Flexibility in the timing of post‐breeding moult in passerines in the UK

Higher temperatures resulting from climate change have led to predictions that the duration of the breeding season of many temperate bird species may be changing. However, the extent to which breeding seasons can be altered will also depend on the degree of flexibility in processes occurring at other points in the annual cycle. In particular, plasticity in the timing of post‐breeding moult (PBM) could facilitate changes in the timing of key events throughout the annual cycle, but little is known about the level of within‐ and between‐species plasticity in PBM. As part of the British Trust for Ornithology (BTO) Ringing Scheme, many ringers routinely record moult scores of flight feathers, and these can be used to provide information on the annual progression of PBM for a range of species. Here we use ringing data to investigate patterns of PBM in 15 passerines, as well as data from the BTO Nest Record Scheme to relate these differences to the timing of breeding of these species across the UK. We find considerable variation in both the mean start (19 May–29 July) and duration (66–111 days) of PBM between species, but find no evidence that species starting PBM later in the season complete it any faster. However, there is considerable within‐species variation in PBM, particularly for multi‐brooded species; PBM starts later and is completed in less time when the duration of the breeding season (difference between first and last nests) is longer. This implies that a later end to breeding can be compensated for by faster PBM, and that advances in breeding could lead to earlier and slower PBM. Our findings suggest that adaptation of PBM in response to climate‐mediated changes in the timing and duration of the breeding season is possible. However, the requirement to complete PBM prior to migration or the onset of winter might constrain the extent to which breeding seasons can lengthen, especially for later nesting species.     

Delayed egg‐laying and shortened incubation duration of Arctic‐breeding shorebirds coincide with climate cooling

Biological impacts of climate change are exemplified by shifts in phenology. As the timing of breeding advances, the within‐season relationships between timing of breeding and reproductive traits may change and cause long‐term changes in the population mean value of reproductive traits. We investigated long‐term changes in the timing of breeding and within‐season patterns of clutch size, egg volume, incubation duration, and daily nest survival of three shorebird species between two decades. Based on previously known within‐season patterns and assuming a warming trend, we hypothesized that the timing of clutch initiation would advance between decades and would be coupled with increases in mean clutch size, egg volume, and daily nest survival rate. We monitored 1,378 nests of western sandpipers, semipalmated sandpipers, and red‐necked phalaropes at a subarctic site during 1993–1996 and 2010–2014. Sandpipers have biparental incubation, whereas phalaropes have uniparental incubation. We found an unexpected long‐term cooling trend during the early part of the breeding season. Three species delayed clutch initiation by 5 days in the 2010s relative to the 1990s. Clutch size and daily nest survival showed strong within‐season declines in sandpipers, but not in phalaropes. Egg volume showed strong within‐season declines in one species of sandpiper, but increased in phalaropes. Despite the within‐season patterns in traits and shifts in phenology, clutch size, egg volume, and daily nest survival were similar between decades. In contrast, incubation duration did not show within‐season variation, but decreased by 2 days in sandpipers and increased by 2 days in phalaropes. Shorebirds demonstrated variable breeding phenology and incubation duration in relation to climate cooling, but little change in nonphenological components of traits. Our results indicate that the breeding phenology of shorebirds is closely associated with the temperature conditions on breeding ground, the effects of which can vary among reproductive traits and among sympatric species.     

Long‐term climate impacts on breeding bird phenology in Pennsylvania,USA

Climate change is influencing bird phenology worldwide, but we still lack information on how many species are responding over long temporal periods. We assessed how climate affected passerine reproductive timing and productivity at a constant effort mist‐netting station in western Pennsylvania using a model comparison approach. Several lines of evidence point to the sensitivity of 21 breeding passerines to climate change over five decades. The trends for temperature and precipitation over 53 years were slightly positive due to intraseasonal variation, with the greatest temperature increases and precipitation declines in early spring. Regardless of broodedness, migration distance, or breeding season, 13 species hatched young earlier over time with most advancing >3 days per decade. Warm springs were associated with earlier captures of juveniles for 14 species, ranging from 1‐ to 3‐day advancement for every 1 °C increase. This timing was less likely to be influenced by spring precipitation; nevertheless, higher rainfall was usually associated with later appearance of juveniles and breeding condition in females. Temperature and precipitation were positively related to productivity for seven and eleven species, respectively, with negative relations evident for six and eight species. We found that birds fledged young earlier with increasing spring temperatures, potentially benefiting some multibrooded species. Indeed, some extended the duration of breeding in these warm years. Yet, a few species fledged fewer juveniles in warmer and wetter seasons, indicating that expected future increases could be detrimental to locally breeding populations. Although there were no clear relationships between life history traits and breeding phenology, species‐specific responses to climate found in our study provide novel insights into phenological flexibility in songbirds. Our research underscores the value of long‐term monitoring studies and the importance of continuing constant effort sampling in the face of climate change.     

Factors influencing plasticity in the arrival‐breeding interval in a migratory species reacting to climate change

Climate change is profoundly affecting the phenology of many species. In migratory birds, there is evidence for advances in their arrival time at the breeding ground and their timing of breeding, yet empirical studies examining the interdependence between arrival and breeding time are lacking. Hence, evidence is scarce regarding how breeding time may be adjusted via the arrival‐breeding interval to help local populations adapt to local conditions or climate change. We used long‐term data from an intensively monitored population of the northern wheatear (Oenanthe oenanthe) to examine the factors related to the length of 734 separate arrival‐to‐breeding events from 549 individual females. From 1993 to 2017, the mean arrival and egg‐laying dates advanced by approximately the same amount (~5–6 days), with considerable between‐individual variation in the arrival‐breeding interval. The arrival‐breeding interval was shorter for: (a) individuals that arrived later in the season compared to early‐arriving birds, (b) for experienced females compared to first‐year breeders, (c) as spring progressed, and (d) in later years compared to earlier ones. The influence of these factors was much larger for birds arriving earlier in the season compared to later arriving birds, with most effects on variation in the arrival‐breeding interval being absent in late‐arriving birds. Thus, in this population it appears that the timing of breeding is not constrained by arrival for early‐ to midarriving birds, but instead is dependent on local conditions after arrival. For late‐arriving birds, however, the timing of breeding appears to be influenced by arrival constraints. Hence, impacts of climate change on arrival dates and local conditions are expected to vary for different parts of the population, with potential negative impacts associated with these factors likely to differ for early‐ versus late‐arriving birds.     

Climate change leads to differential shifts in the timing of annual cycle stages in a migratory bird

Shifts in reproductive phenology due to climate change have been well documented in many species but how, within the same species, other annual cycle stages (e.g. moult, migration) shift relative to the timing of breeding has rarely been studied. When stages shift at different rates, the interval between stages may change resulting in overlaps, and as each stage is energetically demanding, these overlaps may have negative fitness consequences. We used long‐term data of a population of European pied flycatchers (Ficedula hypoleuca) to investigate phenological shifts in three annual cycle stages: spring migration (arrival dates), breeding (egg‐laying and hatching dates) and the onset of postbreeding moult. We found different advancements in the timing of breeding compared with moult (moult advances faster) and no advancement in arrival dates. To understand these differential shifts, we explored which temperatures best explain the year‐to‐year variation in the timing of these stages, and show that they respond differently to temperature increases in the Netherlands, causing the intervals between arrival and breeding and between breeding and moult to decrease. Next, we tested the fitness consequences of these shortened intervals. We found no effect on clutch size, but the probability of a fledged chick to recruit increased with a shorter arrival‐breeding interval (earlier breeding). Finally, mark–recapture analyses did not detect an effect of shortened intervals on adult survival. Our results suggest that the advancement of breeding allows more time for fledgling development, increasing their probability to recruit. This may incur costs to other parts of the annual cycle, but, despite the shorter intervals, there was no effect on adult survival. Our results show that to fully understand the consequences of climate change, it is necessary to look carefully at different annual cycle stages, especially for organisms with complex cycles, such as migratory birds.     

Trophic mismatch and its effects on the growth of young in an Arctic herbivore

In highly seasonal environments, timing of breeding of organisms is typically set to coincide with the period of highest resource availability. However, breeding phenology may not change at a rate sufficient to keep up with rapid changes in the environment in the wake of climate change. The lack of synchrony between the phenology of consumers and that of their resources can lead to a phenomenon called trophic mismatch, which may have important consequences on the reproductive success of herbivores. We analyzed long‐term data (1991–2010) on climate, plant phenology and the reproduction of a long‐distance Arctic migrant, the greater snow goose (Chen caerulescens atlantica), in order to examine the effects of mismatched reproduction on the growth of young. We found that geese are only partially able to adjust their breeding phenology to compensate for annual changes in the timing of high‐quality food plants, leading to mismatches of up to 20 days between the two. The peak of nitrogen concentration in plants, an index of their nutritive quality for goslings, occurred earlier in warm springs with an early snow melt. Likewise, mismatch between hatch dates of young and date of peak nitrogen was more important in years with early snow melt. Gosling body mass and structural size at fledging was reduced when trophic mismatch was high, particularly when the difference between date of peak nitrogen concentration and hatching was >9 days. Our results support the hypothesis that trophic mismatch can negatively affect the fitness of Arctic herbivores and that this is likely to be exacerbated by rising global temperatures.     

Effect of climatic change on the duration of the breeding season in three European thrushes

Capsule Evidence for extension of breeding seasons in Song Thrushes and Blackbirds (multiple‐brooded species) and shortening in Fieldfares (a single‐brooded species).

Aims To analyse breeding data from central Europe during 1964–2006 in relation to climatic conditions operating at breeding and wintering grounds, and to compare breeding phenology in urban and rural habitats.

Methods Using chick ringing data we evaluated long‐term changes in breeding phenology of Fieldfares Turdus pilaris, Song Thrushes T. philomelos and Common Blackbirds T. merula. Changes in phenology were analysed for the 5th percentile, median, 95th percentile, inter‐quantile range (IQR) and brood size.

Results All thrushes showed consistent trends towards earlier onset of breeding. However, there was a contrasting pattern in the later phenological characteristics (median, 95th percentile and IQR) that were associated with single‐ versus multiple‐brooded species. The single‐brooded Fieldfares revealed an advancing trend in all phenological phases, which caused shortening of its breeding period. In contrast, the multiple‐brooded Song Thrushes and Blackbirds delayed both medians and the 95th percentiles of breeding dates leading to extension of their breeding seasons. Temperatures at both wintering and breeding grounds were generally negatively associated with the onset of breeding. Urban birds bred significantly earlier than their rural conspecifics, but brood sizes did not differ.

Conclusion Multiple‐brooded species may respond differently to increasing spring temperatures than single‐brooded species.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Time‐lapse cameras reveal latitude and season influence breeding phenology durations in penguins

Variation in the phenology of avian taxa has long been studied to understand how a species reacts to environmental changes over both space and time. Penguins (Sphenicidae) serve as an important example of how biotic and abiotic factors influence certain stages of seabird phenology because of their large ranges and the extreme, dynamic conditions present in their Southern Ocean habitats. Here, we examined the phenology of gentoo (Pygoscelis papua) and chinstrap penguins (Pygoscelis antarctica) at 17 sites across the Scotia arc, including the first documented monitoring of phenology on the South Sandwich Islands, to determine which breeding phases are intrinsic, or rather vary across a species range and between years. We used a novel method to measure seabird breeding phenology and egg and chick survival: time‐lapse cameras. Contrary to the long‐standing theory that these phases are consistent between colonies, we found that latitude and season had a predominant influence on the length of the nest establishment, incubation, and guard durations. We observe a trend toward longer incubation times occurring farther south, where ambient temperatures are colder, which may indicate that exposure to cold slows embryo growth. Across species, in colonies located farther south, parents abandoned nests later when eggs were lost or chicks died and the latest record of eggs or chicks in the nest occurred earlier during the breeding period. The variation in both space and time observed in penguin phenology provides evidence that the duration of phases within the annual cycle of birds is not fundamental, or genetic, as previously understood. Additionally, the recorded phenology dates should inform field researchers on the best timing to count colonies at the peak of breeding, which is poorly understood.     

The timing of gonadal development and moult in three raptors with different breeding seasons: effects of gender, age and body condition

Differences between species in breeding seasons are thought to be mediated through differences in their reproductive physiology. Little is known about how the timing and duration of gonadal maturation varies between raptor species, how the timing of moult relates to the gonadal cycle, whether the timing and degree of sexual maturation varies between juveniles and adults or whether body condition has a significant effect. To address these questions, data on gonadal size and moult for adults and juveniles of both sexes of three raptor species were extracted from the Predatory Bird Monitoring Scheme (based on birds found dead by members of the public). The three species, Sparrowhawk Accipiter nisus, Kestrel Falco tinnunculus and Barn Owl Tyto alba, have different ecologies – diurnal bird predator, diurnal mammal predator and nocturnal mammal predator, respectively. All are single‐brooded but have different breeding seasons. The duration of gonadal maturation was markedly different between the species. Barn Owls showed the earliest maturation and the latest gonad regression, and Sparrowhawks the latest maturation and earliest gonad regression. Kestrels were intermediate. In males of all species, the testes remained fully mature throughout their respective breeding seasons. In females, the ovaries remained partially mature throughout the breeding season. Moult started slightly earlier in Sparrowhawks than in Kestrels and coincided with gonadal regression in the two species. Although females of the two species started to moult earlier than males, moult duration was similar between the sexes. Barn Owls showed no distinct annual pattern of moult. In juveniles of all three species, the gonads were smaller than in adults throughout spring and started to mature later. Gonad size in birds that had starved tended to be smaller than in birds dying from other causes, but did not influence the difference in gonad mass between adults and juveniles and between seasons. Body condition had no effect on moult. Whilst ecology has led to the evolution of different breeding seasons, differences between species, and between adults and juveniles, are mediated through adaptive differences in their reproductive physiology.     

Diverging phenological responses of Arctic seabirds to an earlier spring

The timing of annual events such as reproduction is a critical component of how free‐living organisms respond to ongoing climate change. This may be especially true in the Arctic, which is disproportionally impacted by climate warming. Here, we show that Arctic seabirds responded to climate change by moving the start of their reproduction earlier, coincident with an advancing onset of spring and that their response is phylogenetically and spatially structured. The phylogenetic signal is likely driven by seabird foraging behavior. Surface‐feeding species advanced their reproduction in the last 35 years while diving species showed remarkably stable breeding timing. The earlier reproduction for Arctic surface‐feeding birds was significant in the Pacific only, where spring advancement was most pronounced. In both the Atlantic and Pacific, seabirds with a long breeding season showed a greater response to the advancement of spring than seabirds with a short breeding season. Our results emphasize that spatial variation, phylogeny, and life history are important considerations in seabird phenological response to climate change and highlight the key role played by the species’ foraging behavior.     

Age‐specific responses to spring temperature in a migratory songbird: older females attempt more broods in warmer springs

Increasing global temperature has led to an interest in plasticity in the timing of annual events; however, little is known about the demographic consequences of changing phenology. Annual reproductive success varies significantly among individuals within a population, and some of that variation has to do with the number of broods attempted by reproducing adults. In birds, female age and the timing of reproduction are often predictors of multiple breeding. We hypothesize that double brooding rates may be affected by spring temperature and that the response may vary with female age. We used a long‐term reproductive data set for a migratory songbird, the prothonotary warbler (Protonotaria citrea) to assess which factors influence (a) an individual female's probability of double brooding and (b) the annual variation in population‐level double brooding rates. We found that older and earlier nesting birds are more likely to double brood, and that there is no evidence for senescence with regard to this trait such that the oldest females were most likely to double brood. Previous experience with double brooding (i.e., whether the female double brooded in the previous year) significantly increased the probability of doing so again. When assessing annual variation in the double brooding rate, we found an interaction between spring temperature and the proportion of older females in the population. Specifically, older females are more likely to double brood in years with warmer springs, but this relationship was not seen for younger females. Previous studies have shown that warmer temperatures lead to earlier and narrower peaks in resources and we hypothesize that these peaks are more available to older and earlier arriving females, enabling them to successfully raise more than one brood in a season. Understanding how different age classes respond to changing environmental conditions will be imperative to managing declining species.     

Effect of climate change on breeding phenology,clutch size and chick survival of an upland bird

Upland birds are predicted to be particularly vulnerable to the effects of climate change, yet few studies have examined these effects on their breeding phenology and productivity. Laying dates of Red Grouse Lagopus lagopus scotica in the Scottish Highlands advanced by 0.5 days/year between 1992 and 2011 and were inversely correlated with pre‐laying temperature, with a near‐significant increase in temperature over this period. Earlier clutches were larger and chick survival was greater in earlier nesting attempts. However, chick survival was also higher in years with lower May temperatures and lower August temperatures in the previous year, the latter probably related to prey abundance in the subsequent breeding season. Although laying dates are advancing, climate change does not currently appear to be having an overall effect on chick survival of Red Grouse within the climate range recorded in this study.     

Individual fitness and the effects of a changing climate on the cessation and length of the breeding period using a 34‐year study of a temperate songbird

Studies of the phenological responses of animals to climate change typically emphasize the initiation of breeding although climatic effects on the cessation and length of the breeding period may be as or more influential of fitness. We quantified links between climate, the cessation and length of the breeding period, and individual survival and reproduction using a 34‐year study of a resident song sparrow (Melospiza melodia) population subject to dramatic variation in climate. We show that the cessation and length of the breeding period varied strongly across years, and predicted female annual fecundity but not survival. Breeding period length was more influential of fecundity than initiation or cessation of breeding alone. Warmer annual temperature and drier winters and summers predicted an earlier cessation of breeding. Population density, the date breeding was initiated, a female's history of breeding success, and the number of breeding attempts initiated previously also predicted the cessation of breeding annually, indicating that climatic, population, and individual factors may interact to affect breeding phenology. Linking climate projections to our model results suggests that females will both initiate and cease breeding earlier in the future; this will have opposite effects on individual reproductive rate because breeding earlier is expected to increase fecundity, whereas ceasing breeding earlier should reduce it. Identifying factors affecting the cessation and length of the breeding period in multiparous species may be essential to predicting individual fitness and population demography. Given a rich history of studies on the initiation of breeding in free‐living species, re‐visiting those data to estimate climatic effects on the cessation and length of breeding should improve our ability to predict the impacts of climate change on multiparous species.     

Amphibian breeding phenology trends under climate change: predicting the past to forecast the future

Global climate warming is predicted to hasten the onset of spring breeding by anuran amphibians in seasonal environments. Previous data had indicated that the breeding phenology of a population of Fowler's Toads (Anaxyrus fowleri) at their northern range limit had been progressively later in spring, contrary to generally observed trends in other species. Although these animals are known to respond to environmental temperature and the lunar cycle to commence breeding, the timing of breeding should also be influenced by the onset of overwintering animals’ prior upward movement through the soil column from beneath the frost line as winter becomes spring. I used recorded weather data to identify four factors of temperature, rainfall and snowfall in late winter and early spring that correlated with the toads’ eventual date of emergence aboveground. Estimated dates of spring emergence of the toads calculated using a predictive model based on these factors, as well as the illumination of the moon, were highly correlated with observed dates of emergence over 24 consecutive years. Using the model to estimate of past dates of spring breeding (i.e. retrodiction) indicated that even three decades of data were insufficient to discern any appreciable phenological trend in these toads. However, by employing weather data dating back to 1876, I detected a significant trend over 140 years towards earlier spring emergence by the toads by less than half a day/decade, while, over the same period of time, average annual air temperature and annual precipitation had both increased. Changes in the springtime breeding phenology for late‐breeding species, such as Fowler's Toads, therefore may conform to expectations of earlier breeding under global warming. Improved understanding of the environmental cues that bring organisms out of winter dormancy will enable better interpretation of long‐term phenological trends.     

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.     

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.     

Mate desertion by male Great Reed Warblers Acrocephalus arundinaceus at the end of the breeding season

Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains.     

Getting the timing right: antler growth phenology and sexual selection in a wild red deer population

There has been growing interest in the determinants of the annual timing of biological phenomena, or phenology, in wild populations, but research on vertebrate taxa has primarily focused on the phenology of reproduction. We present here analyses of the phenology of the annual growth of a secondary sexual characteristic, antlers in red deer (Cervus elaphus) males. The long-term individual-based data from a wild population of red deer on the Isle of Rum, Scotland allow us to consider ecological factors influencing variation in the phenology of growth of antlers, and the implications of variation in antler growth phenology with respect to the phenotype of antler grown (antler mass) and annual breeding success. The phenology of antler growth was influenced by local environmental conditions: higher population density delayed both the start date (during spring) and the relative end date (in late summer) of antler growth, and warmer temperatures in the September and April prior to growth advanced start and end dates, respectively. Furthermore, there was variation between individuals in this phenotypic plasticity of start date, although not in that of end date of growth. The phenology of antler growth impacted on the morphology of antlers grown, with individuals who started and ended growth earliest having the heaviest antlers. The timing of antler growth phenology was associated with breeding success in the following mating season, independently of the mass of antlers grown: an earlier start of antler growth was associated with siring a higher number of the calves born the following spring. Our results suggest that the phenology of traits that are not directly correlated with offspring survival may also regularly show correlations with fitness.     

Phenological delay despite warming in wood frog Rana sylvatica reproductive timing: a 20-year study

Across all taxa, amphibians exhibit some of the strongest phenological shifts in response to climate change. As climates warm, amphibians and other animals are expected to breed earlier in response to temperature cues. However, if species use fixed cues such as daylight, their breeding timing might remain fixed, potentially creating disconnects between their life history and environmental conditions. Wood frogs Rana sylvatica are a cold-adapted species that reproduce in early spring, immediately after breeding ponds are free of ice. We used long-term surveys of wood frog oviposition timing in 64 breeding ponds over 20 yr to show that, despite experiencing a warming of 0.29°C per decade in annual temperature, wood frog breeding phenology has shifted later by 2.8 d since 2000 (1.4 d per decade; 4.8 d per °C). This counterintuitive pattern is likely the result of changes in the timing of snowpack accumulation and melting. Finally, we used relationships between climate and oviposition between 2000 and 2018 to hindcast oviposition dates from climate records to model longer-term trends since 1980. Our study indicates that species can respond to fine-grained seasonal climate heterogeneity within years that is not apparent or counterintuitive when related to annual trends across years.