Peak Food Availability Research Articles

Alpine songbirds at higher elevations are only raised with a slight delay and therefore under harsher environmental conditions.

The breeding phenology of birds is often timed to coincide with a peak in food availability. However, the shortening of the vegetation period with increasing elevation may force bird species at high elevations to breed earlier in relation to optimal environmental conditions due to time constraints. We investigated differences in fledging dates in five Alpine woodland songbird species along an elevational gradient from 1500 to 2200 m in Switzerland. We estimated fledging dates from a nationwide citizen science bird monitoring dataset and used the date when the proportion of observations of 'fledged young' reached 50% among all observations indicating breeding behaviour. This measure had the advantage that we could estimate average timing of the broods across a wide geographic range and over many years without the need to search for individual nests. We then compared differences in timing of the broods with climatic conditions and larch budburst across different elevational bands. The daily mean air temperature of 10-15°C was reached 34-38 days later at 2200 m compared to 1500 m, which is a similar delay as found in previous reports on snow melt-out date. The average delay in larch budburst was 19.2 days at 2200 m compared to 1500 m. In comparison, the average timing of the birds' broods was only 5.4 days later in coal tits and 0.5 days later in Alpine tits at 2200 compared to 1500 m (the two species for which we had the narrowest interval estimates). Also, the estimated delay at higher elevations in the broods of song thrushes, mistle thrushes and Eurasian chaffinches was relatively small. Rather than postponing breeding dates to better environmental conditions later in the season that would match the earlier conditions at low elevation, songbirds breeding at higher elevations may thus have evolved adaptations to cope with the harsher conditions.

The phenology and clutch size of UK Blue Tits does not differ with woodland composition

The deciduous tree‐herbivorous caterpillar‐insectivorous bird food chain is a well‐studied system for investigating the impacts of climate change across trophic levels. To date, across Europe, most attention has focused on the impacts of increasing spring temperature on changes to phenology in Oak‐dominated (Quercus spp.) woodlands. Paridae species and Pied Flycatcher Ficedula hypoleuca are the most studied secondary consumers, all of which demonstrate an advancement in reproductive phenology with increases in spring temperature. Shifts in climate and phenology may also impact on reproductive investment in clutch size, and the effects of climate on phenology and clutch size may vary depending on woodland composition. To date, the effects of among‐habitat variation in phenology and reproductive investment have received little attention. Insectivorous birds inhabiting woodlands that differ in tree composition may differ in the timing of breeding, due to local tree leafing phenology acting as a cue for egg‐laying date and/or clutch size. Moreover, for most insectivorous birds, woodland composition within a territory is likely to be the main determinant of food availability for both adults and chicks. Consequently, if warming springs affect the temporal patterns of food availability differently across different woodland compositions, this may affect the optimal average local phenology for nesting birds. Here, using data from 34 long‐term (mean 15 years) nest monitoring sites across the UK, we investigate the effect of woodland tree composition and temperature on Blue Tit Cyanistes caeruleus first egg date (FED) and clutch size. We supplemented the nest monitoring data by quantifying woodland composition, at a site level, through modified point counts. We predict that birds breeding in woodlands with greater proportions of late‐leafing species, such as Oak and Ash Fraxinus excelsior, will breed later than those breeding in woodlands with greater proportions of early‐leafing species, such as Birch Betula spp. and Beech Fagus sylvatica. We found no evidence for differences in Blue Tit FED or clutch size in relation to the proportion of any of the tree species investigated, after controlling for temperature and latitude (FED: −3.4 and 2.2, clutch size: −0.4 and − 0.2 eggs for one‐unit increase in temperature and latitude, respectively). In recent decades and across all sites, clutch size has decreased as spring temperatures have increased, a strategy which could allow birds flexibly to adjust their breeding phenology such that nestling demand coincides with peak food availability. The lack of an effect of woodland composition on Blue Tit phenology suggests Blue Tits do not fine‐tune their reproductive phenology to the local tree composition. Whether this lack of evidence for phenological divergence is due to an absence of divergent selection on breeding phenology and clutch size or to gene flow is not clear.

Complex relationships between climate and reproduction in a resident montane bird.

Animals use climate-related environmental cues to fine-tune breeding timing and investment to match peak food availability. In birds, spring temperature is a commonly documented cue used to initiate breeding, but with global climate change, organisms are experiencing both directional changes in ambient temperatures and extreme year-to-year precipitation fluctuations. Montane environments exhibit complex climate patterns where temperatures and precipitation change along elevational gradients, and where exacerbated annual variation in precipitation has resulted in extreme swings between heavy snow and drought. We used 10 years of data to investigate how annual variation in climatic conditions is associated with differences in breeding phenology and reproductive performance in resident mountain chickadees (Poecile gambeli) at two elevations in the northern Sierra Nevada mountains, USA. Variation in spring temperature was not associated with differences in breeding phenology across elevations in our system. Greater snow accumulation was associated with later breeding initiation at high, but not low, elevation. Brood size was reduced under drought, but only at low elevation. Our data suggest complex relationships between climate and avian reproduction and point to autumn climate as important for reproductive performance, likely via its effect on phenology and abundance of invertebrates.

Phenology effects on productivity and hatching-asynchrony of American kestrels (Falco sparverius) across a continent

Optimal reproductive performance occurs when birds time reproduction to coincide with peak food availability. Deviation from optimal timing, or mismatch, can affect productivity, though birds may mediate some mismatch effects by altering their incubation behavior. We studied the consequences of nesting timing (i.e., clutch initiation relative to an index of spring start) on productivity across the breeding range of American kestrels ( Falco sparverius ) in the United States and southern Canada, and associations between nesting timing, incubation behavior, and hatching asynchrony. We used observations from long-term nest box monitoring, remote trail cameras, and community-scientist-based programs to obtain data on clutch initiation, productivity, incubation, and hatching synchrony. Kestrels that initiated clutches after the extended spring index (SI-x, start of spring estimate) had higher rates of nest failure and fewer nestlings than earlier nesters, and effects of nesting timing on productivity were strongest in the Northeast. In contrast, kestrels in the Southwest experienced a more gradual decline in productivity across the season. Spatial effects may be the result of regional differences in growing seasons and temporal nesting windows (duration of nesting season). Specifically, resource availability in the Northeast was highly peaked during the breeding season, potentially resulting in shorter nesting windows. Conversely, resource curves were more prolonged in the Southwest, and growing seasons are becoming longer with climate change, potentially resulting in longer nesting windows. We found an inverse relationship between nesting timing and the onset of male incubation. Males from breeding pairs that initiated clutches after SI-x began incubation sooner than males from breeding pairs that initiated clutches before SI-x. Early-onset of male incubation was positively associated with hatching asynchrony, creating increased age variation in developing young. In sum, nesting phenology relative to the SI-x has consequences for American kestrels’ productivity, and these consequences vary across space. The early onset of incubation may act as a potential adaptive behavior to advance the average hatch date and spread out energetic demands. Given the effects of nesting timing on productivity, kestrels are likely to be sensitive to climate-driven advances in growing seasons and vulnerable to phenological mismatch, particularly in the Northeast. Top panel show availability of food (green polygon) in two different types of environments, a highly seasonal environment with peaked productivity and a less seasonal environment with prolonged growing seasons, and the outcome of two nesting attempts, one before the start of spring (yellow) and another after (purple). The later nesting attempt results in fewer young, especially in the seasonal environment. The bottom panel shows that in pairs that nest before the start of spring (yellow), males delay incubation and young hatching at the same time, whereas in later nesting pairs (purple), males may incubate earlier and young hatch asynchronously. • We studied the phenology and productivity of an avian predator across North America. • American kestrels nesting after the start of spring had lower productivity. • Effects of late nesting on productivity were strongest in the Northeast. • Later nesting kestrels altered incubation so that eggs hatched asynchronously. • Spatial patterns of phenology effects reveal populations sensitive to mismatch.

Seasonal trends in adult apparent survival and reproductive trade-offs reveal potential constraints to earlier nesting in a migratory bird.

Birds aim to optimize resources for feeding young and self-maintenance by timing reproduction to coincide with peak food availability. When reproduction is mistimed, birds could incur costs that affect their survival. We studied whether nesting phenology correlated with the apparent survival of American kestrels (Falco sparverius) from two distinct populations and examined trends in clutch-initiation dates. We estimated apparent survival using multi-state mark-recapture models with nesting timing, nesting success, sex, age, and weather covariates. Nesting timing predicted the apparent survival of successful adults; however, the effect differed between populations. Early nesting kestrels had higher apparent survival than later nesters in the western population, where kestrels have a relatively long nesting season. At the eastern site, where kestrels have a relatively short nesting season, the pattern was reversed-later nesters had higher apparent survival than earlier nesters. Nesting timing did not affect the apparent survival of adults with failed nests suggesting that the energetic cost of producing fledglings contributed to the timing effect. Finally, clutch-initiation dates advanced in the western population and remained static in the eastern population. Given that both populations have seasonal declines in productivity, population-specific survival patterns provide insight into seasonal trade-offs. Specifically, nesting timing effects on survival paralleled productivity declines in the western population and inverse patterns of survival and reproduction in the eastern population suggest a condition-dependent trade-off. Concomitant seasonal declines in reproduction and survival may facilitate population-level responses to earlier springs, whereas seasonal trade-offs may constrain phenology shifts and increase vulnerability to mismatch.

Sea ice and local weather affect reproductive phenology of a polar seabird with breeding consequences

Abstract Breeding at the right time is essential for animals living in seasonal environments to ensure that energy requirements for reproduction, especially the nutritional needs for rearing offspring, coincide with peak food availability. Climate change is likely to cause modifications in the timing of maximum food availability, and organisms living in polar environments where the breeding period is heavily contracted may be particularly affected. Here we used a 26-year dataset to study the phenological response of a pagophilic species, the Snow Petrel (Pagodroma nivea), to climate change and its demographic impact. First, we investigated the trends and relationships between climate variables and hatching dates measured in three neighboring colonies. In a second step, we examined the impact of the hatching date and environmental covariates on the fledging probability. Our results showed that sea ice, a climate-related variable, showed a positive temporal trend. We found that hatching date was delayed when sea ice concentration was greater and local weather conditions were worse (i.e., increase in the number of windy days or the number of snow days). Hatching date had a negative effect on fledging probability, and fledging probability showed a bell-shaped temporal trend. We suggest that Snow Petrels can delay breeding phenology in response to environmental conditions. However, this plasticity may be limited as fledging success decreased with delayed hatching, potentially making the Snow Petrel vulnerable to a mismatch between resource availability and nutritional needs.

Artificial light reduces foraging opportunities in wild least horseshoe bats

Artificial light at night has been proposed as a global threat to biodiversity. Insectivorous bats are strictly nocturnal animals that are vulnerable to disruption from artificial light. Given that many light-sensitive bats tend to avoid night light during roost departure, it is often assumed that nighttime light pollution reduces their foraging opportunities, albeit empirical evidence in support of this hypothesis remains elusive. Here, we used least horseshoe bats, Rhinolophus pusillus, to assess whether white artificial light is detrimental for the opportunities of foraging. We manipulated the levels of ambient illumination and perceived predation risk inside the bat roost. We monitored bats' emergence activity using high-speed video and audio recording systems. DNA-based faecal dietary analysis and insect survey were applied to determine activity time of prey in foraging areas. Following experimentally manipulation of white light-emitting diode (LED) lighting 0–15 min after sunset, bat pass, flight duration, and echolocation pulse emission decreased. The mean emergence time of bats flying out was delayed by 14 min under lit treatment compared with the dark control. Only 10% of bats left for foraging during 40 min of light exposure. Aversive effects of LED light on bat emergence were robust regardless of the presence of a potential predator. Insect prey reached a peak of abundance between 30 and 60 min after sunset. These results demonstrate that white artificial light hinders evening emergence behavior in least horseshoe bats, leading to a mismatch between foraging onset and peak food availability. Our findings highlight that light pollution overrides foraging onset, suggesting the importance of improving artificial lighting scheme near the roosts of light-sensitive bats.

Exploring the drivers of variation in trophic mismatches: A systematic review of long-term avian studies.

Many organisms reproduce in seasonal environments, where selection on timing of reproduction is particularly strong as consumers need to synchronize reproduction with the peaked occurrence of their food. When a consumer species changes its phenology at a slower rate than its resources, this may induce a trophic mismatch, that is, offspring growing up after the peak in food availability, potentially leading to reductions in growth and survival. However, there is large variation in the degree of trophic mismatches as well as in its effects on reproductive output.Here, we explore the potential causes for variation in the strength of trophic mismatches in published studies of birds. Specifically, we ask whether the changes in the degree of mismatch that have occurred over time can be explained by a bird's (a) breeding latitude, (b) migration distance, and/or (c) life‐history traits.We found that none of these three factors explain changes in the degree of mismatch over time. Nevertheless, food phenology did advance faster at more northerly latitudes, while shifts in bird phenology did not show a trend with latitude.We argue that the lack of support in our results is attributable to the large variation in the metrics used to describe timing of food availability. We propose a pathway to improve the quantification of trophic mismatches, guided by a more rigorous understanding of links between consumers and their resources.

Seasonal reproduction of northernmost endangered forest musk deer (Moschus berezovskii) in China and the synchronization with climatic conditions

Reproductive rhythms, as a response to environmental seasonality, may maximize the survival possibility of both newborns and parents, which is vital for ex situ conservation and sustainable management in wildlife. Forest musk deer (Moschus berezovskii) is a critically endangered ungulate of China. In order to estimate its reproduction characteristics and performance in northern China, a historical distribution area, the parturition pattern of the northernmost captive forest musk deer in Huailai Musk Deer Farm from 2017 to 2018 was analyzed. We found that the parturition of forest musk deer was highly seasonal with a peak on May 27<sup>th</sup>, and 75% of parturitions were completed within 22 days from May to June, which could be an adaptation to the seasonality of local food resources. The parturition peak was four weeks before the best hydrothermal conditions when food resources were maximally provided. The time lag between the parturition date and the peak of food availability was affected by climatic conditions during lactation as warmer temperatures in the 4<sup>th</sup> and 5<sup>th</sup> week and heavier precipitation in the 4<sup>th</sup> week of lactation shortened the time lag by five days (P < 0.05). The synchronization between parturition rhythms and seasonality of climatic conditions indicated that the forest musk deer could be farmed in Northeastern Taihang Mountains and northern China, where there is a suitable climate for the musk deer reproduction. Sustainable musk production and musk deer release into the wild where there are suitable climatic conditions has been proved to be practicable, which can benefit the ex situ conservation of endangered forest musk deer and musk resources sustainability.

Nestling diet and parental food provisioning in a declining mountain passerine reveal high sensitivity to climate change

Mountain ecosystems naturally experience strong seasonal weather variations leading to a brief peak in food availability that constrains bird reproduction. Climate change accentuates both the intra‐ and interannual weather variability, which in turn can reduce the predictability of food resources and hence impact population demography. Yet, relatively little is known about the influence of environmental factors on the breeding ecology of mountain birds. Here, we quantified the nestling diet and provisioning behaviour of the Alpine ring ouzel Turdus torquatus alpestris, an emblematic and declining thrush species typical of central European treeline ecotones, and relate these parameters to local weather conditions. Nests were monitored with camcorders to assess prey provisioning frequency and identify items delivered by parents to nestlings, as well as to estimate prey biomass. Our results indicate the prominence of earthworms (Lumbricidae) in the nestling diet, both in terms of abundance (80%) and biomass (90%). Elevated ambient temperatures negatively impacted both prey provisioning rates and biomass delivered to chicks by parents, while rainfall had a positive effect on the delivered biomass. The mean prey item biomass decreased throughout the breeding season, as did the proportion of earthworms in nestlings' diet. These findings highlight the key role played by local weather in parental provisioning behaviour, probably reflecting the low availability of the staple food source, earthworms, in warm and dry weather contexts. In particular, they underpin how climate alterations, notably increasing ambient temperatures and changing precipitation regimes, could impact mountain birds. Although effects on reproductive performance and population dynamics still ought to be studied, these results further our understanding of the ecological mechanisms potentially at play in the decline of wildlife inhabiting high‐elevation, climate‐sensitive ecosystems.

Weather conditions explain reproductive success and advancement of the breeding season in Western Capercaillie (Tetrao urogallus)

Climate change and associated changes in weather patterns have globally widespread effects on natural systems. Shifts in phenology can affect the reproductive success of birds by causing a mismatch between the onset of breeding and favourable conditions for reproduction, such as a peak in food availability. Weather conditions and their changes have also long been discussed as affecting the reproductive success of grouse (Tetraoninae) and contributing to rapid declines in many central European populations. We monitored the reproductive success of Western Capercaillie Tetrao urogallus in the Black Forest, Germany, over a 14‐year period to study how weather conditions correlated to reproductive success and to assess whether changing environmental conditions were related to a seasonal shift in reproduction. We observed a temporal shift in peak lekking activity from late to mid‐April, which corresponds to an advance of 0.65 days/year. Furthermore, we show that warm conditions during the reproductive season were positively associated with reproductive success (i.e. larger brood size, more hens with chicks and more chicks per hen) and that precipitation volume during the early weeks after hatching was negatively correlated with brood size. Precipitation before copulation (i.e. peak of lekking) affected the proportion of hens with chicks and the number of chicks per hen. Sex ratio in broods was also affected by weather conditions, with more female than male chicks in years with low temperatures and high precipitation. By analysing spring weather predictions for the period 2020–2039, we found that weather conditions during the reproductive period remained constant when assuming a continued advancement of reproduction onset, although annual variation was high. We show the importance of weather conditions in influencing Western Capercaillie reproductive success. Western Capercaillie in the Black Forest seem to adjust the timing of reproduction according to changes in weather conditions, but this advancement is unlikely to be limitless, making the future impact of climate change on the species uncertain.

Rapid plastic breeding response to rain matches peak prey abundance in a tropical savanna bird.

Changes in climate are shifting the timing of life cycle events in the natural world. Compared to northern temperate areas, these effects are relatively poorly understood in tropical and southern regions, where there is limited information on how timing of breeding and food availability are affected by climatic factors, and where patterns of breeding activity are more unpredictable within and between years. Combining a new statistical modelling approach with 5years of continuous individual-based monitoring of a monsoonal tropical insectivorous bird, we quantified (a) the proximate climatic drivers at two trophic levels: timing of breeding and abundance of arthropod prey; (b) the effect of climate variation on reproductive output and (c) the role of individual plasticity. Rainfall was identified as the main determinant of phenology at both trophic levels. Throughout the year, likelihood of egg laying increased very rapidly in response to even small amounts of rain during the preceding 0-3weeks. Adult body mass and male sperm storage also increased rapidly after rain, suggesting high breeding preparedness. Additionally, females were flexible, since they were more likely to nest whether their previous attempt was longer ago and unsuccessful. Arthropod abundance also increased after rainfall, but more slowly, with a peak around 10weeks. Therefore, the peak food availability coincided with the presence of dependent fledglings. Fitness benefits of nesting after more rain appeared to be linked to offspring quantity rather than quality: nest attempts following higher rainfall produced larger clutches, but showed no improvement in nestling mass or relative fledging success. The response of clutch size to rainfall was plastic, since repeated sampling showed that individual females laid larger clutches after more rain, possibly mediated by improved body mass. Rapid, individually flexible breeding in response to rainfall and slower increase in arthropod abundance also as a response to rainfall, might buffer insectivorous species living in tropical seasonal environments from climate change-induced phenological trophic mismatches.

Snow cover phenology is the main driver of foraging habitat selection for a high-alpine passerine during breeding: implications for species persistence in the face of climate change

High-alpine ecosystems are strongly seasonal and adverse environments. In these ecosystems, the brevity of optimal breeding conditions means species must efficiently track spatiotemporal variations in resources in order to synchronise their reproductive effort with peaks in food availability. Understanding the details of prey-habitat associations and their seasonality in such ecosystems is thus key for deciphering species’ ecological niches and developing sound conservation action. However, the ecological requirements of high-alpine avifauna remain poorly documented. Furthermore, mountain ranges in the Old World are affected not only by profound alterations of climate, but also by changes in land-use, the interaction of which hampers both proper forecasting of species’ resilience to environmental change and delivery of evidence-based conservation guidance. Here, we investigate the prey-habitat associations of a high-alpine passerine, the White-winged Snowfinch (Montifringilla nivalis), by radio-tracking breeding adults in the Swiss Alps. In late spring and early summer, Snowfinches foraged preferentially next to invertebrate-rich, melting snow patches where Tipulidae larvae abound. Later, in mid-summer, they favoured flower-rich alpine meadows. When foraging, they always preferred short ground vegetation while avoiding rock and scree. Their pattern of foraging habitat selection reflects trade-offs between food abundance and accessibility, i.e. prey availability. The reliance of this passerine on a habitat mosaic where snow plays a major role questions its ability to cope with climate change due to future habitat loss and potential phenological mismatches. Targeted grazing could possibly help in habitat management by aiming at maintaining invertebrate-rich meadows with short vegetation. Yet, it remains an open question whether habitat management would suffice to compensate for the potentially detrimental effects of the progressive retreat of snow fields to higher elevations.

The effects of weather, group size and type of nest on the timing of egg-laying in the Southern Ground-hornbill Bucorvus leadbeateri

The timing of breeding in birds has important consequences if food availability varies seasonally. Optimal timing of the most energetically demanding stages of breeding to coincide with peaks in food availability can increase breeding success. Early breeding may be challenging for females producing eggs, but generally birds initiating egg-laying early in the breeding season are more likely to succeed. Initiating egg-laying later may be less costly to females, but raising offspring after the peak in food availability becomes challenging. Balancing these costs and benefits is important for the cooperative breeding Southern Ground-hornbill Bucorvus leadbeateri (hereafter ‘Ground-hornbill’), which typically produces only a single clutch of two eggs and fledges a single chick per breeding season. A previous study found that Ground-hornbill groups that lay early in the breeding season are more likely to fledge a chick than later-breeding groups. We investigated factors associated with the timing of breeding such as group size, the type of nest used, rainfall and temperature in a study site supplemented with nest boxes. Ground-hornbills laid eggs from 9 September to 14 November and the average number of days between the first and the last laying date in a season was 48 ± 32 days (range 10–101 days). Larger groups, particularly those using natural nests compared to those using nest boxes, laid later than smaller groups. Egg-laying was delayed under hot, dry spring conditions and laying dates were earlier following above-average spring rainfall.

No effects of asynchrony between hatching and peak food availability on chick growth in Semipalmated Plovers (Charadrius semipalmatus) near Churchill, Manitoba

Birds rely on consistent patterns of food availability on their breeding grounds to successfully complete their breeding cycle. Due to ongoing warming of the sub-Arctic, there is potential for a mismatch between the peak in available invertebrate biomass and the peak in food demand for shorebird chicks. During the summers of 2010 and 2011, we investigated the relationship between temperature and benthic and terrestrial invertebrate biomass, measured using three sampling techniques in Churchill, Manitoba. We also investigated the relationship between timing of breeding of Semipalmated Plovers (Charadrius semipalmatus) and timing of peaks in invertebrate biomass. In 2011, chick growth rates were also measured to examine whether hatching in synchrony with the peak in invertebrate biomass during the brood rearing period affected growth rates. In 2010, emergent and core invertebrate biomass were negatively related to soil degree days, whereas in 2011, core biomass increased with soil degree days and pitfall biomass increased with air temperature. Total invertebrate biomass (summed over trap types) peaked from 25 to 31 days before the median chick hatch date in 2010 and 10 days after the median chick hatch date in 2011. In 2011 we did not detect any effects of asynchrony on the growth of Semipalmated Plover chicks. These results may indicate that food resources in their environment remain adequate throughout the breeding season, despite inter-annual fluctuations in the timing of invertebrate peaks.

Arctic Geese Tune Migration to a Warming Climate but Still Suffer from a Phenological Mismatch

Climate warming challenges animals to advance their timing of reproduction [1], but many animals appear to be unable to advance at the same rate as their food species [2, 3]. As a result, mismatches can arise between the moment of largest food requirements for their offspring and peak food availability [4-6], with important fitness consequences [7]. For long-distance migrants, adjustment of phenology to climate warming may be hampered by their inability to predict the optimal timing of arrival at the breeding grounds from their wintering grounds [8]. Arrival can be advanced if birds accelerate migration by reducing time on stopover sites [9, 10], but a recent study suggests that most long-distance migrants are on too tight a schedule to do so [11]. This may be different for capital-breeding migrants, which use stopovers not only to fuel migration but also to acquire body stores needed for reproduction [12-14]. By combining multiple years of tracking and reproduction data, we show that a long-distance migratory bird (the barnacle goose, Branta leucopsis) accelerates its 3,000km spring migration to advance arrival on its rapidly warming Arctic breeding grounds. As egg laying has advanced much less than arrival, they still encounter a phenological mismatch that reduces offspring survival. A shift toward using more local resources for reproduction suggests that geese first need to refuel body stores at the breeding grounds after accelerated migration. Although flexibility in body store use allows migrants to accelerate migration, this cannot solve the time constraint they are facing under climate warming.

Impact of urbanization on abundance and phenology of caterpillars and consequences for breeding in an insectivorous bird.

Urbanization can have marked effects on plant and animal populations' phenology, population size, predator-prey, interactions and reproductive success. These aspects are rarely studied simultaneously in a single system, and some are rarely investigated, e.g., how insect phenology responds to urban development. Here, we study a tri-trophic system of trees, phytophagous insects (caterpillars), and insectivorous birds (Great Tits) to assess how urbanization influences (1) the phenology of each component of this system, (2) insect abundance, and (3) avian reproductive success. We use data from two urban and two forest sites in Hungary, central Europe, collected over four consecutive years. Despite a trend of earlier leaf emergence in urban sites, there is no evidence for an earlier peak in caterpillar abundance. Thus, contrary to the frequently stated prediction in the literature, the earlier breeding of urban bird populations is not associated with an earlier peak in caterpillar availability. Despite this the seasonal dynamics of caterpillar biomass exhibited striking differences between habitat types with a single clear peak in forests, and several much smaller peaks in urban sites. Caterpillar biomass was higher in forests than urban areas across the entire sampling period, and between 8.5 and 24 times higher during the first brood's chick-rearing period. This higher biomass was not associated with taller trees in forest sites, or with tree species identity, and occurred despite most of our focal trees being native to the study area. Urban Great Tits laid smaller clutches, experienced more frequent nestling mortality from starvation, reared fewer offspring to fledging age, and their fledglings had lower body mass. Our study strongly indicates that food limitation is responsible for lower avian reproductive success in cities, which is driven by reduced availability of the preferred nestling diet, i.e., caterpillars, rather than phenological shifts in the timing of peak food availability.

Nightjars may adjust breeding phenology to compensate for mismatches between moths and moonlight.

Phenology match–mismatch usually refers to the extent of an organism's ability to match reproduction with peaks in food availability, but when mismatch occurs, it may indicate a response to another selective pressure. We assess the value of matching reproductive timing to multiple selective pressures for a migratory lunarphilic aerial insectivore bird, the whip‐poor‐will (Antrostomus vociferus). We hypothesize that a whip‐poor‐will's response to shifts in local phenology may be constrained by long annual migrations and a foraging mode that is dependent on both benign weather and the availability of moonlight. To test this, we monitored daily nest survival and overall reproductive success relative to food availability and moon phase in the northern part of whip‐poor‐will's breeding range. We found that moth abundance, and potentially temperature and moonlight, may all have a positive influence on daily chick survival rates and that the lowest chick survival rates for the period between hatching and fledging occurred when hatch was mismatched with both moths and moonlight. However, rather than breeding too late for peak moth abundance, the average first brood hatch date actually preceded the peak moth abundance and occurred during a period with slightly higher available moonlight than the period of peak food abundance. As a result, a low individual survival rate was partially compensated for by initiating more nesting attempts. This suggests that nightjars were able to adjust their breeding phenology in such a way that the costs of mismatch with food supply were at least partially balanced by a longer breeding season.

Breeding phenology and performance for four swallows over 57 years: relationships with temperature and precipitation

AbstractClimate change can drive population declines for many species, often through changes to their food supply. These changes can involve a mis‐timing between periods of high food demand and peak food availability, typically from advances in breeding phenology, and/or an overall reduction in food availability. Aerial insectivores, birds that feed on insects caught in flight, are experiencing steep population declines possibly because of shifts in the timing and/or abundance of aerial insects. We determined whether changes in breeding performance over time could account for declines in Bank Riparia riparia, Barn Hirundo rustica, Cliff Petrochelidon pyrrhonota, and Tree Tachycineta bicolor Swallows, and if so, whether changes were related to shifts in breeding phenology and/or climate change. We compared breeding performance and phenology in Maritime Canada before (1962–1972) and after (2006–2016) the onset of steep population declines during the mid‐1980s, to determine whether breeding performance was reduced or phenology was advanced. Then, we modeled relationships between temperature, precipitation, breeding phenology, and performance for Barn and Tree Swallows, the only species with sufficient data, from 1960 to 2016, to determine whether phenology and performance were related to climatic conditions. Between the two time periods, we found significantly lower performance in Bank Swallows, higher performance in Barn and Tree Swallows, and unchanged performance in Cliff Swallows. We also found clutch initiation dates advanced by 8–10 d for all species except Bank Swallows. On the breeding grounds, warmer winter temperatures for Tree Swallows and less winter precipitation for Barn and Tree Swallows in a given year were associated with earlier breeding, and for Tree Swallows, changes in nestling survival. Otherwise, Barn and Tree Swallow breeding performance was unaffected by winter temperature and precipitation. Our results suggest that in this region poorer breeding performance could contribute to population declines for Bank Swallows but not for the other three species.

Advances on the group composition, mating system, roosting and flight behaviour of the European free-tailed bat (Tadarida teniotis)

Abstract We investigated a large colony of European free-tailed bats (Tadarida teniotis) in Spain, using a combination of capture-mark-recapture data and direct observations. Its social and reproductive organisation is complex and the mating system fits a “resource defence polygyny” model. In spring and autumn, aggressive interactions in flight, defence of roosts and mating songs of males to attract females occurred. According to our results, T. teniotis is organised in “harems” consisting of a dominant male and a variable number of females. In addition the sexual cycle displayed a bimodal reproductive pattern (this is unique and remarkable for European bats). The bimodal pattern coincided with peaks in food availability (moths) at high altitudes. Presumably, roost-guarding activities (patrolling, advertising…) make males less prone to move away (e.g. at higher altitudes and longer distances) from roosts, resulting in differences in prey selection and in altitudinal segregation between sexes. This provides a plausible explanation for the differences in diet (predation of more sedentary vs. high-flying migratory moths) between males and females that has been found in published studies.