-lived plants have evolved strategies in response to physiological changes that occur during the development of their hosts. The ability to adapt to host quality changes is necessary particularly if the mobility of an animal is poor or risk-constrained. In the plant-aphid system Centaurea jaceae-Uroleucon jaceae, the responses of the aphid to the seasonal changes in its host plant grown in poor and good quality soil were investigated. Coarse- and fine-tuned physiological reactions were observed in discrete aphid generations which were reared on plants grown either in a growth chamber or in a greenhouse. The number of ovarioles and developmental time depended on extrinsic factors (length of photoperiod) not directly related to plant quality. The reproductive investment of the aphids was also independent of their total dry weight. However, within the gonadal system high correlations were found at the embryonic level (e.g. the number of sclerotized embryos or the length of the oldest embryos per ovariole were highly correlated with gonadal dry weight). Aphids which were living on the 4-leaf stage of high-quality host plants showed a significantly higher investment in their gonads than aphids feeding on senescenting hosts. Factor analysis corroborated that aphids reared on poor-quality hosts revealed no grouping of variables measured, whereas those which were reared on high-quality plants showed clustering in respect of somatic (tibia length, dry weight of soma) and gonadal (number of sclerotized and unsclerotized embryos, length of the oldest embryos, ovariole number, gonad dry weight) factors. Three different levels of adaptive response to plant quality are distinguished: individual response to host quality, maternal influences on offspring and response to changes not specific to habitat (photoperiod). These different levels of regulation are thought to enable the aphids to adapt to a host of a given nutritional quality and anticipate deteriorating habitat quality simultaneously. It is concluded that physiological constraints in aphids are only revealed when aphids are exposed to severe nutritional stress for several generations. [ABSTRACT FROM AUTHOR]

Reproductive output and the growth of captive voles were quantified under high and low avian predation risk in a semi-natural experiment. Voles were exposed to Eurasian kestrels ( Falco tinnunculus), the main avian predator of vole species studied ( Clethrionomys glareolus, Microtus agrestis and M. rossiaemeridionalis). Vole pairs were housed in cages settled under nest-boxes occupied by breeding kestrels or in control cages settled under empty nest-boxes for 2 weeks. The experiment was conducted in mid-summer when kestrels had half-grown nestlings, because in that time hunting adults and begging nestlings produce noise and scats which may indicate significant predation threat to voles housed underneath the nest-boxes. The risk of kestrel predation did not have any obvious impact on pregnancy rates, mean litter sizes, or growth rates of kestrel-exposed voles compared with control voles studied. These results indicate that the risk of avian predation does not depress the reproductive investment of voles. [ABSTRACT FROM AUTHOR]

In an environment fluctuating in a predicatable manner with wide among-year variation in offspring mortality, fitness is largely influenced by the timing of reproductive investment. In vole-eating nocturnal Tengmalm's owls ( Aegolius funereus), within-cycle variation in 1st-year survival of owlets is 3-fold as estimated by the recruitment probability of an offspring. It increases from the peak through the low to the increase phase of the vole cycle. We recorded prey delivery rates of males during a 3-year vole cycle using 4 h continuous recording at night. Males brought significantly more prey items per offspring in a low-vole year than in the increase and peak vole years. In the early night (10 p.m.-12 p.m.), males fed their families equally in the increase and peak vole years, whereas in the late night (0.01-2.00 a.m.) males reduced their feeding rate in the peak year but not in other years. Both prey number and prey mass per offspring were larger in the low and increase vole years than in the peak year, though in the peak phase food is most abundant. We suggest that feeding effort of site-tenacious, long-lived (mean lifespan 3.5 years) male owls culminates in the increase rather than in the peak phase of the vole cycle, because offspring survive better in the former phase. [ABSTRACT FROM AUTHOR]