Survival of Svalbard pink-footed geese Anser brachyrhynchus in relation to winter climate, density and land-use

1. Global change may strongly affect population dynamics, but mechanisms remain elusive. Several Arctic goose species have increased considerably during the last decades. Climate, and land-use changes outside the breeding area have been invoked as causes but have not been tested. We analysed the relationships between conditions on wintering and migration staging areas, and survival in Svalbard pink-footed geese Anser brachyrhynchus. Using mark-recapture data from 14 winters (1989-2002) we estimated survival rates and tested for time trends, and effects of climate, goose density and land-use. 2. Resighting rates differed for males and females, were higher for birds recorded during the previous winter and changed smoothly over time. Survival rates did not differ between sexes, varied over time with a nonsignificant negative trend, and were higher for the first interval after marking (0.88-0.97) than afterwards (0.74-0.93). Average survival estimates were 0.967 (SE 0.026) for the first and 0.861 (SE 0.023) for all later survival intervals. 3. We combined 16 winter and spring climate covariates into two principal components axes. F1 was related to warm/wet winters and an early spring on the Norwegian staging areas and F2 to dry/cold winters. We expected that F1 would be positively related to survival and F2 negatively. F1 explained 23% of survival variation (F1,10=3.24; one-sided P=0.051) when alone in a model and 28% (F1,9=4.50; one-sided P=0.031) in a model that assumed a trend for survival. In contrast, neither F2 nor density, land-use, or scaring practices on important Norwegian spring staging areas had discernible effects on survival. 4. Climate change may thus affect goose population dynamics, with warmer winters and earlier springs enhancing survival and fecundity. A possible mechanism is increased food availability on Danish wintering and Norwegian staging areas. As geese are among the main herbivores in Arctic ecosystems, climate change, by increasing goose populations, may have important indirect effects on Arctic vegetation. Our study also highlights the importance of events outside the breeding area for the population dynamics of migrant species.     

Size‐based ecological interactions drive food web responses to climate warming

Predicting climate change impacts on animal communities requires knowledge of how physiological effects are mediated by ecological interactions. Food‐dependent growth and within‐species size variation depend on temperature and affect community dynamics through feedbacks between individual performance and population size structure. Still, we know little about how warming affects these feedbacks. Using a dynamic stage‐structured biomass model with food‐, size‐ and temperature‐dependent life history processes, we analyse how temperature affects coexistence, stability and size structure in a tri‐trophic food chain, and find that warming effects on community stability depend on ecological interactions. Predator biomass densities generally decline with warming – gradually or through collapses – depending on which consumer life stage predators feed on. Collapses occur when warming induces alternative stable states via Allee effects. This suggests that predator persistence in warmer climates may be lower than previously acknowledged and that effects of warming on food web stability largely depend on species interactions.     

Eutrophication effects on greenhouse gas fluxes from shallow‐lake mesocosms override those of climate warming

Fresh waters make a disproportionately large contribution to greenhouse gas (GHG) emissions, with shallow lakes being particular hot spots. Given their global prevalence, how GHG fluxes from shallow lakes are altered by climate change may have profound implications for the global carbon cycle. Empirical evidence for the temperature dependence of the processes controlling GHG production in natural systems is largely based on the correlation between seasonal temperature variation and seasonal change in GHG fluxes. However, ecosystem‐level GHG fluxes could be influenced by factors, which while varying seasonally with temperature are actually either indirectly related (e.g. primary producer biomass) or largely unrelated to temperature, for instance nutrient loading. Here, we present results from the longest running shallow‐lake mesocosm experiment which demonstrate that nutrient concentrations override temperature as a control of both the total and individual GHG flux. Furthermore, testing for temperature treatment effects at low and high nutrient levels separately showed only one, rather weak, positive effect of temperature (CH₄ flux at high nutrients). In contrast, at low nutrients, the CO₂ efflux was lower in the elevated temperature treatments, with no significant effect on CH₄ or N₂O fluxes. Further analysis identified possible indirect effects of temperature treatment. For example, at low nutrient levels, increased macrophyte abundance was associated with significantly reduced fluxes of both CH₄ and CO₂ for both total annual flux and monthly observation data. As macrophyte abundance was positively related to temperature treatment, this suggests the possibility of indirect temperature effects, via macrophyte abundance, on CH₄ and CO₂ flux. These findings indicate that fluxes of GHGs from shallow lakes may be controlled more by factors indirectly related to temperature, in this case nutrient concentration and the abundance of primary producers. Thus, at ecosystem scale, response to climate change may not follow predictions based on the temperature dependence of metabolic processes.     

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.     

Spatial heterogeneity in climate change effects decouples the long‐term dynamics of wild reindeer populations in the high Arctic

The ‘Moran effect’ predicts that dynamics of populations of a species are synchronized over similar distances as their environmental drivers. Strong population synchrony reduces species viability, but spatial heterogeneity in density dependence, the environment, or its ecological responses may decouple dynamics in space, preventing extinctions. How such heterogeneity buffers impacts of global change on large‐scale population dynamics is not well studied. Here, we show that spatially autocorrelated fluctuations in annual winter weather synchronize wild reindeer dynamics across high‐Arctic Svalbard, while, paradoxically, spatial variation in winter climate trends contribute to diverging local population trajectories. Warmer summers have improved the carrying capacity and apparently led to increased total reindeer abundance. However, fluctuations in population size seem mainly driven by negative effects of stochastic winter rain‐on‐snow (ROS) events causing icing, with strongest effects at high densities. Count data for 10 reindeer populations 8–324 km apart suggested that density‐dependent ROS effects contributed to synchrony in population dynamics, mainly through spatially autocorrelated mortality. By comparing one coastal and one ‘continental’ reindeer population over four decades, we show that locally contrasting abundance trends can arise from spatial differences in climate change and responses to weather. The coastal population experienced a larger increase in ROS, and a stronger density‐dependent ROS effect on population growth rates, than the continental population. In contrast, the latter experienced stronger summer warming and showed the strongest positive response to summer temperatures. Accordingly, contrasting net effects of a recent climate regime shift—with increased ROS and harsher winters, yet higher summer temperatures and improved carrying capacity—led to negative and positive abundance trends in the coastal and continental population respectively. Thus, synchronized population fluctuations by climatic drivers can be buffered by spatial heterogeneity in the same drivers, as well as in the ecological responses, averaging out climate change effects at larger spatial scales.     

Predicting conditions for migration: effects of density dependence and habitat quality

Migration is widespread among animals, but the factors that influence the decision to migrate are poorly understood. Within a single species, populations may be completely migratory, completely sedentary or partially migratory. We use a population model to derive conditions for migration and demonstrate how migratory survival, habitat quality and density dependence on both the breeding and non-breeding grounds influence conditions for migration and the proportion of migrants within a population. Density dependence during the season in which migratory and sedentary individuals use separate sites is necessary for partial migration. High levels of density dependence at the non-shared sites widen the range of survival values within which we predict partial migration, whereas increasing the strength of density dependence at the shared sites narrows the range of survival values within which we predict partial migration. Our results have important implications for predicting how contemporary populations with variable migration strategies may respond to changes in the quality or quantity of habitat.     

Predicting population consequences of ocean climate change for an ecosystem sentinel,the seabird Cassin's auklet

Forecasting the ecological effects of climate change on marine species is critical for informing greenhouse gas mitigation targets and developing marine conservation strategies that remain effective and increase species' resilience under changing climate conditions. Highly productive coastal upwelling systems are predicted to experience substantial effects from climate change, making them priorities for ecological forecasting. We used a population modeling approach to examine the consequences of ocean climate change in the California Current upwelling ecosystem on the population growth rate of the planktivorous seabird Cassin's auklet (Ptychoramphus aleuticus), a demographically sensitive indicator of marine climate change. We use future climate projections for sea surface temperature and upwelling intensity from a regional climate model to forecast changes in the population growth rate of the auklet population at the important Farallon Island colony in central California. Our study projected that the auklet population growth rate will experience an absolute decline of 11–45% by the end of the century, placing this population on a trajectory toward extinction. In addition, future changes in upwelling intensity and timing of peak upwelling are likely to vary across auklet foraging regions in the California Current Ecosystem (CCE), producing a mosaic of climate conditions and ecological impacts across the auklet range. Overall, the Farallon Island Cassin's auklet population has been declining during recent decades, and ocean climate change in this century under a mid‐level emissions scenario is projected to accelerate this decline, leading toward population extinction. Because our study species has proven to be a sensitive indicator of oceanographic conditions in the CCE and a powerful predictor of the abundance of other important predators (i.e. salmon), the significant impacts we predicted for the Cassin's auklet provide insights into the consequences that ocean climate change may have for other plankton predators in this system.     

Effects of climate change and crop planting structure on the abundance of cotton bollworm,Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

The interactions between plants and insects play an important role in ecosystems. Climate change and cropping patterns can affect herbivorous pest insect dynamics. Understanding the reasons for population fluctuations can help improve integrated pest management strategies. Here, a 25‐year dataset on climate, cropping planting structure, and the population dynamics of cotton bollworms (Helicoverpa armigera) from Bachu County, south Xinjiang, China, was analyzed to assess the effects of changes in climate and crop planting structure on the population dynamics of H. armigera. The three generations of H. armigera showed increasing trends in population size with climate warming, especially in the third generation. The relative abundances of the first and second generations decreased, but that of the third generation increased. Rising temperature and precipitation produced different impacts on the development of different generations. The population numbers of H. armigera increased with the increase in the non‐Bacillus thuringiensis (Bt) cotton‐planted area. Asynchrony of abrupt changes existed among climate change, crop flowering dates, and the phenology of H. armigera moths. The asynchronous responses in crop flowering dates and phenology of H. armigera to climate warming would expand in the future. The primary factors affecting the first, second, and third generations of moths were T_mean in June, the last appearance date of the second generation of moths, and the duration of the third generation of moths, respectively. To reduce the harm to crops caused by H. armigera, Bt cotton should be widely planted.     

Synergistic effects of climate and land‐use change influence broad‐scale avian population declines

Climate and land‐use changes are expected to be the primary drivers of future global biodiversity loss. Although theory suggests that these factors impact species synergistically, past studies have either focused on only one in isolation or have substituted space for time, which often results in confounding between drivers. Tests of synergistic effects require congruent time series on animal populations, climate change and land‐use change replicated across landscapes that span the gradient of correlations between the drivers of change. Using a unique time series of high‐resolution climate (measured as temperature and precipitation) and land‐use change (measured as forest change) data, we show that these drivers of global change act synergistically to influence forest bird population declines over 29 years in the Pacific Northwest of the United States. Nearly half of the species examined had declined over this time. Populations declined most in response to loss of early seral and mature forest, with responses to loss of early seral forest amplified in landscapes that had warmed over time. In addition, birds declined more in response to loss of mature forest in areas that had dried over time. Climate change did not appear to impact populations in landscapes with limited habitat loss, except when those landscapes were initially warmer than the average landscape. Our results provide some of the first empirical evidence of synergistic effects of climate and land‐use change on animal population dynamics, suggesting accelerated loss of biodiversity in areas under pressure from multiple global change drivers. Furthermore, our findings suggest strong spatial variability in the impacts of climate change and highlight the need for future studies to evaluate multiple drivers simultaneously to avoid potential misattribution of effects.     

Nonlinear effects of climate on boreal rodent dynamics: mild winters do not negate high‐amplitude cycles

Small rodents are key species in many ecosystems. In boreal and subarctic environments, their importance is heightened by pronounced multiannual population cycles. Alarmingly, the previously regular rodent cycles appear to be collapsing simultaneously in many areas. Climate change, particularly decreasing snow quality or quantity in winter, is hypothesized as a causal factor, but the evidence is contradictory. Reliable analysis of population dynamics and the influence of climate thereon necessitate spatially and temporally extensive data. We combined data on vole abundances and climate, collected at 33 locations throughout Finland from 1970 to 2011, to test the hypothesis that warming winters are causing a disappearance of multiannual vole cycles. We predicted that vole population dynamics exhibit geographic and temporal variation associated with variation in climate; reduced cyclicity should be observed when and where winter weather has become milder. We found that the temporal patterns in cyclicity varied between climatically different regions: a transient reduction in cycle amplitude in the coldest region, low‐amplitude cycles or irregular dynamics in the climatically intermediate regions, and strengthening cyclicity in the warmest region. Our results did not support the hypothesis that mild winters are uniformly leading to irregular dynamics in boreal vole populations. Long and cold winters were neither a prerequisite for high‐amplitude multiannual cycles, nor were mild winters with reduced snow cover associated with reduced winter growth rates. Population dynamics correlated more strongly with growing season than with winter conditions. Cyclicity was weakened by increasing growing season temperatures in the cold, but strengthened in the warm regions. High‐amplitude multiannual vole cycles emerge in two climatic regimes: a winter‐driven cycle in cold, and a summer‐driven cycle in warm climates. Finally, we show that geographic climatic gradients alone may not reliably predict biological responses to climate change.     

Northward range expansion in spring‐staging barnacle geese is a response to climate change and population growth,mediated by individual experience

All long‐distance migrants must cope with changing environments, but species differ greatly in how they do so. In some species, individuals might be able to adjust by learning from individual experiences and by copying others. This could greatly speed up the process of adjustment, but evidence from the wild is scarce. Here, we investigated the processes by which a rapidly growing population of barnacle geese (Branta leucopsis) responded to strong environmental changes on spring‐staging areas in Norway. One area, Helgeland, has been the traditional site. Since the mid‐1990s, an increasing number of geese stage in another area 250 km further north, Vesterålen. We collected data on goose numbers and weather conditions from 1975 to 2017 to explore the extent to which the increase in population size and a warmer climate contributed to this change in staging area use. During the study period, the estimated onset of grass growth advanced on average by 0.54 days/year in each of the two areas. The total production of digestible biomass for barnacle geese during the staging period increased in Vesterålen but remained stable in Helgeland. The goose population has doubled in size during the past 25 years, with most of the growth being accommodated in Vesterålen. The observations suggest that this dramatic increase would not have happened without higher temperatures in Vesterålen. Records of individually marked geese indicate that from the initial years of colonization onwards, especially young geese tended to switch to Vesterålen, thereby predominating in the flocks at Vesterålen. Older birds had a lower probability of switching to Vesterålen, but over the years, the probability increased for all ages. Our findings suggest that barnacle geese integrate socially learned behaviour with adjustments to individual experiences, allowing the population to respond rapidly and accurately to global change.     

Dampening prey cycle overrides the impact of climate change on predator population dynamics: a long‐term demographic study on tawny owls

Predicting the dynamics of animal populations with different life histories requires careful understanding of demographic responses to multifaceted aspects of global changes, such as climate and trophic interactions. Continent‐scale dampening of vole population cycles, keystone herbivores in many ecosystems, has been recently documented across Europe. However, its impact on guilds of vole‐eating predators remains unknown. To quantify this impact, we used a 27‐year study of an avian predator (tawny owl) and its main prey (field vole) collected in Kielder Forest (UK) where vole dynamics shifted from a high‐ to a low‐amplitude fluctuation regime in the mid‐1990s. We measured the functional responses of four demographic rates to changes in prey dynamics and winter climate, characterized by wintertime North Atlantic Oscillation (wNAO). First‐year and adult survival were positively affected by vole density in autumn but relatively insensitive to wNAO. The probability of breeding and number of fledglings were higher in years with high spring vole densities and negative wNAO (i.e. colder and drier winters). These functional responses were incorporated into a stochastic population model. The size of the predator population was projected under scenarios combining prey dynamics and winter climate to test whether climate buffers or alternatively magnifies the impact of changes in prey dynamics. We found the observed dampening vole cycles, characterized by low spring densities, drastically reduced the breeding probability of predators. Our results illustrate that (i) change in trophic interactions can override direct climate change effect; and (ii) the demographic resilience entailed by longevity and the occurrence of a floater stage may be insufficient to buffer hypothesized environmental changes. Ultimately, dampened prey cycles would drive our owl local population towards extinction, with winter climate regimes only altering persistence time. These results suggest that other vole‐eating predators are likely to be threatened by dampening vole cycles throughout Europe.     

Interactions between climate and habitat loss effects on biodiversity: a systematic review and meta‐analysis

Climate change and habitat loss are both key threatening processes driving the global loss in biodiversity. Yet little is known about their synergistic effects on biological populations due to the complexity underlying both processes. If the combined effects of habitat loss and climate change are greater than the effects of each threat individually, current conservation management strategies may be inefficient and at worst ineffective. Therefore, there is a pressing need to identify whether interacting effects between climate change and habitat loss exist and, if so, quantify the magnitude of their impact. In this article, we present a meta‐analysis of studies that quantify the effect of habitat loss on biological populations and examine whether the magnitude of these effects depends on current climatic conditions and historical rates of climate change. We examined 1319 papers on habitat loss and fragmentation, identified from the past 20 years, representing a range of taxa, landscapes, land‐uses, geographic locations and climatic conditions. We find that current climate and climate change are important factors determining the negative effects of habitat loss on species density and/or diversity. The most important determinant of habitat loss and fragmentation effects, averaged across species and geographic regions, was current maximum temperature, with mean precipitation change over the last 100 years of secondary importance. Habitat loss and fragmentation effects were greatest in areas with high maximum temperatures. Conversely, they were lowest in areas where average rainfall has increased over time. To our knowledge, this is the first study to conduct a global terrestrial analysis of existing data to quantify and test for interacting effects between current climate, climatic change and habitat loss on biological populations. Understanding the synergistic effects between climate change and other threatening processes has critical implications for our ability to support and incorporate climate change adaptation measures into policy development and management response.     

A review and meta‐analysis of the effects of climate change on Holarctic mountain and upland bird populations

Mountain regions are globally important areas for biodiversity but are subject to multiple human‐induced threats, including climate change, which has been more severe at higher elevations. We reviewed evidence for impacts of climate change on Holarctic mountain bird populations in terms of physiology, phenology, trophic interactions, demography and observed and projected distribution shifts, including effects of other factors that interact with climate change. We developed an objective classification of high‐elevation, mountain specialist and generalist species, based on the proportion of their breeding range occurring in mountain regions. Our review found evidence of responses of mountain bird populations to climate (extreme weather events, temperature, rainfall and snow) and environmental (i.e. land use) change, but we know little about either the underlying mechanisms or the synergistic effects of climate and land use. Long‐term studies assessing reproductive success or survival of mountain birds in relation to climate change were rare. Few studies have considered shifts in elevational distribution over time and a meta‐analysis did not find a consistent direction in elevation change. A meta‐analysis carried out on future projections of distribution shifts suggested that birds whose breeding distributions are largely restricted to mountains are likely to be more negatively impacted than other species. Adaptation responses to climate change rely mostly on managing and extending current protected areas for both species already present, and for expected colonizing species that are losing habitat and climate space at lower elevation. However, developing effective management actions requires an improvement in the current knowledge of mountain species ecology, in the quality of climate data and in understanding the role of interacting factors. Furthermore, the evidence was mostly based on widespread species rather than mountain specialists. Scientists should provide valuable tools to assess the status of mountain birds, for example through the development of a mountain bird population index, and policy‐makers should influence legislation to develop efficient agri‐environment schemes and forestry practices for mountain birds, as well as to regulate leisure activities at higher elevations.     

Weakening density dependence from climate change and agricultural intensification triggers pest outbreaks: a 37‐year observation of cotton bollworms

Understanding drivers of population fluctuation, especially for agricultural pests, is central to the provision of agro‐ecosystem services. Here, we examine the role of endogenous density dependence and exogenous factors of climate and human activity in regulating the 37‐year population dynamics of an important agricultural insect pest, the cotton bollworm (Helicoverpa armigera), in North China from 1975 to 2011. Quantitative time‐series analysis provided strong evidence explaining long‐term population dynamics of the cotton bollworm and its driving factors. Rising temperature and declining rainfall exacerbated the effect of agricultural intensification on continuously weakening the negative density dependence in regulating the population dynamics of cotton bollworms. Consequently, ongoing climate change and agricultural intensification unleashed the tightly regulated pest population and triggered the regional outbreak of H. armigera in 1992. Although the negative density dependence can effectively regulate the population change rate to fluctuate around zero at stable equilibrium levels before and after outbreak in the 1992, the population equilibrium jumped to a higher density level with apparently larger amplitudes after the outbreak. The results highlight the possibility for exogenous factors to induce pest outbreaks and alter the population regulating mechanism of negative density dependence and, thus, the stable equilibrium of the pest population, often to a higher level, posing considerable risks to the provision of agro‐ecosystem services and regional food security. Efficient and timely measures of pest management in the era of Anthropocene should target the strengthening and revival of weakening density dependence caused by climate change and human activities.     

Relocation,high‐latitude warming and host genetic identity shape the foliar fungal microbiome of poplars

Micro‐organisms associated with plants and animals affect host fitness, shape community structure and influence ecosystem properties. Climate change is expected to influence microbial communities, but their reactions are not well understood. Host‐associated micro‐organisms are influenced by the climate reactions of their hosts, which may undergo range shifts due to climatic niche tracking, or may be actively relocated to mitigate the effects of climate change. We used a common‐garden experiment and rDNA metabarcoding to examine the effect of host relocation and high‐latitude warming on the complex fungal endophytic microbiome associated with leaves of an ecologically dominant boreal forest tree (Populus balsamifera L.). We also considered the potential effects of poplar genetic identity in defining the reactions of the microbiome to the treatments. The relocation of hosts to the north increased the diversity of the microbiome and influenced its structure, with results indicating enemy release from plausible pathogens. High‐latitude warming decreased microbiome diversity in comparison with natural northern conditions. The warming also caused structural changes, which made the fungal communities distinct in comparison with both low‐latitude and high‐latitude natural communities, and increased the abundance of plausible pathogens. The reactions of the microbiome to relocation and warming were strongly dependent on host genetic identity. This suggests that climate change effects on host–microbiome systems may be mediated by the interaction of environmental factors and the population genetic processes of the hosts.