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Climate change affects both terrestrial<ref>Diffenbaugh, N. S. and Field, C. B., 2013. Changes in Ecologically Critical Terrestrial Climate Conditions. Science, 341(6145), pp. 486-492. [https://doi.org/10.1126/science.1237123 doi: 10.1126/science.1237123]</ref> and aquatic biomes<ref>Hoegh-Guldberg, O., and Bruno, J. F., 2010. The Impact of Climate Change on the World’s Marine Ecosystem. Science, 328(5985), pp. 1523-1528. [https://doi.org/10.1126/science.1189930 doi: 10.1126/science.1189930]</ref> causing significant effects on ecosystem functions and biodiversity<ref name=":0">Bellard, C., Berteslsmeier, C., Leadley, P., Thuiller, W., and Courchamp, F., 2012. Impacts of climate change on the future of biodiversity. Ecological Letters, 15(4), pp. 365-377. [https://doi.org/10.1111/j.1461-0248.2011.01736.x doi: 10.1111/j.1461-0248.2011.01736.x] [//www.enviro.wiki/images/a/a4/Bellard2012.pdf Article pdf]</ref>. Climate change is affecting several key ecological processes and patterns that will have cascading impacts on wildlife and habitat<ref name=":1">Inkley, D. B., Anderson, M. G., Blaustein, A. R., Burkett, V. R., Felzer, B., Griffith, B., Price, J., and Root, T. L., 2004. Global Climate Change and Wildlife in North America. Wildlife Society Technical Review 04-2. The Wildlife Society, Bethesda, MD, 26 pp. [//www.enviro.wiki/images/f/f1/Inkley2004.pdf Report pdf]</ref>. For example, sea-level rise, changes in the timing and duration of growing seasons, and changes in primary production are mainly driven by changes to global environmental variables (e.g., temperature and atmospheric CO<sub>2</sub>). Climate-induced changes in the environment ultimately impact wildlife population abundance and distributions.  
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As the global climate continues to warm, changes in local climate conditions put populations of many species at risk of severe decline and even extinction. Predicting which species are most vulnerable to changing conditions is challenging, because climate interacts different life stages in complex ways. Population models allow natural resource managers to integrate the effects of climate across life stages and provide a powerful tool to inform management decisions. However, care must be taken to match model structure to a species’ biology and recognize the limitations of the data used to parameterize models when interpreting predictions.
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<div style="float:right;margin:0 0 2em 2em;">__TOC__</div>
  
 
'''Related Article(s):'''
 
'''Related Article(s):'''
  
 
*[[Climate Change Primer|Climate Change]]
 
*[[Climate Change Primer|Climate Change]]
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*[[Predicting Species Responses to Climate Change with Population Models]]
  
  
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'''Contributor(s):''' Dr. Brian Hudgens
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'''Contributor(s):''' Dr. Breanna F. Powers and Dr. Julie A. Heath
  
  
 
'''Key Resource(s):'''
 
'''Key Resource(s):'''
  
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*Quantitative Conservation Biology<ref>Morris, W.F., and Doak, D.F., 2002. Quantitative Conservation Biology: Theory and Practice of Population Viability Analysis. Sinauer Associates, Inc. Publishers, Sunderland, Massachusetts, USA. ISBN: 978-087893546-8</ref>
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*Global climate change and wildlife in North America. Wildlife Society Technical Review 04-2<ref name=":1" />
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*Evaluating the Use of Spatially Explicit Population Models to Predict Conservation Reliant Species in Nonanalogue Future Environments on DoD Lands, Strategic Environmental Research and Development Program (SERDP)<ref>
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*Impacts of climate change on the future of biodiversity<ref name=":0" />
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Hudgens, B., Abbott, J., Haddad, N., Kiekebusch E., Louthan A., Morris W., Stenzel L., Walters J., 2020. Evaluating the Use of Spatially Explicit Population Models to Predict Conservation Reliant Species in Nonanalogue Future Environments on DoD Lands.  Strategic Environmental Research and Development Program (SERDP), [https://www.serdp-estcp.org/Program-Areas/Resource-Conservation-and-Resiliency/Natural-Resources/Species-Ecology-and-Management/RC-2512 Project RC-2512] [//www.enviro.wiki/images/0/04/RC-2512FinalReport.pdf Final Report pdf]</ref>.
 
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==Climate Change and No-Analogue Environmental Conditions==
 
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The global climate has been changing throughout the past century, with continued changes predicted over coming decades. Generally, temperatures are getting warmer throughout U.S. and worldwide<ref name=":0">
 
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IPCC, 2013. Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. Stocker, T.F., D. Qin, G.-K. Plattner, M. Tignor, S.K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex and P.M. Midgley (eds.). Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA, 1535 pp. [//www.enviro.wiki/images/5/58/2013-IPCC-Climate_Change-The_Physical_science_basis.pdf Report pdf]
 
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</ref>. Precipitation patterns are also changing, with some regions of the U.S.  getting drier, others getting wetter<ref name=":0" /><ref name=":1">
 
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Abatzoglou, J. T., 2013. Development of gridded surface meteorological data for ecological applications and modelling. International Journal of Climatology, 33(1), pp. 121–131. [https://doi.org/10.1002/joc.3413 doi: 10.1002/joc.3413]</ref>.  Further changes are occurring in the timing and duration of precipitation events, with extreme weather events becoming more common<ref name=":0" /><ref name=":1" />.
 
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As a result of these changes, populations, even entire species, are likely to experience novel conditions that impact individual fitness and population viability. In the case of polar bears, extended periods of low sea ice resulting from atmospheric and oceanic warming have led to the species being protected under the Endangered Species Act<ref>U.S. Fish and Wildlife (USFWS), 2016. Polar Bear (Ursus maritimus) Conservation Management Plan, Final. U.S. Fish and Wildlife, Region 7, Anchorage, Alaska. 104 pp. [https://ecos.fws.gov/docs/recovery_plan/PBRT%20Recovery%20Plan%20Book.FINAL.signed.pdf Report pdf]</ref>.
 
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The potential for a changing climate to put populations at risk of extinction creates two imperatives for natural resource managers: 1) detecting climate effects on vital rates such as fecundity, growth and survival, and 2) predicting the overall impact of changing climate on managed species. Importantly, changing climate conditions may have different short and long term effects on populations. For example, a population of eastern tiger salamanders (''Ambystoma tigrinum'') studied at a breeding pond in Fort Bragg, NC, has been observed to suffer consecutive years of no successful offspring when breeding ponds dry out before larvae could undergo metamorphosis<ref>Woodward, D., Hudgens, B., and Haddad, N., 2005. Status and ecology of the Northern Pine Snake, Southern Hognose Snake, Tiger Salamander, and Carolina Gopher Frog on Ft. Bragg, NC. Unpublished report to Ft. Bragg Endangered Species Branch. </ref><ref>Haddad, N., Woodward, D., Hudgens, B., Davidai, N., Fields, W., Chesser, M., 2007. Status and ecology of the Northern Pine Snake, Southern Hognose Snake, Tiger Salamander, and Carolina Gopher Frog on Ft. Bragg, NC. Unpublished report to Ft. Bragg Endangered Species Branch.</ref>. This population would initially benefit from increases in precipitation leading to higher fecundity (reproductive success). However, if higher levels of precipitation convert ephemeral ponds to permanent ponds, many resident amphibians would become vulnerable to increased mortality due to predation by fish and bullfrogs (''Lithobates catesbeianus)''<ref>
 
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Fisher, R.N., and Shaffer, H.B., 1996. The Decline of Amphibians in California’s Great Central Valley. Conservation Biology, 10(5), pp. 1387-1397. [https://doi.org/10.1046/j.1523-1739.1996.10051387.x doi:10.1046/J.1523-1739.1996.10051387.X]
 
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</ref><ref>Cook, M.T., Heppell, S.S., and Garcia, T.S., 2013. Invasive Bullfrog Larvae Lack Developmental Plasticity to Changing Hydroperiod. The Journal of Wildlife Management, 77(4), pp. 655-662. [https://doi.org/10.1002/jwmg.509 doi:10.1002/jwmg.509]  [//www.enviro.wiki/images/e/e4/Cook2013.pdf Article pdf]</ref>.
 
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Previous studies have shown that the effect of changing environmental conditions on a species is best understood by monitoring effects on all life stages<ref>Brown, L.M., Breed, G.A., Severns, P.M. and Crone, E.E., 2017. Losing a battle but winning the war: moving past preference–performance to understand native herbivore–novel host plant interactions. Oecologia, 183(2), pp. 441-453. [https://doi.org/10.1007/s00442-016-3787-y doi: 10.1007/s00442-016-3787-y]</ref>. This is particularly true for understanding how a changing climate may impact a species. Climate change typically involves simultaneous changes in numerous climate variables (e.g. temperature and precipitation), which may interact with a species at different points of its life-cycle. For example, summer temperatures impact young (small) plant growth of the tundra plant moss campion (''Silene acaulis'') while snow cover influences growth and survival of larger plants<ref name=":2">Doak, D.F., and Morris, W.F., 2010. Demographic compensation and tipping points in climate-induced range shifts. Nature, 467, p.p. 959-962. [https://doi.org/10.1038/nature09439 doi:10.1038/nature09439]</ref>.
 
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Even when there is a single, highly dominant climate driver, it is likely to have different effects on different stages of a species life-cycle. For example, drought reduced San Clemente Bell's sparrow fecundity, but not adult survival<ref name=":6">Hudgens, B., Beaudry, F., George, T.L., Kaiser, S., and Munkwitz, N.M., 2011. Shifting threats faced by the San Clemente sage sparrow. The Journal of Wildlife Management, 75(6), pp. 1350-1360. [https://doi.org/10.1002/jwmg.165 doi: 10.1002/jwmg.165] [//www.enviro.wiki/images/a/ae/Hudgens2011.pdf Article pdf]</ref>. Moreover, the variation in the same climate variable can have opposing effects on different life stages. For example, warmer summer temperatures tend to help moss campion plants grow larger, but decrease the number of fruits produced by plants of a given size<ref name=":2" />.
 
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==Population Models Integrating Effects Across Different Life-Stages==
 
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The complex ways in which climate change can impact different species creates a significant challenge to predicting future management needs. Population models can provide a powerful tool for meeting this challenge. Generally, more complex population models capable of integrating climate effects on different life stages are the most useful for predicting species' responses to climate change.  The two types of population models most commonly used for integrating across different life stages are matrix models<ref name=":4">
 
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Ehrlén, J., and Morris, W.F., 2015. Predicting changes in the distribution and abundance of species under environmental change. Ecology Letters, 18(3), pp. 303-314. [https://doi.org/10.1111/ele.12410 doi: 10.1111/ele.12410] [//www.enviro.wiki/images/b/b2/Ehrlen2015.pdf Article pdf]</ref> and individual-based simulations.
 
  
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===Matrix Models===
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==Introduction==
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Matrix models are widely used because there are algebraic tools that facilitate evaluating these models and they have a flexible enough structure to accommodate a wide range of life history strategies<ref>Caswell, H., 2001. Matrix Population Models – Construction, Analysis, and Interpretation. Sinauer Associates, Sunderland, MA, USA, 722 pp. ISBN: 0-87893-096-5</ref><ref>
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[[File:PowersFig1.png|thumb|900px|left | Figure 1. The predicted extinction risk (by percentage with 95% CIs) from climate change by different regions, colors represent a gradient from least to most extinction risks (green to red) based upon the number of relevant studies (n). Figure is from Urban (2015)<ref name=":2" />. Reprinted with permission from AAAS. Any use of this figure requires the prior written permission of AAAS]]
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Cochran, M.E., and Ellner, S., 1992. Simple Methods for Calculating Age‐Based Life History Parameters for Stage‐Structured Populations: Ecological Archives M062-002. Ecological monographs, 62(3), pp. 345-364. [https://doi.org/10.2307/2937115 doi: 10.2307/2937115]</ref>. One of the appeals to matrix models is that the long term growth rate and the proportion of individuals expected to be in each stage class (known as the stable stage distribution) can be calculated directly using matrix algebraic tools, with the long term population growth rate given by the dominant eigenvalue of the matrix model and stable stage distribution given by the corresponding eigenvector<ref>Leslie, P.H., 1945. On the Use of Matrices in Certain Population Mathematics. Biometrika, 33(3), pp.183-212. [https://doi.org/10.2307/2332297 doi: 10.2307/2332297]</ref>. A key assumption in estimating long term growth rates and stable stage distributions from matrix models is that the transition rates from one life stage to the next represented by the matrix elements are constant through time. Differences in observed proportions of a population in different stages from the predicted stable stage distributions can be used to detect differences in past and present transition ratescaused by changing climatic conditions<ref>Doak, D.F., and Morris, W., 1999. Detecting Population‐Level Consequences of Ongoing Environmental Change Without Long‐Term Monitoring. Ecology, 80(5), pp. 1537-1551.  
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Global climate change will affect ecosystem functions and cycles such as nutrient, hydraulic, and carbon cycles, changing aspects of environmental conditions such as temperature, soil moisture, and precipitation<ref>Davidson, E. A. and Janssens, I. A., 2006. Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Nature, 440, pp. 165-173.[https://doi.org/10.1038/nature04514 doi: 10.1038/nature04514]</ref><ref>Melillo, J. M., McGuire, A. D., Kicklighter, D. W., Moore, B., Vorosmarty, C. J., and Schloss, A. L., 1993. Global climate change and terrestrial net primary production. Nature, 363, pp. 234-240. [https://doi.org/10.1038/363234a0 doi: 10.1038/363234a0]</ref>. Wildlife species are adapted to their environments and changes to the environment and habitat conditions will mediate effects, either directly or indirectly, on species survival, fecundity and ultimately population persistence<ref>Alig, R. J., Technical Coordinator, 2011. Effects of Climate Change on Natural Resources and Communities: A Compendium of Briefing Papers. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, General Technical Report, [https://www.fs.usda.gov/treesearch/pubs/37513 PNW-GTR-837], Portland, OR, 169p. [https://doi.org/10.2737/PNW-GTR-837 doi:10.2737/PNW-GTR-837] [//www.enviro.wiki/images/f/f2/Pnwgtr837.pdf Report pdf]</ref><ref>Acevedo-Whitehouse, K., and Duffus, A. L. J., 2009. Effects of environmental change on wildlife health. Philosophical Transactions of the Royal Society B: Biological Sciences, 364(1534), pp. 3429-3438. [https://doi.org/10.1098/rstb.2009.0128 doi: 10.1098/rstb.2009.0128]</ref><ref>Milligan, S. R., Holt, W. V., and Lloyd, R., 2009. Impacts of climate change and environmental factors on reproduction and development in wildlife. Philosophical Transactions of the Royal Society B: Biological Sciences, 364(1534), pp. 3313-3319. [https://doi.org/10.1098/rstb.2009.0175 doi: 10.1098/rstb.2009.0175] [//www.enviro.wiki/images/a/ad/Milligan2009.pdf Article pdf]</ref>. The ability to adapt to changing habitat conditions as a result of climate change will differ across individual species and between populations. Some wildlife species may be more vulnerable to climate change than other species (Figure 1). Vulnerability is often linked to particular life-history traits (e.g., specialized habitat needs or limited dispersal abilities, see Pacifici et al. 2015P<ref>Pacifici, M., Foden, W., and Visconti, P., 2015. Assessing species vulnerability to climate change. Nature Climate Change 5, pp. 215-225. [https://doi.org/10.1038/nclimate2448 doi: 10.1038/nclimate2448]</ref> for a review on species vulnerability to climate change) or genetic composition. For example, grassland birds may be more vulnerable to changing climate than forest birds as forests can buffer change more so than grasslands<ref>Jarzyna, M. A., Zuckerberg, B., Finley, A. O., and Porter, W. F., 2016. Synergistic effects of climate change and land cover: grassland birds are more vulnerable to climate change. Landscape Ecology, 31(10), pp. 2275-2290. [https://doi.org/10.1007/s10980-016-0399-1 doi: 10.1007/s10980-016-0399-1]</ref>. Projected changes in the climate will generally have adverse effects of wildlife populations<ref>IPCC, 2001. Climate Change 2001: Synthesis Report. A Contribution of Working Groups I, II, and III to the Third Assessment Report of the Intergovernmental Panel on Climate Change. R.T. Watson and the Core Writing Team (eds.). Cambridge University Press, Cambridge, United Kingdom, and New York, NY, USA, 398p. [//www.enviro.wiki/images/5/56/IPCC2001.pdf Report pdf]</ref>, though there are some species coping with climate change or benefitting from environmental change.  For example, American kestrels (Falco sparverius) have shifted their breeding phenology to earlier in the year and may now raise two broods of young within a breeding season<ref>Smith, S. H., Steenhof, K., McClure, C. J. W., and Heath, J. A. 2017. Earlier nesting by generalist predatory bird is associated with human responses to climate change. Journal of Animal Ecology, 86(1), pp. 98-107. [https://doi.org/10.1111/1365-2656.12604 doi: 10.1111/1365-2656.12604] [//www.enviro.wiki/images/5/5d/Smith2017.pdf Article pdf]</ref>.
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[https://doi.org/10.2307/176545 doi: 10.2307/176545]</ref>.  
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[[File:PowersFig2.png|thumb|900px|right| Figure 2. Conceptual diagram showing how increased water temperatures and pCO₂ (partial pressure of carbon dioxide) affect the early life stages of fish. Where arrow direction indicated increasing rate ( ↑ ), decreasing rate ( ↓ ), or both directions depending on other environmental variables ( ↕️ ).  Figure is from Pankhurst and Munday (2011)<ref name=":4" />.]]
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==The influence of climate change on wildlife and habitats==
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Climate change effects on wildlife include increases and changes in disease and pathogens distribution, patterns, and outbreaks in wildlife<ref>Bradley, B. A., Wilcove, D. S., and Oppenheimer, M., 2010. Climate change increases risk of plant invasion in the Eastern United States. Biological Invasions, 12, pp.1855-1872. [https://doi.org/10.1007/s10530-009-9597-y doi: 10.1007/s10530-009-9597-y]</ref><ref>Cudmore, T. J., Björklund, N., Carroll, A. L., and Lindgren, B. S., 2010. Climate change and the range expansion of an aggressive bark beetle: evidence of higher beetle reproduction in naïve host tree populations. Journal of Applied Ecology, 47(5), pp. 1036-1043. [https://doi.org/10.1111/j.1365-2664.2010.01848.x doi: 10.1111/j.1365-2664.2010.01848.x] [//www.enviro.wiki/images/b/b3/Cudmore2010.pdf Article pdf]</ref><ref>Price, S. J., Leung, W. T., Owen, C. J., Puschendorf, R., Sergeant, C., Cunningham, A. A., Balloux, F., Garner, T. W., and Nichols, R. A., 2019. Effects of historic and projected climate change on the range and impacts of an emerging wildlife disease. Global Change Biology, 25(8), pp. 2648-2660. [https://doi.org/10.1111/gcb.14651 doi: 10.1111/gcb.14651]</ref>; changes in range distributions and shifts in latitudinal and elevational gradients; changes in phenology or the timing of life cycle events that may create phenological mismatches<ref>Renner, S. S., and Zohner, C. M., 2018. Climate Change and Phenological Mismatch in Trophic Interactions Among Plants, Insects, and Vertebrates. Annual Review of Ecology, Evolution, and Systematics, 49, pp. 165-182. [https://doi.org/10.1146/annurev-ecolsys-110617-062535 doi: 10.1146/annurev-ecolsys-110617-062535]</ref>; and extinction or population reduction<ref name=":2">Urban, M. C., 2015. Accelerating extinction risk from climate change. Science, 348(6234), pp. 571-573. [https://doi.org/ 10.1126/science.aaa4984 doi: 10.1126/science.aaa4984] [//www.enviro.wiki/images/1/15/Urban2015.pdf Article pdf]</ref>. The effects of climate change across a species’ range will most likely not be homogenous, meaning it can vary substantially, especially if a species’ range spans across different continents as exhibited by many migratory birds. Other changes in habitat include shifting vegetation (i.e., tree-lines are shifting to higher elevations), changes in nutrients in plants, earlier snowmelt and run-off, increase in invasive species, warming of streams and rivers, reduction or degradation of habitat (i.e., glacial melt), and an increase in large wildfires<ref>Barbero, R., Abatzoglou, J. T., Larkin, N. K., Kolden, C. A., and Stocks, B., 2015. Climate change presents increased potential for very large fires in the contiguous United States. International Journal of Wildland Fire, 24(7), pp. 892-899. [https://doi.org/10.1071/WF150830128 doi: 10.1071/WF150830128] [//www.enviro.wiki/images/0/08/Barbero2015.pdf Article pdf]</ref>and droughts<ref>Schlaepfer, D. R., Bradford, J. B., Lauenroth, W. K., Munson, S. M., Tietjen, B., Hall, S. A., Wilson, S. D., Duniway, M. C., Jia, G., Pyke, D. A., Lkhagva, A., and Jamiyansharav, K., 2017. Climate change reduces extent of temperate drylands and intensifies drought in deep soils. Nature Communications, 8, pp. 14196. [https://doi.org/10.1038/ncomms14196 doi: 10.1038/ncomms14196]</ref>.
  
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Given that climate does influence vital rates, predicting how climate change will impact population growth requires evaluating matrix models reflecting changing transition rates. One approach is to take advantage of year to year variation in climate conditions and use models fit to data during different periods— perhaps corresponding to wet and dry years, or to cool and warm years<ref name=":3">
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Although climate change effects on wildlife often are linked to species-specific traits, there are general impacts associated with taxonomic groups<ref name=":1" />. For fish it can affect reproduction<ref name=":4">Pankhurst, N. W., and Munday, P. L., 2011. Effects of climate change on fish reproduction and early life history stages. Marine and Freshwater Research, 62(9), pp. 1015-1026. [https://doi.org/10.1071/MF10269 doi: 10.1071/MF10269] [[Special:FilePath/Pankhurst2011.pdf| Article pdf]]</ref>, growth, and recruitment<ref name=":3">Lynch, A. J., Myers, B. J. E., Chu, C., Eby, L. A., Falke, J. A., Kovach, R. P., Krabbenhoft, T. J., Kwak, T. J., Lyons, J., Paukert, C. P., and Whitney, J. E., 2016. Climate Change Effects on North American Inland Fish Populations and Assemblages. Fisheries, 41(7), pp. 346-361. [https://doi.org/10.1080/03632415.2016.1186016 doi: 10.1080/03632415.2016.1186016]</ref>(Figure 2). Cold-water fish such as inland North American species are highly affected with the warming of streams and rivers<ref name=":3" />. Amphibians are highly sensitive to their environment and changes in temperature and moisture can affect development, range, abundance, and phenology<ref>Blaustein, A.R., Walls, S.C., Bancroft, B.A., Lawler, J.J., Searle, C.L., and Gervasi, S.S., 2010. Direct and Indirect Effects of Climate Change on Amphibian Populations. Diversity, 2(2), pp. 281-313.[https://doi.org/10.3390/d2020281 doi: 10.3390/d2020281] [[Media:Blaustein 2010.pdf | Article pdf]]</ref><ref name=":1" /><ref>Ficetola, G. F., and Maiorano, L., 2016. Contrasting effects of temperature and precipitation change on amphibian phenology, abundance and performance. Oecologia, 181(3), pp. 683-693. [https://doi.org/10.1007/s00442-016-3610-9 doi: 10.1007/s00442-016-3610-9]</ref>. In reptiles, climate change effects can alter thermoregulation patterns, affect female reproduction and in some species, change sex ratios with increasing temperature<ref name=":1" />. Furthermore, for many bird species the timing of migration and other phenological events are affected by climate change<ref>Crick, H. Q. P., 2004. The impact of climate change on birds. Ibis, 146(s1), pp. 48-56. [https://doi.org/10.1111/j.1474-919X.2004.00327.x doi: 10.1111/j.1474-919X.2004.00327.x] [//www.enviro.wiki/images/c/c9/Crick2004.pdf Article pdf]</ref>. Range shifts, growth size, and survival are linked to climate change for mammals<ref name=":1" />. Arctic marine mammals are closely linked to sea ice dynamics and a changing climate will affect these dynamics<ref>Kovacs, K. M., Lydersen, C., Overland, J. E., and Moore, S. E., 2011. Impacts of changing sea-ice conditions on Arctic marine mammals. Marine Biodiversity, 41, pp. 81-194. [https://doi.org/10.1007/s12526-010-0061-0 doi: 10.1007/s12526-010-0061-0]</ref>. Therefore, it is increasingly important for conservation and management plans to consider the effects of climate change on wildlife and habitat for the geographic location<ref>Mawdsley, J. R., O’Malley, R., and Ojima, D. S. 2009. A Review of Climate-Change Adaptations Strategies For Wildlife Management and Biodiversity Conservation. Conservation Biology, 23(5), pp. 1080-1089. [https://doi.org/10.1111/j.1523-1739.2009.01264.x doi: 10.1111/j.1523-1739.2009.01264.x]</ref>.
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Gaillard, J.M., Mark Hewison, A.J., Klein, F., Plard, F., Douhard, M., Davison, R., and Bonenfant, C., 2013. How does climate change influence demographic processes of widespread species? Lessons from the comparative analysis of contrasted populations of roe deer. Ecology Letters, 16(1), pp.48-57. [https://doi.org/10.1111/ele.12059 doi: 10.1111/ele.12059] [//www.enviro.wiki/images/5/59/Gaillard2013.pdf Article pdf]</ref>. The same technique can be used to evaluate changes in the timing of seasonal shifts in climate. For example, Gaillard et al.<ref name=":3" /> used matrix models to show that the impact of earlier onset of spring weather on roe deer (''Capreolus capreolus'') was almost entirely due to differences in fecundity between periods of earlier and later spring weather conditions. This observation highlights another key assumption about using sensitivity or elasticity values to determine monitoring or management priorities with respect to climate change: that climate-driven changes in different vital rates are of the same, relatively small, magnitude.
 
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A more general approach better suited to using population models to predict climate change is to make matrix elements functions of climate variables<ref name=":4" /><ref name=":5">Merow, C., Latimer, A.M., Wilson, A.M., McMahon, S.M., Rebelo, A.G., and Silander Jr, J.A., 2014. On using integral projection models to generate demographically driven predictions of species' distributions: development and validation using sparse data. Ecography, 37(12), pp. 1167-1183.[https://doi.org/10.1111/ecog.00839 doi: 10.1111/ecog.00839] [//www.enviro.wiki/images/1/12/Merow2014.pdf Article pdf]</ref>. This approach has additional advantage that other factors influencing vital rates, such as individual size, density dependence, local soil conditions or management activities, can be readily incorporated and corresponding model parameters efficiently estimated from relatively sparse data<ref>Gross, K., Morris, W.F., Wolosin, M.S., and Doak, D.F., 2006. Modeling vital rates improves estimation of population projection matrices. Population Ecology, 48(1), pp. 79-89. [https://doi.org/10.1007/s10144-005-0238-8 doi: 10.1007/s10144-005-0238-8] [//www.enviro.wiki/images/6/6d/Gross2006.pdf Article pdf]</ref> and iterating the population projection forward through time with climate variables changing each time step as predicted by downscaled climate projection models<ref name=":5" />.
 
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===Individual based models===
 
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Individual based simulation models provide an even more general modeling framework. In an individual based model the fate of each individual is tracked through time. Movement and other individual behaviors can be directly incorporated into spatially explicit individual based models, which facilitates looking at interactions between the effects of climate and microhabitat characteristic, on future vital rates. Individual based models are also useful for directly incorporating the effects of demographic stochasticity in small populations. The use of individual based models (also referred to as agent based models) has been facilitated by the development of user-friendly software such as VORTEX <ref>Lacy, R.C., and Pollak, J.P., 2014. Vortex: A Stochastic Simulation of the Extinction Process. Version 10.0. Chicago Zoological Society, Brookfield, Illinois, USA. [https://scti.tools/vortex/ Vortex software]</ref>and NetLogo<ref>Railsback, S.F., and Grimm, V., 2011. Agent-Based and Individual-Based Modeling: A Practical Introduction. Princeton University Press. ISBN: 978-069119083-9</ref>. Hudgens et al.<ref name=":6" /> used VORTEX to simulate San Clemente sage sparrow (since renamed San Clemente Bell's sparrow) population dynamics to highlight the impacts of introduced predators and potential of more frequent drought under different management scenarios. Social interactions often require custom models, such as the model developed to inform management of red-cockaded woodpeckers (''Picoides borealis)''<ref> Walters, J.R., Crowder, L.B., and Priddy, J.A., 2002. Population viability analysis for red‐cockaded woodpeckers using an individual‐based model. Ecological Applications, 12(1), pp. 249-260.[https://doi.org/10.2307/3061150 doi: 10.2307/3061150]</ref><ref>Letcher, B.H., Priddy, J.A., Walters, J.R., and Crowder, L.B., 1998. An individual-based, spatially-explicit simulation model of the population dynamics of the endangered red-cockaded woodpecker, Picoides borealis. Biological Conservation, 86(1), pp.1-14. [https://doi.org/10.1016/S0006-3207(98)00019-6 doi: 10.1016/S0006-3207(98)00019-6]</ref>.
 
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==Promises and Pitfalls of Population Models==
 
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The primary purpose of population models is to integrate our knowledge of a species' ecology. As such, the axiom "garbage-in, garbage-out" applies to population models. Population models that mischaracterize a species' biology are doomed to make inaccurate predictions. Two common modeling mistakes are to oversimplify population structure and failing to account for correlations in how vital rates vary from year to year.
 
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===Oversimplifying population structure===
 
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Oversimplifying population structure often leads to observed variation in population growth rate being misattributed to factors of management interest. Especially in small populations, failing to account for age structure can lead to models that poorly reflect reality.
 
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For example, following removal of golden eagles (''Aquila chrysaetos'') from Santa Cruz Island, island foxes (''Urocyon littoralis'') expanded rapidly from 2000 to 2005. A subsequent decline in population growth rates by 2008 as fox numbers approached 1000 animals<ref>Coonan, T.J., Schwemm, C.A., and Garcelon, D.K., 2010. Decline and Recovery of The Island Fox: A case Study for Population Recovery. Cambridge University Press. eISBN: 9780511781612 [https://doi.org/10.1017/CBO9780511781612 doi: 10.1017/CBO9780511781612]</ref> led to speculation among agencies responsible their recovery that the population was approaching carrying capacity. However, an age-structured matrix model showed that changes in the proportion of foxes in different age classes could also lead to the same reduction in population growth rates<ref>Hudgens, B., Ferrara, F., and Garcelon, D., 2008. Digital radio-telemetry monitoring of San Nicolas Island foxes. Final Report. Department of Defense. December 2008. [//www.enviro.wiki/images/0/07/Hudgens2008.pdf Report pdf]</ref>and subsequent surveys have supported the latter explanation. The near ubiquity of age-related population structure in creatures with lifespans longer than 1-2 years contributes to the widespread use of matrix models in conservation.
 
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===Oversimplifying social structure===
 
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Social structures represent another aspect of the biology of many species that, if not properly accounted for, can lead to model failure. In a dramatic example, Zeigler and Walters<ref>
 
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Zeigler, S.L., and Walters, J.R., 2014. Population models for social species: lessons learned from models of Red‐cockaded Woodpeckers (Picoides borealis). Ecological Applications, 24(8), pp. 2144-2154.
 
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[https://doi.org/10.1890/13-1275.1 doi: 10.1890/13-1275.1]</ref>compared predicted population trajectories for red-cockaded woodpeckers from four population models to observed population dynamics in the Sandhills region of North Carolina. Population projections from the two models that did not incorporate social structure in the form of adult helpers at breeding colonies performed significantly worse than the two models incorporating social structure, even when it was a more complex model.
 
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===Correlations among vital rates===
 
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[[File: HudgensFig1.png|thumb|700px|left | Figure 1. Comparison of population model predictions with and without compensatory breeding. (Left panel) Population trajectory over 12 years (solid line) plotted with predicted trajectories from models with (dashed line, solid circles) and without (dashed line, empty circles) compensatory breeding. Note how model with compensatory breeding captures observed population increase from 2007-2008, while model without does not. (Right panel) Predicted risk of the population dipping below a critical threshold of 500 birds based is higher in simulations without compensatory breeding than in simulations incorporating compensatory breeding.]]
 
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When using population models to predict a species response to climate change, it is particularly important to consider correlations among vital rates. In many species vital rates are inexorably linked such that changes in one are always associated with changes in others<ref>Stearns, S.C., 1989. Trade-offs in life-history evolution. Functional ecology, 3(3), pp.259-268.
 
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[https://doi.org/10.2307/2389364 doi: 10.2307/2389364]</ref>. This kind of tradeoff may also lead to correlations between years in fecundity or growth. A population model presented by Hudgens et al.<ref name=":6" />predicted a high risk of extinction for San Clemente Bell's sparrows associated in part with lack of reproduction during drought years. However, field biologists monitoring the population for the U.S. Navy have subsequently reported extremely high reproductive output in years following drought years. Incorporating this compensatory breeding into the population model substantially lowers both the predicted risk of extinction and predicted potential impact of increased drought frequency on the population (Figure 1).
 
  
 
==References==
 
==References==
 
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==See Also==

Latest revision as of 14:07, 25 April 2022

Climate change affects both terrestrial[1] and aquatic biomes[2] causing significant effects on ecosystem functions and biodiversity[3]. Climate change is affecting several key ecological processes and patterns that will have cascading impacts on wildlife and habitat[4]. For example, sea-level rise, changes in the timing and duration of growing seasons, and changes in primary production are mainly driven by changes to global environmental variables (e.g., temperature and atmospheric CO2). Climate-induced changes in the environment ultimately impact wildlife population abundance and distributions.

Related Article(s):


Contributor(s): Dr. Breanna F. Powers and Dr. Julie A. Heath


Key Resource(s):

  • Global climate change and wildlife in North America. Wildlife Society Technical Review 04-2[4]
  • Impacts of climate change on the future of biodiversity[3]

Introduction

Figure 1. The predicted extinction risk (by percentage with 95% CIs) from climate change by different regions, colors represent a gradient from least to most extinction risks (green to red) based upon the number of relevant studies (n). Figure is from Urban (2015)[5]. Reprinted with permission from AAAS. Any use of this figure requires the prior written permission of AAAS

Global climate change will affect ecosystem functions and cycles such as nutrient, hydraulic, and carbon cycles, changing aspects of environmental conditions such as temperature, soil moisture, and precipitation[6][7]. Wildlife species are adapted to their environments and changes to the environment and habitat conditions will mediate effects, either directly or indirectly, on species survival, fecundity and ultimately population persistence[8][9][10]. The ability to adapt to changing habitat conditions as a result of climate change will differ across individual species and between populations. Some wildlife species may be more vulnerable to climate change than other species (Figure 1). Vulnerability is often linked to particular life-history traits (e.g., specialized habitat needs or limited dispersal abilities, see Pacifici et al. 2015P[11] for a review on species vulnerability to climate change) or genetic composition. For example, grassland birds may be more vulnerable to changing climate than forest birds as forests can buffer change more so than grasslands[12]. Projected changes in the climate will generally have adverse effects of wildlife populations[13], though there are some species coping with climate change or benefitting from environmental change. For example, American kestrels (Falco sparverius) have shifted their breeding phenology to earlier in the year and may now raise two broods of young within a breeding season[14].

Figure 2. Conceptual diagram showing how increased water temperatures and pCO₂ (partial pressure of carbon dioxide) affect the early life stages of fish. Where arrow direction indicated increasing rate ( ↑ ), decreasing rate ( ↓ ), or both directions depending on other environmental variables ( ↕️ ). Figure is from Pankhurst and Munday (2011)[15].

The influence of climate change on wildlife and habitats

Climate change effects on wildlife include increases and changes in disease and pathogens distribution, patterns, and outbreaks in wildlife[16][17][18]; changes in range distributions and shifts in latitudinal and elevational gradients; changes in phenology or the timing of life cycle events that may create phenological mismatches[19]; and extinction or population reduction[5]. The effects of climate change across a species’ range will most likely not be homogenous, meaning it can vary substantially, especially if a species’ range spans across different continents as exhibited by many migratory birds. Other changes in habitat include shifting vegetation (i.e., tree-lines are shifting to higher elevations), changes in nutrients in plants, earlier snowmelt and run-off, increase in invasive species, warming of streams and rivers, reduction or degradation of habitat (i.e., glacial melt), and an increase in large wildfires[20]and droughts[21].

Although climate change effects on wildlife often are linked to species-specific traits, there are general impacts associated with taxonomic groups[4]. For fish it can affect reproduction[15], growth, and recruitment[22](Figure 2). Cold-water fish such as inland North American species are highly affected with the warming of streams and rivers[22]. Amphibians are highly sensitive to their environment and changes in temperature and moisture can affect development, range, abundance, and phenology[23][4][24]. In reptiles, climate change effects can alter thermoregulation patterns, affect female reproduction and in some species, change sex ratios with increasing temperature[4]. Furthermore, for many bird species the timing of migration and other phenological events are affected by climate change[25]. Range shifts, growth size, and survival are linked to climate change for mammals[4]. Arctic marine mammals are closely linked to sea ice dynamics and a changing climate will affect these dynamics[26]. Therefore, it is increasingly important for conservation and management plans to consider the effects of climate change on wildlife and habitat for the geographic location[27].

References

  1. ^ Diffenbaugh, N. S. and Field, C. B., 2013. Changes in Ecologically Critical Terrestrial Climate Conditions. Science, 341(6145), pp. 486-492. doi: 10.1126/science.1237123
  2. ^ Hoegh-Guldberg, O., and Bruno, J. F., 2010. The Impact of Climate Change on the World’s Marine Ecosystem. Science, 328(5985), pp. 1523-1528. doi: 10.1126/science.1189930
  3. ^ 3.0 3.1 Bellard, C., Berteslsmeier, C., Leadley, P., Thuiller, W., and Courchamp, F., 2012. Impacts of climate change on the future of biodiversity. Ecological Letters, 15(4), pp. 365-377. doi: 10.1111/j.1461-0248.2011.01736.x Article pdf
  4. ^ 4.0 4.1 4.2 4.3 4.4 4.5 Inkley, D. B., Anderson, M. G., Blaustein, A. R., Burkett, V. R., Felzer, B., Griffith, B., Price, J., and Root, T. L., 2004. Global Climate Change and Wildlife in North America. Wildlife Society Technical Review 04-2. The Wildlife Society, Bethesda, MD, 26 pp. Report pdf
  5. ^ 5.0 5.1 Urban, M. C., 2015. Accelerating extinction risk from climate change. Science, 348(6234), pp. 571-573. 10.1126/science.aaa4984 doi: 10.1126/science.aaa4984 Article pdf
  6. ^ Davidson, E. A. and Janssens, I. A., 2006. Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Nature, 440, pp. 165-173.doi: 10.1038/nature04514
  7. ^ Melillo, J. M., McGuire, A. D., Kicklighter, D. W., Moore, B., Vorosmarty, C. J., and Schloss, A. L., 1993. Global climate change and terrestrial net primary production. Nature, 363, pp. 234-240. doi: 10.1038/363234a0
  8. ^ Alig, R. J., Technical Coordinator, 2011. Effects of Climate Change on Natural Resources and Communities: A Compendium of Briefing Papers. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, General Technical Report, PNW-GTR-837, Portland, OR, 169p. doi:10.2737/PNW-GTR-837 Report pdf
  9. ^ Acevedo-Whitehouse, K., and Duffus, A. L. J., 2009. Effects of environmental change on wildlife health. Philosophical Transactions of the Royal Society B: Biological Sciences, 364(1534), pp. 3429-3438. doi: 10.1098/rstb.2009.0128
  10. ^ Milligan, S. R., Holt, W. V., and Lloyd, R., 2009. Impacts of climate change and environmental factors on reproduction and development in wildlife. Philosophical Transactions of the Royal Society B: Biological Sciences, 364(1534), pp. 3313-3319. doi: 10.1098/rstb.2009.0175 Article pdf
  11. ^ Pacifici, M., Foden, W., and Visconti, P., 2015. Assessing species vulnerability to climate change. Nature Climate Change 5, pp. 215-225. doi: 10.1038/nclimate2448
  12. ^ Jarzyna, M. A., Zuckerberg, B., Finley, A. O., and Porter, W. F., 2016. Synergistic effects of climate change and land cover: grassland birds are more vulnerable to climate change. Landscape Ecology, 31(10), pp. 2275-2290. doi: 10.1007/s10980-016-0399-1
  13. ^ IPCC, 2001. Climate Change 2001: Synthesis Report. A Contribution of Working Groups I, II, and III to the Third Assessment Report of the Intergovernmental Panel on Climate Change. R.T. Watson and the Core Writing Team (eds.). Cambridge University Press, Cambridge, United Kingdom, and New York, NY, USA, 398p. Report pdf
  14. ^ Smith, S. H., Steenhof, K., McClure, C. J. W., and Heath, J. A. 2017. Earlier nesting by generalist predatory bird is associated with human responses to climate change. Journal of Animal Ecology, 86(1), pp. 98-107. doi: 10.1111/1365-2656.12604 Article pdf
  15. ^ 15.0 15.1 Pankhurst, N. W., and Munday, P. L., 2011. Effects of climate change on fish reproduction and early life history stages. Marine and Freshwater Research, 62(9), pp. 1015-1026. doi: 10.1071/MF10269 Article pdf
  16. ^ Bradley, B. A., Wilcove, D. S., and Oppenheimer, M., 2010. Climate change increases risk of plant invasion in the Eastern United States. Biological Invasions, 12, pp.1855-1872. doi: 10.1007/s10530-009-9597-y
  17. ^ Cudmore, T. J., Björklund, N., Carroll, A. L., and Lindgren, B. S., 2010. Climate change and the range expansion of an aggressive bark beetle: evidence of higher beetle reproduction in naïve host tree populations. Journal of Applied Ecology, 47(5), pp. 1036-1043. doi: 10.1111/j.1365-2664.2010.01848.x Article pdf
  18. ^ Price, S. J., Leung, W. T., Owen, C. J., Puschendorf, R., Sergeant, C., Cunningham, A. A., Balloux, F., Garner, T. W., and Nichols, R. A., 2019. Effects of historic and projected climate change on the range and impacts of an emerging wildlife disease. Global Change Biology, 25(8), pp. 2648-2660. doi: 10.1111/gcb.14651
  19. ^ Renner, S. S., and Zohner, C. M., 2018. Climate Change and Phenological Mismatch in Trophic Interactions Among Plants, Insects, and Vertebrates. Annual Review of Ecology, Evolution, and Systematics, 49, pp. 165-182. doi: 10.1146/annurev-ecolsys-110617-062535
  20. ^ Barbero, R., Abatzoglou, J. T., Larkin, N. K., Kolden, C. A., and Stocks, B., 2015. Climate change presents increased potential for very large fires in the contiguous United States. International Journal of Wildland Fire, 24(7), pp. 892-899. doi: 10.1071/WF150830128 Article pdf
  21. ^ Schlaepfer, D. R., Bradford, J. B., Lauenroth, W. K., Munson, S. M., Tietjen, B., Hall, S. A., Wilson, S. D., Duniway, M. C., Jia, G., Pyke, D. A., Lkhagva, A., and Jamiyansharav, K., 2017. Climate change reduces extent of temperate drylands and intensifies drought in deep soils. Nature Communications, 8, pp. 14196. doi: 10.1038/ncomms14196
  22. ^ 22.0 22.1 Lynch, A. J., Myers, B. J. E., Chu, C., Eby, L. A., Falke, J. A., Kovach, R. P., Krabbenhoft, T. J., Kwak, T. J., Lyons, J., Paukert, C. P., and Whitney, J. E., 2016. Climate Change Effects on North American Inland Fish Populations and Assemblages. Fisheries, 41(7), pp. 346-361. doi: 10.1080/03632415.2016.1186016
  23. ^ Blaustein, A.R., Walls, S.C., Bancroft, B.A., Lawler, J.J., Searle, C.L., and Gervasi, S.S., 2010. Direct and Indirect Effects of Climate Change on Amphibian Populations. Diversity, 2(2), pp. 281-313.doi: 10.3390/d2020281 Article pdf
  24. ^ Ficetola, G. F., and Maiorano, L., 2016. Contrasting effects of temperature and precipitation change on amphibian phenology, abundance and performance. Oecologia, 181(3), pp. 683-693. doi: 10.1007/s00442-016-3610-9
  25. ^ Crick, H. Q. P., 2004. The impact of climate change on birds. Ibis, 146(s1), pp. 48-56. doi: 10.1111/j.1474-919X.2004.00327.x Article pdf
  26. ^ Kovacs, K. M., Lydersen, C., Overland, J. E., and Moore, S. E., 2011. Impacts of changing sea-ice conditions on Arctic marine mammals. Marine Biodiversity, 41, pp. 81-194. doi: 10.1007/s12526-010-0061-0
  27. ^ Mawdsley, J. R., O’Malley, R., and Ojima, D. S. 2009. A Review of Climate-Change Adaptations Strategies For Wildlife Management and Biodiversity Conservation. Conservation Biology, 23(5), pp. 1080-1089. doi: 10.1111/j.1523-1739.2009.01264.x

See Also

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