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with pervasive and sometimes unexpected consequences that propagate through and affect bothbiological and human communities.

CLIMATE CHANGE

Biodiversity redistribution underclimate change: Impacts on
ecosystems and human well-being
Gretta T. Pecl,* Miguel B. Araújo,† Johann D. Bell, Julia Blanchard, Timothy C. Bonebrake,
I-Ching Chen, Timothy D. Clark, Robert K. Colwell, Finn Danielsen, Birgitta Evengård,Lorena Falconi, Simon Ferrier, Stewart Frusher, Raquel A. Garcia, Roger B. Griffis,
Alistair J. Hobday, Charlene Janion-Scheepers, Marta A. Jarzyna, Sarah Jennings,
Jonathan Lenoir, Hlif I. Linnetved, Victoria Y. Martin, Phillipa C. McCormack,
Jan McDonald, Nicola J. Mitchell, Tero Mustonen, John M. Pandolfi, Nathalie Pettorelli,
Ekaterina Popova, Sharon A. Robinson, Brett R. Scheffers, Justine D. Shaw,
Cascade J. B. Sorte, Jan M. Strugnell, Jennifer M. Sunday, Mao-Ning Tuanmu,
Adriana Vergés, Cecilia Villanueva, Thomas Wernberg, Erik Wapstra, Stephen E. Williams
BACKGROUND: The success of human socie-

ties depends intimately on the living components of natural and managed systems. Although
the geographical range limits of species are dynamic and fluctuate over time, climate change
is impelling a universal redistribution of life on
Earth. For marine, freshwater, and terrestrial
species alike, the first response to changing
climate is often a shift in location, to stay within
preferred environmental conditions. At thecooler extremes of their distributions, species

are moving poleward, whereas range limits arecontracting at the warmer range edge, where
temperatures are no longer tolerable. On land,
species are also moving to cooler, higher elevations; in the ocean, they are moving to colder
water at greater depths. Because different species
respond at different rates and to varying degrees,
key interactions among species are often disrupted, and new interactions develop. These
idiosyncrasies can result in novel biotic communities and rapid changes in ecosystem functioning,

ADVANCES: At a time when the world is anticipating unprecedented increases in human population growth and demands, the ability of natural
ecosystems to deliver ecosystem services is being
challenged by the largest climate-driven global
redistribution of species since the Last Glacial
Maximum. We demonstrate the serious consequences of this species redistribution for economic development, livelihoods, food security,
human health, and culture,
ON OUR WEBSITE
and we document feedbacks on climate itself. As
Read the full article
with other impacts of cliat http://dx.doi.
org/10.1126/
mate change, species range
science.aai9214
shifts will leave “winners”..................................................
and “losers” in their wake,radically reshaping the pattern of human wellbeing between regions and different sectors
and potentially leading to substantial conflict.
The pervasive impacts of changes in species
distribution transcend single systems or dimensions, with feedbacks and linkages between multiple interacting scales and throughwhole ecosystems, inclusive of humans. We argue that the negative effects of climate change
cannot be adequately anticipated or preparedfor unless species responses are explicitly included in decision-making and global strategic frameworks.
◥

OUTLOOK: Despite mounting evidence for thepervasive and substantial impacts of a climatedriven redistribution of Earth’s species, current
global goals, policies, and international agreements fail to account for these effects. With thepredicted intensification of species movements
and their diverse societal and environmental impacts, awareness of species “on the move” should
be incorporated into local, regional, and global
assessments as standard practice. This will raise
hope that future targets—whether they be global
sustainability goals, plans for regional biodiversity maintenance, or local fishing or forestry harvest strategies—can be achievable and that society
is prepared for a world of universal ecologicalchange. Human society has yet to appreciate theimplications of unprecedented species redistribution for life on Earth, including for human
lives. Even if greenhouse gas emissions stopped
today, the responses required in human systems
to adapt to the most serious effects of climatedriven species redistribution would be massive.
Meeting these challenges requires governance that
can anticipate and adapt to changing conditions,as well as minimize negative consequences.
As the global climate changes, human well-being, ecosystem function, and even climate
itself are increasingly affected by the shifting geography of life. Climate-driven changes in species
distributions, or range shifts, affect human well-being both directly (for example, through emerging
diseases and changes in food supply) and indirectly (by degrading ecosystem health). Some range shifts
even create feedbacks (positive or negative) on the climate system, altering the pace of climate change.
Pecl et al., Science 355, 1389 (2017)31 March 2017▪

The list of author affiliations is available in the full article online.
*Corresponding author. Email: gretta.pecl@utas.edu.au†All authors after the first author are listed alphabetically.
Cite this article as G. T. Pecl et al., Science 355, eaai9214(2017). DOI: 10.1126/science.aai9214

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REVIEW SUMMARYREVIEW◥

CLIMATE CHANGE

Biodiversity redistribution underclimate change: Impacts on
ecosystems and human well-being
Gretta T. Pecl,1,2* Miguel B. Araújo,3,4,5† Johann D. Bell,6,7 Julia Blanchard,1,2
Timothy C. Bonebrake,8 I-Ching Chen,9 Timothy D. Clark,1,10 Robert K. Colwell,5,11,12,13
Finn Danielsen,14 Birgitta Evengård,15 Lorena Falconi,16 Simon Ferrier,17
Stewart Frusher,1,2 Raquel A. Garcia,18,19 Roger B. Griffis,20 Alistair J. Hobday,2,21
Charlene Janion-Scheepers,22 Marta A. Jarzyna,23 Sarah Jennings,2,24
Jonathan Lenoir,25 Hlif I. Linnetved,26 Victoria Y. Martin,27 Phillipa C. McCormack,28
Jan McDonald,2,28 Nicola J. Mitchell,29 Tero Mustonen,30 John M. Pandolfi,31Nathalie Pettorelli,32 Ekaterina Popova,33 Sharon A. Robinson,34 Brett R. Scheffers,35
Justine D. Shaw,36 Cascade J. B. Sorte,37 Jan M. Strugnell,38,39 Jennifer M. Sunday,40
Mao-Ning Tuanmu,41 Adriana Vergés,42 Cecilia Villanueva,1,2 Thomas Wernberg,29,43
Erik Wapstra,44 Stephen E. Williams16
Distributions of Earth’s species are changing at accelerating rates, increasingly driven by humanmediated climate change. Such changes are already altering the composition of ecologicalcommunities, but beyond conservation of natural systems, how and why does this matter? We
review evidence that climate-driven species redistribution at regional to global scales affects
ecosystem functioning, human well-being, and the dynamics of climate change itself. Productionof natural resources required for food security, patterns of disease transmission, and processesof carbon sequestration are all altered by changes in species distribution. Consideration ofthese effects of biodiversity redistribution is critical yet lacking in most mitigation andadaptation strategies, including the United Nation’s Sustainable Development Goals.

The history of life on Earth is closely associated with environmental change on multiple spatial and temporal scales (1). A criticalcomponent of this association is the capacity
for species to shift their distributions in response to tectonic, oceanographic, or climatic events(2). Observed and projected climatic changes forthe 21st century, most notably global warming,
are comparable in magnitude to the largest globalchanges in the past 65 million years (3, 4). The
combined rate and magnitude of climate change
is already resulting in a global-scale biological re-sponse. Marine, freshwater, and terrestrial organisms are altering distributions to stay within theirpreferred environmental conditions (5–8), and species are likely changing distributions more rapidly
than they have in the past (9). Unlike the introduction of non-native species, which tends to be
idiosyncratic and usually depends on humanmediated transport, climate-driven redistribution is
ubiquitous, follows repeated patterns, and is poised
to influence a greater proportion of Earth’s biota.
This redistribution of the planet’s living organisms
is a substantial challenge for human society.
Despite agreements to curb greenhouse gas
emissions, the climate will continue to change forat least the next several hundred years, given theinertia of the oceanic and atmospheric circulation
systems (10), and species will continue to respond,
often with unpredictable consequences. Since 1880,
there has been an average warming of 0.85°C globally (10), resulting in well-documented shifts in species distributions with far-reaching implications
for human societies, yet governments have agreed
to accept more than double this amount of warming in the future (e.g., the 2°C target from the Paris
Conference of Parties 21). Moreover, current global commitments will only limit warming to 2.7° to
3.7°C, more than three to four times the warming
already experienced (11). To date, all key international discussions and agreements regarding climatechange have focused on the direct socioeconomic
implications of emissions reduction and on funding
mechanisms; shifting natural ecosystems have notyet been considered in detail.
Here we review the consequences of climatedriven species redistribution for economic development and the provision of ecosystem services,
including livelihoods, food security, and culture,
as well as for feedbacks on the climate itself (Fig.
1 and table S1). We start by examining the impactsof climate-driven species redistribution on ecosystem health, human well-being, and the climate
system, before highlighting the governance challenges these impacts individually and collectively

1

Institute for Marine and Antarctic Studies, Hobart, Tasmania 7001, Australia. 2Centre for Marine Socioecology, Hobart, Tasmania 7001, Australia. 3Department of Biogeography and Global
Change, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas, 28006 Madrid, Spain. 4Centro de Investigação em Biodiversidade e Recursos Geneticos,
Universidade de Évora, 7000-890 Évora, Portugal. 5Department of Biology, Center for Macroecology, Evolution and Climate, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen
O, Denmark. 6Australian National Centre for Ocean Resources and Security, University of Wollongong, New South Wales 2522, Australia. 7Betty and Gordon Moore Center for Science andOceans, Conservation International, Arlington, VA 22202, USA. 8School of Biological Sciences, The University of Hong Kong, Hong Kong SAR, China. 9Department of Life Sciences, National Cheng
Kung University, Tainan 701, Taiwan, Republic of China. 10Commonwealth Scientific and Industrial Research Organization (CSIRO) Agriculture and Food, Hobart, Tasmania 7000, Australia.
11
Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269, USA. 12University of Colorado Museum of Natural History, Boulder, CO 80309, USA.
13
Departmento de Ecologia, Universidade Federal de Goiás, CP 131, 74.001-970 Goiânia, Goiás, Brazil. 14NORDECO, Copenhagen DK-1159, Denmark. 15Division of Infectious Diseases, Departmentof Clinical Microbiology, Umea University, 90187 Umea, Sweden. 16College of Marine and Environmental Science, James Cook University, Townsville, Queensland 4811, Australia. 17CSIRO Land andWater, Canberra, Australian Capital Territory 2601, Australia. 18Centre for Statistics in Ecology, the Environment and Conservation, Department of Statistical Sciences, University of Cape Town,
Rondebosch 7701, Cape Town, South Africa. 19Centre for Invasion Biology, Department of Botany and Zoology, Faculty of Science, Stellenbosch University, Matieland 7602, South Africa.
20
National Oceanic and Atmospheric Administration (NOAA) Fisheries Service, Silver Spring, MD 20912, USA. 21CSIRO Oceans and Atmosphere, Hobart, Tasmania 7000, Australia. 22Monash
University, School of Biological Sciences, Clayton, Victoria 3800, Australia. 23Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06520, USA. 24Tasmanian School ofBusiness and Economics, University of Tasmania, Hobart, Tasmania 7001, Australia. 25EDYSAN (FRE 3498 CNRS-UPJV), Université de Picardie Jules Verne, 80037 Amiens Cedex 1, France.
26
Institute of Food and Resource Economics, Faculty of Science, University of Copenhagen, Rolighedsvej 25, DK-1958 Frederiksberg C, Denmark. 27School of Environment, Science andEngineering, Southern Cross University, Lismore, New South Wales 2480, Australia. 28Faculty of Law, University of Tasmania, Hobart, Tasmania 7001, Australia. 29School of Biological Sciences,
The University of Western Australia, Crawley, Western Australia 6009, Australia. 30Snowchange Cooperative, University of Eastern Finland, Joensuu, FIN 80100 Finland. 31School of Biological
Sciences, Autralian Research Council (ARC) Centre of Excellence for Coral Reef Studies, The University of Queensland, Brisbane, Queensland 4072, Australia. 32Institute of Zoology, ZoologicalSociety of London, Regent’s Park, NW1 4RY London, UK. 33National Oceanography Centre, University of Southampton Waterfront Campus, European Way, Southampton, SO14 3ZH, UK. 34Centre
for Sustainable Ecosystem Solutions, School of Biological Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia. 35Department of Wildlife Ecology and Conservation,
Institute of Food and Agricultural Sciences, University of Florida, Gainesville, FL 32611, USA. 36Centre for Biodiversity and Conservation Science, School of Biological Sciences, The University of
Queensland, St Lucia, Queensland 4072, Australia. 37Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA. 38Centre for Sustainable Tropical Fisheries
and Aquaculture, College of Science and Engineering, James Cook University, Townsville, 4811 Queensland, Australia. 39Department of Ecology, Environment and Evolution, School of Life
Sciences, La Trobe University, Melbourne, Victoria 3086, Australia. 40Biodiversity Research Centre, University of British Columbia, Vancouver, British Columbia V6T 1Z4, Canada. 41Biodiversity
Research Center, Academia Sinica, Taipei 115, Taiwan, Republic of China. 42Centre for Marine Bio-Innovation and Evolution and Ecology Research Centre, School of Biological, Earth andEnvironmental Sciences, University of New South Wales, Sydney, New South Wales 2052, Australia. 43Oceans Institute, The University of Western Australia, Perth, Western Australia 6009,
Australia. 44School of Biological Sciences, University of Tasmania, Hobart, Tasmania 7001, Australia.
*Corresponding author. Email: gretta.pecl@utas.edu.au †All authors after the first author are listed alphabetically.

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create. Critically, the pervasive effects of changes
in species distribution transcend single systems
or dimensions, with feedbacks and linkages among
multiple interacting spatial and temporal scales
and through entire ecosystems, inclusive of humans (Figs. 2 and 3). We conclude by considering
species redistribution in the context of Earth systems and sustainable development. Our Review
suggests that the negative effects of climate change
cannot be adequately mitigated or minimized unless species responses are explicitly included indecision-making and strategic frameworks.
Biological responses andecosystem healthSpecies are affected by climate in many ways, including range shifts, changes in relative abundance within species ranges, and subtler changes
in activity timing and microhabitat use (12, 13). The
geographic distribution of any species depends
upon its environmental tolerance, dispersal constraints, and biological interactions with otherspecies (14). As climate changes, species must
either tolerate the change, move, adapt, or facewhen factors other than temperature drive species
redistribution. For example, precipitation changes
or interspecific interactions can cause downward
elevation shifts as climate warms (20). Althoughspecies may adapt to changing climates, either
through phenotypic plasticity or natural selection(21), all species have limits to their capacity foradaptive response to changing environments (12),and these limits are unlikely to increase for species already experiencing warm temperatures close
to their tolerance limits (22).
The idiosyncrasies of species responses to climate change can result in discordant range shifts,
leading to novel biotic communities as species separate or come into contact in new ways (23). In turn,
altered biotic interactions hinder or facilitate further range shifts, often with cascading effects (24).
Changes in predation dynamics, herbivory, hostplant associations, competition, and mutualisms
can all have substantial impacts at the community level (16, 25). A case in point involves the expected effects of crabs invading the continental shelfhabitat of Antarctic seafloor echinoderms andmollusks—species that have evolved in the absence

extinction (15). Surviving species may thus haveincreased capacity to live in new locations or decreased ability to persist where they are currently
situated (13).
Shifts in species distributions across latitude,
elevation, and with depth in the ocean have been
extensively documented (Fig. 1). Meta-analyses
show that, on average, terrestrial taxa move poleward by 17 km per decade (5) and marine taxa by
72 km per decade (6, 16). Just as terrestrial species
on mountainsides are moving upslope to escapewarming lowlands (17), some fish species are driven deeper as the sea surface warms (18).
The distributional responses of some species
lag behind climate change (6, 8). Such lags can arise
from a range of factors, including species-specificphysiological, behavioral, ecological, and evolutionary responses (12). Lack of adequate habitat connectivity and access to microhabitats and associatedmicroclimates are expected to be critical in increasing exposure to macroclimatic warming and extreme heat events, thus delaying shifts of somespecies (19). Furthermore, distribution shifts areoften heterogeneous across geographic gradients

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Ecosystem

Governance

Climate feedbacks

Fig. 1. Climate-driven changes in the distribution of life on Earth are affecting ecosystem health, human well-being, and the dynamics of climate change, challenging local and regional systems of governance. Examples of documented and predicted climate-driven changes in the
distribution of species throughout marine, terrestrial, and freshwater systems of the globe in tropical, temperate, and polar regions are shown. Details of
the impacts associated with each of these changes in distribution are provided in table S1, according to the numbered key, and the links to specificSustainable Development Goals are given in table S2.
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of skeleton-crushing predators (26). The community impacts of shifting species can be of the sameor greater magnitude as the introduction of nonnative species (16), itself recognized as one ofthe primary drivers of biodiversity loss (27).
When species range shifts occur in foundationor habitat-forming species, they can have pervasive effects that propagate through entire communities (28). In some cases, the impacts are so severethat species redistribution alters ecosystem productivity and carbon storage. For example, climatedriven range expansion of mangroves worldwide,at the expense of saltmarsh habitat, is changing
local rates of carbon sequestration (29). The loss ofkelp-forest ecosystems in Australia and their replacement by seaweed turfs have been linked toincreases in herbivory by the influx of tropicalfishes, exacerbated by increases in water temperature beyond the kelp’s physiological tolerance
limits (30, 31). Diverse disruptions from theredistribution of species include effects on terrestrial productivity (32), impacts on marine community assembly (33), and threats to the healthof freshwater systems from widespread cyanobacteria blooms (34).
The effects on ecosystem functioning and condition arising from species turnover and changes
in the diversity of species within entire communities are less well understood. The redistributionof species may alter the community compositionin space and time (beta diversity), the number of
species co-occurring at any given location (alphadiversity), and/or the number of species found within a larger region (gamma diversity) (35). The diversity and composition of functional traits withincommunities may also change as a result of species range shifts (36), although changes in functional traits may occur through alterations in relative
abundance or community composition, withoutchanges in species richness. Increasingly, evidence
indicates that species diversity, which underlies
functional diversity, has a positive effect on themean level and stability of ecosystem functioning
at local and regional scales (37). It therefore appears likely that any changes in diversity resulting
from the redistribution of species will have indirectconsequences for ecosystem condition.
Extinction risk from climate change has been
widely discussed and contested (38–40), and predictions of extinction risk for the 21st century are
considerable (41). In some cases, upslope migration allows mountain-dwelling species to track
suitable climate, but topography and range loss
can sometimes trap species in isolated and eventually unsuitable habitats (42). The American pika
(Ochotona princeps) has been extirpated or severely diminished in some localities, signaling climateinduced extinction or at least local extirpation (43).
Complicated synergistic drivers or “extinction debt”—
a process in which functional extinction precedes
physical extinction—may make climate-induced
extinction seem a distant threat. However, the disappearance of the Bramble Cay melomys (Melomys
rubicola), an Australian rodent declared extinct due
to sea level rise (44), shows that anthropogenic climate change has already caused irreversible species loss.
Pecl et al., Science 355, eaai9214 (2017)31 March 2017

Notwithstanding the rich body of evidence fromthe response to climate change of species and ecosystems in the fossil record (45), understanding
more recent, persistent responses to climate change
usually requires several decades of data to rigorously assess pre- and postclimate change trends
at the level of species and ecosystems (46). Suchlong-term data sets for biological systems are rare,
and recent trends of declining funding undermine the viability of monitoring programs required
to document and respond to climate change.
Human well-being
The well-being of human societies is tied to thecapacity of natural and altered ecosystems to produce a wide range of goods and services. Humanwell-being, survival, and geographical distribution have always depended on the ability to respond to environmental change. The emergenceof early humans was likely conditioned by a capacity to switch prey and diets as changing climatic conditions made new resources available (47).
However, recent technological changes in agriculture, forestry, and fisheries have weakened the direct link between human migration and survival.
Now, human societies rely more on technologicaland behavioral innovation to accommodate human demography, trade and economics, and foodproduction to changing species distribution patterns. The redistributions of species are expected
to affect the availability and distribution of goods
and services for human well-being in a number ofways, and the relative immobility of many humansocieties, largely imposed by jurisdictional borders,has limited capacity to respond to environmental
change by migration.
Redistributions of species are likely to drive major changes in the supply of food and other products.
For example, the relative abundance of skipjack
tuna in the tropical Pacific, which underpins government revenue and food security for many small
island states, is expected to become progressively
greater in eastern areas of the western and central
Pacific Ocean, helping to offset the projected ubiquitous decline in the supply of fish from degradedcoral reefs in that region (48). Conversely, it is estimated that an average of 34% of European forest
lands, currently covered with valuable timber trees,such as Norway spruce, will be suitable only for
Mediterranean oak forest vegetation by 2100, resulting in much lower economic returns for forest
owners and the timber industry (49).
The indirect effects of climate change on foodwebs are also expected to compound the direct effects on crops. For example, the distribution andabundance of vertebrate species that control crop
pests are predicted to decline in European states,
where agriculture makes important contributions
to the gross domestic product (50). Shifts in thespatial distribution of agriculture will be required
to counter the impact of these combined direct
and indirect effects of changing climate. Geographic shifts in natural resource endowments and insystems supporting agriculture, forestry, fisheries,
and aquaculture will result in winners and losers,with many of the negative effects likely to occur indeveloping countries (51). A prime example is the

projected effect of climate change on the supply ofcoffee, with principal coffee-growing regions expected to shift (52).
Species range shifts are also affecting the intrinsic and economic values of recreation and tourism, in both negative and positive ways (53). The
buildup of jellyfish due to warmer temperatures in
a Mediterranean lagoon has had a negative effecton local economies linked to recreation, tourism,
and fishing (54). In southeast Australia, a rangeextending sea urchin has overgrazed macroalgae,
resulting in localized loss of up to 150 associatedtaxa and contributing to reduced catch limits for
popular recreational fisheries species dependenton large seaweed (55). Impacts have been positivein some contexts, such as the recent emergenceof highly prized species in recreational fishingareas (53).
Indirect effects from changes in species distributions that underpin society and culture can bedramatic. In the Arctic, changes in distributionsof fish, wild reindeer, and caribou are affecting thefood security, traditional knowledge systems, andendemic cosmologies of indigenous societies (Figs.
1 and 2) (7). In partial response, the Skolt Sámi in
Finland have introduced adaptation measures toaid survival of Atlantic salmon stocks faced withwarming waters and to maintain their spiritual
relationship with the species. These measures include increasing the catch of pike to reduce predation pressure on salmon. In the East Siberian
tundra, faced with melting permafrost, the Chukchi
people are struggling to maintain their traditional nomadic reindeer-herding practices (56)(Fig. 2). Citizen-recording of climate-induced changes
to complement assessments based on scientificsampling and remote sensing forms part of theirstrategy to maintain traditional practices.
Human health is also likely to be seriously affected by changes in the distribution and virulence of animal-borne pathogens, which alreadyaccount for 70% of emerging infections (57, 58).
Movement of mosquitoes in response to global
warming is a threat to health in many countries
through predicted increases in the number of known
and potentially new diseases (Fig. 3). Malaria, themost prevalent mosquito-borne disease, has long
been a risk for almost half of the world’s population, with more than 200 million cases recorded
in 2014 (59). Malaria is expected to reach new areas
with the poleward and elevational migration ofAnopheles mosquito vectors (60). Climate-relatedtransmission of malaria can result in epidemics
due to lack of immunity among local residents (59)and will challenge health systems at national andinternational scales, diverting public- and privatesector resources from other uses.
The winners and losers arising from the redistributions of species will reshape patterns of human well-being among regions and sectors ofindustry and communities (61). Those regionswith the strongest climate drivers, with the mostsensitive species, and where humans have the leastcapacity to respond will be among the most affected. Developing nations, particularly those near
the equator, are likely to experience greater climaterelated local extinctions due to poleward and3 of 9Downloaded from http://science.sciencemag.org/ on March 30, 2017

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in snow cover but also through a northward shiftof coniferous trees (Fig. 2). Pearson et al. (68)projected that by 2050, vegetation in the Arctic
will mostly shift from tundra (dominated bylichens and mosses with high albedo) to boreal
forest (dominated by coniferous trees with low
albedo). Additionally, the greenhouse effect may
be amplified by top-of-atmosphere radiative

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elevational range shifts (62) and will face greater
economic constraints. In some cases, species redistribution will also lead to substantial conflict—the recent expansion of mackerel into Icelandic
waters is a case in point (Fig.
1 and table S1). The mackerel
fishery in Iceland increased
from 1700 metric tons in 2006
to 120,000 metric tons in 2010,
resulting in “mackerel wars”
between Iceland and competing countries that have traditionally been allocated mackerel
quotas (63). Likewise, with upslope shift of climate zones in
the Italian Alps, intensified
conflict is anticipated between
recreation and biodiversity sectors. For example, climate-driven
contractions in the most valuable habitat for high-elevation
threatened bird species and forski trails are predicted to increase, along with an increasein the degree of overlap between the bird habitat and theareas most suitable for future
ski trail construction (64).
Climate feedbacks
Species redistributions are expected to influence climate
feedbacks via changes in albedo, biologically driven sequestration of carbon fromthe atmosphere to the deepsea (the “biological pump”),
and the release of greenhousegases (65). For instance, terrestrial plants affect albedo via
leaf area and color and regulate the global carbon cycle
through CO2 atmosphere-landexchanges. Similarly, CO 2
atmosphere-ocean exchanges
are biologically modulated by
CO2-fixing photosynthetic phytoplankton and by the biological pump that exports carbon
into deep ocean reservoirs (66).
The climate-driven shifts inspecies distributions most likely to affect biosphere feedbacks
involve redistribution of vegetation on land (Figs. 2 and 4)
and phytoplankton in the ocean.
Decreased albedo, arising fromthe combined effect of earlier
snowmelt and increasing shrub
density at high latitudes, already contributes to increased
net radiation and atmospheric heating, amplifying highlatitude warming (67). Thus,
continued warming will decrease the albedo in the Arctic, not only through a decline

Fig. 2. Species on the move drive greening of the Arctic. Changes in species distribution can lead to climate
feedbacks, changes in ecosystem services, and impacts on human societies, with feedbacks and linkages between eachof these dimensions, illustrated here through climate-driven changes in Arctic vegetation. See Fig. 4 for a more comprehensive description of the direct and indirect climate feedbacks. See also (10, 68, 69, 75, 106–110).

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tree dieback. These processes, in turn, increase
fuel loads and fire frequency (73), ultimately driving additional feedback through massive biomassburning and CO2 release. Finally, increased shrub
canopy cover at high latitudes may locally reduce
soil temperatures through a buffering effect (74),slowing the release of CO2 from permafrost degradation, thus potentially mitigating warming(75) (Fig. 2).
Redistribution of marine phytoplankton is expected to affect the ocean’s biological and carbonate pumps and the production of atmosphericaerosols. The subpolar North Atlantic, which isalready highly productive and stores ~25% of theocean’s anthropogenic CO2 (76), may experience
phytoplankton changes due to retreat of the

Arctic sea ice and strengthening of ocean stratification. These changes are expected to lead, respectively, to northward movement of productive areasand suppression of the spring bloom, substantially altering CO2 exchanges between the ocean andthe atmosphere at high latitudes (77), although
the net effect is uncertain. Rising temperatures
may also lead to changes in the composition ofdifferent plankton functional groups (78). Expectedchanges in the relative dominance of diatoms andcalcareous plankton can strongly affect the biological cycling of carbon. Such a change was a possible
contributor to CO2 differences between Pleistocene glacial and interglacial periods (79). Similarly, shifts from diatom- to flagellate-dominatedsystems in temperate latitudes and increased

Downloaded from http://science.sciencemag.org/ on March 30, 2017imbalance from enhanced evapotranspiration
associated with the greening of the Arctic (69).
At low latitudes, ongoing plant redistribution [e.
g., mangrove expansion and forest dieback (29)]potentially amplifies climate warming throughcarbon-cycle feedbacks (70). However, future
projections in the tropics are uncertain because ofa lack of close climatic analogs from which to
extrapolate (71).
Species redistribution at high latitudes also affects vegetation state indirectly through pests likedefoliators and bark beetles that are moving northward and upslope in boreal forests (72) (Figs. 1, 2,
and 4). The combined effects of increasing temperatures and droughts increase plant stress, thus
contributing to the severity of pest outbreaks andFig. 3. Mosquito species on the move as vectors of disease. Climate change has facilitated an increase in the distribution of disease vectors, withconsiderable human cost and associated governance challenges. The bars in the human well-being graphs represent the minimum and maximum ranges;
the boxes depict the 25th, 50th, and 75th percentiles of the distribution; and the circles represent outliers. See also (60, 111–117).
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microbial remineralization, both associated withwarming, are expected to reduce the efficiency ofthe biological pump and therefore affect atmospheric CO2 (80).
Temperature-related changes in phytoplankton distributions will also affect production ofdimethyl sulfide (DMS), which contributes sulfur
particles to the atmosphere and seeds cloud formation (81). These particles are expected to decrease surface temperature, but they may also act
as a greenhouse gas, so the net effect on climate
warming is not yet clear. There is no simple relationship between DMS production and phytoplankton biomass, chlorophyll concentration, or
primary production, which suggests a complex
regulation of DMS production by the whole marine planktonic ecosystem and the physical environment controlling it. Hence, current climate
models cannot give an estimate of the strengthor even the direction of the phytoplankton-DMSclimate feedback.
Climate-influenced links between terrestrial
and marine regions may also lead to species redistribution and climate feedbacks. For example,
episodic land-atmosphere-ocean deposition of iron(e.g., pulses of Sahara dust) produces phytoplankton blooms (82) and enhances carbon export via
the biological pump. Changes to the phytoplanktondriven drawdown of atmospheric CO2 may therefore
arise through changes in the spatial distribution ofiron deposition, which may be affected by changes indrought conditions, agricultural practices, and largescale atmospheric circulation (83). These complex
processes—not only driven by climate-induced species redistribution but also affecting the climate
system itself—need to be incorporated into climate models to improve future projections (65).
Governance challenges
The impacts of the global redistribution of species
on human welfare and ecosystem services require
new governance mechanisms for biodiversity conservation and management. A dynamic and multilevel legal and policy approach is needed to address
the effects of species range limits moving acrosslocal, national, and international jurisdictionalboundaries. The development of international
guidance where laws do not yet exist will need toaccount for different legal regimes, resources, andnational capacities.
Shifts in species distributions will require changes
in the objectives of conservation law, which havetraditionally emphasized in situ conservation andretention of historical conditions. Objectives should
acknowledge that species will move beyond theirtraditional ranges, that novel ecosystems will inevitably be created and that historic ecosystems
may disappear, as a consequence of such movements (84). The experience of transjurisdictional
managed relocations (conservation introductions
outside of historical ranges) may inform the development of risk assessment processes that mustnavigate the complex ethical challenges arisingfrom novel interactions (85) and risks of collateral damage (86). Moreover, communication among
relevant agencies throughout the new and former
ranges of shifting species is essential to avoid inPecl et al., Science 355, eaai9214 (2017)31 March 2017

Fig. 4. Climate feedbacks and processes driven by the redistribution of plant species at highlatitudes. Climate affects vegetation at high latitudes directly through climatic processes but also
indirectly through pests like defoliators and bark beetles that are moving northward and upslope inboreal forests. Some processes increase warming (red arrows), whereas others may serve todecrease warming (blue arrows). Increasing shrub canopy cover in the Arctic at high latitudes may
reduce soil temperatures locally through a buffering effect, potentially slowing down CO2 carbon
release due to permafrost degradation, thus acting to slow climate warming. However, greening of
the Arctic also decreases albedo, which accelerates warming.

vesting in protecting species in locations where
they are no longer viable and yet failing to manage them appropriately in their new ranges.
Legal instruments are typically slow to change
and often privilege the protection of property anddevelopment rights. Although this inertia provides certainty and stability, it underscores theneed for flexible approaches that can respondquickly to novel threats arising from species movement or to capitalize on new opportunities. For
example, the Landscape Resilience Program ofAustralia’s Queensland government identified
priority locations for new protected areas thatwould maximize available habitat for range-shifting
species (87). Some jurisdictions with well-developed
land use and development processes have moved
toward adaptive development approvals, andAustralia’s fisheries management regime uses decision rules that automatically trigger new arrangements when predetermined environmental
conditions are reached (88). Mechanisms of thissort could be used more widely to implement
adaptive management for broader conservation
purposes, such as management plans with preset
increases in protective strategies that are triggered, or the automatic expansion of protection
for habitat outside protected areas when certain
climatic indicators are observed.
The changing distribution of species withincountries, between countries, and between national borders and the global commons will require increased cooperation and governance acrossmultiple scales among new stakeholders. The
European Union’s Habitats Directive [European
Commission (EC), 1992] and Birds Directive (EC,1979) are early examples of a cooperative approach to identifying and protecting networks ofhabitat across national borders. Initiatives suchas the Transfrontier Conservation Areas in Southern Africa (Southern African Development Community Protocol, 1999) also provide useful insights
to guide future multiscale and cross-border initiatives. Some challenges may also be addressedby increased use of dynamic management techniques. Several countries are already implementing
dynamic ocean management practices for bycatchprotection (89), though equivalent applications ina terrestrial context are more limited. Collaborative initiatives with indigenous communities may
also offer new opportunities for conservation ofrange-shifting species. Indigenous communities
can provide traditional ecological knowledge thatcomplements remote sensing and field data andprovides historical context (56), and new management arrangements may incentivize conservation
activities.
Earth systems andsustainable development
Human survival, for urban and rural communities, depends on other life on Earth. The biological components of natural systems are “on themove,” changing local abundances and geographical distributions of species. At the same time, the6 of 9Downloaded from http://science.sciencemag.org/ on March 30, 2017

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Managing for movement
Under extensive reshuffling of the world’s biota,
how should conservation goals and strategies forpolicy and implementation be developed to maximize long-term resilience of biodiversity andhuman systems? How should natural resource
management across diverse, multiuse, multiscaleland and seascapes be integrated to maximize resilience of both human and natural systems? Howshould specific threats and stressors (including
their interactions) be managed while minimizing impacts on valued ecosystem assets? For thescientific community to help develop mitigation
and adaptation strategies in the face of widespread change in species distribution and ecosystem functioning, a better understanding of themechanisms underlying such changes is needed.
Scientists also need access to real-time data streams,
as well as to integrate this information into decisionsupport frameworks. Moreover, scientists andtheir institutions need to rapidly communicate
Pecl et al., Science 355, eaai9214 (2017)31 March 2017

advances and outcomes to the broader public andto policy-makers. However, the natural world responds in dynamic and unpredictable ways, andthe phenomenon of species redistribution is not,nor will it ever be, fully understood or completely
predictable. This uncertainty necessitates flexible
and dynamic governance so adaptation to changing conditions can be rapid, maximizing opportunities and minimizing negative consequences.

monitoring system that successfully integrates field
and remote-sensing data could substantially
improve our ability to manage the changes tocome while potentially driving faster mitigation
measures (100).
Incorporating species on the move into
integrated assessment models

Because knowledge of the biological and ecological processes underlying resilience of organisms
to predicted average and extreme environmental
conditions is limited, the traits on which naturaland anthropogenic selection will act are uncertain. For example, specific physiological mechanisms have been hypothesized to underlie thethermal ranges of ectothermic organisms (94), yeta lack of universality in the proposed mechanisms
highlights a need for novel, multidisciplinary investigations (95). Large-scale, multigenerational
experimental research programs are required toprovide a robust understanding of the adaptive responses of organisms to environmental change
and to determine the heritability of key traits,
as recently has been achieved for sea turtles(96). Modeling approaches, lab and field-based
experimental manipulations, and field-based monitoring programs need to be combined with more
effective policy communication to understand andimplement responses to species redistributions.

Knowledge of underlying biological processes
and access to real-time data are necessary but not
sufficient for informed responses. Improved capacity to model linkages and feedbacks between
species range shifts and ecosystem functioning,food security, human health, and the climate isrequired. Modeling is essential to reliably project
the potential impacts of alternative scenarios andpolicy options on human well-being, as the basis
for evidence-based policy and decision support(101). One avenue forward is to incorporate species redistribution and its associated effects into
integrated assessment models (IAMs) (102), which
are used widely within the climate science community and are now being rapidly mobilized andextended to address synergies and trade-offs between multiple SDGs (103). IAMs offer a promising approach for connecting processes, existing
data, and scenarios of demographic, social, andeconomic change and governance. Although species distribution models are commonplace, advances are needed to connect species redistribution
with ecosystem integrity (104) and feedbacks between humans and the biosphere.

Monitoring programs

Communication for public and policy

To best adjust to species redistributions, gaps inunderstanding need to be acknowledged andfilled through hypothesis testing. Our understanding is weakest in poorly surveyed regions
such as the tropics and Antarctica (8). As range
shifts continue to unfold, there will be opportunities to refine our understanding of the process, but taking advantage of these opportunities
requires access to consistent, high-quality, nearreal-time data on a series of environmental andbiological parameters (97).
The current absence of a global, comprehensive, coordinated biodiversity monitoring system
is a major obstacle to our understanding of climate change implications for natural systems.
Thus far, extensive global cooperation and progress have been achieved in terms of coordinating
the collection and distribution of physical andchemical environmental monitoring data. For example, the Global Climate Observing System facilitated international agreement and a global
commitment toward consistent monitoring of climate variables, ultimately supporting the development of spatiotemporally-explicit and uncertaintyexplicit predictions about changes in our climate(98). Ongoing efforts through the Group on Earth
Observation Biodiversity Observation Networkand the Intergovernmental Oceanographic Commission Global Ocean Observing System are beginning to implement the use of Essential Biodiversity
Variables (41) and ecosystem Essential Ocean Variables (99), respectively, but the process is slow
and underresourced. A global, robust biodiversity

How does the scientific community engage effectively with the public on the issue of species
redistribution and its far-reaching impacts? Partof the answer could be citizen science and participatory observing approaches, in which community members are directly involved in datacollection and interpretation (105). These tools
can help to address gaps in both data and communication (100). When properly designed andcarefully tailored to local issues, such approaches
can provide quality data, cost-effectively and sustainably, while simultaneously building capacity
among local constituents and prompting practical and effective management interventions (106).

Underlying biological processes

Conclusions
The breadth and complexity of the issues associated with the global redistribution of species
driven by changing climate are creating profoundchallenges, with species movements already affecting societies and regional economies from thetropics to polar regions. Despite mounting evidence for these impacts, current global goals,
policies, and international agreements do not
sufficiently consider species range shifts in theirformulation or targets. Enhanced awareness, supported by appropriate governance, will provide
the best chance of minimizing negative consequences while maximizing opportunities arisingfrom species movements—movements that, with orwithout effective emission reduction, will continue
for the foreseeable future, owing to the inertia
in the climate system.
7 of 9Downloaded from http://science.sciencemag.org/ on March 30, 2017ability of people and communities to track thesepervasive species redistributions and adapt to themis increasingly constrained by geopolitical boundaries, institutional rigidities, and inertias at all
temporal and spatial scales.
In the coming century, all people and societies
will face diverse challenges associated with development and sustainability, many of which will
be exacerbated by the redistribution of specieson the planet (Figs. 2 and 3). The impacts ofspecies redistribution will intersect with at least
11 of the United Nations’ Sustainable Development Goals (SDGs) (table S2) and will be particularly prominent for several of these SDGs.
SDG2 (Zero Hunger) requires feeding morethan 9 billion people by 2050 (90). However, theability to deliver food through agriculture will be
altered via the direct effects of climate change, as
the distributions and abundances of pollinatorschange and as plant pathogens and pests become more prevalent or emerge in new places asa result of global warming (91, 92). SDG3 (Good
Health and Well-Being) is made more challenging by tropical illnesses spreading to new areas(58) and changes in food security and the distribution of economic wealth on local, regional, andglobal scales. Moreover, human well-being is also
related to many other facets of society and culture, including attachment to place (56, 93) andthe living environment found around us. The
mental health of indigenous and rural communities, in particular, may be affected as species
redistribution alters the capacity for traditional
practices, subsistence, or local industries. The
success of SDG13 (Climate Action) will depend on
accounting for the direct and indirect influencesof shifting organisms and associated feedbacks
on our biosphere, yet these processes and feedbacks are rarely accounted for in projections of
future climate. Sustainable management and theconservation of SDGs 14 and 15 (Life Below Water and Life on Land) are unlikely to be effective
unless climate-driven alterations in species ranges
and their profound ecosystem consequences aretaken into consideration.

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AC KNOWLED GME NTS

We thank the attendees of the international Species on the Move
conference held in Hobart, Tasmania, Australia, in February 2016.
G.T.P., E.W., and T.W. were supported by ARC Future Fellowships(FT140100596, FT110100597, and FT110100174, respectively). R.A.G.’s
participation was made possible by the South African National
Research Foundation (KIC 98457 and Blue Skies 449888). M.A.J.
was funded by Yale Climate and Energy Institute. T.M.’s participation
was supported by the WAPEAT (Finnish Academy 263465) Project.
J.M.P. was funded by the ARC Centre of Excellence for Coral Reef
Studies and ARC DP130100250, J.M.St. was supported by ARC
DP150101491, and S.A.R. was funded by ARC DP110101714. B.E. was
supported by Nordforsk. A. Cooper [Institute for Marine and Antarctic
Studies (IMAS)] assisted with the figures. The workshop andconference leading to this paper were supported by the Universityof Tasmania, IMAS, NOAA Fisheries Service, CSIRO, National
Climate Change Adaptation Research Facility Natural Ecosystems
Network, the Ian Potter Foundation, the Antarctic Climate andEcosystems Cooperative Research Centre, and the ARC Centre of
Excellence for Environmental Decisions.
SUPPLEMENTARY MATERIALS

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Tables S1 and S2References (118–135)
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Science 355 (6332), . [doi: 10.1126/science.aai9214]
Editor's Summary
Consequences of shifting species distributions
Climate change is causing geographical redistribution of plant and animal species globally. These
distributional shifts are leading to new ecosystems and ecological communities, changes that will affecthuman society. Pecl et al. review these current and future impacts and assess their implications forsustainable development goals.
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