The response of the Great Reed Warbler Acrocephalus arundinaceus to climate change

We examined long-term responses in the breeding performance of the Great Reed Warblers Acrocephalus arundinaceus to climate change. The study took place in various years from 1970 to 2007. During the study period, mean temperatures in the breeding season of the species increased and precipitation decreased significantly. We found evidence for the significant advancement in both earliest and annual median first-egg-laying dates. This advancement correlated with temperature increases early in the season. The latest first-egg laying dates, however, remained unchanged. Other breeding statistics: clutch size, nest losses, and production of young per nest, did not change significantly over the study period. Precipitation did not affect any of the analysed measures. It is important to note, though, that during dry seasons, the production of young per successful nest was higher. In contrast to some woodland species, the Great Reed Warbler seems to adapt well to climate change by shifting laying dates. The reason for this is probably to optimise food resources.     

Variable effects of climate change on six species of North American birds

Many recent studies have shown that birds are advancing their laying date in response to long-term increases in spring temperatures. These studies have been conducted primarily in Europe and at local scales. If climate change is a large-scale phenomenon, then we should see responses at larger scales and in other regions. We examined the effects of long-term temperature change on the laying dates and clutch sizes of six ecologically diverse species of North American birds using 50 years of nest record data. As predicted, laying dates for most (four of six) species were earlier when spring temperatures were warmer. Over the long-term, laying dates advanced over time for two species (red-winged blackbirds, Agelaius phoeniceus and eastern bluebirds, Sialia sialis). Laying date of song sparrows (Melospiza melodia) also advanced with increasing temperature when the analysis was restricted to eastern populations. Neither laying date nor clutch sizes changed significantly over time in the remaining species (American coot, Fulica americana, killdeer, Charadrius vociferous, and American robin, Turdus migratorius), an unsurprising result given the lack of increase in temperatures over time at nest locations of these species. This study indicates that the relationship between climate change and breeding in birds is variable within and among species. In large-scale analyses of North American birds, four of seven species have shown advances in laying dates with increasing temperature (including song sparrows in the east). These variable responses within and among species highlight the need for more detailed studies across large spatial scales.     

Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus,Parus major)

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first‐egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak‐beech forest, and found that both advanced their mean first‐egg dates by 11–12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2–3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70–75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within‐individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.     

The breeding biology of the Chiffchaff Phylloscopus collybita in Britain: a comparison of an intensive study with records of the BTO Nest Record Scheme

Most known aspects of the breeding biology of the Chiffchaff come from studies carried out in Central Europe. This study documents aspects of its breeding biology in Britain by comparing data gathered during an intensive study in Wytham Woods (Oxford) from 1992 to 1994, and records from 1933 to 1993 held by the BTO's Nest Record Scheme. Comparison of these two data sets showed close similarities in parameters such as: (1) laying dates; (2) length of the breeding season; (3) nest site usage; (4) clutch size; (5) length of the nestling period; and (6) the relative importance of causes of nest failure. First clutches are laid in the second half of April, and early May, with second clutches in June. Nests are built close to the ground, usually in Bramble bushes (Rubus spp.). Average clutch size decreases from 6 to 4 eggs through the season. Incubation and nestling periods last 13–14 days. Nest losses are mainly due to predation, which accounted for approximately 75% of losses in both data sets, and the Weasel (Mustela nivalis) appears to be the main predator in Wytham Woods.     

9. LIST OF MEMBERS (as at date of publication)

We studied the breeding ecology of the Chestnut-backed Sparrowlark Eremopterix leucotis over three years between 2008 and 2010. The breeding season was bimodal with a main peak in laying in autumn (March–April) and another smaller peak in spring (September–October). Nest microhabitat analyses showed they prefer nesting in open areas with lots of bare ground (median 67.5%). Nest entrance directions were biased towards the south (mean vector (µ) = 186.44°). The majority of nests (78.2%) had an apron at the nest entrance. The mean clutch size was 1.88 but there was geographic variation in clutch size between northern and southern races of the species. The mean incubation and nestling periods were 10.33 d (range 10–11 d) and 9.20 d (range 8–10 d), respectively. The results suggest that parental contributions during incubation are almost equal, but females made significantly more food deliveries during the nestling period compared to males. The diet of nestlings comprised mainly of invertebrates (50.2%), seeds (34.4%) and unidentified food items (15.4%). Breeding success was low, averaging 16.1% (range 8.1–20.6%), and the average number of fledged young per pair was 0.36 ± 0.71. Replacement broods were common and we also recorded repeat brooding attempts.     

Effect of food abundance on laying date and clutch size in the White Stork Ciconia ciconia

CapsuleFood independently affects both laying date and clutch size, suggesting that seasonal decline in clutch size is related to a decrease in food availability.

Aim To test the effect of food abundance on laying date and clutch size of the White Stork and identify the cause of seasonal decline in the number of eggs laid.

Methods During 1991 and 1996 we recorded clutch size and laying date of pairs breeding next to rubbish dumps (food abundant and constant throughout the breeding season) and birds breeding far from rubbish dumps (using natural food sources).

Results In 1991 there was no difference in mean laying date between pairs nesting at rubbish dumps and control pairs. Clutch size was significantly larger at rubbish dump nests. In contrast, mean laying date was earlier in control pairs in 1996 and there was no significant differences in clutch sizes, even when controlling for laying date effect.

Conclusion The results support the hypothesis that food availability independently affects both laying date and clutch size. The seasonal decline in clutch size close to rubbish dumps was negligible (1991) or much smaller than in the control zone (1996) suggesting that a progressive deterioration of natural food sources is the most probable reason for a decline in clutch size as the season advances.     

THE EVOLUTION OF CLUTCH SIZE. I. AN EQUATION FOR PREDICTING CLUTCH SIZE

We derive an equation for calculating the clutch sizes of birds and other long-lived animals from Murray's (1979) theory on the evolution of clutch size. For the Prairie Warbler (Dendroica discolor) in Indiana, this equation predicts an average clutch size of 3.49, less than half an egg smaller than the recorded average clutch size of 3.89. We attribute the discrepancy to sampling error and suggest that the equation satisfactorily identifies the important factors affecting the evolution of clutch size. The success of the equation in predicting clutch size of the Prairie Warbler provides additional support for Murray's theory on the evolution of clutch size.     

Breeding phenology,reproductive traits and apparent survival of sympatric Marsh Warbler Acrocephalus palustris and Blyth’s Reed Warbler Acrocephalus dumetorum

Capsule: Sympatric Marsh Warblers Acrocephalus palustris and Blyth’s Reed Warblers Acrocephalus dumetorum differ significantly in their life history traits.

Aims: To provide a direct comparison of demographic parameters among two sympatric populations of the closely related Marsh Warbler and Blyth’s Reed Warbler.

Methods: We examined breeding phenology and reproductive traits at a 25?ha study plot. We use program MARK to estimate daily nest survival and adult apparent survival rates.

Results: On average, Marsh Warblers laid the first eggs 3 days later than Blyth’s Reed Warblers. Mean clutch size in the Marsh Warbler was significantly lower than in the Blyth’s Reed Warbler. There are no significant differences between the two species for nest daily survival, duration of incubation and nestling periods. Apparent survival of adults was slightly higher in Marsh Warblers than in Blyth’s Reed Warblers.

Conclusion: Our findings suggest that two ecologically similar sympatric species differ significantly in terms of life history traits. We assume that observed differences could be the result of adaptations to environmental factors in the central parts of the species’ ranges or due to differences in mortality on migratory pathways or wintering grounds.     

Latitudinal variation in clutch size–lay date regressions in Tachycineta swallows: effects of food supply or demography?

In a study of almost 16 000 nest records from seven swallow species across the entire Western Hemisphere, clutch sizes decline with relative laying date in each population, but the slope of this decline grows steeper with increasing distance from the equator. Late‐laying birds at all latitudes lay clutches of similar sizes, suggesting that latitudinal differences may be driven primarily by earlier‐laying birds. Focused comparisons of site‐years in North America with qualitatively different food availability indicate that food supply significantly affects mean clutch size but not the clutch size–lay date regression. Other studies on the seasonality of swallow food also indicate that steeper clutch size–lay date declines in the North are not caused by steeper earlier food peaks there. The distribution of lay dates grows increasingly right‐skewed with increasing latitude. This variation in lay‐date distributions could be due to the predominance of higher quality, early‐laying (and large‐clutched) individuals among populations at higher latitudes, resulting from latitudinal variation in mortality rates and the intensity of sexual selection. Our results underscore the importance of studying clutch size and lay date in tandem and suggest new research into the causes of their joint geographic variation.     

Effect of interannual variations in sea-surface temperature on egg-laying parameters of black-tailed gulls (Larus crassirostris) at Teuri Island, Japan

Sea-surface temperature (SST) directly and indirectly affects the distribution and abundance of prey species for seabirds, so we expect variation in SST to be associated with variation in seabird life history traits. In black-tailed gulls (Larus crassirostris) at Teuri Island in northern Hokkaido, Japan, we investigated the diet of the gulls prior to egg laying in 2004 and 2005, and examined the influences of SST in March or April, when the gulls congregate in the colony, on egg-laying parameters using 13 years of data (1992–2004). The gulls fed on krill (Thysanoessa inermis) and fish prior to the egg laying. Mean first egg dates and clutch sizes were significantly and quadratically related to SST anomalies in March, but were not influenced by SST anomalies in April. There was no significant effect of SSTs in either March or April on egg volume. Sea-surface temperature anomalies in March of the years of early laying (−1 to 1°C) were higher than those in 2001 (−2.2°C), but lower than those in 1992 (+1.2°C) and 2004 (+1.1°C). Thysanoessa inermis congregates to spawn at the sea surface, when SSTs rise 3−4°C. Thus, a mismatch between food availability and the timing of egg production in the gulls could have occurred in these 3 years. This study suggests that SST fluctuations prior to laying are important in breeding success of black-tailed gulls.     

Climate change, migratory connctivity and changes in laying date and clutch size of the pied flycatcher

We examined long-term (1943–2003) variability in laying dates and clutch sizes in a Finnish population of the pied flycatcher Ficedula hypoleuca Pallas, and analysed whether potential changes were explained by changes in climatic factors at the wintering area in Africa, at migration route or at breeding grounds. Among-year variation in both mean and skewness of laying dates increased, which for mean laying date appeared to be explained by variability of temperatures at the breeding grounds and for skewness by variable temperature trends along the migration route. Pied flycatchers bred earlier in warm springs, but despite a warming trend in pre-laying temperatures, the laying dates tended to delay. Laying dates became continuously later in relation to the phenology of the environment. Mean clutch size decreased with time when mean laying date was controlled for, but the climatic factors did not appear to explain the decrease. The advancement of spring phenology may have shifted some food sources needed for egg-laying, thus leading to later laying and smaller clutches. Variation in clutch size increased when wintering conditions were favourable so that clutch size distribution was skewed with a tail of small clutches when there had been lot of rainfall (more vegetation and insects) in the wintering area. We suggest that when ecological conditions during winter were good, the tail of small clutches represented low-quality individuals that were not able to breed after bad winters. Our analyses demonstrate that measures of spread and symmetry give different information about population level changes than means, and thus complement the understanding of the potential influences of climate change on populations.     

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.     

Temperature but not rainfall influences timing of breeding in a desert bird, the trumpeter finch ( Bucanetes githagineus )

Reproductive performance in birds depends on several factors, one of the most important being the time of breeding. Birds try to fit offspring birth and growth to peak vegetative production in order to assure fledgling survival. In arid environments, where weather conditions are often extreme, birds must face unpredictable abiotic conditions. This study uses a border population of the trumpeter finch (Bucanetes githagineus) as a model to test whether climate variables (rainfall and temperature) influence breeding parameters by comparing 2 years with very different weather. The study was carried out in the Tabernas desert (southeastern Spain) in 2004 and 2005. A comparison of laying dates in the 2 years shows a 40-day delay in the date of the first clutch in the coldest year (mean minimum temperature 3°C lower in 2005 than in 2004). However, once the breeding season started, the number of clutches, clutch size, duration of the incubation period, nestling phase, fledgling rates and productivity were similar. One likely explanation for this delay is that low temperatures did not allow the germination of Diplotaxis sp., a plant forming the bulk of the trumpeter finch diet during spring. Its absence could prevent onset of breeding, although other temperature-related factors could also be involved. Although rainfall has frequently been reported as a limiting factor for arid bird species, our 2-year study shows that temperature can also influence the breeding biology of arid bird species, by affecting its timing.     

Use of statistical models based on radiographic measurements to predict oviposition date and clutch size in rock iguanas (Cyclura nubila)

The ability to noninvasively estimate clutch size and predict oviposition date in reptiles can be useful not only to veterinary clinicians but also to managers of captive collections and field researchers. Measurements of egg size and shape, as well as position of the clutch within the coelomic cavity, were taken from diagnostic radiographs of 20 female Cuban rock iguanas, Cyclura nubila, 81 to 18 days prior to laying. Combined with data on maternal body size, these variables were entered into multiple regression models to predict clutch size and timing of egg laying. The model for clutch size was accurate to 0.53 ± 0.08 eggs, while the model for oviposition date was accurate to 6.22 ± 0.81 days. Equations were generated that should be applicable to this and other large Cyclura species. © 1995 Wiley-Liss, Inc.     

Breeding ground temperature rises,more than habitat change,are associated with spatially variable population trends in two species of migratory bird

Habitat loss and climate change are key drivers of global biodiversity declines but their relative importance has rarely been examined. We attempted to attribute spatially divergent population trends of two Afro-Palaearctic migrant warbler species, Willow Warbler Phylloscopus trochilus and Common Chiffchaff Phylloscopus collybita, to changes in breeding grounds climate or habitat. We used bird counts from over 4000 sites across the UK between 1994 and 2017, monitored as part of the BTO/JNCC/RSPB Breeding Bird Survey. We modelled Willow Warbler and Common Chiffchaff population size and growth in relation to habitat, climate and weather. We then used the abundance model coefficients and observed environmental changes to determine the extent to which spatially varying population trends in England and Scotland were consistent with attribution to climate and habitat changes. Both species' population size and growth correlated with habitat, climate and weather on their breeding grounds. Changes in habitat, in particular woodland expansion, could be linked to small population increases for both species in England and Scotland. Both species' populations correlated more strongly with climate than weather, and both had an optimum breeding season temperature: 11°C for Willow Warbler and around 13.5°C for Common Chiffchaff (with marginally different predictions from population size and growth models). Breeding ground temperature increases, therefore, had the potential to have caused some of the observed Willow Warbler declines in England (where the mean breeding season temperature was 12.7°C) and increases in Scotland (mean breeding season temperature was 10.2°C), and some of the differential rates of increase for Common Chiffchaff. However, much of the variation in species' population abundance and trends were not well predicted by our models and could be due to other factors, such as species interactions, habitat and climate change in their wintering grounds and on migration. This study provides evidence that the effect of climate change on a species may vary spatially and may switch from being beneficial to being detrimental if a temperature threshold is exceeded.     

Testing for biases in selection on avian reproductive traits and partitioning direct and indirect selection using quantitative genetic models

Key life history traits such as breeding time and clutch size are frequently both heritable and under directional selection, yet many studies fail to document microevolutionary responses. One general explanation is that selection estimates are biased by the omission of correlated traits that have causal effects on fitness, but few valid tests of this exist. Here, we show, using a quantitative genetic framework and six decades of life‐history data on two free‐living populations of great tits Parus major, that selection estimates for egg‐laying date and clutch size are relatively unbiased. Predicted responses to selection based on the Robertson–Price Identity were similar to those based on the multivariate breeder's equation (MVBE), indicating that unmeasured covarying traits were not missing from the analysis. Changing patterns of phenotypic selection on these traits (for laying date, linked to climate change) therefore reflect changing selection on breeding values, and genetic constraints appear not to limit their independent evolution. Quantitative genetic analysis of correlational data from pedigreed populations can be a valuable complement to experimental approaches to help identify whether apparent associations between traits and fitness are biased by missing traits, and to parse the roles of direct versus indirect selection across a range of environments.     

Variation in laying date in relation to spring temperature in three species of tits (Paridae) and pied flycatchers Ficedula hypoleuca in southernmost Sweden

This study documents the advancement of laying dates in three species of tits (Paridae) in southernmost Sweden during recent decades, and the absence of a similar response in the pied flycatcher Ficedula hypoleuca. It is based on several different nestbox studies; the oldest one starting in 1969. During 1969 to 2012, mean spring temperatures in the study area increased by between 0.06 and 0.08°C per year, depending on the period considered. Great tits Parus major, blue tits Cyanistes caeruleus and marsh tits Poecile palustris, which generally start egg laying between the last week of April and the first week of May, all advanced laying date at a similar rate during the study period (0.25 d yr^–1). This indicates that these species were similarly affected by increasing temperatures. When accounting for mean spring temperature variation, we still found an advancement of laying date over the study period, mostly due to such relationships among marsh and blue tits. This result could reflect ongoing microevolution favouring earlier laying, but could also be a result of other factors such as increased intra‐ or inter‐specific competition for early breeding. Pied flycatchers, which generally lay during the third week of May, did not significantly advance the date of egg laying despite that the long‐term trend in the increase in ambient temperature during the 30‐d period preceding the start of egg laying was similar for pied flycatchers compared to the tit species.     

Breeding biology of the Little Egret Egretta garzetta on the southern coast of the Bay of Biscay

ABSTRACT

Capsule: Annual reproductive parameters of Little Egrets Egretta garzetta in an Atlantic coastal colony showed strong variation in a 20-year study, mainly due to extreme events.

Aims: To describe the breeding biology of Little Egrets in the Bay of Biscay and compare it with that of the Mediterranean basin. Also, to explore relationships between breeding parameters and colony size with some climatic indicators.

Methods: Phenology, number of nests, clutch size, number of hatchlings, brood size, and hatching and breeding success were recorded over 20 years.

Results: Median laying date of 214 nests was 1 May (range: 1 April–25 June) and 54% of the clutches were laid in the second half of April and the first half of May. Over the 20-year study period mean clutch size of 270 nests was 4.0, mean number of hatchlings was 3.1, and mean brood size was 2.3. Hatching success ranged from 46.1% to 100% and breeding success from zero to 100%. Number of nests was negatively associated with clutch and brood size. The highest clutch size and lowest brood size were recorded at the beginning of the season. Significant relationships were found between the number of hatchlings and the rainfall during the pre-breeding season, and between brood size and the summer rainfall.

Conclusions: Reproductive parameters showed significant variation over the study period, which highlights the importance of using long-term data sets. Breeding occurred one month later than in natural colonies of the Mediterranean basin. Negative relationships between the number of nests and clutch and brood size suggest some degree of density-dependent effects. Ranges of clutch size, number of hatchlings, and brood size were within those reported in Mediterranean populations. The effect of rainfall on reproductive parameters was weak. Extreme weather and predation events caused low rates of hatching and breeding success that affected the growth of the colony.     

Changing climate and the phenological response of great tit and collared flycatcher populations in floodplain forest ecosystems in Central Europe

This study is based on 47 years of observations (1961–2007) on two common bird species, the Great Tit (Parus major) and the Collared Flycatcher (Ficedula albicollis), and a dominant tree species in their habitat, the English Oak (Quercus robur). The study took place at four research sites in the Czech Republic located in full-grown, multi-aged floodplain forests with no forestry management. An increase in air temperature over the evaluated period clearly influenced the length of phenological phases. The full foliage date of English Oak has advanced by 8.7 days during the past 47 years. Great Tit and Collared Flycatcher populations have reacted to the changing climate in the same way, with first laying date and mean laying date advancing by between 6.0 and 9.0 days. In all cases, the trends are highly significant and consistent over all sites. Despite the ongoing shift in phenological stages toward the beginning of the year, the change does not appear to have led to mistiming in the trophic food chain. Overall, this study shows almost identical rates of change in egg laying dates for both bird species in all the floodplain forests studied, and these trends are coherent with those of English Oak and peak herbivorous caterpillar activity.     

Estimating fat and protein fuel from fat and muscle scores in passerines

Fat is the prime energy source for birds during prolonged exercise, but protein is also catabolized. Estimates of the amount of catabolizable fat and protein (termed fat and protein fuel) are therefore important for studying energetics of birds. As fat and protein fuel can only be measured by sacrificing individuals or by use of technically complex methods, scoring systems were invented to estimate fat and protein fuel of birds in the field. The visible subcutaneous fat deposits and the thickness of the flight muscles are each scored on an ordinal scale but these scales do not correspond linearly to fat and protein fuel within species, which is needed for analyses such as flight range estimates. We developed an anova ‐type model to estimate fat and protein fuel from fat scores (FS) and muscle scores (MS) along with total mass and a size measurement. Using data from 11 337 individuals of eight passerine species (Common Nightingale Luscinia megarhynchos, Eurasian Reed Warbler Acrocephalus scirpaceus, Melodious Warbler Hippolais polyglotta, Willow Warbler Phylloscopus trochilus, Orphean Warbler Sylvia hortensis, Garden Warbler Sylvia borin, Common Whitethroat Sylvia communis, Subalpine Warbler Sylvia cantillans) mist‐netted in Mauritania, West Africa, we tested for independence of FS and MS and for variation in the relationship between scores and associated mass in response to physiological state. FS, MS and third primary length (size) explained variation in body mass of all eight species analysed (R²: 0.56–0.77). The parameter estimates of the model showed that fat and protein fuel increased monotonically with increasing fat and muscle scores. In two species we found small differences in the estimates between physiological states (seasons). We evaluated our model by comparing the predicted body mass of birds with both FS and MS equal to 0 with the mean body mass of individuals mist‐netted with both scores equal to zero. The values were very close. The amount of fat extracted from dead Garden and Willow Warblers was within the range of predicted fat fuel derived from the model. We conclude that our model is a useful non‐invasive method to estimate simultaneously mean fat and protein fuel of small passerines and we provide recommendations on its use.