Plasticity in laying dates of Canada Geese in response to spring phenology

Observed phenological changes can be explained either by individual phenotypic plasticity or by evolutionary changes, but there is more evidence pointing towards phenotypic plasticity to explain the mechanism behind changes in bird phenology. However, most studies on phenology have been conducted on insectivorous bird species for which breeding is closely tied to temperature and insect emergence. In this study, we examined the consequences of climatic conditions on the nesting phenology of temperate breeding Canada Geese Branta canadensis maxima, which rely on a continuous food supply, during a 14‐year period (2003–16). We determined whether laying dates were plastically adjusted to spring environmental conditions, and whether this adjustment resulted in a laying date advancement. We further estimated the strength and shape of selection acting on breeding timing, by looking at the effect of laying date on the relative number of young successfully hatched in a nest. We found that Geese plastically adjusted their laying date to spring maximum temperature (and not to precipitation or ice break‐up), resulting in a 9‐day advancement of laying date in the population for that period. Laying date was also moderately repeatable (r = 0.23) and subject to directional selection, but stabilizing selection was negligible. We thus demonstrate how Canada Geese plastically adjust laying dates to temperature, which may further be beneficial to nesting success. Evolutionary change of laying date to selection related to climate change, however, is still possible.     

Climatic effects on arrival and laying dates in a long-distance migrant, the Collared Flycatcher Ficedula albicollis

Long‐distance migrants may respond to climate change in breeding, wintering or staging area by changing their phenology. The geographical variation in such responses (e.g. coastal vs. continental Europe) and the relative importance of climate at different spatial scales remain unclear. Here we analysed variation in first arrival dates (FADs) and laying dates of the Collared Flycatcher Ficedula albicollis in a central European population, from 1973 to 2002. The North Atlantic Oscillation (NAO) index correlated weakly with local temperature during the laying period. Decreasing spring temperatures until 1980 were associated with a trend towards later laying. The rate of warming (0.2 °C per year) and laying advancement (0.4 days per year) since 1980 are amongst the highest values reported elsewhere. This long‐term trend in laying date was largely explained by the change in climatic factors. The negative effect of local spring temperature on laying was relatively stronger than that of NAO. The number of clutches initiated on a particular day was marginally affected by the temperature 3 days prior to laying and the response of females to daily variation in temperature did not change over years. Correspondence between the average population‐level and the individual‐level responses of laying date to climate variation suggests that the advancement of laying was due to phenotypic plasticity. Despite warmer springs and advanced laying, FADs did not change over years and were not correlated with local spring temperature. Marginal evidence suggests later departure from wintering grounds and faster migration across staging areas in warmer conditions. Advancement of arrival was probably constrained by low local temperatures in early spring just before arrival that have not changed over years. The interval between first arrival and laying has declined since 1980 (0.5 days per year), but the increasing temperature during that period may have kept the food supply approximately unchanged.     

Plasticity results in delayed breeding in a long‐distant migrant seabird

A major question for conservationists and evolutionary biologists is whether natural populations can adapt to rapid environmental change through micro‐evolution or phenotypic plasticity. Making use of 17 years of data from a colony of a long‐distant migratory seabird, the common tern (Sterna hirundo), we examined phenotypic plasticity and the evolutionary potential of breeding phenology, a key reproductive trait. We found that laying date was strongly heritable (0.27 ± 0.09) and under significant fecundity selection for earlier laying. Paradoxically, and in contrast to patterns observed in most songbird populations, laying date became delayed over the study period, by about 5 days. The discrepancy between the observed changes and those predicted from selection on laying date was explained by substantial phenotypic plasticity. The plastic response in laying date did not vary significantly among individuals. Exploration of climatic factors showed individual responses to the mean sea surface temperature in Senegal in December prior to breeding: Common terns laid later following warmer winters in Senegal. For each 1°C of warming of the sea surface in Senegal, common terns delayed their laying date in northern Germany by 6.7 days. This suggests that warmer waters provide poorer wintering resources. We therefore found that substantial plastic response to wintering conditions can oppose natural selection, perhaps constraining adaptation.     

Consistent within‐individual plasticity is sufficient to explain temperature responses in red deer reproductive traits

Warming global temperatures are affecting a range of aspects of wild populations, but the exact mechanisms driving associations between temperature and phenotypic traits may be difficult to identify. Here, we use a 36‐year data set on a wild population of red deer to investigate the causes of associations between temperature and two important components of female reproduction: timing of breeding and offspring size. By separating within‐ versus between‐individual associations with temperature for each trait, we show that within‐individual phenotypic plasticity (changes within a female's lifetime) was entirely sufficient to generate the observed population‐level association with temperature at key times of year. However, despite apparently adequate statistical power, we found no evidence of any variation between females in their responses (i.e. no “IxE” interactions). Our results suggest that female deer show plasticity in reproductive traits in response to temperatures in the year leading up to calving and that this response is consistent across individuals, implying no potential for either selection or heritability of plasticity. We estimate that the plastic response to rising temperatures explained 24% of the observed advance in mean calving date over the study period. We highlight the need for comparable analyses of other systems to determine the contribution of within‐individual plasticity to population‐level responses to climate change.     

Life history of breeding partners alters age‐related changes of reproductive traits in a natural population of blue tits

Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.     

Phenological mismatch drives selection on elevation,but not on slope,of breeding time plasticity in a wild songbird

Phenotypic plasticity is an important mechanism for populations to respond to fluctuating environments, yet may be insufficient to adapt to a directionally changing environment. To study whether plasticity can evolve under current climate change, we quantified selection and genetic variation in both the elevation (RN_E) and slope (RN_S) of the breeding time reaction norm in a long‐term (1973–2016) study population of great tits (Parus major). The optimal RN_E (the caterpillar biomass peak date regressed against the temperature used as cue by great tits) changed over time, whereas the optimal RN_S did not. Concordantly, we found strong directional selection on RN_E, but not RN_S, of egg‐laying date in the second third of the study period; this selection subsequently waned, potentially due to increased between‐year variability in optimal laying dates. We found individual and additive genetic variation in RN_E but, contrary to previous studies on our population, not in RN_S. The predicted and observed evolutionary change in RN_E was, however, marginal, due to low heritability and the sex limitation of laying date. We conclude that adaptation to climate change can only occur via micro‐evolution of RN_E, but this will necessarily be slow and potentially hampered by increased variability in phenotypic optima.     

CONTRASTING PATTERNS OF PHENOTYPIC PLASTICITY IN REPRODUCTIVE TRAITS IN TWO GREAT TIT (PARUS MAJOR) POPULATIONS

Phenotypic plasticity is an important mechanism via which populations can respond to changing environmental conditions, but we know very little about how natural populations vary with respect to plasticity. Here we use random‐regression animal models to understand the multivariate phenotypic and genetic patterns of plasticity variation in two key life‐history traits, laying date and clutch size, using data from long‐term studies of great tits in The Netherlands (Hoge Veluwe [HV]) and UK (Wytham Woods [WW]). We show that, while population‐level responses of laying date and clutch size to temperature were similar in the two populations, between‐individual variation in plasticity differed markedly. Both populations showed significant variation in phenotypic plasticity (IxE) for laying date, but IxE was significantly higher in HV than in WW. There were no significant genotype‐by‐environment interactions (GxE) for laying date, yet differences in GxE were marginally nonsignificant between HV and WW. For clutch size, we only found significant IxE and GxE in WW but no significant difference between populations. From a multivariate perspective, plasticity in laying date was not correlated with plasticity in clutch size in either population. Our results suggest that generalizations about the form and cause of any response to changing environmental conditions across populations may be difficult.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

Immense plasticity of timing of breeding in a sedentary forest passerine,Poecile palustris

Numerous bird species have advanced their breeding seasons in response to climate warming. These changes were mostly brought about by phenotypic plasticity, i.e. flexible reactions of individual birds, rather than by microevolutionary change. Knowing the limits of plasticity is thus of paramount importance in any attempt to predict possible reactions of birds to climate warming. However, the breeding performance of the same individuals in contrasting environmental conditions, necessary to answer this question, is rarely observed. Here, we provide data on the flexibility in timing of egg‐laying of individual marsh tit Poecile palustris females breeding in an extremely late (2013) and early (2014) spring in Bia?owie?a National Park (Poland). In both years the birds stayed in the same places in the primeval old‐growth forest, free of direct human influences (no nest‐boxes, no additional food). The weather variation was within the range of conditions observed during 40 yr in the study area, and no climate warming occurred in the marsh tit's pre‐breeding period. Females (n = 16) shifted the onset of laying by 13–23 (median = 20) days between the seasons. This range of individual flexibility encompasses almost the whole latitudinal range of the breeding dates found across Europe. Such a buffer of plasticity would probably be sufficient for marsh tits to adjust the onset of egg‐laying to the forecasted range of climate change. A combination of temperature and photoperiod appears to be involved in fine tuning of the birds’ breeding times with spring conditions, but how the birds asses and integrate this information remains poorly understood.     

Effects of extrinsic and intrinsic factors on breeding success in a long lived seabird

There is growing concern over the impacts of climate change on animal species. Many studies have demonstrated impacts of climate change at the population level, and density dependent effects of climate are frequently reported. However, there is an increasing recognition of the differential impact of such factors on individuals since there is marked variation in individual performance. We investigated the relationships between breeding success and environmental conditions (winter NAO and one year lagged winter NAO) and intrinsic effects (colony size, pair bond duration, past breeding success rate) in the northern fulmar Fulmarus glacialis , using data from a long-term study commenced in 1950. There was a negative trend in breeding success over time, and a negative relationship with winter NAO and lagged winter NAO, which themselves had shown positive increases over the study period. The effects of lagged winter NAO remained after accounting for the linear trend. There was no evidence of density dependence, with breeding success positively related to colony size. We found strong evidence that breeding success was negatively related to pair bond duration but positively related to past breeding success rate. There was also an interaction between these two intrinsic effects such that those pairs that had historically been successful maintained success with increasing pair bond duration, whereas less successful pairs showed a decline. The prediction that there would be a differential impact of extrinsic factors among pairs was supported by an interaction between past breeding success rate and winter NAO, such that pairs with low past success rate exhibited a sharp decline in breeding success with increasing winter NAO, whereas more successful pairs did not. It is critically important to understand interactions between extrinsic factors and individual heterogeneity since a differential impact on individuals will affect population structure, and hence population dynamics.     

No effect of partner age and lifespan on female age‐specific reproductive performance in blue tits

Studies of age‐specific reproductive performance are fundamental to our understanding of population dynamics and the evolution of life‐history strategies. In species with bi‐parental care, reproductive ageing trajectories of either parent may be influenced by their partner's age, but this has rarely been investigated. We investigated within‐individual age‐specific performance (laying date and number of eggs laid) in wild female blue tits Cyanistes caeruleus and evaluated how the age and longevity of their male partner indirectly influenced the females’ reproductive performance. Females showed clear age‐dependence in both laying date and number of eggs laid. We found that female reproductive performance improved in early life, before showing a decline. Longer‐lived females had an earlier laying date throughout their lives than shorter‐lived females, but there was no difference in number of eggs laid between longer‐ and shorter‐lived females. Within breeding pairs, the female's (age‐specific) reproductive performance was not dependent on the age and longevity of the male partner. We conclude that the age and quality of the male partner may be of little importance for traits that are under direct female control.     

Nesting outcomes under anthropogenic change – effects of changing climate and nestbox provision on the reproduction of Great Tits Parus major

Global anthropogenic changes are significantly impacting the ecology and evolution of many species. Among temperate taxa, changes to reproductive phenology as a result of warming springs are apparent. However, how such responses to abiotic change interact with biotic impacts resulting from human management interventions are less clear. Here we examine the response of a range of breeding metrics (laying date, clutch size, hatching and fledging success) to interactions between climatic variables and changes in conspecific density (and hence intraspecific competition) resulting from changes in nestbox provision. Using a 37‐year dataset on the Great Tit Parus major we found little evidence for interacting effects of these two drivers. Instead we found that either climatic or competitive effects were the key influence on different metrics. Annual mean laying date substantially advanced with a warming climate, whereas clutch size, hatching success and fledging success were significantly inversely associated with conspecific density. Annual variance in clutch size and hatching success increased weakly with measures of conspecific density, but there was no association between either climatic or density measures and the annual variance in laying date or fledging success. Increasing conspecific densities therefore resulted in years with lower, but more variable, reproductive rates. These results highlight the importance of adaptive provisioning of nestboxes to enhance reproductive output of hole‐nesting birds regardless of climate‐induced phenological change. Such management is likely to be appropriate for assisting with sustaining populations responding to a changing climate.     

Consequences of a changing environment on the breeding phenology and reproductive success components in a long-distance migratory bird

Migratory birds have a narrow time window to breed, especially in the Arctic, where early nesting typically yields the highest reproductive success. We assessed temporal changes (1991–2015) in reproductive success components in relation to timing of breeding in greater snow geese (Chen caerulescens atlantica). This species breeds in the Canadian Arctic, a region that has experienced a strong warming trend. We tested the effect of laying or hatching date, year and their interaction on six reproductive components: Total clutch laid, nesting success, egg survival, hatching success, prefledging, and postfledging survival. Over 25 years, mean laying date changed little, even though it advanced 1.8 days in early breeders and was delayed 3 days in late breeders. Likewise, the number of eggs in nests initiated early in the season decreased by 0.6 egg, whereas in late nests it increased by 0.3 egg. Success of nests initiated early and late in the season was lower than nests initiated near the population mean, and consistently increased over time. The proportion of eggs surviving to partial predation and postfledging survival decreased with laying date but the pattern did not change over time. In contrast, prefledging survival was not affected by laying date initially but declined in nests initiated late in the season toward the end of the study. Overall, nests initiated close to the population mean showed little temporal change for most components of reproductive success and seem to be less affected by environmental change than nests initiated early and late in the season.     

Melanin coloration has temperature‐dependent effects on breeding performance that may maintain phenotypic variation in a passerine bird

Fluctuating selection pressure may maintain phenotypic variation because of different types of individuals being adapted to different environmental conditions. We show that the extensive variation in the coloration of male pied flycatchers (Ficedula hypoleuca) can be maintained through differences in the reproductive success of male phenotypes under different conditions. The effects of weather conditions on the relative success of different male phenotypes varied between different phases of breeding. The reproductive output of black males was the highest when it was cold during egg‐laying but warm during the nestling period, whereas the fledgling production of brown males was highest when it was continuously warm. In addition, male forehead and wing patch sizes had context‐dependent effects on timing of breeding and nestling mortality, respectively. These results indicate that environmental heterogeneity plays a role in maintaining phenotypic variation. As melanin‐based coloration is heritable, climate change may alter phenotype frequencies depending on the patterns of warming.     

Influence of Snowmelt Timing on the Diet Quality of Pyrenean Rock Ptarmigan (Lagopus muta pyrenaica): Implications for Reproductive Success

The Pyrenean rock ptarmigan (Lagopus muta pyrenaica) is the southernmost subspecies of the species in Europe and is considered threatened as a consequence of changes in landscape, human pressure, climate change, and low genetic diversity. Previous studies have shown a relationship between the date of snowmelt and reproductive success in the Pyrenean ptarmigan. It is well established that birds laying early in the breeding season have higher reproductive success, but the specific mechanism for this relationship is debated. We present an explicative model of the relationship between snowmelt date and breeding success mediated by food quality for grouse in alpine environments. From microhistological analyses of 121 faecal samples collected during three years in the Canigou Massif (Eastern Pyrenees), and the assessment of the chemical composition of the main dietary components, we estimated the potential quality of individual diets. Potential dietary quality was correlated with free-urate faecal N, a proxy of the digestible protein content ingested by ptarmigan, and both were correlated with phenological stage of consumed plants, which in turn depends on snowmelt date. Our findings suggest that the average snowmelt date is subject to a strong interannual variability influencing laying date. In years of early snowmelt, hens benefit from a longer period of high quality food resources potentially leading to a higher breeding success. On the contrary, in years of late snowmelt, hens begin their breeding period in poorer nutrient condition because the peaks of protein content of their main food items are delayed with respect to laying date, hence reducing breeding performance. We discuss the possible mismatch between breeding and snowmelt timing.     

Early springs and breeding performance in two sympatric duck species with different migration strategies

The capacity of migratory species to adapt to climate change may depend on their migratory and reproductive strategies. For example, reproductive output is likely to be influenced by how well migration and nesting are timed to temporal patterns of food abundance, or by temperature variations during the brood rearing phase. Based on two decades (1988–2009) of waterfowl counts from a boreal catchment in southern Finland we assessed how variation in ice break‐up date affected nesting phenology and breeding success in two sympatric duck species, Mallard Anas platyrhynchos and Eurasian Teal Anas crecca. In Fennoscandia these species have similar breeding habitat requirements but differ in migration distance; Teal migrate roughly seven times as far as do Mallard. Annual ice break‐up date was used as a proxy of spring ‘earliness’ to test the potential effect of climate change on hatching timing and breeding performance. Both species were capable of adapting their nesting phenology, and bred earlier in years when spring was early. However, the interval from ice break‐up to hatching tended to be longer in early springs in both species, so that broods hatched relatively later than in late springs. Ice break‐up date did not appear to influence annual number of broods per pair or annual mean brood size in either species. Our study therefore does not suggest that breeding performance in Teal and Mallard is negatively affected by advancement of ice break‐up at the population level. However, both species showed a within‐season decline in brood size with increasing interval between ice break‐up and hatching. Our study therefore highlights a disparity between individuals in their capacity to adjust to ice break‐up date, late breeders having a lower breeding success than early breeders. We speculate that breeding success of both species may therefore decline should a consistent trend towards earlier springs occur.     

Annual global mean temperature explains reproductive success in a marine vertebrate from 1955 to 2010

The salient feature of anthropogenic climate change over the last century has been the rise in global mean temperature. However, global mean temperature is not used as an explanatory variable in studies of population‐level response to climate change, perhaps because the signal‐to‐noise ratio of this gross measure makes its effect difficult to detect in any but the longest of datasets. Using a population of Leach's storm‐petrels breeding in the Bay of Fundy, we tested whether local, regional, or global temperature measures are the best index of reproductive success in the face of climate change in species that travel widely between and within seasons. With a 56‐year dataset, we found that annual global mean temperature (AGMT) was the single most important predictor of hatching success, more so than regional sea surface temperatures (breeding season or winter) and local air temperatures at the nesting colony. Storm‐petrel reproductive success showed a quadratic response to rising temperatures, in that hatching success increased up to some critical temperature, and then declined when AGMT exceeded that temperature. The year at which AGMT began to consistently exceed that critical temperature was 1988. Importantly, in this population of known‐age individuals, the impact of changing climate was greatest on inexperienced breeders: reproductive success of inexperienced birds increased more rapidly as temperatures rose and declined more rapidly after the tipping point than did reproductive success of experienced individuals. The generality of our finding that AGMT is the best predictor of reproductive success in this system may hinge on two things. First, an integrative global measure may be best for species in which individuals move across an enormous spatial range, especially within seasons. Second, the length of our dataset and our capacity to account for individual‐ and age‐based variation in reproductive success increase our ability to detect a noisy signal.     

Phenological shifts assist colonisation of a novel environment in a range‐expanding raptor

In a rapidly changing world understanding the capacity of populations to adapt to novel environments is increasingly urgent. Timing of breeding can be a highly flexible trait and adjustments in this trait can potentially buffer populations from climate change and facilitate the colonisation of new environments. Recent range‐expansions into novel climatic regimes provide a valuable opportunity to investigate the implications of plasticity in timing of breeding for population processes. Black sparrowhawks have recently colonised the Cape Peninsula of South Africa where they experience dramatically different weather patterns to those in their historical range. These include a total reversal in the rainfall regime, with the majority of rain falling in the winter as opposed to the summer months. We investigate the breeding phenology of black sparrowhawks in relation to both regional and local climate variation and, using a long‐term dataset, explore the implications of phenological shifts for reproductive success and population growth following colonisation. In the recently colonised Cape Peninsula the breeding season began up to three months earlier than within their historical range and these early breeding attempts produced more offspring. Population models suggested that this adjustment assisted the colonisation of the Cape Peninsula, reducing the probability of extinction by 23%. Contrary to expectations, we found little support for the hypothesis that black sparrowhawks were responding to local variation in rainfall. We suggest that shifts in breeding phenology may be driven in part by other novel processes, such as interspecific competition for nest sites and lower temperatures during late summer. These results provide insight into the processes that facilitated the colonisation of a novel climatic regime highlighting the potential role of a diverse range of factors.     

Reduced parental effort in relation to laying date in house sparrows (Passer domesticus): a study under controlled conditions

It is known that breeding success in birds declines with an advance in the laying date. In this paper, breeding success diminished with laying date in a house sparrow (Passer domesticus) population breeding in captivity. The principal cause of the cline in breeding success with date was a decrease in the female feeding rate. The parental effort in females decreased with the advance of the laying date, although food access was ad libitum throughout the breeding season. Consequently, this cline in breeding success may be due to a reduced parental investment with the advance of the laying date or to exhaustion due to previous reproductive events in the season. Hypotheses usually used to explain the cline in breeding success with date can not explain results in this study.     

Phenological shifts in North American red squirrels: disentangling the roles of phenotypic plasticity and microevolution

Phenological shifts are the most widely reported ecological responses to climate change, but the requirements to distinguish their causes (i.e. phenotypic plasticity vs. microevolution) are rarely met. To do so, we analysed almost two decades of parturition data from a wild population of North American red squirrels (Tamiasciurus hudsonicus). Although an observed advance in parturition date during the first decade provided putative support for climate change‐driven microevolution, a closer look revealed a more complex pattern. Parturition date was heritable [h² = 0.14 (0.07–0.21 (HPD interval)] and under phenotypic selection [β = ?0.14 ± 0.06 (SE)] across the full study duration. However, the early advance reversed in the second decade. Further, selection did not act on the genetic contribution to variation in parturition date, and observed changes in predicted breeding values did not exceed those expected due to genetic drift. Instead, individuals responded plastically to environmental variation, and high food [white spruce (Picea glauca) seed] production in the first decade appears to have produced a plastic advance. In addition, there was little evidence of climate change affecting the advance, as there was neither a significant influence of spring temperature on parturition date or evidence of a change in spring temperatures across the study duration. Heritable traits not responding to selection in accordance with quantitative genetic predictions have long presented a puzzle to evolutionary ecologists. Our results on red squirrels provide empirical support for one potential solution: phenotypic selection arising from an environmental, as opposed to genetic, covariance between the phenotypic trait and annual fitness.