Why climate change will invariably alter selection pressures on phenology

The seasonal timing of lifecycle events is closely linked to individual fitness and hence, maladaptation in phenological traits may impact population dynamics. However, few studies have analysed whether and why climate change will alter selection pressures and hence possibly induce maladaptation in phenology. To fill this gap, we here use a theoretical modelling approach. In our models, the phenologies of consumer and resource are (potentially) environmentally sensitive and depend on two different but correlated environmental variables. Fitness of the consumer depends on the phenological match with the resource. Because we explicitly model the dependence of the phenologies on environmental variables, we can test how differential (heterogeneous) versus equal (homogeneous) rates of change in the environmental variables affect selection on consumer phenology. As expected, under heterogeneous change, phenotypic plasticity is insufficient and thus selection on consumer phenology arises. However, even homogeneous change leads to directional selection on consumer phenology. This is because the consumer reaction norm has historically evolved to be flatter than the resource reaction norm, owing to time lags and imperfect cue reliability. Climate change will therefore lead to increased selection on consumer phenology across a broad range of situations.     

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.     

Influence of climate change on the abundance, distribution and phenology of woodland bird species in temperate regions

There is now overwhelming evidence that an increase in the concentration of greenhouse gases in the Earth's atmosphere has caused global temperatures to increase by 0.6 °C since 1900 and further increases of between 1.4 and 5.8 °C are predicted over the next century. Changes in climatic conditions have already influenced the demography, phenology and distribution of a wide range of plant and animal taxa. This review focuses on the impacts, both observed and potential, of climate change on birds breeding in temperate woodlands of the Western Palaearctic, a significant proportion of which are currently declining. Changes in ambient temperatures and patterns of precipitation may have direct and indirect effects on the survival rates and productivity of bird species, thus influencing population sizes. For some species or populations, the timing of events such as egg-laying and return from the wintering grounds is also changing in relation to shifts in the peak of food availability during the breeding season. The degree to which different individuals are able to track these temporal changes will have a significant bearing on population sizes and distributions in the future. Unless active management steps are taken, the relatively low dispersal rates of tree species may lead to a decrease in the total area of some woodland habitat types as losses at the southern edge of the range are likely to occur much more quickly than expansion at the northern edge. In addition, the dispersal rates of many woodland birds are themselves low, which could affect their ability to move to new habitat patches if currently occupied areas become unsuitable. Thus, woodland birds may be particularly susceptible to the impacts of climate change.     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.     

Reproduction and survival of Glaucous Gulls breeding in an Arctic seabird colony

ABSTRACT.   Despite being widespread and easily observed, little is known about the life history of Glaucous Gulls ( Larus hyperboreus ). From 1984 to 2007, we examined their breeding biology and demography at Coats Island, Nunavut, Canada, where they nest alongside a colony of 30,000 pairs of Thick-billed Murres ( Uria lomvia ). The gulls fed mainly on murre eggs and chicks and by scavenging adult carcasses. The median age at first breeding was 5 yr, and the mean age was 4.8 ± 0.9 yr. Adult survival was estimated as 0.84 ± 0.03 (SE). The mean clutch size was 2.56 eggs and the mean number of young reared per year was 1.6 (range = 0.9–2.2). Birds reared at the colony provided 40% of recruits. Assuming that survival of locally reared chicks that emigrated was similar to that of chicks that returned to the colony, about 22% of the young gulls survived to breeding age. The timing of breeding by Glaucous Gulls appeared related to the timing of laying by murres. Although the demographic characteristics of Glaucous Gulls in our study were similar to those of populations of other large gulls, adult survival was at the lower end of the range for populations of large Larus gulls. There is some evidence that Glaucous Gulls exhibit lower survival than large gulls breeding in temperate areas, possibly because of contaminant burdens. In general, however, the demographic characteristics of large gulls show little variation and are probably a product of their common phylogeny.     

Severe recent decrease of adult body mass in a declining insectivorous bird population

Migratory bird species that feed on air-borne insects are experiencing widespread regional declines, but these remain poorly understood. Agricultural intensification in the breeding range is often regarded as one of the main drivers of these declines. Here, we tested the hypothesis that body mass in breeding individuals should reflect habitat quality in an aerial insectivore, the tree swallow (Tachycineta bicolor), along a gradient of agricultural intensity. Our dataset was collected over 7 years (2005–2011) and included 2918 swallow captures and 1483 broods. Analyses revealed a substantial decline of the population over the course of the study (−19% occupancy rate), mirrored by decreasing body mass. This trend was especially severe in females, representing a total loss of 8% of their mass. Reproductive success was negatively influenced by intensive agriculture, but did not decrease over time. Interestingly, variation in body mass was independent of breeding habitat quality, leading us to suggest that this decline in body mass may result from carry-over effects from non-breeding areas and affect population dynamics through reduced survival. This work contributes to the growing body of evidence suggesting that declines in migratory aerial insectivores are driven by multiple, complex factors requiring better knowledge of year-round habitat use.     

Phenotypic plasticity in breeding plumage signals in both sexes of a migratory bird: responses to breeding conditions

Adaptive phenotypic plasticity may respond to present ambient conditions. Sexual and social signals in both sexes may express phenotype performance. Plumage signals that change discontinuously allow relating discrete variation to previous performance. Both sexes of the pied flycatcher Ficedula hypoleuca present white patches on the wings and on the forehead, which constitute sexual and social signals. Forehead patches are moulted together with body plumage in Africa, while wing patches are partly moulted in Africa and partly in the breeding area soon after breeding. We studied individual inter‐year changes (corrected for regression to the mean) in the size of forehead and wing patches of both sexes in seven years for females or six years for males in two nearby study areas in central Spain. We found that initial signal extent strongly delimits the possible subsequent changes negatively. There is a negative association of male age with forehead patch changes. Cold and rainy springs are associated in females with decreases in both patch areas and vice versa, while no association with climate is observed in male wing patch changes. Cold pre‐breeding conditions predict positive changes in female wing and male forehead patches. Breeding success is positively associated with forehead patch changes in females. Late‐breeding males experience more positive changes in forehead patch size than early‐breeding males. Some of these trends can be explained by variable costs of breeding in certain conditions for subsequent signal production and/or maintenance, while absence of trends in some cases may be explained by sex differences in costs of breeding and interactions with phenotypic quality of breeders.     

Nesting activity,breeding success and colony size for the Wedge-tailed Shearwater Puffinus pacificus on Heron Island

Abstract The Capricorn Group of islands in Australia's Great Barrier Reef Marine Park sustains one of the world's largest breeding populations of the Wedge-tailed Shearwater Puffinus pacificus. Heron Island, a 13.5 ha coral cay which supports tourist and research station leases as well as a national park, is the third largest nesting island in the group. Sample censuses of breeding burrows were conducted each year between 1985 and 1990 and a further survey was completed in 1993. These returned estimates of between 13 264±1387 and 16 337±1545 active burrows (Y±SE). Burrow densities within each of the habitats monitored showed no significant trends between years, although there were large differences in burrow density between habitats. There were roughly the same number of burrows in the developed (west) and national park (east) halves of the cay. A miniature video camera system (burrowscope), which allowed nesting chambers at the ends of burrows to be inspected, was used in 1989, 1990 and 1993. This demonstrated that around half the burrows were occupied by incubating birds. Variations were found in the distribution of incubating birds between habitats, although this did not remain constant between the years. In the 1993 season, breeding activity was traced from the burrow establishment to fledging stage. Fifty-one per cent of burrows were used for breeding (eggs laid), 77% of eggs hatched and 80% of chicks produced a fledgling. Overall breeding success for the island was estimated at 61%. In 1993 the area designated as Buildings was found to have significantly lower hatching success compared with natural habitats. Most mortality occurred at the egg stage; however, in the Fringe habitat, mortality was highest at the chick stage. Previous surveys have estimated the breeding population from burrow counts. It now appears that only about 30% of such burrows produce fledglings.     

Effects of supplementary feeding on territoriality, breeding success and survival of pheasants

1. The effects of supplementary food in spring on subsequent pheasant breeding in an intensively farmed area in southern England were assessed by a large-scale, replicated field experiment.
2. Territorial cock pheasants were counted in April, and the breeding success and survival of radio-tagged hens were monitored in six 1-km² plots during 1994 and 1995. Total numbers of young reared and post-breeding pheasant densities were found by August counts. In 1994, three randomly selected plots were supplied with wheat grains via hoppers along woodland edges and hedgerows. The other three plots acted as controls and the treatments were reversed in 1995.
3. The density of cock territories increased significantly in food-supplemented plots (44 ± 8 km−2) in relation to control plots (29 ± 8 km−2), and the presence of hoppers significantly affected the locations of territories. However, similar proportions of territorial males acquired harems in the control and food-supplemented plots. Hen density did not increase and, consequently, the mean harem size was significantly lower with supplementary feeding.
4. Hens given supplementary food did not nest earlier and the number of nests initiated, clutch sizes and the proportion of successful nests did not differ significantly from those of controls. However, hens supplied with supplementary food re-nested significantly more quickly following the loss of a nest or brood.
5. Radio-tagged hens did not rear significantly more young with supplementary feeding. Hen survival was unchanged and post-breeding pheasant densities were no higher.
6. On present evidence, spring feeding cannot be advocated as a management technique to improve the breeding success of pheasants surviving the winter.     

Age and clutch size variation in Dusky Flycatcher nest survival

Examination of spatial and temporal factors that influence nest survival can provide insight into habitat selection, reproductive decisions (e.g., clutch size), population dynamics, and conservation requirements for species. We used nest survival data for the Dusky Flycatcher Empidonax oberholseri to examine several factors that may influence nesting success. Our prediction was that the number of nest initiations would be positively associated with period nest survival. We used a model selection framework and found that nesting success was a function of clutch size and a cubic effect of age. Clutches with one, two, three, and four eggs had period survival rates of 0, 0.05, 0.33, and 0.49, respectively. Daily survival rates decreased from the onset of egg-laying and increased during the later stages of incubation before remaining relatively constant through the later portions of the nestling stage. Model-selection criterion provided support for a date effect on daily survival (i.e., daily nest survival declined across the nesting season) although the 95% confidence interval for the estimate included zero. We found that the majority of nest initiations occurred early in the nest season and declined across the season as period nest survival declined. Our prediction concerning nest survival was partially supported. In addition, we found substantial positive associations between clutch size and nest survival. While low daily survival rates for clutches with one or two eggs suggested that individuals may have reduced reproductive effort in response to nest predation risk, we did not find strong evidence that individuals reduced their clutch sizes in subsequent nest attempts. Alternative predictions, including the preferential settlement of higher quality individuals (e.g., those with the ability to lay full clutches to replace depredated nests) into high-quality habitat and differences in behavior patterns (e.g., number of visits to provision nestlings), may provide more consistent explanations for these patterns.     

The Norwegian sheep breeding scheme: description, genetic and phenotypic change

The developments in Norwegian sheep breeding since the early 1990s are reviewed. For the largest breeding population, the Norwegian White Sheep, results are presented for both genetic and phenotypic changes. Of the nine traits that make up the aggregate genotype, the largest gain per year, in per cent of the corresponding phenotypic average, was found for carcass grade (1.66%) and carcass weight (0.99%), number of lambs born at 1, 2 and 3 years of age (0.32% to 0.60%) and the maternal effect on weaning weight (0.26%). For fat grade, a genetic deterioration was estimated. This may be due to the too small weighting of this trait in the aggregate genotype and the true genetic parameters being somewhat different from the estimates in the prediction of breeding values. For lamb as well as ewe fleece weight, genetic change was close to zero - interpreted as mainly a correlated response to other traits in the aggregate genotype. Data for the two traits of fleece weight were, respectively, selected and few. Thus, phenotypic change was calculated for all traits except for fleece weight, and in addition for number of lambs at weaning, being indirectly selected for through number of lambs born. For all traits, with the exception of fat grade, advantageous phenotypic change was estimated. For weaning and carcass weight, the phenotypic change was less than the genetic change, while the opposite was observed for carcass and fat grade and number of lambs born. The latter traits can be more easily controlled by environmental actions, and the results thus exemplify the interdependency between environmental and genetic change.     

Dispersal, interpatch movements, and survival in a shrubland breeding bird community

ABSTRACT Dispersal events can affect the distribution, abundance, population structure, and gene flow of animal populations, but little is known about long‐distance movements due to the difficulty of tracking individuals across space. We documented the natal and breeding dispersal of shrubland birds among 13 study sites in a 1000 km² area in southeastern Ohio. In addition, we radio‐marked and tracked 37 adult males of one shrubland specialist, the Yellow‐breasted Chat (Icteria virens). We banded 1925 juveniles and 2112 adults of nine shrubland species from 2002 to 2005. Of these, 33 (1.7%) juveniles were encountered in subsequent years (2003–2006) as adults (natal dispersal) and 442 (20.9%) birds initially banded as breeding adults were re‐encountered in subsequent years (breeding dispersal). Apparent survival of juvenile shrubland birds on their natal patches was 0.024 (95% CI 0.016–0.036). After accounting for the probability of detection, we found that 21% of birds banded as juveniles and recaptured as adults returned to their natal patches, whereas 78% of adult birds showed fidelity to the patch where they were originally captured. Moreover, natal dispersers tended to move farther than breeding dispersers (corrected natal median = 1.7 km ± 0.37; corrected breeding median = 0.23 km ± 0.10). We used our estimates of natal dispersal and annual apparent survival to estimate true survival at 0.11 (95% CI 0.07–0.18) for juveniles in their first year. However, this estimate was only applicable for birds dispersing within 7 km of their natal patches. Interpatch movements of radio‐marked Yellow‐breasted Chats were not uncommon, with 13 of 37 males located in more than one habitat patch. Overall, we observed low natal philopatry, but high adult site fidelity for shrubland birds in our study area. Considering the frequency of short‐distance movements observed (median = 531 m, range = 88–1045 m), clustering of patches within 1 km might facilitate use of shrubland habitat.     

Effects of clutch size manipulations on reproductive behaviour and nesting success in the cooperatively breeding Bell Miner (Manorina melanophrys)

Summary An experimental manipulation of clutch size was carried out on a wild population of the cooperatively breeding Bell Miner (Manorina melanophrys, Meliphagidae) to assess which factor(s) limit clutch size in this species. Results provide some support for the trade-off hypothesis since there is a cost of reproduction for the breeding female in terms of loss of body mass. The breeding female performs most of the nestling care. Clutches of three eggs are also laid during the mid-breeding season which is the period most favourable for breeding (i.e. nestlings grow faster). This evidence also supports the intrabrood competition hypothesis. Clutches that have lost an egg were more likely to be deserted; this may be an antipredator strategy since partial clutch predation has been recorded in the field. Nest predation was high in this study (64.9%), suggesting that many small clutches may be a strategy to decrease the effect of nest predation on reproductive success over the whole breeding season (nest predation hypothesis). Both the trade-off hypothesis and the nest predation hypothesis may apply in this case since they are not mutually exclusive. The size of the attending group did not greatly affect reproductive success in the short term, although if both age structure and size of the group are taken into account, reproductive success can be better predicted.     

The demands of incubation and avian clutch size

We reviewed information on the demands of incubation to examine whether these could influence the optimal clutch size of birds. The results indicate that appreciable metabolic costs of incubation commonly exist, and that the incubation of enlarged clutches can impose penalties on birds. In 23 studies on 19 species, incubation metabolic rate (IMR) was not elevated above the metabolic rate of resting non-incubating birds (RMR), but contrary to the physiological predictions of King and others, IMR was greater than RMR in 15 studies on 15 species. Across species, IMR was substantially above basal metabolic rate (BMR), averaging 1.606 × BMR. Of six studies on three species performed under thermo-neutral conditions, none found IMR to be in excess of RMR. IMRs measured exclusively within the thermo-neutral zone averaged only 1.08 × BMR contrasting with the significantly higher figure of 1.72 × BMR under wider conditions. 16 of 17 studies on procellariiforms found IMR below RMR, indicating a significant difference between this and other orders. We could find no other taxonomic, or ecological factors which had clear effects on IMR. Where clutch size was adjusted experimentally during incubation, larger clutches were associated with: significantly lower percentage hatching success in 11 of 19 studies; longer incubation periods in eight of ten studies; greater loss of adult body condition in two of five studies; and higher adult energy expenditure in eight of nine studies. Given that incubation does involve metabolic costs and given that the demands of incubation increase sufficiently with clutch size to affect breeding performance, we propose that the optimal clutch size of birds may in part by shaped by the number of eggs the parents can afford to incubate.     

The association between the emergence of cooperative breeding and clutch size

Previous theoretical work has suggested that smaller brood sizes helped facilitate the emergence of cooperative breeding in birds. However, recent empirical evidence has found no statistically significant difference between the clutch sizes of cooperative breeders and that of noncooperative breeders. One explanation for this finding is that while small clutch sizes may predispose species to cooperative breeding, the emergence of cooperative breeding itself may influence the evolution of clutch size. Here, we develop a set of models using population dynamics to describe how the emergence of cooperative breeding influences clutch size. We find, in contrast to previous theoretical work, that the emergence of cooperative breeding does not necessarily decrease (and under certain conditions may actually increase) clutch size. In particular, clutch size may increase after the emergence of cooperative breeding if helpers – philopatric individuals that assist their breeding relatives – are able to substantially improve breeder fecundity at low costs to their own survival, and if the association between breeder and helper is brief. In many cases, clutch size increases following the emergence of cooperative breeding not because it is optimal for the breeder, but as the result of breeder–helper conflict over resource allocation.     

Synergistic effects of the invasive Chinese tallow (Triadica sebifera) and climate change on aquatic amphibian survival

Changes in climate and the introduction of invasive species are two major stressors to amphibians, although little is known about the interaction between these two factors with regard to impacts on amphibians. We focused our study on an invasive tree species, the Chinese tallow (Triadica sebifera), that annually sheds its leaves and produces leaf litter that is known to negatively impact aquatic amphibian survival. The purpose of our research was to determine whether the timing of leaf fall from Chinese tallow and the timing of amphibian breeding (determined by weather) influence survival of amphibian larvae. We simulated a range of winter weather scenarios, ranging from cold to warm, by altering the relative timing of when leaf litter and amphibian larvae were introduced into aquatic mesocosms. Our results indicate that amphibian larvae survival was greatly affected by the length of time Chinese tallow leaf litter decomposes in water prior to the introduction of the larvae. Larvae in treatments simulating warm winters (early amphibian breeding) were introduced to the mesocosms early in the aquatic decomposition process of the leaf litter and had significantly lower survival compared with cold winters (late amphibian breeding), likely due to significantly lower dissolved oxygen levels. Shifts to earlier breeding phenology, linked to warming climate, have already been observed in many amphibian taxa, and with most climate models predicting a significant warming trend over the next century, the trend toward earlier breeding should continue if not increase. Our results strongly suggest that a warming climate can interact with the effects of invasive plant species, in ways we have not previously considered, to reduce the survival of an already declining group of organisms.     

Natural selection and inheritance of breeding time and clutch size in the collared flycatcher

Many characteristics of organisms in free-living populations appear to be under directional selection, possess additive genetic variance, and yet show no evolutionary response to selection. Avian breeding time and clutch size are often-cited examples of such characters. We report analyses of inheritance of, and selection on, these traits in a long-term study of a wild population of the collared flycatcher Ficedula albicollis. We used mixed model analysis with REML estimation ("animal models") to make full use of the information in complex multigenerational pedigrees. Heritability of laying date, but not clutch size, was lower than that estimated previously using parent-offspring regressions, although for both traits there was evidence of substantial additive genetic variance (h2 = 0.19 and 0.29, respectively). Laying date and clutch size were negatively genetically correlated (rA = -0.41 +/- 0.09), implying that selection on one of the traits would cause a correlated response in the other, but there was little evidence to suggest that evolution of either trait would be constrained by correlations with other phenotypic characters. Analysis of selection on these traits in females revealed consistent strong directional fecundity selection for earlier breeding at the level of the phenotype (beta = -0.28 +/- 0.03), but little evidence for stabilising selection on breeding time. We found no evidence that clutch size was independently under selection. Analysis of fecundity selection on breeding values for laying date, estimated from an animal model, indicated that selection acts directly on additive genetic variance underlying breeding time (beta = -0.20 +/- 0.04), but not on clutch size (beta = 0.03 +/- 0.05). In contrast, selection on laying date via adult female survival fluctuated in sign between years, and was opposite in sign for selection on phenotypes (negative) and breeding values (positive). Our data thus suggest that any evolutionary response to selection on laying date is partially constrained by underlying life-history trade-offs, and illustrate the difficulties in using purely phenotypic measures and incomplete fitness estimates to assess evolution of life-history trade-offs. We discuss some of the difficulties associated with understanding the evolution of laying date and clutch size in natural populations.