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Climate Change and the Ecology and Evolution of Arctic Vertebrates

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Climate Change and the Ecology and Evolution of Arctic Vertebrates Ann. N.Y. Acad. Sci. ISSN 0077-8923 ANNALS OF THE NEW YORK ACADEMY OF SCIENCES Issue: The Year in Ecology and Conservation Biology Climate change and the ecology and evolution of Arctic vertebrates Olivier Gilg,1,2,3 Kit M. Kovacs,4 Jon Aars,4 Jer´ omeˆ Fort,5 Gilles Gauthier,6 David Gremillet,´ 7 Rolf A. Ims,8 Hans Meltofte,5 Jer´ omeˆ Moreau,1 Eric Post,9 Niels Martin Schmidt,5 Glenn Yannic,3,6 and Lo¨ıc Bollache1 1Universite´ de Bourgogne, Laboratoire Biogeosciences,´ UMR CNRS 5561, Equipe Ecologie Evolutive, Dijon, France. 2Division of Population Biology, Department of Biological and Environmental Sciences, University of Helsinki, Finland. 3Groupe de Recherche en Ecologie Arctique (GREA), Francheville, France. 4Norwegian Polar Institute, FRAM Centre, Tromsø, Norway. 5Department of Bioscience, Aarhus University, Roskilde, Denmark. 6Departement´ de Biology and Centre d’Etudes´ Nordiques, Universite´ Laval, Quebec,´ Quebec,´ G1V 0A6, Canada. 7FRAM Centre d’Ecologie Fonctionnelle et Evolutive, UMR CNRS 5175, Montpellier, France. 8Department of Arctic and Marine Biology, University of Tromsø, Tromsø, Norway. 9Department of Biology, Penn State University, University Park, Pennsylvania Address for correspondence: Olivier Gilg, Laboratoire Biogeosciences,´ UMR CNRS 5561, Equipe Ecologie Evolutive, 6 Boulevard Gabriel, 21000 Dijon, France. [email protected] Climate change is taking place more rapidly and severely in the Arctic than anywhere on the globe, exposing Arctic vertebrates to a host of impacts. Changes in the cryosphere dominate the physical changes that already affect these animals, but increasing air temperatures, changes in precipitation, and ocean acidification will also affect Arctic ecosystems in the future. Adaptation via natural selection is problematic in such a rapidly changing environment. Adjustment via phenotypic plasticity is therefore likely to dominate Arctic vertebrate responses in the short term, and many such adjustments have already been documented. Changes in phenology and range will occur for most species but will only partly mitigate climate change impacts, which are particularly difficult to forecast due to the many interactions within and between trophic levels. Even though Arctic species richness is increasing via immigration from the South, many Arctic vertebrates are expected to become increasingly threatened during this century. Keywords: impacts; phenological changes; plasticity; range shifts; adaptations; threat; trophic interactions; mis- matches; sea ice; tundra; parasites; geese; shorebirds; rodents; lemmings; large herbivores; seabirds; marine mammals; polar bear ing: in Africa, water stress is expected to become the Introduction main constraint in the future, with arid and semi- Several studies have already reviewed the ongoing arid land projected to increase; in eastern South and forecasted impacts of climate change on the America, drought and increasing temperatures will ecology and evolution of vertebrates on Earth.1,2 induce a gradual replacement of tropical forests by Overall, changes in phenologies, in distributional savanna; in boreal regions, increasing rainfall and range, in evolutionary adaptations—on the long temperature will in many places allow the forest term—and, in the structure and the dynamics of to expand in altitude and latitude, replacing the communities, are forecast to be the main biological Arctic and Alpine tundras; and in many coastal consequences of climate change. However, due to lowlands, the elevation of sea level will replace ter- regional differences in climate change and in biotic restrial ecosystems by marine ones. In the Arctic, characteristics, climate-induced impacts on species physical changes will be dominated by changes in the will differ in their nature and magnitude between re- cryosphere (glaciers, permafrost, sea ice, and snow gionsandecosystems.3 Examples include the follow- layers), primarily due to increasing temperatures. doi: 10.1111/j.1749-6632.2011.06412.x 166 Ann. N.Y. Acad. Sci. 1249 (2012) 166–190 c 2012 New York Academy of Sciences. Gilg et al. Climate change impacts on Arctic vertebrates To understand how Arctic vertebrates will respond type in the Arctic. If these trends continue as pre- to these changes, we have to consider their particu- dicted by models, the Arctic Ocean should become lar history, the extreme physical environment they ice free in summer well before the end of this cen- currently inhabit, and the relatively simple structure tury,7 and this would be a first for Arctic marine and composition of the communities to which they systems during the last 5+ million years.8 belong. Arctic vertebrates possess many adaptations that Of course, changes in climate are not entirely new, have enabled their persistence in Arctic environ- and Arctic vertebrates have already lived through ments. These traits include behavioral (e.g., day tor- several periods of “historical climate changes”— por, seasonal migration), physiological (e.g., sea- although the speed and the extent of the current sonal metabolic adjustments, fasting ability), and changes are probably unprecedented. However, un- morphological (e.g., white coats, short extremities, less we enter the realm of paleoecology, we know extensive blubber layers, thick fur, lack of dorsal little about ecological responses of vertebrates to fins among ice whales) mechanisms. But despite past changes in climate. Vibe’s seminal study4 is their extreme and varied adaptations to coping with one notable exception. He clearly showed, nearly the harsh and highly variable Arctic climate, these 50 years ago, the importance of climate, and more species are considered to be particularly vulnera- precisely the availability of specific ice types, in de- ble to ongoing climate changes. Their peculiar life- termining the distribution of marine mammals on history strategies have served them well in a low- the west coast of Greenland. More recent analyses competition situation but leave them vulnerable to of decadal patterns of sea ice and their influences on competitive stress from temperate species that are marine mammals can also be used to predict that already invading their ranges. Arctic endemics have changes in recent years are likely to impact resident little potential for northward retraction to avoid marine mammal populations at both regional and such overlap. In addition, they may have limited hemispheric scales.5 capacity to deal with exposure to disease because The Arctic biome is already changing, and this of their limited exposure in the past. Furthermore, change is rapid and severe compared to other re- the heavy lipid-based dietary dependence of some gions of the globe. It is expected that the changes that Arctic species leaves them disproportionately vul- will take place in the Arctic in the coming decades nerable to the effects of lipophilic contaminants. will surpass what many Arctic vertebrates have ex- Arctic vertebrates can adjust to cope with climate- perienced in the past, despite the extreme variabil- related changes (i.e., through their phenotypic plas- ity that is normal in the climate of the Arctic. In- ticity) or adapt (via the differential selection of some deed, while the global mean surface temperature has genotypes) in many different ways. The aim of our warmed by 0.6–0.8◦ Cduringthe20thcentury,ithas paper is to review these different responses and, increased ca. twice as much in the Arctic during the whenever possible, to present examples of the im- same period.3 Theconsequencesofthiswarmingin- pacts of climate change at the community level, clude decreases in snow, ice, and frozen ground lay- explicitly taking into account interspecific interac- ers, while precipitation has increased overall in the tions. Given the wide range of possible interactions, Northern hemisphere.3 The strong environmental predictionsmadeatthislevelusuallyremainspecu- constraints that prevail in the Arctic—including the lative. On the other hand, predictions that overlook harsh climate, strong seasonality, low productivity, interaction processes would lack realism. To avoid and numerous ecological barriers—are likely to ex- these pitfalls, the examples supporting our text are acerbate the impacts that are expected due to direct primarily chosen from the few biological models influences of climate change.6 that have already been studied comprehensively in Documented declines in Arctic sea ice are symp- the Arctic in recent years—that is, rodents and their tomatic of these ongoing changes. Its overall extent, predators, geese and shorebirds, large herbivores, thickness, seasonal duration, and the proportion of seabirds, and marine mammals and their prey. multiyear ice have all declined. Multiyear sea ice Our paper is structured in nine sections and has disappeared three times faster during the past follows a process orientation, rather than the tra- decade than during the previous three decades, and ditional species by species approach. In the first seasonal ice is thus becoming the dominant sea ice section, we present a general outline of how Ann. N.Y. Acad. Sci. 1249 (2012) 166–190 c 2012 New York Academy of Sciences. 167 Climate change impacts on Arctic vertebrates Gilg et al. vertebrates will respond to changes in the The winter is a key period in the annual cycle cryosphere, because most species living in the Arctic of Arctic small mammals as some—for example, biome are exceedingly dependent on snow and ice the Arctic lemmings—live under the snow for up regimes for their reproduction
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