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1.  Birds are tracking climate warming, but not fast enough 
Range shifts of many species are now documented as a response to global warming. But whether these observed changes are occurring fast enough remains uncertain and hardly quantifiable. Here, we developed a simple framework to measure change in community composition in response to climate warming. This framework is based on a community temperature index (CTI) that directly reflects, for a given species assemblage, the balance between low- and high-temperature dwelling species. Using data from the French breeding bird survey, we first found a strong increase in CTI over the last two decades revealing that birds are rapidly tracking climate warming. This increase corresponds to a 91 km northward shift in bird community composition, which is much higher than previous estimates based on changes in species range edges. During the same period, temperature increase corresponds to a 273 km northward shift in temperature. Change in community composition was thus insufficient to keep up with temperature increase: birds are lagging approximately 182 km behind climate warming. Our method is applicable to any taxa with large-scale survey data, using either abundance or occurrence data. This approach can be further used to test whether different delays are found across groups or in different land-use contexts.
PMCID: PMC2605823  PMID: 18713715
birds; breeding bird survey; climate warming; community composition; global changes; range edges
2.  Large-Scale Changes in Community Composition: Determining Land Use and Climate Change Signals 
PLoS ONE  2012;7(4):e35272.
Human land use and climate change are regarded as the main driving forces of present-day and future species extinction. They may potentially lead to a profound reorganisation of the composition and structure of natural communities throughout the world. However, studies that explicitly investigate both forms of impact—land use and climate change—are uncommon. Here, we quantify community change of Dutch breeding bird communities over the past 25 years using time lag analysis. We evaluate the chronological sequence of the community temperature index (CTI) which reflects community response to temperature increase (increasing CTI indicates an increase in relative abundance of more southerly species), and the temporal trend of the community specialisation index (CSI) which reflects community response to land use change (declining CSI indicates an increase of generalist species). We show that the breeding bird fauna underwent distinct directional change accompanied by significant changes both in CTI and CSI which suggests a causal connection between climate and land use change and bird community change. The assemblages of particular breeding habitats neither changed at the same speed and nor were they equally affected by climate versus land use changes. In the rapidly changing farmland community, CTI and CSI both declined slightly. In contrast, CTI increased in the more slowly changing forest and heath communities, while CSI remained stable. Coastal assemblages experienced both an increase in CTI and a decline in CSI. Wetland birds experienced the fastest community change of all breeding habitat assemblages but neither CTI nor CSI showed a significant trend. Overall, our results suggest that the interaction between climate and land use changes differs between habitats, and that comparing trends in CSI and CTI may be useful in tracking the impact of each determinant.
PMCID: PMC3327650  PMID: 22523579
3.  Photoperiodic response may facilitate adaptation to climatic change in long-distance migratory birds. 
Recent climatic change is causing spring events in northern temperate regions to occur earlier in the year. As a result, migratory birds returning from tropical wintering sites may arrive too late to take full advantage of the food resources on their breeding grounds. Under these conditions, selection will favour earlier spring arrival that could be achieved by overwintering closer to the breeding grounds. However, it is unknown how daylength conditions at higher latitudes will affect the timing of life cycle stages. Here, we show in three species of Palaearctic-African migratory songbirds that a shortening of migration distance induces an advancement of springtime activities. Birds exposed to daylengths simulating migration to and wintering in southern Europe considerably advanced their spring migratory activity and testicular development. This response to the novel photoperiodic environment will enable birds wintering further north to advance spring arrival and to start breeding earlier. Thus, phenotypic flexibility in response to the photoperiod may reinforce selection for shorter migration distance if spring temperatures continue to rise.
PMCID: PMC1698020  PMID: 12952632
4.  Regional climatic warming drives long-term community changes of British marine fish. 
Climatic change has been implicated as the cause of abundance fluctuations in marine fish populations worldwide, but the effects on whole communities are poorly understood. We examined the effects of regional climatic change on two fish assemblages using independent datasets from inshore marine (English Channel, 1913-2002) and estuarine environments (Bristol Channel, 1981-2001). Our results show that climatic change has had dramatic effects on community composition. Each assemblage contained a subset of dominant species whose abundances were strongly linked to annual mean sea-surface temperature. Species' latitudinal ranges were not good predictors of species-level responses, however, and the same species did not show congruent trends between sites. This suggests that within a region, populations of the same species may respond differently to climatic change, possibly owing to additional local environmental determinants, interspecific ecological interactions and dispersal capacity. This will make species-level responses difficult to predict within geographically differentiated communities.
PMCID: PMC1691641  PMID: 15156925
5.  Climate change and the demographic demise of a hoarding bird living on the edge 
Population declines along the lower-latitude edge of a species' range may be diagnostic of climate change. We report evidence that climate change has contributed to deteriorating reproductive success in a rapidly declining population of the grey jay (Perisoreus canadensis) at the southern edge of its range. This non-migratory bird of boreal and subalpine forest lives on permanent territories, where it hoards enormous amounts of food for winter and then breeds very early, under still-wintry conditions. We hypothesized that warmer autumns have increased the perishability of hoards and compromised subsequent breeding attempts. Our analysis confirmed that warm autumns, especially when followed by cold late winters, have led to delayed breeding and reduced reproductive success. Our findings uniquely show that weather months before the breeding season impact the timing and success of breeding. Warm autumns apparently represent hostile conditions for this species, because it relies on cold storage. Our study population may be especially vulnerable, because it is situated at the southern edge of the range, where the potential for hoard rot is most pronounced. This population's demise may signal a climate-driven range contraction through local extinctions along the trailing edge.
PMCID: PMC1664634  PMID: 17015367
climate change; demography; grey jay; Perisoreus canadensis; hoarding
6.  Re-Shuffling of Species with Climate Disruption: A No-Analog Future for California Birds? 
PLoS ONE  2009;4(9):e6825.
By facilitating independent shifts in species' distributions, climate disruption may result in the rapid development of novel species assemblages that challenge the capacity of species to co-exist and adapt. We used a multivariate approach borrowed from paleoecology to quantify the potential change in California terrestrial breeding bird communities based on current and future species-distribution models for 60 focal species. Projections of future no-analog communities based on two climate models and two species-distribution-model algorithms indicate that by 2070 over half of California could be occupied by novel assemblages of bird species, implying the potential for dramatic community reshuffling and altered patterns of species interactions. The expected percentage of no-analog bird communities was dependent on the community scale examined, but consistent geographic patterns indicated several locations that are particularly likely to host novel bird communities in the future. These no-analog areas did not always coincide with areas of greatest projected species turnover. Efforts to conserve and manage biodiversity could be substantially improved by considering not just future changes in the distribution of individual species, but including the potential for unprecedented changes in community composition and unanticipated consequences of novel species assemblages.
PMCID: PMC2730567  PMID: 19724641
7.  Avian migrants adjust migration in response to environmental conditions en route 
Biology Letters  2008;4(6):685-688.
The onset of migration in birds is assumed to be primarily under endogenous control in long-distance migrants. Recently, climate changes appear to have been driving a rapid change in breeding area arrival. However, little is known about the climatic factors affecting migratory birds during the migration cycle, or whether recently reported phenological changes are caused by plastic behavioural responses or evolutionary change. Here, we investigate how environmental conditions in the wintering areas as well as en route towards breeding areas affect timing of migration. Using data from 1984 to 2004 covering the entire migration period every year from observatories located in the Middle East and northern Europe, we show that passage of the Sahara Desert is delayed and correlated with improved conditions in the wintering areas. By contrast, migrants travel more rapidly through Europe, and adjust their breeding area arrival time in response to improved environmental conditions en route. Previous studies have reported opposing results from a different migration route through the Mediterranean region (Italy). We argue that the simplest explanation for different phenological patterns at different latitudes and between migratory routes appears to be phenotypic responses to spatial variability in conditions en route.
PMCID: PMC2614143  PMID: 18700199
birds; migration; phenology; climate change; normalized difference vegetation index
8.  Gray Wolves as Climate Change Buffers in Yellowstone 
PLoS Biology  2005;3(4):e92.
Understanding the mechanisms by which climate and predation patterns by top predators co-vary to affect community structure accrues added importance as humans exert growing influence over both climate and regional predator assemblages. In Yellowstone National Park, winter conditions and reintroduced gray wolves (Canis lupus) together determine the availability of winter carrion on which numerous scavenger species depend for survival and reproduction. As climate changes in Yellowstone, therefore, scavenger species may experience a dramatic reshuffling of food resources. As such, we analyzed 55 y of weather data from Yellowstone in order to determine trends in winter conditions. We found that winters are getting shorter, as measured by the number of days with snow on the ground, due to decreased snowfall and increased number of days with temperatures above freezing. To investigate synergistic effects of human and climatic alterations of species interactions, we used an empirically derived model to show that in the absence of wolves, early snow thaw leads to a substantial reduction in late-winter carrion, causing potential food bottlenecks for scavengers. In addition, by narrowing the window of time over which carrion is available and thereby creating a resource pulse, climate change likely favors scavengers that can quickly track food sources over great distances. Wolves, however, largely mitigate late-winter reduction in carrion due to earlier snow thaws. By buffering the effects of climate change on carrion availability, wolves allow scavengers to adapt to a changing environment over a longer time scale more commensurate with natural processes. This study illustrates the importance of restoring and maintaining intact food chains in the face of large-scale environmental perturbations such as climate change.
Reintroducing wolves can help ameliorate the negative effects of warmer winters on other species and reveals the importance of maintaining intact food chains in the face of climate change
PMCID: PMC1064850  PMID: 15757363
9.  Non-breeding habitat preference affects ecological speciation in migratory waders 
Die Naturwissenschaften  2007;95(4):347-354.
Models of ecological speciation predict that certain types of habitat should be more conducive to species diversification than others. In this study, I test this hypothesis in waders of the sub-order Charadrii using the number of morphological sub-species per species as an index of diversity. I classified all members of this clade as spending the non-breeding season either coastally or inland and argue that these represent fundamentally different environments. Coastal mudflats are characterised by high predictability and patchy worldwide distribution, whilst inland wetlands are widespread but unpredictable. The results show that migratory species that winter coastally are sub-divided into more sub-species than those that winter inland. This was not the case for non-migratory species. I argue that coastal environments select for more rigid migratory pathways, whilst inland wetlands favour more flexible movement patterns. Population sub-division could then result from the passive segregation of breeding sites or from the active selection for assortative mating of ecomorphs.
PMCID: PMC2270370  PMID: 18087687
Speciation; Waders; Non-breeding habitat; Sub-species
10.  Demersal Fish Assemblages and Spatial Diversity Patterns in the Arctic-Atlantic Transition Zone in the Barents Sea 
PLoS ONE  2012;7(4):e34924.
Direct and indirect effects of global warming are expected to be pronounced and fast in the Arctic, impacting terrestrial, freshwater and marine ecosystems. The Barents Sea is a high latitude shelf Sea and a boundary area between arctic and boreal faunas. These faunas are likely to respond differently to changes in climate. In addition, the Barents Sea is highly impacted by fisheries and other human activities. This strong human presence places great demands on scientific investigation and advisory capacity. In order to identify basic community structures against which future climate related or other human induced changes could be evaluated, we analyzed species composition and diversity of demersal fish in the Barents Sea. We found six main assemblages that were separated along depth and temperature gradients. There are indications that climate driven changes have already taken place, since boreal species were found in large parts of the Barents Sea shelf, including also the northern Arctic area. When modelling diversity as a function of depth and temperature, we found that two of the assemblages in the eastern Barents Sea showed lower diversity than expected from their depth and temperature. This is probably caused by low habitat complexity and the distance to the pool of boreal species in the western Barents Sea. In contrast coastal assemblages in south western Barents Sea and along Novaya Zemlya archipelago in the Eastern Barents Sea can be described as diversity “hotspots”; the South-western area had high density of species, abundance and biomass, and here some species have their northern distribution limit, whereas the Novaya Zemlya area has unique fauna of Arctic, coastal demersal fish. (see Information S1 for abstract in Russian).
PMCID: PMC3328492  PMID: 22545093
11.  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.
PMCID: PMC3049050  PMID: 20861045
climate change; conservation; ecological mismatch; migration; population trend
12.  Future Climate Change Will Favour Non-Specialist Mammals in the (Sub)Arctics 
PLoS ONE  2012;7(12):e52574.
Arctic and subarctic (i.e., [sub]arctic) ecosystems are predicted to be particularly susceptible to climate change. The area of tundra is expected to decrease and temperate climates will extend further north, affecting species inhabiting northern environments. Consequently, species at high latitudes should be especially susceptible to climate change, likely experiencing significant range contractions. Contrary to these expectations, our modelling of species distributions suggests that predicted climate change up to 2080 will favour most mammals presently inhabiting (sub)arctic Europe. Assuming full dispersal ability, most species will benefit from climate change, except for a few cold-climate specialists. However, most resident species will contract their ranges if they are not able to track their climatic niches, but no species is predicted to go extinct. If climate would change far beyond current predictions, however, species might disappear. The reason for the relative stability of mammalian presence might be that arctic regions have experienced large climatic shifts in the past, filtering out sensitive and range-restricted taxa. We also provide evidence that for most (sub)arctic mammals it is not climate change per se that will threaten them, but possible constraints on their dispersal ability and changes in community composition. Such impacts of future changes in species communities should receive more attention in literature.
PMCID: PMC3527567  PMID: 23285098
13.  Disparities between observed and predicted impacts of climate change on winter bird assemblages 
Understanding how climate change affects the structure and function of communities is critical for gauging its full impact on biodiversity. To date, community-level changes have been poorly documented, owing, in part, to the paucity of long-term datasets. To circumvent this, the use of ‘space-for-time’ substitution—the forecasting of temporal trends from spatial climatic gradients—has increasingly been adopted, often with little empirical support. Here we examine changes from 1975 to 2001 in three community attributes (species richness, body mass and occupancy) for 404 assemblages of terrestrial winter avifauna in North America containing a total of 227 species. We examine the accuracy of space-for-time substitution and assess causal associations between community attributes and observed changes in annual temperature using a longitudinal study design. Annual temperature and all three community attributes increased over time. The trends for the three community attributes differed significantly from the spatially derived predictions, although richness showed broad congruence. Correlations with trends in temperature were found with richness and body mass. In the face of rapid climate change, applying space-for-time substitution as a predictive tool could be problematic with communities developing patterns not reflected by spatial ecological associations.
PMCID: PMC2817117  PMID: 19520804
avian communities; body-size distributions; climate change; longitudinal analysis; space-for-time substitution; species richness
14.  Rainfall-induced changes in food availability modify the spring departure programme of a migratory bird 
Climatic warming has intensified selection for earlier reproduction in many organisms, but potential constraints imposed by climate change outside the breeding period have received little attention. Migratory birds provide an ideal model for exploring such constraints because they face warming temperatures on temperate breeding grounds and declining rainfall on many tropical non-breeding areas. Here, we use longitudinal data on spring departure dates of American redstarts (Setophaga ruticilla) to show that annual variation in tropical rainfall and food resources are associated with marked change in the timing of spring departure of the same individuals among years. This finding challenges the idea that photoperiod alone regulates the onset of migration, providing evidence that intensifying drought in the tropical winter could hinder adaptive responses to climatic warming in the temperate zone.
PMCID: PMC3177634  PMID: 21450737
American redstart; climate change; ecological constraint; microevolution; phenotypic plasticity; Setophaga ruticilla
15.  Timing of autumn bird migration under climate change: advances in long-distance migrants, delays in short-distance migrants. 
As a response to increasing spring temperature in temperate regions in recent years, populations of many plant and animal species, including migratory birds, have advanced the seasonal start of their reproduction or growth. However, the effects of climate changes on subsequent events of the annual cycle remain poorly understood. We investigated long-term changes in the timing of autumn migration in birds, a key event in the annual cycle limiting the reproductive period. Using data spanning a 42-year period, we analysed long-term changes in the passage of 65 species of migratory birds through Western Europe. The autumn passage of migrants wintering south of the Sahara has advanced in recent years, presumably as a result of selection pressure to cross the Sahel before its seasonal dry period. In contrast, migrants wintering north of the Sahara have delayed autumn passage. In addition, species with a variable rather than a fixed number of broods per year have delayed passage, possibly because they are free to attempt more broods. Recent climate changes seem to have a simple unidirectional effect on the seasonal onset of reproduction, but complex and opposing effects on the timing of subsequent events in the annual cycle, depending on the ecology and life history of a species. This complicates predictions of overall effects of global warming on avian communities.
PMCID: PMC1691393  PMID: 12965011
16.  Woody plants and the prediction of climate-change impacts on bird diversity 
Current methods of assessing climate-induced shifts of species distributions rarely account for species interactions and usually ignore potential differences in response times of interacting taxa to climate change. Here, we used species-richness data from 1005 breeding bird and 1417 woody plant species in Kenya and employed model-averaged coefficients from regression models and median climatic forecasts assembled across 15 climate-change scenarios to predict bird species richness under climate change. Forecasts assuming an instantaneous response of woody plants and birds to climate change suggested increases in future bird species richness across most of Kenya whereas forecasts assuming strongly lagged woody plant responses to climate change indicated a reversed trend, i.e. reduced bird species richness. Uncertainties in predictions of future bird species richness were geographically structured, mainly owing to uncertainties in projected precipitation changes. We conclude that assessments of future species responses to climate change are very sensitive to current uncertainties in regional climate-change projections, and to the inclusion or not of time-lagged interacting taxa. We expect even stronger effects for more specialized plant–animal associations. Given the slow response time of woody plant distributions to climate change, current estimates of future biodiversity of many animal taxa may be both biased and too optimistic.
PMCID: PMC2880125  PMID: 20513712
biodiversity change; biotic interactions; climate-change impacts; general circulation model; predictive model; uncertainty
17.  Hybridization in a warmer world 
Ecology and Evolution  2014;4(10):2019-2031.
Climate change is profoundly affecting the evolutionary trajectory of individual species and ecological communities, in part through the creation of novel species assemblages. How climate change will influence competitive interactions has been an active area of research. Far less attention, however, has been given to altered reproductive interactions. Yet, reproductive interactions between formerly isolated species are inevitable as populations shift geographically and temporally as a result of climate change, potentially resulting in introgression, speciation, or even extinction. The susceptibility of hybridization rates to anthropogenic disturbance was first recognized in the 1930s. To date, work on anthropogenically mediated hybridization has focused primarily on either physical habitat disturbance or species invasion. Here, I review recent literature on hybridization to identify how ecological responses to climate change will increase the likelihood of hybridization via the dissolution of species barriers maintained by habitat, time, or behavior. Using this literature, I identify several cases where novel hybrid zones have recently formed, likely as a result of changing climate. Future research should focus on identifying areas and taxonomic groups where reproductive species interactions are most likely to be influenced by climate change. Furthermore, a better understanding of the evolutionary consequences of climate-mediated secondary contact is urgently needed. Paradoxically, hybridization is both a major conservation concern and an important source of novel genetic and phenotypic variation. Hybridization may therefore both contribute to increasing rates of extinction and stimulate the creation of novel phenotypes that will speed adaptation to novel climates. Predicting which result will occur following secondary contact will be an important contribution to conservation for many species.
PMCID: PMC4063493  PMID: 24963394
Global change; hybrid zone dynamics; mate choice; phenology; species distribution
18.  Species richness changes lag behind climate change 
Species-energy theory indicates that recent climate warming should have driven increases in species richness in cool and species-poor parts of the Northern Hemisphere. We confirm that the average species richness of British butterflies has increased since 1970–82, but much more slowly than predicted from changes of climate: on average, only one-third of the predicted increase has taken place. The resultant species assemblages are increasingly dominated by generalist species that were able to respond quickly. The time lag is confirmed by the successful introduction of many species to climatically suitable areas beyond their ranges. Our results imply that it may be decades or centuries before the species richness and composition of biological communities adjusts to the current climate.
PMCID: PMC1560312  PMID: 16777739
biodiversity; butterflies; climate change; species richness
19.  Range-Wide Latitudinal and Elevational Temperature Gradients for the World's Terrestrial Birds: Implications under Global Climate Change 
PLoS ONE  2014;9(5):e98361.
Species' geographical distributions are tracking latitudinal and elevational surface temperature gradients under global climate change. To evaluate the opportunities to track these gradients across space, we provide a first baseline assessment of the steepness of these gradients for the world's terrestrial birds. Within the breeding ranges of 9,014 bird species, we characterized the spatial gradients in temperature along latitude and elevation for all and a subset of bird species, respectively. We summarized these temperature gradients globally for threatened and non-threatened species and determined how their steepness varied based on species' geography (range size, shape, and orientation) and projected changes in temperature under climate change. Elevational temperature gradients were steepest for species in Africa, western North and South America, and central Asia and shallowest in Australasia, insular IndoMalaya, and the Neotropical lowlands. Latitudinal temperature gradients were steepest for extratropical species, especially in the Northern Hemisphere. Threatened species had shallower elevational gradients whereas latitudinal gradients differed little between threatened and non-threatened species. The strength of elevational gradients was positively correlated with projected changes in temperature. For latitudinal gradients, this relationship only held for extratropical species. The strength of latitudinal gradients was better predicted by species' geography, but primarily for extratropical species. Our findings suggest threatened species are associated with shallower elevational temperature gradients, whereas steep latitudinal gradients are most prevalent outside the tropics where fewer bird species occur year-round. Future modeling and mitigation efforts would benefit from the development of finer grain distributional data to ascertain how these gradients are structured within species' ranges, how and why these gradients vary among species, and the capacity of species to utilize these gradients under climate change.
PMCID: PMC4031198  PMID: 24852009
20.  Climate-Induced Elevational Range Shifts and Increase in Plant Species Richness in a Himalayan Biodiversity Epicentre 
PLoS ONE  2013;8(2):e57103.
Global average temperature increase during the last century has induced species geographic range shifts and extinctions. Montane floras, in particular, are highly sensitive to climate change and mountains serve as suitable observation sites for tracing climate-induced biological response. The Himalaya constitute an important global biodiversity hotspot, yet studies on species’ response to climate change from this region are lacking. Here we use historical (1849–50) and the recent (2007–2010) data on temperature and endemic species’ elevational ranges to perform a correlative study in the two alpine valleys of Sikkim. We show that the ongoing warming in the alpine Sikkim Himalaya has transformed the plant assemblages. This study lends support to the hypothesis that changing climate is causing species distribution changes. We provide first evidence of warmer winters in the region compared to the last two centuries, with mean temperatures of the warmest and the coldest months may have increased by 0.76±0.25°C and 3.65±2°C, respectively. Warming-driven geographical range shifts were recorded in 87% of 124 endemic plant species studied in the region; upper range extensions of species have resulted in increased species richness in the upper alpine zone, compared to the 19th century. We recorded a shift of 23–998 m in species’ upper elevation limit and a mean upward displacement rate of 27.53±22.04 m/decade in the present study. We infer that the present-day plant assemblages and community structure in the Himalaya is substantially different from the last century and is, therefore, in a state of flux under the impact of warming. The continued trend of warming is likely to result in ongoing elevational range contractions and eventually, species extinctions, particularly at mountaintops.
PMCID: PMC3577782  PMID: 23437322
21.  Seasonal Variations in the Diet and Foraging Behaviour of Dunlins Calidris alpina in a South European Estuary: Improved Feeding Conditions for Northward Migrants 
PLoS ONE  2013;8(12):e81174.
During the annual cycle, migratory waders may face strikingly different feeding conditions as they move between breeding areas and wintering grounds. Thus, it is of crucial importance that they rapidly adjust their behaviour and diet to benefit from peaks of prey abundance, in particular during migration, when they need to accumulate energy at a fast pace. In this study, we compared foraging behaviour and diet of wintering and northward migrating dunlins in the Tagus estuary, Portugal, by video-recording foraging birds and analysing their droppings. We also estimated energy intake rates and analysed variations in prey availability, including those that were active at the sediment surface. Wintering and northward migrating dunlins showed clearly different foraging behaviour and diet. In winter, birds predominantly adopted a tactile foraging technique (probing), mainly used to search for small buried bivalves, with some visual surface pecking to collect gastropods and crop bivalve siphons. Contrastingly, in spring dunlins generally used a visual foraging strategy, mostly to consume worms, but also bivalve siphons and shrimps. From winter to spring, we found a marked increase both in the biomass of invertebrate prey in the sediment and in the surface activity of worms and siphons. The combination of these two factors, together with the availability of shrimps in spring, most likely explains the changes in the diet and foraging behaviour of dunlins. Northward migrating birds took advantage from the improved feeding conditions in spring, achieving 65% higher energy intake rates as compared with wintering birds. Building on these results and on known daily activity budgets for this species, our results suggest that Tagus estuary provides high-quality feeding conditions for birds during their stopovers, enabling high fattening rates. These findings show that this large wetland plays a key role as a stopover site for migratory waders within the East Atlantic Flyway.
PMCID: PMC3849249  PMID: 24312532
22.  Phylogenetic patterns of climatic, habitat and trophic niches in a European avian assemblage 
The origins of ecological diversity in continental species assemblages have long intrigued biogeographers. We apply phylogenetic comparative analyses to disentangle the evolutionary patterns of ecological niches in an assemblage of European birds. We compare phylogenetic patterns in trophic, habitat and climatic niche components.
From polygon range maps and handbook data we inferred the realized climatic, habitat and trophic niches of 405 species of breeding birds in Europe. We fitted Pagel’s lambda and kappa statistics, and conducted analyses of disparity through time to compare temporal patterns of ecological diversification on all niche axes together. All observed patterns were compared with expectations based on neutral (Brownian) models of niche divergence.
In this assemblage, patterns of phylogenetic signal (lambda) suggest that related species resemble each other less in regard to their climatic and habitat niches than they do in their trophic niche. Kappa estimates show that ecological divergence does not gradually increase with divergence time, and that this punctualism is stronger in climatic niches than in habitat and trophic niches. Observed niche disparity markedly exceeds levels expected from a Brownian model of ecological diversification, thus providing no evidence for past phylogenetic niche conservatism in these multivariate niches. Levels of multivariate disparity are greatest for the climatic niche, followed by disparity of the habitat and the trophic niches.
Main conclusions
Phylogenetic patterns in the three niche components differ within this avian assemblage. Variation in evolutionary rates (degree of gradualism, constancy through the tree) and/or non-random macroecological sampling probably lead here to differences in the phylogenetic structure of niche components. Testing hypotheses on the origin of these patterns requires more complete phylogenetic trees of the birds, and extended ecological data on different niche components for all bird species.
PMCID: PMC4001463  PMID: 24790525
Birds; Brownian motion; comparative method; disparity through time; ecological diversification; Europe; macroecological sampling; niche conservatism; phylogenetic signal
23.  Ecophysiological constraints shape autumn migratory response to climate change in the North American field sparrow 
Biology Letters  2008;4(5):595-598.
Our ability to accurately forecast species' geographical responses to climate change requires knowledge of the proximate and ultimate drivers of their distribution. Here, we consider the ecophysiological and demographic determinants of the distribution of a partial migrant, the North American field sparrow, Spizella pusilla. From 1940 to 1963, the field sparrow extended its winter northern range margin 222 km polewards. Such expansion was coincident with not only a geographical expansion into suitable breeding habitats, but also a decrease in mean abundance across sites occupied during the winter surveys. Combined, these trends suggest that declining populations along the expansion front either stopped migrating or altered their autumn migration. The poleward expansion was not coincident with climatically induced decreases in peak metabolic energy demand, but it did track increases in ecosystem net primary productivity. After 1963, the species' lower lethal temperature prevented further poleward movement. These findings show how different ecophysiological constraints can interact to change migration and distribution in a demographically declining species.
PMCID: PMC2610066  PMID: 18664419
climate change; demographic decline; ecophysiological niche tracking; geographical range expansion; migration
24.  Is breeding of farmland wading birds depressed by a combination of predator abundance and grazing? 
Biology Letters  2008;4(3):256-258.
Agri-environment schemes have been implemented across Europe to counter biodiversity loss in agricultural landscapes and halt the continual decline of farmland birds, including waders. Such schemes provide financial compensation for changes in agricultural practice, including livestock grazing regimes. Scheme uptake has been variable, partly because farmers believe that other factors, notably predation, are key to wader population declines. On the basis of wader breeding surveys across Shetland, UK, we show that predator density and livestock grazing, through reducing sward height, interact to influence territoriality and thereby are likely to affect wader breeding success. Our results appear to reflect views of both farmers and government agencies, which indicates that future agri-environment schemes would benefit from genuine stakeholder participation to maximize scheme uptake, implementation and beneficial effects on biodiversity. Our findings also imply that agri-environment schemes will reap the greatest benefits for waders through reducing stocking rate where avian predators are abundant.
PMCID: PMC2610041  PMID: 18381262
agri-environment scheme; farmers; lapwing; livestock grazing; oystercatcher; predation
25.  Climate Change or Land Use Dynamics: Do We Know What Climate Change Indicators Indicate? 
PLoS ONE  2011;6(4):e18581.
Different components of global change can have interacting effects on biodiversity and this may influence our ability to detect the specific consequences of climate change through biodiversity indicators. Here, we analyze whether climate change indicators can be affected by land use dynamics that are not directly determined by climate change. To this aim, we analyzed three community-level indicators of climate change impacts that are based on the optimal thermal environment and average latitude of the distribution of bird species present at local communities. We used multiple regression models to relate the variation in climate change indicators to: i) environmental temperature; and ii) three landscape gradients reflecting important current land use change processes (land abandonment, fire impacts and urbanization), all of them having forest areas at their positive extremes. We found that, with few exceptions, landscape gradients determined the figures of climate change indicators as strongly as temperature. Bird communities in forest habitats had colder-dwelling bird species with more northern distributions than farmland, burnt or urban areas. Our results show that land use changes can reverse, hide or exacerbate our perception of climate change impacts when measured through community-level climate change indicators. We stress the need of an explicit incorporation of the interactions between climate change and land use dynamics to understand what are current climate change indicators indicating and be able to isolate real climate change impacts.
PMCID: PMC3080866  PMID: 21533025

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