Rapid change in host use of the common cuckoo Cuculus canorus linked to climate change

Parasites require synchrony with their hosts so if host timing changes with climate change, some parasites may decline and eventually go extinct. Residents and short-distance migrant hosts of the brood parasitic common cuckoo, Cuculus canorus, have advanced their phenology in response to climate change more than long-distance migrants, including the cuckoo itself. Because different parts of Europe show different degrees of climate change, we predicted that use of residents or short-distance migrants as hosts should have declined in areas with greater increase in spring temperature. Comparing relative frequency of parasitism of the two host categories in 23 European countries before and after 1990, when spring temperatures in many areas had started to increase, we found that relative parasitism of residents and short-distance migrants decreased. This change in host use was positively related to increase in spring temperature, consistent with the prediction that relative change in phenology for different migrant classes drives host-use patterns. These findings are consistent with the hypothesis that climate change affects the relative abundance of different host races of the common cuckoo.     

Climate and the complexity of migratory phenology: sexes,migratory distance,and arrival distributions

The intra- and inter-season complexity of bird migration has received limited attention in climatic change research. Our phenological analysis of 22 species collected in Chicago, USA, (1979–2002) evaluates the relationship between multi-scalar climate variables and differences (1) in arrival timing between sexes, (2) in arrival distributions among species, and (3) between spring and fall migration. The early migratory period for earliest arriving species (i.e., short-distance migrants) and earliest arriving individuals of a species (i.e., males) most frequently correlate with climate variables. Compared to long-distance migrant species, four times as many short-distance migrants correlate with spring temperature, while 8 of 11 (73%) of long-distance migrant species’ arrival is correlated with the North Atlantic Oscillation (NAO). While migratory phenology has been correlated with NAO in Europe, we believe that this is the first documentation of a significant association in North America. Geographically proximate conditions apparently influence migratory timing for short-distance migrants while continental-scale climate (e.g., NAO) seemingly influences the phenology of Neotropical migrants. The preponderance of climate correlations is with the early migratory period, not the median of arrival, suggesting that early spring conditions constrain the onset or rate of migration for some species. The seasonal arrival distribution provides considerable information about migratory passage beyond what is apparent from statistical analyses of phenology. A relationship between climate and fall phenology is not detected at this location. Analysis of the within-season complexity of migration, including multiple metrics of arrival, is essential to detect species’ responses to changing climate as well as evaluate the underlying biological mechanisms.     

Avian population consequences of climate change are most severe for long-distance migrants in seasonal habitats

One consequence of climate change is an increasing mismatch between timing of food requirements and food availability. Such a mismatch is primarily expected in avian long-distance migrants because of their complex annual cycle, and in habitats with a seasonal food peak. Here we show that insectivorous long-distance migrant species in The Netherlands declined strongly (1984–2004) in forests, a habitat characterized by a short spring food peak, but that they did not decline in less seasonal marshes. Also, within generalist long-distance migrant species, populations declined more strongly in forests than in marshes. Forest-inhabiting migrant species arriving latest in spring declined most sharply, probably because their mismatch with the peak in food supply is greatest. Residents and short-distance migrants had non-declining populations in both habitats, suggesting that habitat quality did not deteriorate. Habitat-related differences in trends were most probably caused by climate change because at a European scale, long-distance migrants in forests declined more severely in western Europe, where springs have become considerably warmer, when compared with northern Europe, where temperatures during spring arrival and breeding have increased less. Our results suggest that trophic mismatches may have become a major cause for population declines in long-distance migrants in highly seasonal habitats.     

Climatic effects and phenological mismatch in cuckoo–host interactions: a role for host phenotypic plasticity in laying date?

Climatic effects on breeding phenology vary across organisms and therefore might promote a phenological mismatch in ecologically interacting species, including those engaged in coevolutionary interactions such as brood parasites and their hosts. Recent studies suggest that climatic induced changes in migration phenology may have mismatched cuckoos and their hosts in Europe. However, it is currently unknown whether cuckoo–host phenological mismatch results from different degrees of phenotypic plasticity or to different speeds of microevolutionary processes affecting hosts and parasites. Here we performed 1) cross‐sectional correlations between climate conditions and population level of phenological mismatch between the migratory brood parasite great spotted cuckoo Clamator glandarius and its main resident host in Europe, the magpie Pica pica; and 2) a longitudinal analysis to study within‐individual variation in breeding phenology for individual hosts experiencing different climate conditions over a period of nine years (2005–2013). Cross‐sectional analyses revealed independent and contrary effects of winter and spring temperature on magpie phenology: magpie hosts tend to breed earlier those years with lower February temperatures, however, high temperature in the first half of April spur individuals to lay eggs. Breeding phenology of cuckoos was tuned to that of their magpie host in time and duration. However, annual phenological mismatch between cuckoos and magpie hosts increased with NAO index and January temperature. Longitudinal analyses revealed high individual consistency in magpie host phenology, but a low influence of climate, suggesting that the climatic‐driven phenological mismatch between cuckoos and magpies at the population‐level cannot be explained by a host plastic response to climatic conditions.     

Winter rainfall predicts phenology in widely separated populations of a migrant songbird

Climate change is affecting behaviour and phenology in many animals. In migratory birds, weather patterns both at breeding and at non-breeding sites can influence the timing of spring migration and breeding. However, variation in responses to weather across a species range has rarely been studied, particularly among populations that may winter in different locations. We used prior knowledge of migratory connectivity to test the influence of weather from predicted non-breeding sites on bird phenology in two breeding populations of a long-distance migratory bird species separated by 3,000 km. We found that winter rainfall showed similar associations with arrival and egg-laying dates in separate breeding populations on an east–west axis: greater rainfall in Jamaica and eastern Mexico was generally associated with advanced American redstart (Setophaga ruticilla) phenology in Ontario and Alberta, respectively. In Ontario, these patterns of response could largely be explained by changes in the behaviour of individual birds, i.e., phenotypic plasticity. By explicitly incorporating migratory connectivity into responses to climate, our data suggest that widely separated breeding populations can show independent and geographically specific associations with changing weather conditions. The tendency of individuals to delay migration and breeding following dry winters could result in population declines due to predicted drying trends in tropical areas and the tight linkage between early arrival/breeding and reproductive success in long-distance migrants.     

Bird migration times, climate change, and changing population sizes

Past studies of bird migration times have shown great variation in migratory responses to climate change. We used 33 years of bird capture data (1970–2002) from Manomet, Massachusetts to examine variation in spring migration times for 32 species of North American passerines. We found that changes in first arrival dates – the unit of observation used in most studies of bird migration times – often differ dramatically from changes in the mean arrival date of the migration cohort as a whole. In our study, the earliest recorded springtime arrival date for each species occurred 0.20 days later each decade. In contrast, the mean arrival dates for birds of each species occurred 0.78 days earlier each decade. The difference in the two trends was largely explained by declining migration cohort sizes, a factor not examined in many previous studies. We found that changes in migration cohort or population sizes may account for a substantial amount of the variation in previously documented changes in migration times. After controlling for changes in migration cohort size, we found that climate variables, migration distance, and date of migration explained portions of the variation in migratory changes over time. In particular, short-distance migrants appeared to respond to changes in temperature, while mid-distance migrants responded particularly strongly to changes in the Southern Oscillation Index. The migration times of long-distance migrants tended not to change over time. Our findings suggest that previously reported changes in migration times may need to be reinterpreted to incorporate changes in migration cohort sizes.     

Advanced departure dates in long-distance migratory raptors

Evidences for phenological changes in response to climate change are now numerous. One of the most documented changes has been the advance of spring arrival dates in migratory birds. However, the effects of climate change on subsequent events of the annual cycle remain poorly studied and understood. Moreover, the rare studies on autumn migration have mainly concerned passerines. Here, we investigated whether raptor species have changed their autumn migratory phenology during the past 30 years at one of the most important convergent points of western European migration routes in France, the Organbidexka pass, in the Western Pyrenees. Eight out of the 14 studied raptor species showed significant phenological shifts during 1981–2008. Long-distance migrants displayed stronger phenological responses than short-distance migrants, and advanced their mean passage dates significantly. As only some short-distance migrants were found to delay their autumn migration and as their trends in breeding and migrating numbers were not significantly negative, we were not able to show any possible settling process of raptor populations. Negative trends in numbers of migrating raptors were found to be related to weaker phenological responses. Further studies using data from other migration sites are necessary to investigate eventual changes in migration routes and possible settling process.     

Divergent patterns of impact of environmental conditions on life history traits in two populations of a long-distance migratory bird

Some areas have experienced recent dramatic warming due to climate change, while others have shown no change at all, or even recent cooling. We predicted that patterns of selection on life history would differ between southern and northern European populations of a long-distance migratory bird, the barn swallow Hirundo rustica, because global patterns of weather as reflected by large-scale weather phenomena such as the North Atlantic Oscillation (NAO) and the El Niño-Southern Oscillation (ENSO) have different effects on environmental conditions in different parts of the world frequented during the annual cycle. We investigated relationships between mean arrival date, dispersal rate and yearling survival rate among years, using two long-term population studies in Spain and Denmark. We found evidence of a difference in the effects of normalized difference vegetation index in North and West Africa on mean arrival date of male barn swallows, with the effect differing significantly between populations. Second, there was a significant interaction between ENSO and population on dispersal rate, showing that conditions in Africa during winter differentially affected dispersal in the two populations. Finally, the NAO index in winter had an effect on yearling survival that differed between populations. These findings highlight the divergent patterns of response to climate change among populations, and they suggest that climate change can differentially affect important life history traits with potential implications for maintenance of viable populations and gene flow among populations.     

Climate warming,ecological mismatch at arrival and population decline in migratory birds

Climate is changing at a fast pace, causing widespread, profound consequences for living organisms. Failure to adjust the timing of life-cycle events to climate may jeopardize populations by causing ecological mismatches to the life cycle of other species and abiotic factors. Population declines of some migratory birds breeding in Europe have been suggested to depend on their inability to adjust migration phenology so as to keep track of advancement of spring events at their breeding grounds. In fact, several migrants have advanced their spring arrival date, but whether such advancement has been sufficient to compensate for temporal shift in spring phenophases or, conversely, birds have become ecologically mismatched, is still an unanswered question, with very few exceptions. We used a novel approach based on accumulated winter and spring temperatures (degree-days) as a proxy for timing of spring biological events to test if the progress of spring at arrival to the breeding areas by 117 European migratory bird species has changed over the past five decades. Migrants, and particularly those wintering in sub-Saharan Africa, now arrive at higher degree-days and may have therefore accumulated a ‘thermal delay’, thus possibly becoming increasingly mismatched to spring phenology. Species with greater ‘thermal delay’ have shown larger population decline, and this evidence was not confounded by concomitant ecological factors or by phylogenetic effects. These findings provide general support to the largely untested hypotheses that migratory birds are becoming ecologically mismatched and that failure to respond to climate change can have severe negative impacts on their populations. The novel approach we adopted can be extended to the analysis of ecological consequences of phenological response to climate change by other taxa.     

Spring arrival response to climate change in birds: a case study from eastern Europe

This paper analyses the dependence of the first spring arrival dates of short/medium- and long-distance migrant bird species on climate warming in eastern Europe. The timing of arrival of the selected species at the observation site correlates with the North Atlantic Oscillation (NAO) index, air temperature, atmospheric pressure, precipitation and wind characteristics. A positive correlation of fluctuations in winter and spring air temperatures with variations in the NAO index has been established in eastern Europe. Positive winter NAO index values are related to earlier spring arrival of birds in the eastern Baltic region and vice versa—arrival is late when the NAO index is negative. The impact of climate warming on the bird’s life cycle depends on local or regional climate characteristics. We tested the hypothesis that differences in climate indices between North Africa and Europe can influence the timing of spring arrival. Our results support the hypothesis that differences in first spring arrival dates between European populations occur after individuals cross the Sahara. We assume that the endogenous programme of migration control in short/medium-distance migrants synchronises with the changing environment on their wintering grounds and along their migration routes, whereas in long-distance migrants it is rather with environmental changes in the second part of their migratory route in Europe. Our results strongly indicate that the mechanism of dynamic balance in the interaction between the endogenous regulatory programme and environmental factors determines the pattern of spring arrival, as well as migration timing.     

Reduced compensatory growth capacity in mistimed broods of a migratory passerine

Phenotypic plasticity has recently been proposed to increase population viability when rapid anthropogenic environmental changes cannot be tracked by means of evolution. This assumes that environmental changes do not constrain phenotypic plasticity itself, which has rarely been examined in natural populations. In areas of climate warming, many long-distance migratory birds breed increasingly late relative to the period of peak food supply, and the temporal mismatch may constrain plastic life-history traits such as nestling growth. We combined 23 years of food availability and breeding data with a 3-year experimental manipulation of nestling growth trajectories in a Central European population of collared flycatchers (Ficedula albicollis) to examine the potential impact of climate-related mistimed breeding on nestling developmental plasticity. Timing of the food peak was predicted by winter climate, and the median hatching date of broods was earlier in springs with earlier food peaks. However, the adjustment of hatching date was incomplete and the population largely missed the food peak in years with very early food peaks. After imposing a temporary, experimental food shortage on nestlings, the extent of compensatory growth in body mass differed among years, and this difference was apparently related to the distance of hatching dates from the yearly food peak. Growth compensation declined with distance from the peak. These results suggest that mistimed phenology may not only create permanently adverse conditions for migratory species but it may also constrain the plastic responses of individuals to temporary disturbances. Therefore, climate change may not only favour but also restrict phenotypic plasticity.     

Predicting the effects of climate change on natural enemies of agricultural pests

Climate change can have diverse effects on natural enemies of pest species. Here we review these effects and their likely impacts on pest control. The fitness of natural enemies can be altered in response to changes in herbivore quality and size induced by temperature and CO₂ effects on plants. The susceptibility of herbivores to predation and parasitism could be decreased through the production of additional plant foliage or altered timing of herbivore life cycles in response to plant phenological changes. The effectiveness of natural enemies in controlling pests will decrease if pest distributions shift into regions outside the distribution of their natural enemies, although a new community of enemies might then provide some level of control. As well as being affected by climate through host plants and associated herbivores, the abundance and activity of natural enemies will be altered through adaptive management strategies adopted by farmers to cope with climate change. These strategies may lead to a mismatch between pests and enemies in space and time, decreasing their effectiveness for biocontrol. Because of the diverse and often indirect effects of climate change on natural enemies, predictions will be difficult unless there is a good understanding of the way environmental effects impact on tritrophic interactions. In addition, evolutionary changes in both hosts and natural enemies might have unexpected consequences on levels of biocontrol exerted by enemies. We consider interactions between the pest light brown apple moth and its natural enemies to illustrate the type of data that needs to be collected to make useful predictions.     

The disproportionate effect of global warming on the arrival dates of short-distance migratory birds in North America

Recent studies have shown that, in response to global climate change, diverse avian taxa are now nesting measurably earlier (< 10 days) in both the United States and Britain. Similarly, several studies on European birds have now demonstrated that a variety of species (although not all) are arriving increasingly early. However, surprisingly, widespread changes in North American migrant phenology have not been demonstrated. It is hypothesized that short-distance migrants (birds that winter in the southern United States) may be quicker to adapt to climate change than long-distance migrants (birds that winter south of the United States), as short-distance migrants can respond to meteorological cues indicating weather conditions to the north whereas long-distance migrants must rely on photoperiod. This study examined the first arrival dates of 103 migrant birds in New York and Massachusetts and found that, on average, all migrants arrived significantly earlier during the period 1951–1993 than the period 1903–1950. From 1951–1993 birds wintering in the southern United States arrived on average 13 days earlier while birds wintering in South America arrived 4 days earlier. Although a change in observer effort cannot be quantified and may be a source of bias, a comparison of the numbers of reporting observers during the 1930s and the 1980s revealed no significant difference. These results are consistent with those expected under a scenario of global warming.     

Delayed herbivory by migratory geese increases summer‐long CO2 uptake in coastal western Alaska

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to “phenological mismatches” as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three‐year experiment to examine the consequences for CO₂ exchange of advanced spring green‐up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green‐up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO₂ uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green‐up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m^?2 s^?1 toward a greater CO₂ sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO₂ uptake. Importantly, while the direct effect of climate warming on phenology of green‐up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.     

Advanced autumn migration of sparrowhawk has increased the predation risk of long-distance migrants in Finland

Predation affects life history traits of nearly all organisms and the population consequences of predator avoidance are often larger than predation itself. Climate change has been shown to cause phenological changes. These changes are not necessarily similar between species and may cause mismatches between prey and predator. Eurasian sparrowhawk Accipiter nisus, the main predator of passerines, has advanced its autumn phenology by about ten days in 30 years due to climate change. However, we do not know if sparrowhawk migrate earlier in response to earlier migration by its prey or if earlier sparrowhawk migration results in changes to predation risk on its prey. By using the median departure date of 41 passerine species I was able to show that early migrating passerines tend to advance, and late migrating species delay their departure, but none of the species have advanced their departure times as much as the sparrowhawk. This has lead to a situation of increased predation risk on early migrating long-distance migrants (LDM) and decreased the overlap of migration season with later departing short-distance migrants (SDM). Findings highlight the growing list of problems of declining LDM populations caused by climate change. On the other hand it seems that the autumn migration may become safer for SDM whose populations are growing. Results demonstrate that passerines show very conservative response in autumn phenology to climate change, and thus phenological mismatches caused by global warming are not necessarily increasing towards the higher trophic levels.     

Consequences of the genetic threshold model for observing partial migration under climate change scenarios

Migration is a widespread phenomenon across the animal kingdom as a response to seasonality in environmental conditions. Partially migratory populations are populations that consist of both migratory and residential individuals. Such populations are very common, yet their stability has long been debated. The inheritance of migratory activity is currently best described by the threshold model of quantitative genetics. The inclusion of such a genetic threshold model for migratory behavior leads to a stable zone in time and space of partially migratory populations under a wide range of demographic parameter values, when assuming stable environmental conditions and unlimited genetic diversity. Migratory species are expected to be particularly sensitive to global warming, as arrival at the breeding grounds might be increasingly mistimed as a result of the uncoupling of long‐used cues and actual environmental conditions, with decreasing reproduction as a consequence. Here, we investigate the consequences for migratory behavior and the stability of partially migratory populations under five climate change scenarios and the assumption of a genetic threshold value for migratory behavior in an individual‐based model. The results show a spatially and temporally stable zone of partially migratory populations after different lengths of time in all scenarios. In the scenarios in which the species expands its range from a particular set of starting populations, the genetic diversity and location at initialization determine the species’ colonization speed across the zone of partial migration and therefore across the entire landscape. Abruptly changing environmental conditions after model initialization never caused a qualitative change in phenotype distributions, or complete extinction. This suggests that climate change‐induced shifts in species’ ranges as well as changes in survival probabilities and reproductive success can be met with flexibility in migratory behavior at the species level, which will reduce the risk of extinction.     

Patterns of change in timing of spring migration in North European songbird populations

From 1976 to 1997 passerines were mist-netted and ringed on the island of Christiansø, in the Baltic Sea. Here we present analyses of phenological changes (i.e. time of arrival) for 25 species based on the entire populations of mist-netted songbirds during spring migration. We used two approaches (least square and quantile regression) to test for changes in arrival time of first individuals and three different parts of the songbird populations (i.e. first 5%, 50% and 95% of the total number of trapped individuals corrected for trapping effort). Our results generally confirm earlier spring arrival of migratory passerines with an overall earlier arrival of 0.26 days per year. Changes in the arrival time of first individuals are often the only data available. They are typically analysed on the assumption that they are representative of their respective population. We found a unidirectional, significant change towards earlier arrival for all four measures of arrival timing which seem to support this. However, the four measures of arrival are changing at different rates. First individuals changed arrival time more rapidly than the first 5%, 50% and 95% of the spring total. Such differences are likely to be important for our understanding of population-dynamic changes in relation to climate change. These differences may also have long-term evolutionary consequences. Migration distance seems to affect the degree of change in arrival time, but we found no difference between species wintering in different regions of Africa.     

Patterns of change in timing of spring migration in North European songbird populations

From 1976 to 1997 passerines were mist‐netted and ringed on the island of Christiansø, in the Baltic Sea. Here we present analyses of phenological changes (i.e. time of arrival) for 25 species based on the entire populations of mist‐netted songbirds during spring migration. We used two approaches (least square and quantile regression) to test for changes in arrival time of first individuals and three different parts of the songbird populations (i.e. first 5%, 50% and 95% of the total number of trapped individuals corrected for trapping effort). Our results generally confirm earlier spring arrival of migratory passerines with an overall earlier arrival of 0.26 days per year. Changes in the arrival time of first individuals are often the only data available. They are typically analysed on the assumption that they are representative of their respective population. We found a unidirectional, significant change towards earlier arrival for all four measures of arrival timing which seem to support this. However, the four measures of arrival are changing at different rates. First individuals changed arrival time more rapidly than the first 5%, 50% and 95% of the spring total. Such differences are likely to be important for our understanding of population‐dynamic changes in relation to climate change. These differences may also have long‐term evolutionary consequences. Migration distance seems to affect the degree of change in arrival time, but we found no difference between species wintering in different regions of Africa.     

Differences in timing of migration and response to sexual signalling drive asymmetric hybridization across a migratory divide

Ecological traits and sexual signals may both contribute to the process of ecological speciation. Here we investigate the roles of an ecological trait, seasonal migratory behaviour and a sexual trait, song, in restricting or directing gene flow across a migratory divide in the Swainson's thrush (Catharus ustulatus). We show that short-distance migratory ecotypes wintering in Central America arrive earlier at the breeding grounds than long-distance migratory ecotypes wintering primarily in South America, providing the potential for some premating isolation. Playback experiments suggest that early- and late-arriving forms recognize each other as competitors, but that the early-arriving form responds more aggressively to a broader spectrum of stimuli. Genetic analysis suggests that hybridization occurs more often between males of the early-arriving ecotype and females of the late-arriving ecotype. Together our results suggest that differences in arrival times may reduce the temporal coincidence of mate choice, but asymmetry in response to heterotypic song may hinder complete divergence. These data provide further insight into the roles of ecological traits and sexual signals during the incipient stages of speciation.     

Quantitative genetics of migration syndromes: a study of two barn swallow populations

Migration is a complex trait although little is known about genetic correlations between traits involved in such migration syndromes. To assess the migratory responses to climate change, we need information on genetic constraints on evolutionary potential of arrival dates in migratory birds. Using two long-term data sets on barn swallows Hirundo rustica (from Spain and Denmark), we show for the first time in wild populations that spring arrival dates are phenotypically and genetically correlated with morphological and life history traits. In the Danish population, length of outermost tail feathers and wing length were negatively genetically correlated with arrival date. In the Spanish population, we found a negative genetic correlation between arrival date and time elapsed between arrival date and laying date, constraining response to selection that favours both early arrival and shorter delays. This results in a decreased rate of adaptation, not because of constraints on arrival date, but constraints on delay before breeding, that is, a trait that can be equally important in the context of climate change.