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https://doi.org/10.1038/s41558-020-0768-2
1Department of Ecology and Evolutionary Biology, University of California, Irvine, Irvine, CA, USA. 2Northeast Climate Adaptation Science Center, US
Geological Survey, Amherst, MA, USA. 3Department of Environmental Conservation, University of Massachusetts, Amherst, MA, USA. 4Organismic
and Evolutionary Biology, University of Massachusetts, Amherst, MA, USA. 5Department of Natural Resources and the Environment, University of New
Hampshire, Durham, NH, USA. 6Miller Worley Center for the Environment, Mount Holyoke College, South Hadley, MA, USA. 7USDA Agricultural Research
Service, Rangeland Resources & Systems Research Unit, Fort Collins, CO, USA. 8Department of Forestry and Natural Resources, Purdue University, West
Lafayette, IN, USA. 9Department of Biological Sciences, Purdue University, West Lafayette, IN, USA. 10Centre for Ecology and Conservation, Penryn
Campus, University of Exeter, Cornwall, UK. 11Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI, USA. 12School
for Environment and Sustainability, University of Michigan, Ann Arbor, MI, USA. 13Estación Biológica de Doñana–Consejo Superior de Investigaciones
Científicas (EBD–CSIC), Sevilla, Spain. 14Department of Plant Biology and Ecology, University of Sevilla, Sevilla, Spain. 15These authors contributed equally:
Piper D. Wallingford, Toni Lyn Morelli. ✉e-mail: tmorelli@usgs.gov
Anthropogenic climate change is increasingly affecting spe-
cies and ecosystems across the globe, threatening biodi-
versity at both local and broad scales1. In response, species
from many taxonomic groups and ecosystems are undergoing redis-
tribution towards higher latitudes and elevations due to both the
direct (for example, physical limitations) and indirect (for example,
altered species interactions) effects of climate change2–6. Because
colonizing new habitats helps species persist both regionally and
globally7,8, range shifts are seen as overwhelmingly beneficial to
biodiversity conservation9. With the exception of some problematic
species (for example, forest pests)10, as well as translocations and
assisted migrations11–13, few studies (although see ref. 3 for an exam-
ple) have assessed the community and ecosystem impacts of spe-
cies tracking their climate niche into new areas. This research gap
remains despite theoretical literature that recognizes the potential
for impacts and the need for such research14–17. The lack of stud-
ies on range shift impacts is surprising given that the introduction
and spread of new species is often viewed by ecologists through the
lens of invasion biology, where the primary concern is the potential
for negative impacts on the recipient community. This dichotomy
underscores the importance of considering the ecological impacts
of range-shifting species in terms of both the benefits, particularly
to their persistence, as well as the potential costs to recipient com-
munities and ecosystem processes.
There are important ecological differences between introduced
and range-shifting species (see Table 1 for definitions) that result
in different levels of risk. For example, synthesis work considering
a broad range of introduced species suggests that 10–50% become
invasive and have negative impacts18–20. In contrast, results from
analyses of range shift impacts are mixed, with some showing mag-
nitudes of impacts similar to introduced species3 and others indicat-
ing that native species are less likely to be problematic when shifting
to nearby recipient communities20. Potential differences in impact
could be driven by range shifters’ shared evolutionary history with
some species in the recipient community; however, understanding
which species are likely to have a large negative impact is critical for
conservation of species in the many communities globally that are
being joined by range shifters. Invasion ecology, therefore, provides
insight for considering these interactions and assessing risk on a
species-by-species basis.
The movement of populations in response to climate change is,
in many ways, similar to the invasion of introduced species: it cre-
ates the potential for novel species interactions15. Both introduced
and range-shifting species have been shown to impact recipi-
ent communities by consuming, parasitizing or competing with
native species that lack the ability or defences to overcome them3,10.
Nevertheless, range shifters frequently share an evolutionary his-
tory with some species in the recipient community, so interactions
will not be completely novel, decreasing their potential for harmful
impacts due to established niches and community roles21. As more
species shift in response to climate change, methods for assessing
potential impacts on recipient communities, and thus prioritizing
which species to facilitate, become more valuable. Here, we lever-
age our understanding of biological invasions to describe a frame-
work for assessing the likelihood and degree to which range shifters
could impact recipient communities.
Adjusting the lens of invasion biology to focus
on the impacts of climate-driven range shifts
Piper D. Wallingford 1,15, Toni Lyn Morelli 2,3,4,15 ✉ , Jenica M. Allen2,5,6, Evelyn M. Beaury 4,
Dana M. Blumenthal7, Bethany A. Bradley3,4, Jeffrey S. Dukes 8,9, Regan Early10, Emily J. Fusco 3,
Deborah E. Goldberg11, Inés Ibáñez 12, Brittany B. Laginhas4, Montserrat Vilà 13,14 and
Cascade J. B. Sorte1
As Earth’s climate rapidly changes, species range shifts are considered key to species persistence. However, some range-shifting
species will alter community structure and ecosystem processes. By adapting existing invasion risk assessment frameworks,
we can identify characteristics shared with high-impact introductions and thus predict potential impacts. There are fundamen-
tal differences between introduced and range-shifting species, primarily shared evolutionary histories between range shifters
and their new community. Nevertheless, impacts can occur via analogous mechanisms, such as wide dispersal, community
disturbance and low biotic resistance. As ranges shift in response to climate change, we have an opportunity to develop plans
to facilitate advantageous movements and limit those that are problematic.
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Assessing the impacts of climate-driven range shifts
Invasion ecologists have invested considerable effort in developing
rubrics for predicting which introduced species are likely to become
problematic22. Catford et al.23 proposed a holistic framework that
broadly grouped these hypotheses into categories of propagule pres-
sure, abiotic characteristics of the recipient community and biotic
characteristics of both the recipient community and introduced
species. Many (but not all) of the factors influencing invasion suc-
cess, as identified in the Catford etal. framework, might also trans-
late to impacts of range-shifting species. We can, therefore, use this
framework to help assess the potential impacts of range shifters as
well as to identify vulnerable recipient communities (Fig. 1).
Propagule pressure. Propagule or dispersal pressure is critical to
the establishment of any introduced species24,25. Most invasive spe-
cies experience a lag period between the initial introduction and
the time at which they become invasive. This lag can last from
3–140 years in plants and 10–38 years in birds26, and is attributed
to a founder effect of the initial established population. Increased
propagule pressure can reduce this lag time by increasing genetic
diversity and adaptability of spreading populations27,28. Unlike with
introduced species, for which genetic diversity is strongly limited by
propagule pressure and number of introduction events, propagules
of range shifters are likely to have been arriving sporadically into
the recipient community at locations near the range margin. Thus,
the existence of nearby source populations of range shifters could
reduce time lags and increase the rate of population growth and
range expansion, especially for species that are prolific propagule
producers (Fig. 1)29. For example, marine organisms are expanding
by an order of magnitude faster than terrestrial species, likely due
to higher connectivity between communities, which translates to
fewer barriers to widespread dispersal3. Higher propagule pressure
at range margins makes it more likely that a range-shifting species
will establish and spread into a novel recipient ecosystem.
Abiotic effects on impacts. Introduced species can establish in
new communities when they have a competitive advantage or they
occupy an empty niche; for example, anthropogenic disturbances
can provide a window of opportunity for non-natives30. As the
climate continues to change, recipient communities are likely to
experience more frequent and acute abiotic stresses, which might
lead to decreased population sizes and extirpations (even extinc-
tions) in these communities4,31. This may enable the establishment
of range shifters as they track their optimal climate conditions.
For example, shorter winters and higher minimum temperatures
are allowing many range-shifting insect pests (such as spruce and
pine beetles) to colonize forests that were previously outside their
ranges32–34, leading to profound impacts on these ecosystems35. As
these fast-growing insect pests shift into novel forest communities,
drought conditions increase trees’ vulnerability and exacerbate the
pests’ impacts36,37.
Similarly, some of the most problematic introduced woody
plant species host nitrogen-fixing microorganisms in their roots,
thus allowing them to outcompete native species in an otherwise
stressful, low-nutrient environment. Myrica faya in Hawaii, Lupinus
arboreus in California grasslands and Acacia spp. in South Africa
are examples of highly invasive shrubs and trees that benefit from
greater access to nitrogen in nitrogen-poor soils38. Black locust
(Robinia pseudoacacia) is a fast-growing, nitrogen-fixing native
tree of southeastern North America that is currently undergo-
ing a climate-mediated range shift39. As black locust moves north
of its current range in response to climate change40, it is likely to
have a competitive advantage over native vegetation, especially in
nitrogen-deficient soils. Thus, recipient ecosystems that are heavily
disturbed or have low nutrient availability may incur larger impacts
from fast-growing and nitrogen-fixing range shifters.
Biotic characteristics. As with introductions, biotic characteristics
of shifting species and recipient communities influence potential
impacts (Fig. 1). Traits that make introduced species successful
(for example, high fecundity, fast growth, generalist feeding habits,
ability to engineer ecosystem conditions, and so on) will also facili-
tate the spread of range shifters41–46. Yet, because of the differences
in shared evolutionary history with species in the recipient com-
munity, impacts on the recipient community are likely to differ47.
Introduced species often benefit from interacting with new species
(naïve prey)48 and leaving old enemies behind (enemy release)21.
In contrast, species undergoing climate-induced range shifts settle
in an adjacent community, which is likely to have some overlap of
species composition and interactions with the donor community3,49.
Such overlaps make it less likely that range shifters will leave ene-
mies behind or encounter naïve prey, reducing the likelihood of
novel impacts.
Yet there is evidence that range-shifting species can also experi-
ence enemy release50,51, especially when a range shift occurs over a
long distance or crosses a biogeographic boundary that previously
limited dispersal52. The probability that a range shifter will experi-
ence release from natural enemies and encounter naïve species in
the recipient community is higher at ecotone edges, where dissimi-
lar adjacent communities meet53. For example, the movement of
tropical fish species to temperate communities has been facilitated
by ‘naïve’ temperate algae with weaker chemical defences. In the
southeastern Mediterranean Sea, the expansion of tropical herbi-
vores led to a 60% loss in benthic biomass, a 40% decrease in species
richness and, ultimately, a shift from a temperate reef system to one
that more closely resembles adjacent tropical communities54. Such
tropicalization of marine systems has become widespread as a result
of range shifts55,56.
By applying an invasive ecology framework, we hypothesize
that range shifters will be less likely to impact communities if some
species have co-existed and interacted within the donor commu-
nity. As with introduced species, the strongest impacts will likely
be seen in recipient communities with high concentrations of spe-
cialist species57, populations with low genetic variability7, species
that are already threatened by exploitation58 or species with low
population sizes58. Communities with traits that confer high biotic
resistance, such as high rates of predation, herbivory or strong
competitive interactions59, will be most resistant to impacts of
range shifters (Fig. 1)60.
Table 1 | Definitions of terms as used in this manuscript
Range shifter or
range-shifting species A species tracking its environmental niche
through a range expansion or relocation beyond
its historical native range.
Introduced species A non-native species transported to a new
ecosystem by humans, whether intentionally or
unintentionally.
Invasive species An introduced species that causes negative
ecological, economic or environmental impacts.
Recipient community The community into which an introduced or
range-shifting species arrives.
Donor community The community from which an introduced or
range-shifting species originates.
Establishment The process by which a founding population
increases in size and becomes self-sustaining.
Spread The process by which a species’ range expands
into new locations at an increasing distance from
the original area of establishment.
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Impacts of range shifting can parallel introductions
In contrast to introduced species, research on the effects of known
range shifters has been relatively scarce despite several studies show-
ing that the ecological and economic impacts can be equivalent61. In
marine systems, for example, range-shifting and introduced species
can cause community-level effects of the same direction and magni-
tude, but these impacts have been studied in fewer than 10% of doc-
umented marine range shifts3. Here, we present several examples
that illustrate how impacts of range shifters could have been pre-
dicted by applying an invasive species risk assessment framework
based on the traits and associated impacts reviewed above.
Range shifters benefit from novel interactions. Range shifters
encountering new species can have significant impacts on recipient
communities through changes to biotic processes, such as predation,
competition and the transmission of new parasites or pathogens. In
North American forests, the southern flying squirrel (Glaucomys sab-
rinus) is displacing the smaller northern flying squirrel (Glaucomys
volans) as the southern species expands its range in response to
increasing temperatures62. In addition to being superior competitors,
southern flying squirrels are carriers of an intestinal parasite that is
deadly to northern flying squirrels63,64. In the same forests, white-tailed
deer and their associated parasites are expanding northward in
response to climate change65. Due to the introduction of these para-
sites as well as increasing thermal stress, moose, boreal specialists, are
ultimately predicted to be extirpated from sites along their southern
range edge66. Conversely, moose are expanding at the northern end of
their range in response to the encroachment of deciduous forest into
the tundra, leading to a decline in native caribou populations67 (Fig.
2). In addition to highlighting the complexity of the impacts of cli-
mate change, these examples show how some range shifters will cause
localized extirpations, similar to introduced species. Risk assessments
can be used to identify range-shifting pathogen carriers and commu-
nities with vulnerable species or naïve prey before such impacts occur.
Invasive traits in range-shifting species. As with introduced spe-
cies that become invasive, range shifters with certain traits are more
likely to have negative impacts. For example, many shifting tree
populations are composed of conifers, which often have biologi-
cal traits that make them good colonizers. Most notably, many pine
species have relatively high growth rates, are resistant to environ-
mental stresses and develop monospecific stands that provide high
propagule pressure68.
Many invasive species that become dominant are also generalists,
able to utilize a variety of different resources. In marine systems, ocean
warming has allowed the long-spined sea urchin Centrostephanus
rodgersii, previously limited by juvenile growth, to redistribute pole-
ward from mainland Australia to Tasmania69. This urchin consumes
a wide range of macroalgal species, leading to significant declines in
kelp forest habitat70. Additionally, the long-spined urchin, a general-
ist herbivore, consumes many of the same prey species as the blacklip
abalone (Haliotis rubra), a specialist. Reduction in resource availabil-
ity has led to decreased abundance, fitness and survival among aba-
lone populations71. As with introduced species, range shifters that
are generalist consumers and possess ‘weedy’ traits are more likely to
impact a recipient community.
Community changes by range shifters can scale up to alter eco-
systems. The abundance, role and trophic level of a species in its
donor community can be indicative of its success in a recipient com-
munity72,73. These effects on populations and communities can ulti-
mately scale up to alter ecosystem states and processes. Ecosystem
alterations can be observed as trees shift into new areas, leading to
increases in aboveground and belowground biomass and shifts in
nutrient cycling through litter decomposition6,74–76. Climate-related
shifts of tree lines have been described in many parts of the world,
and grasses are expanding in the tropics as fire frequency increases77.
Yet the ecosystem impacts of these changes have been less explored
than those that occur after invasions by introduced trees and
grasses. Community and ecosystem effects have been observed in
aquatic and marine systems as well. For example, herbivory by the
long-spined sea urchin C. rodgersii has resulted in the collapse of
kelp forests, leading to decreased biodiversity, a simplified food web
and—at the ecosystem level—lower primary productivity78.
Another ecosystem shift occurring in tropical and subtropi-
cal regions is via the poleward expansion of mangroves into salt
Propagule or dispersal
pressure (species)
Abiotic effects
(community)
Biotic characteristics
Species Community
High risk
of impacts
Low risk
of impacts
High fecundity
Wide dispersal
Continuous propagules
High genetic diversity
Low fecundity
Limited dispersal
History of disturbance
Increasing environmental
stress
Breach of biogeographic
barriers
Resilient or resistant
to disturbance
Similar environmental
conditions
Invasive elsewhere
Abundant in home range
Fast growth
Generalists
Foundation species or
ecosystem engineers
Pathogen carriers
Threatened or endangered
Endemic
Obligate mutualist
Specialists
Rare community
Naïve prey
Enemy release
Shared evolutionary
history
Biotic resistance
Fig. 1 | Risk assessments for biological introductions focus on the importance of three main components that lead to the successful establishment and
spread of species: the introduction of propagules, the abiotic environment and biotic interactions. If characteristics that lead to negative impacts of
introduced species on recipient communities (indicated in bold) are shared with species undergoing range shifts, there is greater risk (shown in orange) to
recipient ecosystems from range shifters, which can inform management strategies.
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marshes79. In Florida, mangrove forests have doubled their area at
the northern end of their historical range due to reduced frequency
of cold-weather extremes80. Both mangrove trees and salt marsh
grasses are foundation species in their respective ecosystems and
play an important role in structuring communities by providing
habitat and altering nutrient cycling80. Because mangroves have
greater aboveground biomass and, therefore, structural complexity
than native salt marsh vegetation, their expansion has broad impli-
cations for coastal wetland ecosystems. The establishment of intro-
duced mangroves in sandflats has increased the concentration of
fine sediments and organic matter, leading to a higher abundance
and diversity of non-native macrofauna79.
The lack of defences of temperate species to tropical herbi-
vores81,82, general patterns of increased nutrient content with lati-
tude83 and increased disease due to increased herbivory84,85 might
accelerate the tropicalization of these temperate wetland regions
under future climate change. Previous research on the impacts of
biological invasions on ecosystem properties and processes has
shown that these impacts are highly context-dependent, as the
magnitude and direction can vary both within and between types
of impacts depending on taxa and ecosystems86. As with introduc-
tions, species that can alter the physical properties of the commu-
nity (for example, ecosystem engineers) are most likely to have
ecosystem-level impacts.
Balancing conservation with risk
Conservation options for promoting persistence (and prevent-
ing extinction) of species threatened by climate change include
increasing habitat connectivity between communities to facilitate
species movement and actively moving species—that is, assisted
migration11,12. In the context of assisted migration to facilitate
climate change adaptation, conservation organizations, such as the
International Union for the Conservation of Nature (IUCN)87, are
already considering invasion risk. However, increasing habitat con-
nectivity to facilitate the movement of range-shifting species is gen-
erally considered an unmitigated good with little consideration of
the full range of impacts on the recipient community.
Rather than placing a value on all species movements, we suggest
using a risk–benefit analysis framework to assess potential impacts
on a case-by-case basis. In some contexts, increasing habitat con-
nectivity might best be based on analyses of donor and recipient
communities with a focus on providing connectivity for low risk,
nearby natives (Fig. 1). While there are inherent value judgements
in assigning worth to species, we suggest that management should
generally (1) facilitate range shifts that promote ecosystem services
and biodiversity88 and (2) discourage range shifts of species with the
potential to negatively impact sensitive or rare species and com-
munities89. In some cases, the analyses will be straightforward. For
example, when range-shifting species are both locally and region-
ally uncommon, they could pose little risk to recipient communities
(Fig. 1) and would benefit from opportunities to shift their ranges.
This is unlikely to be true for species that have large impacts on their
donor communities. Keystone predators (species with a dispropor-
tionate impact relative to their abundance) and foundation species
(species that facilitate diversity by providing habitat and modify-
ing their environment) might lead to management conundrums, as
such species could pose great risk to recipient communities but also
support the colonization of other range-shifting species with which
they interact90.
Even range-shifting species with small impacts in their donor
communities can have large impacts in recipient communities
because of ecological surprises, such as novel interactions with spe-
cies in the community91. A single invasion can be devastating to a
community92, and risk assessments are a useful and often-applied
tool for identifying species of concern. Therefore, like others who
warn about the potential dangers of assisted migration93, we pro-
pose that, before facilitating range shifts, the ecological, economic
and societal impacts associated with these management actions
be considered88.
There are many assessment tools to evaluate the potential
impacts of introduced species94. Notably, the Environmental Impact
Classification for Alien Taxa (EICAT) framework is a standardized,
objective and transparent approach adopted by the IUCN in 2016
that identifies the mechanisms through which introduced taxa can
impact recipient communities73,87. Although this assessment was
developed for introduced species, the mechanisms of impact out-
lined in EICAT apply to the interactions between range shifters and
recipient communities as well. Identified mechanisms primarily fall
into the biotic characteristics of the Catford etal.23 framework and
consist of competition, herbivory and predation (including para-
sites and pathogens), hybridization, poisoning/toxicity, biofouling,
ability to alter the ecosystem and interactions with other non-native
species. These mechanisms are scored based on their magnitude of
impact to the community, ranging from minimal (that is, negligible
impacts, but no reductions in fitness for native species) to massive
(that is, irreversible impacts through local, population or global
extinctions; Fig. 3).
We suggest applying EICAT to rank and prioritize range-shifting
species based on their potential impacts on recipient communities
and to develop monitoring or control plans to limit those impacts.
For example, communities receiving range-shifting species with
the lowest potential to experience impacts (minimal and minor)
are likely to benefit most from passive management actions, such
as monitoring. Such range shifts could maintain or even increase
community diversity and allow for persistence of populations
under increasingly stressful environmental conditions. Although
minor and moderate impacts lead to reductions of fitness in
individuals or declines in population abundances, respectively, the
Fig. 2 | Range shifters can impact recipient communities. For example, as
white-tailed deer expand their range (yellow) northward (arrows pointing
upwards) in response to climate change, moose at the southern edge of
their range (green) are experiencing greater rates of parasitism and are
projected to undergo population declines66. In contrast, moose populations
at the northern range edge are increasing and replacing caribou67 (blue).
Smaller icons indicate range contractions. Ranges based on IUCN Red List
of Threatened Species 2016 (refs. 112–114).
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recipient community structure and functioning are preserved.
Future communities might not be analogous to existing recipient
communities, but shifts are likely to have a net positive impact on
global biodiversity.
Species with major or massive impacts, however, might need to
be actively managed through facilitating or restricting movement,
as their impacts could alter community structure and composition
and cause local extinctions in the recipient community. While such
changes, by definition within the EICAT framework, are revers-
ible for species with major impacts, those with massive impacts
are likely to cause irreversible changes as the community passes a
threshold from which it can no longer recover. In the cases of spe-
cies with major or massive impacts, serious consideration should
be given to whether the benefits of promoting the persistence of
the range-shifting species or populations justify the impacts on the
recipient communities. Based on approaches traditionally used to
manage invasive species, we suggest the following specific strategies
for range shifters:
• Involve stakeholders early: work closely with natural resource
managers, conservation practitioners and decision-makers at
every step of the process, including to identify priority ecologi-
cal and cultural species95,96 and important ecological services5
associated with both range shiers and recipient communities.
• Identify management priorities for range-shiing species
and recipient communities: what is the conservation status of
the species? How important is the range shi for its persistence?
How unique is the recipient ecosystem? How important are its
constituent species and associated services for stakeholders?
Supporting range shis for species of conservation concern will
remain a key climate change adaptation tactic for conservation
practitioners and natural resource managers.
• Incorporate species distribution model forecasts: use the best
available data and models to anticipate the movement of range
shiers and identify high priority conservation areas, hotspots
of biodiversity97 and hotspots of high impacts98,99. Additionally,
triaging which species are most likely to persist under projected
climatic conditions can inform where resources can be most
eectively allocated. Regularly revise management proposals to
incorporate updates to these forecasts.
• Use tools to assess invasion risk: consider the parallels between
traits common in successful introduced invasives22 (Fig. 1) and
their potential impacts (EICAT73) to identify high- and low-risk
range-shiing species.
• Monitor changes in recipient communities: monitor for shis
in abundance of target species and the arrival of new species,
especially following disturbance and extreme climatic events24.
Challenges and potential limitations
Important knowledge gaps related to range-shifting species must
be addressed to better understand the impacts that these species
might have while also promoting persistence of species as their
climate zones move. While range shifts have been documented
for hundreds of species across taxa and ecosystems3–5, they can
be difficult to detect, as the historical ranges for many species are
Recipient community
Donor community
Climate warming through time
Massive
Major
Moderate
Minimal or minor
Potential impact scenarios
Fig. 3 | As climate change alters environmental conditions, range shifts can lead to new species interactions and changes to community structures
depending on the magnitude of associated impacts. For example, as individuals from a nearby donor community (blue birds in grey circle) shift into a
novel recipient community (green and black birds in black circle) in response to climate change, they might have minimal or minor impacts (few blue
birds in a community of primarily green and black birds) up to major or massive impacts where the shifting species predominates. This range of impacts
can be seen in the examples discussed here, including cases of southern flying squirrels displacing northern flying squirrels (moderate due to effects on
populations) to tropicalization (massive, irreversible shifts in ecosystems). Photographs courtesy of Alexej Sirén and the U.S. National Park Service.
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unknown or imprecise and our understanding of a ‘native range’
is not well-defined100. This is especially true for systems that are
not as well-studied, such as deep-water marine systems that are
difficult to access, and incorporating different spatial or tempo-
ral scales could further alter our definition of what constitutes a
range-shifting species.
The impacts of range shifters, which might accrue more slowly
than impacts of introduced species, have received less attention
than invasion impacts; thus, our ability to predict future outcomes
is limited. Range-shifting species could potentially undergo hybrid-
ization, experience toxicity, or evolve or adapt; an increased under-
standing of potential interactions in new environments is needed
to evaluate these possible outcomes. Additionally, effects may differ
across scales. Addressing these knowledge gaps will require working
across broad stakeholder groups to leverage and continue existing
monitoring programs and incorporate diverse resources, such as
local and traditional ecological knowledge101.
Predicting potential shifts is further complicated by our limited
understanding of the abiotic and biotic limits to species’ ranges.
Predicting which species are likely to undergo shifts requires a
knowledge of organismal physiology and thermal limits and how
these contribute to ability to disperse as well as to adapt in place.
Additionally, while temperature is a primary driver of distribu-
tion patterns102,103, biotic resistance also plays a critical role49. Yet
biotic interactions are, themselves, often altered by abiotic con-
ditions16,104. Traits can act synergistically (for example, a drought
causes reduced propagules), creating feedbacks that alter the mag-
nitude of impacts. To detect species interactions and community
impacts, manipulative insitu experiments are likely a necessary
and important focus for climate change researchers. However,
these experiments can be time-consuming or expensive, and a
lack of experimental studies does not preclude using general risk
assessment frameworks (Fig. 1) and knowledge from invasion
biology to inform decision making. Additionally, risk assessments
that are continually updated as new empirical data accrue can be
used by practitioners, providing an accessible resource for those
making management decisions.
Finally, we must acknowledge that anthropogenic climate change
has led to unprecedented disruptions to global environments at a
level rarely experienced before the Anthropocene105,106. Many spe-
cies’ ranges have already been dramatically altered by human
impacts, which raises questions about how to classify species that
expand into their historical habitat following extirpation and which
incarnation of a community should be protected107,108. The rapid rate
of anthropogenic climate change is likely to outpace species’ ability
to adapt. Range shifts, therefore, have been viewed as an alternative
means to promoting global biodiversity. Yet, the potential feedbacks
and consequences need to be considered as conservation goals may
conflict with one another depending on the individual case. For
example, mangroves and salt marshes provide similar ecosystem
services. However, salt marsh systems have experienced significant
area loss109, and range-shifting mangroves could further impact
these vulnerable communities, highlighting the important of hav-
ing clear objectives for management actions. At the same time, as
range shifters are altering recipient communities, those communi-
ties themselves are responding to climate change; disentangling the
causes and effects of these drivers will be a continuing challenge.
Conclusions
Although the impacts of range-shifting species can vary from minor
to massive, considerations of species movements in the context of
climate change has almost entirely focused on positive impacts7,28,86.
As species shift to track a changing climate, we have a unique
opportunity to facilitate advantageous, and discourage potentially
problematic, movement of species in real time. However, both
researchers and managers will likely need to adopt a more fluid
and dynamic view of what constitutes a community, as differences
in species’ responses could result in communities with no current
analogue110. Communities are unlikely to shift as a whole, and par-
tial shifts will disrupt species interactions and lead to trophic mis-
matches111. Rather than developing new strategies to evaluate the
potential impacts of range-shifting species, we suggest leveraging
invasion ecology theory and risk assessment tools (for example,
EICAT) to quantify the magnitude of the potential impacts of range
shifters and define specific conservation goals in response. This will
allow us to maintain biodiversity and ecosystem functioning most
effectively despite a rapidly changing climate.
Received: 21 May 2019; Accepted: 30 March 2020;
Published: xx xx xxxx
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Acknowledgements
This work was initiated at a working group led by C.J.B.S., B.A.B., A.E.B., and R.E. and
was supported by the Albert and Elaine Borchard Foundation. We thank R. Whitlock
for his insight during initial discussions, V. Pasquarella for her comments on an early
draft, and C. Millar and J. McMullen who provided valuable feedback. Funding for this
project was provided in the form of a University of Michigan catalyst grant to I.I., and
from the National Institute of Food and Agriculture, U.S. Department of Agriculture, the
Massachusetts Agricultural Experiment Station, the U.S. Geological Survey Northeast
Climate Adaptation Science Center and the Department of Environmental Conservation
under Project Number MAS00033 to B.A.B. Any use of trade, firm or product names is
for descriptive purposes only and does not imply endorsement by the U.S. Government.
Author contributions
T.L.M., C.J.B.S. and P.D.W. conceptualized the idea for this Review independently.
T.L.M. and P.D.W. proposed the project, led breakout sessions during the working
group and managed the project throughout its development, including writing,
reviewing and editing all manuscript versions. B.A.B. and C.J.B.S. provided invaluable
feedback throughout the project and contributed though mentoring and supervision,
as well as writing and in-depth review. B.A.B., B.B.L., T.L.M. and P.D.W. created figure
visualizations. J.M.A., E.M.B., D.M.B., J.S.D., R.E., E.J.F., D.E.G., I.I., B.B.L. and M.V.
contributed equally to writing and providing feedback.
Competing interests
The authors declare no competing interests.
Additional information
Correspondence should be addressed to T.L.M.
Peer review information Nature Climate Change thanks I-Ching Chen, Jorge E. Ramos and
the other, anonymous, reviewer(s) for their contribution to the peer review of this work.
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