Climate change and the optimal arrival of migratory birds

Recent climate change has sparked an interest in the timing of biological events, which is a general problem in life-history evolution. Reproduction in many organisms breeding in seasonal environments, e.g. migratory birds, is dependent on the exploitation of a short but rich food supply. If the seasonal timing of the food peak advances owing to climate change, then one would expect the bird to track those changes, hence, initiate migration and breeding earlier. However, when there is competition for territories and a risk of pre-breeding mortality, the optimal response to a shifting food distribution is no longer obvious. We develop a theoretical model to study how the optimal arrival time depends on the mean and variance of the food distribution, the degree of competition for territories and the risk of mortality. In general, the optimal shift in arrival date should never be as extreme as the shift in food peak date. Our results also show that we should expect the high variation of trends in arrival date observed among migratory birds, even if migration and information about climate change were unconstrained.     

Climate change can alter competitive relationships between resident and migratory birds

Climate change could affect resource competition between resident and migratory bird species by changing the interval between their onsets of breeding or by altering their population densities. We studied interspecific nest-hole competition between resident great tits and migrant pied flycatchers in South-Western Finland over the past five decades (1953-2005). We found that appearance of fatal take-over trials, the cases where a pied flycatcher tried to take over a great tit nest but was killed by the tit, increased with a reduced interspecific laying date interval and with increasing densities of both tits and flycatchers. The probability of pied flycatchers taking over great tit nests increased with the density of pied flycatchers. Laying dates of the great tit and pied flycatcher are affected by the temperatures of different time periods, and divergent changes in these temperatures could consequently modify their competitive interactions. Densities are a result of reproductive success and survival, which can be affected by separate climatic factors in the resident great tit and trans-Saharan migrant pied flycatcher. On these bases we conclude that climate change has a great potential to alter the competitive balance between these two species.     

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.     

Estimating partial observability and nonlinear climate effects on stochastic community dynamics of migratory waterfowl

1. Understanding the impact of environmental variability on migrating species requires the estimation of sequential abiotic effects in different geographic areas across the life cycle. For instance, waterfowl (ducks, geese and swans) usually breed widely dispersed throughout their breeding range and gather in large numbers in their wintering headquarters, but there is a lack of knowledge on the effects of the sequential environmental conditions experienced by migrating birds on the long-term community dynamics at their wintering sites. 2. Here, we analyse multidecadal time-series data of 10 waterfowl species wintering in the Guadalquivir Marshes (SW Spain), the single most important wintering site for waterfowl breeding in Europe. We use a multivariate state-space approach to estimate the effects of biotic interactions, local environmental forcing during winter and large-scale climate during breeding and migration on wintering multispecies abundance fluctuations, while accounting for partial observability (observation error and missing data) in both population and environmental data. 3. The joint effect of local weather and large-scale climate explained 31·6% of variance at the community level, while the variability explained by interspecific interactions was negligible (<5%). In general, abiotic conditions during winter prevailed over conditions experienced during breeding and migration. Across species, a pervasive and coherent nonlinear signal of environmental variability on population dynamics suggests weaker forcing at extreme values of abiotic variables. 4. Modelling missing observations through data augmentation increased the estimated magnitude of environmental forcing by an average 30·1% and reduced the impact of stochasticity by 39·3% when accounting for observation error. Interestingly however, the impact of environmental forcing on community dynamics was underestimated by an average 15·3% and environmental stochasticity overestimated by 14·1% when ignoring both observation error and data augmentation. 5. These results provide a salient example of sequential multiscale environmental forcing in a major migratory bird community, which suggests a demographic link between the breeding and wintering grounds operating through nonlinear environmental effects. Remarkably, this study highlights that modelling observation error in the environmental covariates of an ecological model can be proportionally more important than modelling this source of variance in the population data.     

Examining the total arrival distribution of migratory birds

This paper reports on the total distribution of spring migration timing of willow warbler, chiffchaff and pied flycatcher at locations in the UK, Germany, Russia and Finland. This is the first time that high‐quality data based on known‐effort monitoring has been examined on a continental scale. First arrival dates, commonly reported in the literature, were positively correlated with mean arrival dates although they would not make good predictors of the latter. At all locations, at least one aspect of the arrival distribution of each species had got significantly earlier in recent years. The trend towards earliness was associated with warmer local temperatures and more positive winter North Atlantic Oscillation index. In years that were early, the arrival distribution became more elongated and skewed. Researchers should now investigate the consequences of earlier arrival on current and future bird populations.     

Radionuclide transfer by migratory birds]

Evaluation of the zoogenic transfer of radionuclides from the 30-km zone around the Chernobyl NPP was necessary because of the enormous heavily polluted territory and mighty flow of migratory birds who tended to large rivers, the Dnieper and Pripyat. The integral estimate of the transferred amount was obtained as a product of three variables: the transfer factor (0.0077 m2/kg for 137Cs; 0.0107 m2/kg for 90Sr), the density of birds (0.002 kg/m2, at the mass of migrants about 5000 t per year), and the total fund of radionuclides throughout the territory. The upper estimated limit of the annual transfer rate was 5.5 Ci (2E + 11 Bq) for 137 Cs and 1.8 Ci (6.7E + 10 Bq) for 90 Sr. Restrictions of hunting are recommended within the northern part of the Kiev reservoir.     

Optimal moult strategies in migratory birds

Avian migration, which involves billions of birds flying vast distances, is known to influence all aspects of avian life. Here we investigate how birds fit moult into an annual cycle determined by the need to migrate. Large variation exists in moulting patterns in relation to migration: for instance, moult can occur after breeding in the summer or after arrival in the wintering quarters. Here we use an optimal annual routine model to investigate why this variation exists. The modelled bird's decisions depend on the time of year, its energy reserves, breeding status, experience, flight feather quality and location. Our results suggest that the temporal and spatial variations in food are an important influence on a migratory bird's annual cycle. Summer moult occurs when food has a high peak on the breeding site in the summer, but it is less seasonal elsewhere. Winter moult occurs if there is a short period of high food availability in summer and a strong winter peak at different locations (i.e. the food is very seasonal but in opposite phase on these areas). This finding might explain why only long-distance migrants have a winter moult.     

The scheduling of molt in migratory birds

Summary We model the yearly cycle of small migratory birds to explain the variation in scheduling of complete molt, in particular why some birds molt immediately after breeding on the breeding grounds (summer molt) whereas others migrate to their wintering grounds before molt is initiated (winter molt). We employ the method of dynamic programming, because of its suitability for modelling life history traits. Feather quality and latitude entered the model as state variables and were assumed to affect survival rate and reproductive success. Migration and molt were assumed to be associated with increased mortality risks. By changing the parameters in the model we were able to generate most existing molt patterns, including summer and winter molt, biannual (summer and winter) molt, and molt migration. Our model suggests that the scheduling of molt is basically a result of a trade-off between having a high feather quality during breeding versus during the non-breeding period. A high impact of feather quality on survival rate in combination with low costs of molt resulted in biannual molt. Winter molt became more likely as the survival rateper se increased. A low seasonal amplitude in survival rate is a prerequisite for the occurrence of molt migration. Molt duration, migration costs and reproductive successper se were found to have no impact on the timing of molt. We also investigated the effect of benefits from prior occupancy at breeding and winter grounds.     

Orientation of migratory birds under ultraviolet light

In view of the finding that cryptochrome 1a, the putative receptor molecule for the avian magnetic compass, is restricted to the ultraviolet single cones in European Robins, we studied the orientation behaviour of robins and Australian Silvereyes under monochromatic ultraviolet (UV) light. At low intensity UV light of 0.3 mW/m², birds showed normal migratory orientation by their inclination compass, with the directional information originating in radical pair processes in the eye. At 2.8 mW/m², robins showed an axial preference in the east–west axis, whereas silvereyes preferred an easterly direction. At 5.7 mW/m², robins changed direction to a north–south axis. When UV light was combined with yellow light, robins showed easterly ‘fixed direction’ responses, which changed to disorientation when their upper beak was locally anaesthetised with xylocaine, indicating that they were controlled by the magnetite-based receptors in the beak. Orientation under UV light thus appears to be similar to that observed under blue, turquoise and green light, albeit the UV responses occur at lower light levels, probably because of the greater light sensitivity of the UV cones. The orientation under UV light and green light suggests that at least at the level of the retina, magnetoreception and vision are largely independent of each other.     

Protandry and sexual dimorphism in trans-Saharan migratory birds

Earlier arrival to reproductive sites of males relative to females(protandry) is widespread among migratory organisms. Diversemechanisms have been proposed that may select for protandry,including competition for limiting resources (e.g., territories)or mates. In species with large variation in male reproductivesuccess, such as polygamous species and those with intense spermcompetition, early arriving males may accrue a fitness advantagebecause they acquire more mates or have larger chances of paternity.Comparative studies of birds have shown that sexual size dimorphism(SSD) is positively associated with the level of polygyny, whereasintense sperm competition is associated with sexual dichromatism(SD). Positive correlations between protandry and SSD or SDcan therefore be expected to exist across avian species. Becauselarge males are predicted to be better able to cope with adverseecological conditions early in the breeding season, selectionfor protandry, in turn, may have a correlated response on SSDamong migratory species breeding in boreal latitudes. Althoughprevious studies of birds have analyzed the association betweenSSD and protandry, none has analyzed SD in relation to protandry.Here we analyze the association between protandry during springmigration, SSD, and SD in 21 trans-Saharan monogamous migratorybird species. The difference in median migration dates betweenfemales and males, reflecting protandry, was positively associatedwith SD but not with SSD. Because dichromatism is positivelyrelated to sperm competition across species, present resultsare consistent with predictions derived from sexual selectionhypotheses for the evolution of protandry mediated by spermcompetition.     

Genetic control of migratory behaviour in birds

Although it has long been suspected that biannual migration in birds has a direct genetic basis, only in the last decade have details of the inheritance of behavioural traits such as migratory activity and directional preferences been demonstrated. A model has now been developed to estimate how inexperienced first-time migrants manage to reach their unknown winter quarters on the basis of inherited spatio-temporal programs. Furthermore, in obligate partial migrants the decision to migrate or not has been shown to have a strong genetic base. Migratoriness and sedentariness in partial migrants have been shown to have a high potential for rapid evolution. A recent set of results has suggested that novel migratory habits can evolve in less than 25 years. A possible consequence is that environmental changes, including 'greenhouse' effects, might considerably alter avian migration systems by acting on genetic variation for migratory tendencies.     

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.     

Macroecology of habitat choice in long-distance migratory birds

Patterns of habitat choice in ecological communities are not only influenced by present-day selective forces but also by historical processes, such as the biogeographical history of the lineages they are composed of. Nevertheless, it has been very difficult to test historical factors. The possible tropical origin of long-distance migratory birds provides an opportunity for such a test. If habitat choice of long-distance migrants is inherited from their tropical ancestors then Nearctic long-distance migrants might have acquired their habitat choice from Neotropical forest species and European long-distance migrants from African savannah species. Here we use a macroecological approach to show that this hypothesis can be confirmed. Long-distance migrants in the Nearctic are found in forested habitat types, while those in Europe are found in open ones. In comparison, the habitat choice of residents and short-distance migrants (in genera without long-distance migration) does not differ between the Nearctic and Europe. These results demonstrate that habitat choice in temperate bird communities can be explained by the tropical history of long-distance migrants. Thus, habitat choice seems to be shaped not only by local mechanisms, but also by processes acting on much larger spatial and temporal scales.     

Identifying type I excitability using dynamics of stochastic neural firing patterns

The stochastic firing patterns are simulated near saddle-node bifurcation on an invariant cycle corresponding to type I excitability in stochastic Morris–Lecar model. In absence of external periodic signal, the stochastic firing manifests continuous distribution in ISI histogram (ISIH), whose amplitude at first increases sharply and then decreases exponentially. In presence of the external periodic signal, stochastic firing patterns appear as two cases of integer multiple firing with multiple discrete peaks in ISIH. One manifests perfect exponential decay in all peaks and the other imperfect exponential decay except a lower first peak. These stochastic firing patterns simulated with or without external periodic signal can be demonstrated in the experiments on rat hippocampal CA1 pyramidal neurons. The exponential decay laws in the multiple peaks are also acquired using probability analysis method. The perfect decay law is determined by the independent characteristic within the firing while the imperfect decay law is from the inhibitory effect. In addition, the stochastic firing patterns corresponding to type I excitability are compared to those of type II excitability. The results not only reveal the dynamics of stochastic firing patterns with or without external signal corresponding to type I excitability, but also provide practical indicators to availably identify type I excitability.     

Zugunruhe of migratory and non-migratory birds in a circannual context

Recent findings in non-migratory birds have reopened questions about the interpretation and seasonal organization of Zugunruhe . I address the relationship between Zugunruhe and migration by comparing underlying circannual patterns in captive populations of migratory and non-migratory stonechats. Zugunruhe was highly variable and lacked clear periodicity, indicating its sensitivity to external cues. Patterns of Zugunruhe were similar in African residents, European short-distance migrants, and Siberian long-distance migrants, revealing no major difference in circannual organization. Moult was regulated independently of Zugunruhe and timed more rigidly, particularly in stonechats from equatorial Africa. The persistent and variable circannual patterns of Zugunruhe suggest that non-migratory and migratory stonechats have similar underlying programs but have modified the expression of actual migration. The findings, together with published observations from other species, emphasize the importance of considering programs for migration in a wide range of species, without losing sight of its environmental context.     

Population dynamics in migratory networks

Migratory animals are comprised of a complex series of interconnected breeding and nonbreeding populations. Because individuals in any given population can arrive from a variety of sites the previous season, predicting how different populations will respond to environmental change can be challenging. In this study, we develop a population model composed of a network of breeding and wintering sites to show how habitat loss affects patterns of connectivity and species abundance. When the costs of migration are evenly distributed, habitat loss at a single site can increase the degree of connectivity (mixing) within the entire network, which then acts to buffer global populations from declines. However, the degree to which populations are buffered depends on where habitat loss occurs within the network: a site that has the potential to receive individuals from multiple populations in the opposite season will lead to smaller declines than a site that is more isolated. In other cases when there are equal costs of migration to two or more sites in the opposite season, habitat loss can result in some populations becoming segregated (disconnected) from the rest of the network. The geographic structure of the network can have a significant influence on relative population sizes of sites in the same season and can also affect the overall degree of mixing in the network, even when sites are of equal intrinsic quality. When a migratory network is widely spaced and migration costs are high, an equivalent habitat loss will lead to a larger decline in global population size than will occur in a network where the overall costs of migration are low. Our model provides an important foundation to test predictions related to habitat loss in real-world migratory networks and demonstrates that migratory networks will likely produce different dynamics from traditional metapopulations. Our results provide strong evidence that estimating population connectivity is a prerequisite for successfully predicting changes in migratory populations.     

Passive internal dispersal of insect larvae by migratory birds

It has long been assumed that the resistant eggs of many zooplankton are able to survive passage through the gut of migratory waterbirds, thus facilitating their dispersal between isolated aquatic habitats. We present the first evidence that such passive internal transport within birds may be relevant for insect populations. In three out of six faecal samples from black-tailed Godwits on autumn migration in southwest Spain, we found larvae of the chironomid Chironomus salinarius which had survived gut passage. Although adult chironomids can fly, they are likely to disperse greater distances when transported as larvae via birds. In insects with discrete generations, such passive transport also enables colonization of new habitats at times when flight by adults is not an option.     

Influenza in migratory birds and evidence of limited intercontinental virus exchange

Migratory waterfowl of the world are the natural reservoirs of influenza viruses of all known subtypes. However, it is unknown whether these waterfowl perpetuate highly pathogenic (HP) H5 and H7 avian influenza viruses. Here we report influenza virus surveillance from 2001 to 2006 in wild ducks in Alberta, Canada, and in shorebirds and gulls at Delaware Bay (New Jersey), United States, and examine the frequency of exchange of influenza viruses between the Eurasian and American virus clades, or superfamilies. Influenza viruses belonging to each of the subtypes H1 through H13 and N1 through N9 were detected in these waterfowl, but H14 and H15 were not found. Viruses of the HP Asian H5N1 subtypes were not detected, and serologic studies in adult mallard ducks provided no evidence of their circulation. The recently described H16 subtype of influenza viruses was detected in American shorebirds and gulls but not in ducks. We also found an unusual cluster of H7N3 influenza viruses in shorebirds and gulls that was able to replicate well in chickens and kill chicken embryos. Genetic analysis of 6,767 avian influenza gene segments and 248 complete avian influenza viruses supported the notion that the exchange of entire influenza viruses between the Eurasian and American clades does not occur frequently. Overall, the available evidence does not support the perpetuation of HP H5N1 influenza in migratory birds and suggests that the introduction of HP Asian H5N1 to the Americas by migratory birds is likely to be a rare event.