The breeding biology of the White-spectacled Bulbul,Pycnonotus xanthopygos,at the northwestern edge of its distribution range

The White-spectacled Bulbul, Pycnonotus xanthopygos, is an abundant and possibly invasive species in Turkey, where it has gradually expanded its distribution and breeding range in both western and southeastern directions. This study focused on its breeding biology, which is still poorly known. The breeding activity extends from February until September. The preferred nesting areas are mainly gardens and maquis groves, where 24 different nesting tree species have been identified. The clutch size is 3.3 ± 0.8 eggs per pair, nesting success 68%, hatching success 94%, fledgling success 95%, and overall breeding success 89%. While nesting success differs significantly between the years, we found no significant differences in hatching, fledging, and overall breeding success between the years studied. Despite favourable climatic conditions in the Mediterranean region, the species makes only one brood per year in a relatively extended breeding season extending over seven months, and has a relatively a high reproduction rate per nest.     

Reduced reproductive performance associated with warmer ambient temperatures during incubation in a winter‐breeding,food‐storing passerine

Timing of reproduction can influence individual fitness whereby early breeders tend to have higher reproductive success than late breeders. However, the fitness consequences of timing of breeding may also be influenced by environmental conditions after the commencement of breeding. We tested whether ambient temperatures during the incubation and early nestling periods modulated the effect of laying date on brood size and dominant juvenile survival in gray jays (Perisoreus canadensis), a sedentary boreal species whose late winter nesting depends, in part, on caches of perishable food. Previous evidence has suggested that warmer temperatures degrade the quality of these food hoards, and we asked whether warmer ambient temperatures during the incubation and early nestling periods would be associated with smaller brood sizes and lower summer survival of dominant juveniles. We used 38 years of data from a range‐edge population of gray jays in Algonquin Provincial Park, Ontario, where the population has declined over 50% since the study began. Consistent with the “hoard‐rot” hypothesis, we found that cold temperatures during incubation were associated with larger brood sizes in later breeding attempts, but temperatures had little effect on brood size for females breeding early in the season. This is the first evidence that laying date and temperature during incubation interactively influence brood size in any bird species. We did not find evidence that ambient temperatures during the incubation period or early part of the nestling period influenced summer survival of dominant juveniles. Our findings provide evidence that warming temperatures are associated with some aspects of reduced reproductive performance in a species that is reliant on cold temperatures to store perishable food caches, some of which are later consumed during the reproductive period.     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

Reproductive success and helper effects in the cooperatively breeding grey-crowned babbler

Cooperative breeding, where some individuals help to raise offspring that are not their own, is a relatively rare social system in birds. We studied the breeding biology of a declining cooperative breeder, the grey-crowned babbler Pomatostomus temporalis , with the aim of isolating the social factors that affect its reproductive success. Most breeding pairs were assisted by philopatric offspring, although pairs could breed successfully without helpers. Females laid up to four clutches (usually three eggs per clutch) per season. Male (but not female) helpers increased the number of young fledged from individual nests and the likelihood of re-nesting, resulting in higher seasonal fledgling production. Helper effects on brood size and fledgling production were greater in the second year of the study, which was also characterized by higher nest failure. This suggests that helpers enhance reproduction more in poor conditions. Our study demonstrates the interacting effects of social and ecological factors on reproductive success, and that retention of offspring is not always beneficial for the breeders in cooperative species.     

Reproductive performance of the Bald Ibis Geronticus calvus in relation to rainfall and grass-burning

Reproductive performance of the Bald Ibis Geronticus calvus was studied during three consecutive nesting seasons (1978–80) at a single large breeding colony in Natal, South Africa. A focal sample of 22 nest sites (out of a total of 60 in the colony) was inspected at frequent intervals to determine clutch sizes, egg survival, hatching success, nestling mortality and fledgling production in each year. Mean clutch size was slightly lower in 1980, compared to the two previous years, although survival and hatching success of eggs were similar in all three years. However, nestling mortality was significantly higher in 1980, largely as a result of parental nest desertion and nestling starvation, so that the proportion of breeding pairs producing at least one fledged offspring, and the average brood size at fledging, were lower compared to 1978 and 1979.
The 1978 and 1979breeding seasons were both preceded by 12 months of above average rainfall, and Bald Ibises fed extensively on small invertebrates obtained in burnt grassland in the study area during the nesting period. By contrast, the onset of nesting in 1980 followed 11 consecutive months of sub-normal rainfall. As a result of the drought, and the consequent lack of controlled burning by landowners in the study area in 1980, very little burnt grassland was available during the Bald Ibis' breeding season, and the birds fed primarily in cultivated pastures during the customary burning period. The lower reproductive performance of Bald Ibises at the study site in 1980 is discussed in relation to the possible effects of drought on the abundance of invertebrate prey and the extent of controlled burning in the species' grassland biotope.     

Factors affecting nesting success of a bird assembly in the central Monte Desert, Argentina

We estimated nesting success of a breeding bird assemblage in the central Monte Desert in four breeding seasons (1995–1999). We analyzed the effect of year, period within the breeding season, nest type, and nest location on daily survival rates (DSR). Averaged nesting success of passeriforms in natural habitats was very low (<20% on average, n=12 species). DSR tended to be lower during the egg-laying stage in Furnariidae, Rhinocryptidae, Mimidae, and Emberizidae. Tyrannidae had the highest DSR for all nesting stages and Emberizidae the lowest. Predation was the main cause of nest failure (>90% on average, n=12 species). DSR differed among years, being higher in the driest year (1995) and lower in the most humid year (1997). Nests initiated in the first half of the breeding season had higher DSR than those initiated in the second half. Closed nests had similar survival rates to open nests. DSR differed according to species of plant that supported the nest. Nest predation in the central Monte Desert may be an important selective pressure in birds' life history, which is in accordance with other results for South American temperate birds.     

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

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

Genetic and environmental effects on morphology and fluctuating asymmetry in nestling barn swallows

A barn swallow Hirundo rustica partial cross‐fostering experiment with simultaneous brood size manipulation was conducted in two years with contrasting weather conditions, to estimate heritable variation in tarsus, tail and wing size and fluctuating asymmetry. Environmental stress had contrasting effects depending on trait type. Significant heritabilities for tarsus, tail and wing size were found only in enlarged broods irrespective of year effects, while tarsus asymmetry was significantly heritable in the year with benign weather conditions irrespective of brood size manipulation effects. Tail, wing and composite (multicharacter) asymmetry were never significantly heritable. The environment with the higher heritability generally had higher additive genetic variance and lower environmental variance, irrespective of trait type. Heritability was larger for trait size than for trait asymmetry. Patterns of genetic variation in nestlings do not necessarily translate to the juvenile or adult stage, as indicated by lack of correlation between nestling and fledgling traits.     

Is the blue tit falling into an ecological trap in Argentine ant invaded forests?

Because insectivorous birds must evaluate resources for reproduction before settling into a breeding habitat, they can fall into an ecological trap if informative cues about habitat suitability become dissociated from their actual yield. Given their potential to affect ecological networks, invasive ant species are potential candidates for triggering such ecological traps. We combined observational and experimental approaches to examine whether the variation in food supply for nestlings resulting from the invasion of the Argentine ant, Linepithema humile, had any influence on the breeding ecology of the blue tit, Cyanistes caeruleus, an insectivorous foliage-gleaner. We investigated the effects of the ant invasion on breeding performance (nesting success, clutch size, brood size and breeding success) and offspring quality (body size and condition, developmental stability and plumage colour) in replicated Mediterranean forest areas over a period of 3 years. There was no evidence that the reduction in caterpillar availability resulting from the invasion had a concurrent negative effect on the blue tit’s ability to successfully rear nestlings in optimal conditions, at least as measured here. Although the raw figures suggest an increased level of nutritional stress in blue tits breeding in invaded forests, the data analyses showed no significant alterations in terms of productivity or offspring fitness. The reproductive performance of the blue tit has been shown to be remarkably resilient to the Argentine ant-mediated food shortage, either because the prey reduction following the invasion did not reach a critical threshold or because of compensatory activity by the progenitors. We cannot conclusively reject an ecological trap triggered by the ant invasion on blue tits, since neither fledgling recruitment nor the prospective survival of parents were assessed. Even though we could not confirm short-term consequences of the Argentine ant invasion on blue tit reproductive fitness, the long-term bottom-up effects of the invasion remain unknown and should not be ruled out.     

Density-dependent decline of breeding success in an introduced, increasing mute swan Cygus olor population

An increasing population of the mute swan Cygnus olor , was studied from its very establishment in 1976 until 1998. As the number of pairs increased, there was a decline in all production parameters, including the average number of clutches per pair, the average number of broods per clutch, and the average number of fledged young per brood. In a multiple regression analysis covering the whole breeding season, the number of pairs and the average number of clutches per pair explained 71% of the variation in the average number of fledglings per pair. During the brood season, the average number of fledglings per brood was an additionally important parameter in explaining fledgling production. Habitat quality seemed to affect breeding: in places occupied earlier more cygnets fledged from a clutch than in habitats inhabited later. The decreased production of young very likely reflects density-dependent effects on reproduction. This density dependence seems to operate on the breeding grounds, since winter harshness did not affect breeding success. Density-dependent processes started acting already when pairs were beginning to nest, and continued during the brood period. Density dependence has apparently not been detected at the pair stage in earlier studies of bird populations.     

Long-term trends in the timing of breeding and brood size in the Red-Backed Shrike Lanius collurio in the Czech Republic, 1964–2004

Climate change has affected breeding dates and clutch sizes in many bird species. To date, most of the studies aimed at assessing the effects of climate change on these phenological events in birds have been on hole-nesting species and the changes linked either to local climate variation or to some large-scale composite variables, such as the North Atlantic Oscillation (NAO). Relatively less is known about the climate responses of open-nesting birds and on the relative roles of climate variables at different scales. Using bird ringing records covering a time span of 41 years, we documented shifts in the timing of breeding and brood size in a long-distance migrant, the Red-Backed Shrike (Lanius collurio) from a central European population. We found a 3- to 4-day shift towards earlier breeding and an increase in brood size by approximately 0.3 nestlings since 1964. The Red-Backed Shrikes start to breed in May and rear the first nestlings in June. During the period 1964–2004, temperatures in May significantly increased, while the increase in June temperatures was not significant. Simultaneous tests on the influence of local climate variables and the NAO index revealed a better performance of local climate. The increasing temperature in May was positively associated with the advancement of breeding. Similarly, at a local scale, higher May temperatures were followed by larger brood sizes, while a high amount of rainfall had a strong negative effect.     

Control of invasive predators improves breeding success of an endangered alpine passerine

Birds living in alpine environments are becoming increasingly impacted by human‐induced threats. We investigated the impacts of introduced mammalian predators on an endangered alpine species, the New Zealand Rockwren Xenicus gilviventris, and assessed whether predator control improved its breeding success. Nest monitoring revealed that the primary cause of nest failure was predation by invasive mammals, primarily Stoats Mustela erminea and House Mice Mus musculus. Daily survival rates (DSR) decreased with nest age, and nests were at their most vulnerable to predators just prior to fledging. DSR, egg‐hatching and fledgling rates were all improved by predator trapping, demonstrating the significant impacts that even low numbers of invasive predators can have on sensitive alpine and upland species.     

Cold snaps,snowfall and sudden death from ischemic heart disease.

The short-term effect of low temperature on the incidence of ischemic heart disease over a 15-year period was examined. To reduce confounding by other seasonal factors the analysis was restricted to the winter months and was based on the change in the daily rate of sudden death at the time of cold snaps (arbitrarily defined as days on which the mean temperature was at least 4.4 degrees C lower than the day before) and around the time of heavy snowfalls. A statistically significant increase in the daily rate of sudden death at the time of cold snaps occurred only in men under 65 years of age, and even in this group the effect was of relatively small magnitude (+16%) compared with the large change in rate following heavy snowfalls (+88%). Among persons aged 65 years or over cold snaps had virtually no effect, and only the men in this group showed an increased daily rate of sudden death following a snowfall. These results suggest that much of the increased frequency of death from ischemic heart disease in winter, particularly among the elderly, must be due to factors other than short-term cold stress.     

Effects of site fidelity and breeding performance on mate retention in a short‐lived passerine,the tree swallow Thachycineta bicolor

We studied the relationship among re‐mating, site fidelity and breeding performance in the tree swallow Tachycineta bicolor using 16 y of data on reproductive biology in a population breeding in nest boxes near Ithaca, New York. Of 217 pairs for which both members survived the non‐breeding season, 76% mated with a new partner and 24% reunited with their previous mate. Pairs did not increase their breeding success by breeding together for more than one breeding season. Males produced fewer fledglings after breeding with a new partner, but females neither increased nor decreased their success when breeding with a new mate. Females who bred with a new partner were younger than females that reunited with their previous mates, and they were more likely to move to a different nest box. Males that bred with a new mate were of similar age to males that reunited, and they did not move more often. The probability of breeding with a new partner was better predicted by female age than by previous breeding success, suggesting that re‐mating was not strongly affected by past breeding performance. Because younger females change breeding sites more frequently than do older females and females that mated with a new partner were younger than females that reunited with their previous mates, we suggest that the tendency of tree swallows to change partners between years is a by‐product of lower site fidelity of younger females rather than a strategy for increasing breeding success.     

Climate change could alter the distribution of mountain pine beetle outbreaks in western Canada

Climate change can markedly impact biology, population ecology, and spatial patterns of eruptive insects due to the direct influence of temperature on insect development and population success. The mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae), is a landscape‐altering insect that infests forests of North America. Abundant availability of host trees due to altered disturbance regimes has facilitated an unprecedented, landscape‐wide outbreak of this pest in British Columbia and Alberta, Canada, during the past decade. A previous outbreak in the 1980s, in central British Columbia, collapsed due to host depletion and extreme cold weather events. Despite the importance of such extreme weather events and other temperature‐related signals in modulating an outbreak, few landscape‐level models have studied the associations of extreme cold events with outbreak occurrences. We studied the individual associations of several biologically‐relevant cold temperature variables, and other temperature/degree‐day terms, with outbreak occurrences in a spatial‐temporal logistic regression model using data from the current outbreak. Timing, frequency, and duration of cold snaps had a severe negative association with occurrence of an outbreak in a given area. Large drops in temperature (>10°C) or extreme winter minimum temperatures reduced the outbreak probability. We then used the model to apply eight different climate change scenarios to the peak year of the current outbreak. Our scenarios involved combinations of increasing annual temperature and different variances about this trend. Our goal was to examine how spatial outbreak pattern would have changed in the face of changing thermal regime if the underlying outbreak behaviour remained consistent. We demonstrate that increases in mean temperature by 1°C to 4°C profoundly increased the risk of outbreaks with effects first being manifested at higher elevations and then at increasing latitudes. However, increasing the variance associated with a mean temperature increase did not change the overall trend in outbreak potential.     

Great tits growing old: selective disappearance and the partitioning of senescence to stages within the breeding cycle

Deterioration of reproductive traits with age is observed in an increasing number of species. Although such deterioration is often attributed to reproductive senescence, a within-individual decline in reproductive success with age, few studies on wild animals have focused on direct fitness measures while accounting for selective disappearance and terminal effects, and to our knowledge none have determined how senescence effects arise from underlying reproductive traits. We show for female great tits that such an approach helps understanding of the onset, impact and architecture of senescence. Cross-sectional analysis of 49 years of breeding data shows annual recruit production to decline from 3.5 years of age, this decline affecting 9 per cent of females each year. Longitudinal analyses, however, show that selective disappearance of poor-quality breeders partly masks senescence, which in fact starts at 2.8 years and affects 21 per cent of females each year. There is no evidence for abrupt terminal effects. Analyses of underlying traits show no deterioration in clutch size, but significant declines in brood size and fledgling number. Furthermore, these traits contribute −9, 12 and 39 per cent to the senescent decline in recruit production, respectively. Besides providing detailed knowledge of the patterns and architecture of senescence in a natural population, these results illustrate the importance of modelling individual variation, and facilitate study of the underlying mechanisms of senescence.     

Within-season sequential density dependence regulates breeding success in mallards Anas platyrhynchos

Density dependence in vital rates is a key issue in population ecology but remains largely unexplored experimentally. We studied breeding success, lake use, and prey availability in wild mallards Anas platyrhynchos on small nemoral lakes in a replicated, two-year cross-over experiment in which pair density was increased. The number of wild mallards that settled on lakes prior to introductions of extra pairs did not differ between control and introduction years. Introductions led to a lake-level reduction in the number of broods observed. However, the number of stage 2+ (almost fledged) ducklings did not differ between treatments, nor did lake utilization by nonbreeding adults, broods and ducklings. Prey resource availability differed greatly among lakes, but it did not correlate with breeding success. Partialling out the possible effect of food competition from wild adult nonbreeding mallards did not change this conclusion. Our study demonstrates sequential density dependence in breeding success; introductions caused a decrease in brood number, but despite fewer broods a similar number of nearly fledged ducklings were produced. We suggest that predation and/or lake change of broods soon after hatching created these patterns. We conclude that using a single and late measure of breeding success such as fledged birds can mask regulatory processes. Implications of density dependence and its relation to individual reproductive success are understood better if breeding success is decomposed into nest success, duckling survival and fledgling survival.     

Nest survival of piping plovers at a dynamic reservoir indicates an ecological trap for a threatened population

In the past 60 years, reservoirs have reshaped riverine ecosystems and transformed breeding habitats used by the threatened piping plover (Charadrius melodus; hereafter plover). Currently, 29 % of the Northern Great Plains plover population nests at reservoirs that might function as ecological traps because reservoirs have more diverse habitat features and greater dynamics in water levels than habitats historically used by breeding plovers. We examined factors influencing daily survival rates (DSR) of 346 plover nests at Lake Sakakawea (SAK; reservoir) during 2006–2009 by evaluating multiple a priori models, and we used our best model to hindcast nest success of plovers during 1985–2009. Our observed and hindcast estimates of nest success were low compared to published estimates. Previous findings indicate that plovers prefer nest sites that are low relative to water level. We found that elevation of nests above the water level had a strong positive correlation with DSR because water levels of SAK typically increased throughout the nesting period. Habitat characteristics on the reservoir differ from those that shaped nest-site selection for plovers. Accordingly, extraordinary nest loss occurs there in many years, largely due to inundation of nests, and based on low fledging rates those losses were not compensated by potential changes in chick survival. Therefore, our example supports the concept of ecological traps in birds because it addresses quantitative assessments of habitat preference and productivity over 25 years (since species listing) and affects a large portion of the population.     

Hungry chicks and mortal parents: A state-variable approach to the breeding seasons of birds

We here report the results of a dynamic programming model of the breeding season of a bird, patterned after the Black-capped Chickadee (Parus atricapillus) nesting near Ithaca, New York. Optimal breeding seasons fall largely before the peak in food distribution, and this shift appears to be because peak metabolic demands of the brood are coordinated with the peak in food density and because the risk of adult mortality selects for earlier breeding than would otherwise be favored. This latter effect, first proposed by Fisher, may explain the nearly universal observation that early breeding appears to be favored in birds from a wide range of habitats and phylogenies.     

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.