Investing in a nest egg: intraspecific variation in the timing of egg laying across a latitudinal gradient.

Avian reproductive strategies vary widely, and many studies of life-history variation have focused on the incubation and hatching stages of nesting. Birds make proximate decisions regarding reproductive investment during the laying stage, and these decisions likely constrain and tradeoff with other traits and subsequent behaviors. However, we know relatively little about egg-laying stage behaviors given the difficulty of locating and monitoring nest sites from the onset of laying. We used non-invasive continuous video recording to quantify variation in the egg-laying behaviors of burrowing owls (Athene cunicularia) along a 1400-km latitudinal gradient in western North America. Burrowing owls laid eggs disproportionately in the morning hours, and that tendency was strongest among first eggs in a clutch. However, selection appeared to act more strongly on laying intervals (the time between laying of consecutive eggs) than on the diel time of laying, and laying intervals varied widely among and within clutches. Laying intervals declined seasonally and with increasing clutch size but increased with increasing burrow temperature and as a function of laying stage nest attentiveness, which may be a strategy to preserve egg viability. Laying interval was positively correlated with the duration of hatching intervals, suggesting that laying interval duration is one mechanism (along with timing of incubation onset) that generates variation in hatching asynchrony. Our results lend support to two general hypotheses to explain laying schedules; selection favors laying eggs in the morning, but other selective pressures may override that pattern. These conclusions indicate that allocation decisions during laying are an important part of avian life-history strategies which are subject to energetic constraints and tradeoffs with other traits.

Nest microclimate and limits to egg viability explain avian life-history variation across latitudinal gradients.

Variation in life-history strategies is central to our understanding of population dynamics and how organisms adapt to their environments. Yet we lack consensus regarding the ecological processes that drive variation in traits related to reproduction and survival. For example, we still do not understand the cause of two widespread inter- and intraspecific patterns: (1) the ubiquitous positive association between avian clutch size and latitude; and (2) variation in the extent of asynchronous hatching of eggs within a single clutch. Well-known hypotheses to explain each pattern have largely focused on biotic processes related to food availability and predation risk. However, local adaptation to maintain egg viability could explain both patterns with a single abiotic mechanism. The egg viability hypothesis was initially proposed to explain the cause of asynchronous hatching and suggests that asynchronous hatching results from early incubation onset in response to unfavorable nest microclimatic conditions, which otherwise reduce egg viability. However, allocation of resources to early incubation, prior to clutch completion, may energetically constrain clutch size and help explain the positive association between clutch size and latitude. We measured intraspecific variation in five functionally linked life-history traits of burrowing owls at five study sites spanning a 1,400-km latitudinal transect in western North America: clutch size, the timing of incubation onset, the degree of hatching asynchrony, the probability of hatching failure, and nestling survival. We found that most traits varied clinally with latitude, but all the traits were more strongly associated with individual nest microclimates than with latitude, and all varied with nest microclimate in the directions predicted by the egg viability hypothesis. Furthermore, incubation began earlier, hatching asynchrony increased, and clutch size declined across the breeding season. These results suggest that nest microclimate drives an important life-history trade-off and that thermal gradients are often sufficient to account for observed biogeographic and seasonal patterns in life-history strategies. Furthermore, our results reveal a potentially important indirect mechanism by which reproductive success and recruitment could be affected by climate change.

Ashmole's hypothesis and the latitudinal gradient in clutch size.

One enduring priority for ecologists has been to understand the cause(s) of variation in reproductive effort among species and localities. Avian clutch size generally increases with increasing latitude, both within and across species, but the mechanism(s) driving that pattern continue to generate hypotheses and debate. In 1961, a Ph.D. student at Oxford University, N. Philip Ashmole, proposed the influential hypothesis that clutch size varies in direct proportion to the seasonality of resources available to a population. Ashmole's hypothesis has been widely cited and discussed in the ecological literature. However, misinterpretation and confusion has been common regarding the mechanism that underlies Ashmole's hypothesis and the testable predictions it generates. We review the development of well-known hypotheses to explain clutch size variation with an emphasis on Ashmole's hypothesis. We then discuss and clarify sources of confusion about Ashmole's hypothesis in the literature, summarise existing evidence in support and refutation of the hypothesis, and suggest some under-utilised and novel approaches to test Ashmole's hypothesis and gain an improved understanding of the mechanisms responsible for variation in avian clutch size and fecundity, and life-history evolution in general.

Variation in selective regimes drives intraspecific variation in life-history traits and migratory behaviour along an elevational gradient.

Comparative studies, across and within taxa, have made important contributions to our understanding of the evolutionary processes that promote phenotypic diversity. Trait variation along geographic gradients provides a convenient heuristic for understanding what drives and maintains diversity. Intraspecific trait variation along latitudinal gradients is well-known, but elevational variation in the same traits is rarely documented. Trait variation along continuous elevational gradients, however, provides compelling evidence that individuals within a breeding population may experience different selective pressures. Our objectives were to quantify variation in a suite of traits along a continuous elevational gradient, evaluate whether individuals in the population experience different selective pressures along that gradient and quantify variation in migratory tendency along that gradient. We examined variation in a suite of 14 life-history, morphological and behavioural traits, including migratory tendency, of yellow-eyed juncos along a continuous 1000-m elevational gradient in the Santa Catalina Mountains of Arizona. Many traits we examined varied along the elevational gradient. Nest survival and nestling growth rates increased, while breeding season length, renesting propensity and adult survival declined, with increasing elevation. We documented the migratory phenotype of juncos (partial altitudinal migrants) and show that individual migratory tendency is higher among females than males and increases with breeding elevation. Our data support the paradigm that variation in breeding season length is a major selective pressure driving life-history variation along elevational gradients and that individuals breeding at high elevation pursue strategies that favour offspring quality over offspring quantity. Furthermore, a negative association between adult survival and breeding elevation and a positive association between nest survival and breeding elevation help explain both the downslope and reciprocal upslope seasonal migratory movements that characterize altitudinal migration in many birds. Our results demonstrate how detailed studies of intraspecific variation in suites of traits along environmental gradients can lend new insights into the evolutionary processes that promote diversification and speciation, the causes of migratory behaviour, and how animal populations will likely respond to climate change.