Latitudinal and Elevational Range Shifts under Contemporary Climate Change

Jonathan Lenoir and Jens-Christian Svenning, Aarhus University, Aarhus, Denmark

r 2013 Elsevier Inc. All rights reserved.

GlossaryFront edge Stable margin of a species’ range where

population size is growing.

Leading edge Dynamic margin of a species’ range where it

is expanding into neighboring unoccupied areas.

Range core Central part of the distribution of a species.

Range periphery Marginal parts of the distribution of a

species.

Rear edge Stable margin of a species’ range where

population size is declining.

Species’ niche The scope of environmental conditions

under which a species is able to maintain stable or

increasing populations.

Species’ optimum The value of a given environmental

variable at which a species reaches its maximum probability

of occurrence or abundance.

Species’ range The spatial distribution of a species, that is,

the sum of all geographic locations of its individuals over a

given period in time.

Species’ response curve The relationship between a

species’ probability of occurrence or its density (number of

individuals per unit area) and the values of a given

environmental variable.

Trailing edge Dynamic margin of a species’ range where

it is becoming locally extinct, causing the range to

retract.

The concept of range shifts refers to the dynamic process

whereby species shift their distribution over time, tracking

geographic shifts in the climate and habitat conditions that they

require. Range shifts are also commonly defined as migration

events that occur on decadal to millennial or longer time

scales, usually beyond the life span of individuals. In contrast,

the concept does not include migration events occurring

over the life span of individuals, that is, on a seasonal to annual

or shorter time scales, as a result of recurring movements

of individuals between spatially separate habitats in order

to avoid temporarily unfavorable conditions. It also does

not include essentially stochastic long-distance dispersal events

of a single or few individuals that fail to establish new popu-

lations. Recently, species range shifts have attracted a great deal

of interest from the scientific community as a direct con-

sequence of the increasing focus on current global environ-

mental changes, notably the changing climate (IPCC, 2007a).

There is now a growing body of studies reporting and fore-

casting species range shifts across the main geographic dimen-

sions (latitude, longitude, and elevation) as a response to both

contemporary and future changes in climatic conditions. Here,

the focus is on realized range shifts in response to contemporary

climate change, a phenomenon that is of key importance for

developing well-founded predictive models of range shifts

under expected substantial future climate change (IPCC,

2007b). Because dramatic range shifts have also occurred in the

geological past, species range shifts, until recently, have been

described in textbooks through paleontological examples, but

contemporary range shifts are now becoming even more

evident.

Species’ Range: Geographic Dimensions andEcological Drivers

As a species’ range can be defined by the geographic locations

of all its individuals over a given period of time (Brown and

Lomolino, 1998), it is often difficult and challenging to cap-

ture and represent it. Several abstractions of the real distri-

bution of a single species have been used, with the most

frequent approaches being to represent a species’ range as

outline, dot, or contour maps in a two-dimensional geo-

graphic space, most often across the latitudinal and longi-

tudinal dimensions. One could also represent a species’ range

in a three-dimensional geographic space by adding occurrence

along elevational gradients to the representation (Figure 1).

Another more simplistic abstraction is to project the known or

estimated occurrences of the species against one of these three

dimensions. Hence, latitude, longitude, and elevation consti-

tute the geographic dimensions over which a species’ range is

estimated, and a species’ range shift is defined as changes in

these geographic dimensions over the fourth dimension: time

(Figure 1).

A species’ range is generally understood to be at least partly

determined by the species’ environmental requirements and

tolerances, that is, its ecological niche (Hutchinson, 1957;

Soberon and Nakamura, 2009). The ecological niche is typi-

cally considered a multidimensional space (or ‘‘hypervo-

lume’’), in which the different dimensions represent different

environmental conditions (Hutchinson, 1957). Especially at

large scales, climate is often thought to constitute the niche

axes that are most relevant for species ranges (Pearson and

Dawson, 2003). Therefore, one should be able to deduce a

species’ climatic niche from its range. Unfortunately, the

geographic distribution of a species is unlikely to fully repre-

sent its ecological niche for at least three reasons (Colwell and

Rangel, 2009): (1) geographic unavailability of some portions

of its ecological niche; (2) geographic inaccessibility due to its

limited dispersal abilities (Svenning and Skov, 2004; Blach-

Overgaard et al., 2010); and (3) geographic control by con-

straining species interactions (Case et al., 2005). Throughout

this article, therefore, the species’ range is solely being con-

sidered as an incomplete geographic reflection of the species’

niche. Still, it is useful to relate ranges to the niche concept in

Encyclopedia of Biodiversity, Volume 4 http://dx.doi.org/10.1016/B978-0-12-384719-5.00375-0 599

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order to understand the mechanics of range shifts under cli-

mate change. Given this conceptual framework, a given species

may respond differently to the environmental changes over

time. It may (1) relocate toward newly suitable geographic

locations, completely or partially tracking its environmental

requirements (range shift) (Lenoir et al., 2008); (2) remain

within the same locations and acclimate or adapt to its new

environment (niche shift) (Davis and Shaw, 2001); (3) go

extinct (Svenning, 2003); or some combination of these re-

sponses. In short: move, adapt, or die (Maggini et al., 2011).

How Do Species Shift Their Ranges?

Species’ ranges are not static over time, but constantly shift

either more or less stochastically by means of population

dynamics within a steady-state environment or directionally

through population dynamics within a shifting environment

that is spatially autocorrelated (Figure 2). Although many

human-induced changes in the global environment, such as

land-use change, climate change, nitrogen deposition, bio-

logical invasions, and atmospheric CO2 increase (Sala et al.,

2000) may trigger species range shifts, here the focus is on the

effect of contemporary climate change.

The Mechanics of Range Shifts: Distinction between RangeCore and Range Periphery

In a steady-state environment, stochastic range shifts are the

net result of growth, decline, colonization, and extinction

processes that are tuned to different temporal scales (Breshears

et al., 2008). Whereas local populations grow and decline over

relatively short time periods, local populations colonize and

go extinct over much longer time periods, especially for long-

lived species. Additionally, growth, decline, colonization, and

extinction processes are tuned differently across a species’

range (Figure 2(a)). Notably, there may be a strong

core–periphery distinction leading to lower population dens-

ities near the peripheral area in comparison to the core area

(Brown and Lomolino, 1998). Here the term ‘‘core’’ instead of

‘‘center’’ is explicitly used when referring to the geographic

location of the abundance peak within a species’ range. In-

deed, the geographic location of the highest population

density or abundance of a given species within its range is not

necessarily located at the geometric center or centroid of its

range (Murphy et al., 2006), even though the core area of a

species’ range may be represented at the range center for

simplicity (Figure 2(a)). At the core area of a species’ range,

sometimes called species’ optimum to avoid confusion with

the geometric center, local populations are generally thought

to experience favorable conditions leading to enhanced fitness

and population density. In other words, at the core, birth and

emigration rates may exceed death and immigration rates,

respectively, leading to sink populations (Pulliam, 2000).

Such populations not only are likely to grow within the

suitable localities already occupied, but are also likely to col-

onize and establish populations in suitable localities not

previously occupied by the species in the core area of its range

and even beyond, if dispersal and biotic interactions are not

constraints. Decline and extinction processes might occur

there as well, but to a lesser extent, unless there is a drastic

shift in environmental conditions. At the peripheral area of a

species’ range, local populations may experience unfavorable

conditions, leading to reduced fitness and reduced population

density. In other words, at the periphery, death and immi-

gration rates may exceed birth and emigration rates, respect-

ively, leading to sink populations (Pulliam, 2000). Such range-

edge sink populations will tend to decline and eventually go

extinct, except when maintained by a ‘‘rescue effect’’ through

constant immigration from source populations, typically lo-

cated close to the core area of the species range. Growth and

colonization processes might occur there as well, but to a

lesser extent, unless there is a drastic shift in environmental

conditions.

Directional Range Shifts under Climate Change

Let us now consider shifting conditions in the environment,

such as the temperature and precipitation changes observed

during contemporary climate change (IPCC, 2007b). Any shift

in climatic conditions in a given location will impact the in-

dividuals living there and thus affect population dynamics

locally. For instance, species sensitive to temperature may re-

spond to a warmer climate through local changes in growth,

decline, colonization, and extinction rates. On the one hand,

Latitude

Latitude

(a)

(b)

Longitude

Longitude

Figure 1 Representation of a species’ range (red dots) across the

three geographic dimensions (latitude, longitude, and elevation) of a

given landscape in the Northern Hemisphere ( the ‘‘volcano’’ data set

available in R has been used (R Development Core Team, 2010) for

illustrative purpose) and at two different time periods: t1 (a) and t2(b). Note that the illustrative species shifted its geographic range

upward and poleward between t1 and t2.

600 Latitudinal and Elevational Range Shifts under Contemporary Climate Change

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locations within the peripheral area of a species’ range that

were too cold and therefore less suitable might become more

suitable in a warmer climate, thus turning sink populations

into source populations with higher growth and colonization

rates. On the other hand, locations within the core area of a

species’ range that were highly suitable might become too

warm and therefore less suitable, turning source populations

into sink populations with higher decline and extinction rates.

Across species ranges, the net result of these changes in

growth, decline, colonization, and extinction processes will

affect their geographic distribution in different ways. However,

an increase in temperature is likely to have an overall dir-

ectional impact on species range shifts, because temperatures

are autocorrelated in space, linking warmer conditions at

lower latitudes and elevations with cooler conditions at higher

latitudes and elevations. Therefore, one expects to observe

poleward and upward range shifts as climate warms, even after

accounting for dispersal limitations (Engler et al., 2009) or

biotic interactions (Araujo and Luoto, 2007). Accordingly,

poleward and upward range shifts in the warming climates

following the glacial recessions of the past interglacial periods

have been widely reported for plants, birds, and mammals

(Brown and Lomolino, 1998). For instance, Pleistocene fossils

of several species of rodents have been found several thousand

kilometers southward of the southern limit of their modern

distribution in northern America (Graham, 1986), suggesting

strong poleward shifts during the Holocene. Similarly, Pleis-

tocene macrofossils of Podocarpus have been found c. 1000 m

below the lower limit of their contemporary distribution on

the Andean flank in western Amazonia (Cardenas et al., 2011),

thus suggesting a large upward shift during the current inter-

glacial period.

Although the expectations given temperature increase

alone are relatively straightforward, it is difficult to predict

how concurrent changes in other climatic factors, especially

precipitation, will affect species ranges. Temperatures are

negatively correlated along the latitudinal and elevational

gradients and have risen globally over the past decades (IPCC,

2007b). In contrast, precipitation changes are more hetero-

geneous across space and time (IPCC, 2007b), leading to

strong regional differences. Such regional variation in climate

change, mainly due to the effect of precipitation changes on

the water balance equation (balance between evapo-

transpiration and precipitation), may affect species ranges as

well, leading to unexpected regional range shifts as climate

warms globally (Crimmins et al., 2011). Additionally, biotic

interactions may also affect the magnitude of species range

shifts as climate warms (Araujo and Luoto, 2007) with some

suggestions that interactions could explain unexpected range

shifts as well (Lenoir et al., 2010a). Therefore, at global to

continental scales, poleward and upward range shifts are likely

(a)

(b)

(c)

(d)

Abundance

Abundance

Marc

hS

tochasticity

Latitude/altitude

Latitude/altitude

Abundance

Lean

Abundance

Cra

sh

Latitude/altitude

Latitude/altitude

Figure 2 Conceptual representations of latitudinal and elevational

range shifts and their mechanics under a steady-state environment

(a) and under climate warming (b)–(d). In each case, the

figure shows the relative importance of growth, decline, colonization,

and extinction processes across the species range. For the march-,

lean-, and crash-range shifts, gray shades represent conditions

before a climate warming event whereas overlaying transparent

colors represent shifting conditions under a climate warming event.

Latitudinal and Elevational Range Shifts under Contemporary Climate Change 601

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to be the main responses to the global increase in temperature

conditions, albeit counterintuitive shifts should be expected as

well at more regional scales due to regional variation in cli-

mate change and biotic interactions.

Types of Range Shifts

There are different types of range shifts. Many different terms

have been used to refer to the many ways species may shift

poleward or upward as climate warms, among others: leading-

edge expansion, trailing-edge retraction, optimum shift, and

abundance changes. Recently, a catalog of the possible species

range shifts observable along a given gradient has been laid

out, suggesting that observed species distribution changes are

rarely complete shifts, but intermediate states in an ongoing

shifting process (Maggini et al., 2011). More generally, three

main categories of observed range shifts can be distinguished

as climate warms (Breshears et al., 2008): (1) ‘‘march’’; (2)

‘‘lean’’; and (3) ‘‘crash’’ (Figures 2(b)–(d)).

• March-range shifts represent the most dynamic category of

range shifts and involve dynamic edges with establishment

from enhanced colonization and growth at the leading

edge and mortality from enhanced decline and extinction

at the trailing edge. Such dynamics driven by the edges

usually lead to expansion toward higher latitudes or ele-

vations (leading-edge expansion) or retractions from lower

latitudes or elevations (trailing-edge retraction) (Maggini

et al., 2011). Therefore, out of this first category, one can

develop several kinds of range shifts involving any com-

binations of expansion at the leading edge or retraction at

the trailing edge, the most dynamic being a combination of

both or march (Figure 2(b)). Short-lived, small-sized, and

mobile species for which colonization and extinction

processes occur on a relatively short time scale are likely to

exhibit such dynamic range shifts. For instance, short-lived

and small-sized butterfly species not strongly limited by

their dispersal abilities have been reported to shift north-

ward at both their leading and trailing edges (Parmesan

et al., 1999).

• Lean-range shifts involve stable edges with the optimum

shifting within the existing range (Figure 2(c)). This rep-

resents the net result of enhanced growth at the highest

latitudes or elevations (front edge) and decline at the

lowest latitudes or elevations (rear edge) of the existing

range, resulting in a shifting abundance pattern. This kind

of shift might be transient and might correspond to early

stages of a full shifting process (Maggini et al., 2011). Long-

lived, large-sized, and immobile species for which colon-

ization and extinction processes occur on a much longer

time scale than the period of observation are likely to ex-

hibit such transient stages (time lag) across their ranges.

For instance, long-lived and large-sized tree species, which

tend to be strongly limited by their dispersal abilities, have

been reported to shift their optimum elevation upward

without shifting their range edges: persistence at the rear

edge without establishment success beyond the front edge

(Kelly and Goulden, 2008).

• Crash-range shifts involve stable edges and a stable opti-

mum, but overall declines across the existing range

(Figure 2(d)). Rare species having a restricted distribution

for which the degree of climate warming far exceeds their

ability to shift toward suitable conditions (because of dis-

persal limitation and natural or anthropogenic habitat

fragmentation) are likely to exhibit such changes across

their ranges. Similarly, there is no possible escape for en-

demic species having a distribution restricted to a specific

mountain summit and facing contemporary climate

change. In such situations, these threatened populations

may simply crash. For instance, in 1987, a multipopulation

crash of the endemic golden toad of Costa Rica’s Cordillera

de Tilaran has been reported to have led to the dis-

appearance of this species (Pounds et al., 1997). Add-

itionally, species range shifts at the bottom of a food web

may trigger trophic mismatch leading to multipopulation

crash within the higher levels of the food web. For instance,

fluctuations in plankton have resulted in a general decline

in cod biomass and recruitment throughout a bottom-up

control in the North Sea (Beaugrand et al., 2003).

Of course, these three categories are not mutually exclusive

and can be combined to develop all possible patterns of

poleward or upward shifts as climate warms (Breshears et al.,

2008). Using the species response curve representation along

one of the three geographic dimensions (Figure 2), the dif-

ferent combinations may affect the edges, optimum, level of

abundance, or skewness of each species’ response curve, thus

leading to many possible patterns of range shifts (Maggini

et al., 2011).

Contemporary Evidence for Latitudinal andElevational Range Shifts

An overwhelming number of reports of latitudinal and ele-

vational range shifts under contemporary climate change have

recently appeared in the scientific literature (IPCC, 2007a).

The expected poleward and upward range shifts in warming

regions have been reported for many taxonomic groups across

the plant and animal kingdoms (Parmesan et al., 1999; Sturm

et al., 2001; Lenoir et al., 2008; Moritz et al., 2008). A recent

meta-analysis (Chen et al., 2011a) concluded that the distri-

butions of species have recently shifted poleward at a median

rate of 16.9 km per decade and upward at a median rate of

11.0 m per decade, which is two to three times faster than

reported in a previous meta-analysis (Parmesan and Yohe,

2003). However, these global trends represent averages across

considerable variation in range dynamics under climate

change (e.g., Chen et al., 2011a). Indeed, most of the studies

that have reported the expected range shifts toward higher

latitudes or elevations as climate warms globally have also

detected, at a smaller frequency and to a lesser extent, species

moving toward lower latitudes or elevations (Parmesan et al.,

1999; La Sorte and Thompson, 2007; Lenoir et al., 2008,

2010a; Moritz et al., 2008). In most cases, modern latitudinal

and elevational range shifts have been documented by using

and sometimes resurveying historical occurrence data such as

museum collections and field notes (Tingley and Beissinger,

2009). These historically based studies can be divided into two

main groups (Parmesan, 2006): (1) studies that directly assess

602 Latitudinal and Elevational Range Shifts under Contemporary Climate Change

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range shifts by comparing historical and modern data on a

species’ distribution across its entire ranges or sections of the

range; and (2) studies that infer range shifts by comparing

historical and modern data on species composition or abun-

dance within local communities. The following sections cover

both types of studies.

Latitudinal Range Shifts

We carried out a selective review of historically based studies

documenting contemporary latitudinal range shifts. Many

species across many organism groups ranging from the most

simple such as plankton, algae, and lichens to the most

complex such as flowering plants, birds, and mammals, across

both marine and terrestrial ecosystems, have shifted their

latitudinal ranges over recent decades (Table 1). Most of these

contemporary latitudinal range shifts are coherent with con-

temporary climate change, even though some studies reported

the absence of latitudinal range shifts or even unexpected

latitudinal range shifts, in addition to the expected and em-

pirically predominant poleward range shifts (Thomas

and Lennon, 1999; Warren et al., 2001; Hill et al., 2002;

Brommer, 2004; Hitch and Leberg, 2007; La Sorte and

Thompson, 2007).

Most latitudinal range shifts have been documented within

the animal kingdom (invertebrates, fishes, amphibians, rep-

tiles, birds, and mammals), whereas only a handful of cases

have so far been documented for plants (Table 1). Among

animals, there is clear evidence of poleward range shifts

through either expansion at the leading edge or retraction at

the trailing edge (Parmesan et al., 1999; Hickling et al., 2005),

or increasing and decreasing abundance at the front and rear

edges, respectively (Sagarin et al., 1999; Myers et al., 2009). For

plants, however, reported changes seem to occur principally

at the leading/front edge (Smith, 1994; Sturm et al., 2001;

Walther et al., 2005b) without much clear evidence for chan-

ges at the trailing/rear edge (Lesica and McCune, 2004), in line

with conclusions from an earlier review (Jump et al., 2009) on

the apparent lack of latitudinal range shifts at the trailing/rear

edge of plant species ranges, especially for long-lived plants

such as woody species (Jump et al., 2009).

Hitherto, most latitudinal range shifts have been docu-

mented in the Northern Hemisphere, with few studies focus-

ing on the Southern Hemisphere (including Antarctica)

(Table 1), consistent with previously suggested differences in

observed climate-induced changes in natural and managed

ecosystems between the Northern and Southern Hemispheres

(IPCC, 2007a). Additionally, in the Northern Hemisphere,

most latitudinal range shifts have been documented within

the Arctic, boreal, temperate, and Mediterranean biomes, that

is, from middle to high latitudes (Table 1). Only few studies of

continental extent have covered tropical and subtropical areas

below 301 latitude (Beaugrand et al., 2002; Hitch and Leberg,

2007; La Sorte and Thompson, 2007). Hence, there is a

striking dearth of evidence for contemporary latitudinal range

shifts from areas between –601 and 301 latitude (Table 1). The

paucity of evidence from the Tropics and the Southern

Hemisphere (except Antarctica) may have several explanations

(IPCC, 2007a): (1) a marked scarcity of good historical data

from developing countries that are suitable for resampling

along the latitudinal gradient between –601 and 301 latitude;

(2) a lack of taxonomical knowledge, especially in the Tropics,

making it difficult to study multispecies latitudinal range

shifts; (3) logistic problems, making it even more difficult to

collect, access, and manage ecological data in these regions of

the world; (4) the fact that mean annual temperature within

the Tropics is approximately constant between –211 and 211

latitude, at any given elevation, so that little opportunity for

latitudinal escape from warming exists (Colwell et al., 2008);

and (5) a huge imbalance of the Earth’s land mass distribution

with relatively little terrestrial area between –601 and 01 lati-

tude, which probably contributes to the paucity of evidence

for contemporary latitudinal range shifts for terrestrial eco-

systems at these latitudes.

Elevational Range Shifts

A selective review of historically based studies documenting

modern elevational range shifts was also carried out. Although

most studies have reported range shifts toward higher ele-

vations for a majority of taxa (Table 2), these studies have also

reported, to a lesser extent, unexpected range shifts toward

lower elevations (Lenoir et al., 2010a). A few studies have even

reported a general directional shift toward lower elevations for

some taxonomic groups (Hickling et al., 2006; Crimmins et al.,

2011). In Northern California, for instance, most plant species

shifted downward to track regional changes in climatic water

balance rather than temperature (Crimmins et al., 2011). This

shows that range shifts may differ not only in extent, but also

in direction due to regional variation in climate change.

Contrary to latitudinal range shifts, as many elevational

range shifts have been reported for plants as for animals

(Table 2). Across all taxonomic groups, elevational range shifts

toward higher elevations have been documented through

changes either at the trailing/rear edge (Wilson et al., 2005;

Frei et al., 2010), the leading/front edge (Hickling et al., 2006;

Frei et al., 2010), or the core (Konvicka et al., 2003; Lenoir

et al., 2008) of species ranges. Therefore, all possible patterns

of upward shift between march- and lean-range shifts have

been documented along the elevational gradient (Kelly and

Goulden, 2008; Moritz et al., 2008; Raxworthy et al., 2008;

Bergamini et al., 2009). However, it has been suggested that

plants, especially long-lived plants such as woody species, are

more likely to lean rather than march their elevational range

upward (Breshears et al., 2008).

Just as for latitudinal range shifts, there is a strong im-

balance of evidence for observed elevational range shifts under

contemporary climate change between the Northern and

Southern Hemispheres, most likely due to the higher density

of mountainous area in the Northern Hemisphere relative to

the Southern Hemisphere. Indeed, most elevational range

shifts reports across taxonomic groups have been documented

in the Northern Hemisphere, especially between 301 and 601

latitude (Table 2). Although there is a strong focus on tem-

perate ecosystems in the Northern Hemisphere, most of the

examples have been reported for subalpine, alpine, and nival

ecosystems. Therefore, research efforts on elevational range

shifts have been focusing on the coldest biomes (Antarctic,

Latitudinal and Elevational Range Shifts under Contemporary Climate Change 603

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Arctic, and alpine) with much less research on elevational

range shifts in hotter areas such as Mediterranean and tropical

biomes (Table 2). However, on the contrary to the latitudinal

range shifts, there is relatively many studies documenting

elevational range shifts in the Tropics, for example, for plants,

insects, amphibians, and reptiles (Pounds et al., 1997; Rax-

worthy et al., 2008; Chen et al., 2009; Feeley et al., 2011; Juvik

et al., 2011). Continental-scale studies of elevational range

shifts under contemporary climate change are still lacking

(Table 2).

Table 1 Examples of observed latitudinal range shifts under contemporary climate change

Taxonomic

group

No. taxa Latitudinal

zone (deg.)

Spatial

extentaNo. years T/R C L/F Main changes Reference

Crustaceans 1 [ÿ 90; –60] C 75 x x x Decreasing abundance Atkinson et al. (2004)

Chordates 1 [ÿ 90; –60] C 75 x x x Increasing abundance Atkinson et al. (2004)

Moths 1 [30; 60] R 30 x Northward range shift Battisti et al. (2005)

Fishes 2 [30; 60] R 80 x Increasing abundance Beare et al. (2004)

Plankton 25 [30; 90] C 40 x Increasing richness Beaugrand et al. (2002)

Plankton 11 [30; 90] C 40 x Decreasing richness Beaugrand et al. (2002)

Birds 116 [60; 90] R 10 x Northward range shift Brommer (2004)

Birds 34 [60; 90] R 10 x Absence of range shift Brommer (2004)

Butterflies 4 [30; 60] R 35 x Northward range shift Franco et al. (2006)

Odonates 24 [30; 60] R 35 x Northward range shift Hickling et al. (2005)

Odonates 4 [30; 60] R 35 x Northward range shift Hickling et al. (2005)

Arthropods 280 [30; 60] R 25 x Northward range shift Hickling et al. (2006)

Herptiles 3 [30; 60] R 25 x Southward range shift Hickling et al. (2006)

Fishes 15 [30; 60] R 25 x Northward range shift Hickling et al. (2006)

Birds 22 [30; 60] R 20 x Absence of range shift Hickling et al. (2006)

Mammals 9 [30; 60] R 25 x Northward range shift Hickling et al. (2006)

Butterflies 46 [30; 60] R 35 x Absence of range shift Hill et al. (2002)

Butterflies 4 [30; 60] R 35 x Absence of range shift Hill et al. (2002)

Birds 27 [30; 60] C 25 x Northward range shift Hitch and Leberg (2007)

Birds 29 [30; 60] C 25 x Absence of range shift Hitch and Leberg (2007)

Fishes 75 [30; 60] L 20 x x x Decreasing abundance Holbrook et al. (1997)

Birds 254 [30; 60] C 30 x x Northward range shift La Sorte and Thompson

(2007)

Plants 7 [30; 60] L 15 x Decreasing abundance Lesica and McCune (2004)

Algae 13 [30; 60] R 55 x Northward range shift Lima et al. (2007)

Algae 26 [30; 60] R 55 x Northward range shift Lima et al. (2007)

Acarids 1 [60; 90] R 15 x Increasing abundance Lindgren et al. (2000)

Mammals 4 [30; 60] R 125 x Increasing abundance Myers et al. (2009)

Mammals 5 [30; 60] R 125 x Decreasing abundance Myers et al. (2009)

Butterflies 35 [30; 90] C 50 x x Northward range shift Parmesan et al. (1999)

Fishes 43 [30; 60] R 25 x Northward range shift Perry et al. (2005)

Plankton B400 [30; 60] C 45 x x x Northward range shift Richardson and Schoeman

(2004)

Invertebrates 11 [30; 60] L 65 x Increasing abundance Sagarin et al. (1999)

Invertebrates 7 [30; 60] L 65 x Decreasing abundance Sagarin et al. (1999)

Plants 2 [ÿ 90; –60] R 25 x Increasing abundance Smith (1994)

Plants 4 [60; 90] R 50 x Increasing abundance Sturm et al. (2001)

Birds 59 [30; 60] R 20 x Northward range shift Thomas and Lennon (1999)

Birds 42 [30; 60] R 20 x Absence of range shift Thomas and Lennon (1999)

Lichens 12 [30; 60] R 20 x Increasing richness van Herk et al. (2002)

Lichens 66 [30; 60] R 20 x Decreasing richness van Herk et al. (2002)

Plants 1 [30; 60] R 50 x Northward range shift Walther et al. (2005a, b)

Butterflies 46 [30; 60] R 30 x Decreasing abundance Warren et al. (2001)

Birds 41 [30; 60] R 20 x Absence of range shift Zuckerberg et al. (2009)

Birds 129 [30; 60] R 20 x Northward range shift Zuckerberg et al. (2009)

Birds 43 [30; 60] R 20 x Northward range shift Zuckerberg et al. (2009)

aC: continental; L: local; R: regional.

Examples of recent studies reporting, for both marine and terrestrial ecosystems, modern latitudinal range shifts of organisms at the trailing/rear edge (T/R), the core (C), or the

leading/front edge (L/F) of their latitudinal range. We restricted our literature survey to studies that have explicitly used historical occurrence data to assess range shifts under

modern climate change. We focused on the following two types of historically based studies (Parmesan, 2006): (1) those that directly assess range shifts by comparing historical

and modern data on species distribution across their entire ranges or sections of ranges; and (2) those that infer range shifts by comparing historical and modern data on species

composition or abundance within local communities. Shaded cells represent reports for marine ecosystems, whereas nonshaded cells represent reports for terrestrial ecosystems.

604 Latitudinal and Elevational Range Shifts under Contemporary Climate Change

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Latitudinal and Elevational Range Shifts are Laggingbehind Climate Change

It is clear that contemporary climate change is driving lati-

tudinal and elevational range shifts in species distribution

worldwide (Tables 1 and 2). However, the magnitude of these

biotic responses is unlikely to match the magnitude of con-

temporary climate change due to differences in speed,

magnitude, and direction between the climatic changes and

the biotic responses as well as differences among species. Time

lags between geographical range shifts and past climate

changes have already been reported for plants and animals

(Davis, 1986; Svenning and Skov, 2007; Araujo et al., 2008).

One way to assess the importance of such time lags in the

biotic responses to contemporary climate change is to com-

pare the magnitude of observed latitudinal and elevational

Table 2 Examples of observed elevational range shifts under contemporary climate change

Taxonomic

group

No. taxa Latitudinal

zone (deg.)

Spatial

extentaNo. years T/R C L/F Main changes References

Birds 41 [30; 60] L 30 x Absence of range shift Archaux (2004)

Moths 1 [30; 60] L 30 x Upward range shift Battisti et al. (2005)

Trees 3 [30; 60] L 40 x Increasing abundance Beckage et al. (2008)

Trees 3 [30; 60] L 40 x Decreasing abundance Beckage et al. (2008)

Bryophytes 61 [30; 60] R 125 x x x Upward range shift Bergamini et al. (2009)

Moths 102 [0; 30] L 40 x Upward range shift Chen et al. (2009)

Moths 28 [0; 30] L 40 x Downward range shift Chen et al. (2011b)

Moths 28 [0; 30] L 40 x Upward range shift Chen et al. (2011b)

Moths 109 [0; 30] L 40 x Upward range shift Chen et al. (2011b)

Moths 109 [0; 30] L 40 x Upward range shift Chen et al. (2011b)

Plants 64 [30; 60] R 80 x Downward range shift Crimmins et al. (2011)

Epiphyte 1 [30; 60] R 95 x Upward range shift Dobbertin et al. (2005)

Trees 38 [ÿ 30; 0] L 5 x Upward range shift Feeley et al. (2011)

Butterflies 4 [30; 60] R 35 x Upward range shift Franco et al. (2006)

Plants 125 [30; 60] L 95 x x Upward range shift Frei et al. (2010)

Plants 60 [30; 60] L 95 x Increasing richness Grabherr et al. (1994)

Arthropods 280 [30; 60] R 25 x Upward range shift Hickling et al. (2006)

Herptiles 3 [30; 60] R 25 x Downward range shift Hickling et al. (2006)

Fishes 15 [30; 60] R 25 x Upward range shift Hickling et al. (2006)

Birds 22 [30; 60] R 20 x Absence of range shift Hickling et al. (2006)

Mammals 9 [30; 60] R 25 x Upward range shift Hickling et al. (2006)

Butterflies 15 [30; 60] R 35 x Upward range shift Hill et al. (2002)

Plants 140 [30; 60] L 120 x Upward range shift Holzinger et al. (2008)

Plants 22 [0; 30] L 50 x Increasing richness Juvik et al. (2011)

Plants 10 [30; 60] L 30 x x x Upward range shift Kelly and Goulden (2008)

Butterflies 119 [30; 60] R 50 x Upward range shift Konvicka et al. (2003)

Trees 7 [60; 90] L 60 x Upward range shift Kullman (2002)

Plants 171 [30; 60] R 100 x Upward range shift Lenoir et al. (2008)

Plants 60 [30; 60] R 20 x x x Upward range shift Lenoir et al. (2010b)

Plants 22 [ÿ 90; –60] L 40 x x Upward range shift le Roux and McGeoch (2008)

Mammals 28 [30; 60] L 90 x x x Upward range shift Moritz et al. (2008)

Plants 78 [60; 90] R 40 x Upward range shift Odland et al. (2010)

Plants 93 [30; 60] L 50 x Upward range shift Parolo and Rossi (2008)

Plants 10 [30; 60] L 10 x Increasing abundance Pauli et al. (2007)

Plants 10 [30; 60] L 10 x Decreasing abundance Pauli et al. (2007)

Birds 94 [0; 30] 30 x Upward range shift Peh (2007)

Birds 94 [0; 30] 30 x Absence of range shift (Peh 2007)

Amphibians 50 [0; 30] L 5 x x x Decreasing abundance Pounds et al. (1997)

Reptiles 30 [ÿ 30; 0] R 10 x x x Upward range shift Raxworthy et al. (2008)

Birds 1 [30; 60] R 30 x x Upward range shift Tryjanowski et al. (2005)

Plants 31 [30; 60] L 100 x Increasing richness Vittoz et al. (2008)

Plants 18 [30; 60] L 100 x Increasing richness Walther et al. (2005a)

Butterflies 16 [30; 60] R 35 x Upward range shift Wilson et al. (2005)

Birds 41 [30; 60] R 20 x Absence of range shift Zuckerberg et al. (2009)

Birds 129 [30; 60] R 20 x Downward range shift Zuckerberg et al. (2009)

Birds 43 [30; 60] R 20 x Absence of range shift Zuckerberg et al. (2009)

aC: continental; L: local; R: regional.

Examples of recent studies reporting, for terrestrial ecosystems, modern elevational range shifts of organisms at the trailing/rear edge (T/R), the core (C), or the leading/front edge

(L/F) of their elevational range. We restricted our literature survey to studies that have explicitly used historical occurrence data to assess range shifts under modern climate change

(see Table 1 for further explanations).

Latitudinal and Elevational Range Shifts under Contemporary Climate Change 605

Author's personal copy

range shifts of different taxonomic groups with the magnitude

of expected range shifts given climate change alone. In the

French mountain forests, for instance, the core of the ele-

vational range for plants (n¼171 species) shifted toward

higher elevations (mean: 29 m per decade B0.17 1C per

decade, considering an adiabatic lapse rate of 0.6 1C per 100 m

in that region) only 40% of that expected simply from the

observed temperature increase in that region between 1965

and 2005 (0.45 1C per decade B75 m per decade) (Lenoir

et al., 2008). For purposes of comparison, the core of the

elevational range for reptiles and amphibians (n¼30 species)

in Madagascar shifted toward higher elevations (mean: 65 m

per decade B 0.32 1C per decade, considering an adiabatic

lapse rate of 0.5 1C per 100 m in that region) approximately

85% of that expected from the observed temperature increase

in that region between 1984–1993 and 1994–2003 (0.37 1C

per decade B74 m per decade) (Raxworthy et al., 2008). This

suggests that the magnitude of lags in range-shift responses to

climate change may vary considerably among taxonomic

groups (Davis, 1986).

Many different factors might be responsible for such

delayed responses. Life span is obviously one of those factors.

Species with shorter life spans such as insects and small

mammals should be able to respond relatively fast to climate

change, being delayed only a few years after a climatic event,

whereas species with longer life spans, including species with

slow maturation such as large mammals and many trees,

should show much longer time lags, only responding to long-

term climatic changes (Davis, 1986). Accordingly, it has been

documented under contemporary climate change that species

with shorter life cycles have shifted poleward and upward with

greater magnitude than species with longer life cycles (Perry

et al., 2005; Lenoir et al., 2008) (Figure 3). Time lags may also

result from low dispersal ability. Species with limited dispersal

ability, such as many plants, may not be able to rapidly col-

onize newly suitable climatic areas, even in the vicinity of their

geographic ranges, causing time lags in their colonization of

these areas (Davis, 1986). Suggestive of this mechanism, larger

elevational range shifts have been reported for species with

lighter diaspores as compared to species with heavier dia-

spores in the Italian Alps (Parolo and Rossi, 2008). Colon-

ization of newly suitable habitats may also be delayed by

competition from resident species (Davis, 1986). For instance,

recruitment of Fagus sylvatica in southern France is delayed in

grassland habitats due to its low tolerance to competition

from herbaceous species (Kunstler et al., 2007). Time lags in

range-shift responses to climate change is furthermore likely to

be most pronounced for species with small geographic ranges,

as such species are often concentrated in long-term

stable areas (Svenning and Skov, 2007; Araujo et al., 2008),

and thus likely to be particularly dispersal-limited. Linked to

this idea that species with small geographic ranges might have

pronounced time lags, range-shifts gaps – that is, lacking

overlap between the current range and a future suitable range

– are likely to occur and to increase the magnitude of these

time lags if the geographic range of a species is smaller than

the projected range shift under climate change (Colwell et al.,

2008). Crash-range shifts are a likely expectation for such

species facing range-shift gaps under contemporary climate

change. Finally, physiological adaptation to changing climate

conditions through either evolutionary response or pheno-

typic plasticity may in some cases also cause ranges to shift less

than expected from climate change (Davis and Shaw, 2001).

In addition to these biotic factors, spatial features may also

affect time lags in range-shift responses to climate change.

Considering contemporary climate warming, closeness among

spatial isotherms is probably the most obvious factor. Indeed,

time lags in range-shift responses to contemporary climate

change are likely to be longer in flatland areas, which offer no

short-distance escapes for species facing climate change and

where the velocity of climate change is expected to be much

stronger, compared to mountainous terrain (Loarie et al.,

2009). Therefore, time lags in latitudinal-range-shift responses

to climate change should be longer than time lags in ele-

vational-range-shift responses to climate change for a given

species, particularly within the tropics, where latitudinal gra-

dients in temperature are virtually absent (Colwell et al.,

2008). However, Chen et al. (2011a) found greater elevation

lags than latitudinal lags in their meta-analysis, arguing that

relatively strong elevation lags may arise from topographic and

topoclimatic complexity in mountainous areas. Additionally,

habitat fragmentation is likely to lengthen time lags in range-

shift responses to climate change. In the context of future

climate change, it has been simulated that the range of forest

plant species in central Belgium will lag behind climate

change, especially within the most fragmented landscapes

(Honnay et al., 2002).

Ecological Consequences

Ecological consequences of latitudinal and elevational range

shifts under contemporary climate change are numerous, af-

fecting biodiversity, communities, and ecosystem functioning.

The often mentioned potential polar and mountaintop ex-

tinctions constitute an obvious example of the likely bio-

diversity consequences of climate-change-induced range shifts.

Indeed, species restricted to these cold extremes have nowhere

mosses

and

shrubs

Trees

lllustration: P. Mouche/Terre Sauvage

and

5m

8+

ferns,

Herbs,

+ 20

m

Figure 3 Differential range shifts between short- (herbs, ferns, and

mosses) and long-lived (trees and shrubs) plant species along the

elevational gradient of the French mountain forests under

contemporary climate change (Lenoir et al., 2008). The drawing is

from P Mouche/Terre Sauvage. Reproduced from Etienne Hurault.

606 Latitudinal and Elevational Range Shifts under Contemporary Climate Change

Author's personal copy

else to go to escape climate warming. Therefore, their potential

habitat is shrinking through a general reduction in surface

area. Such range-restricted species may experience severe range

contractions and may be the first groups in which species have

already been driven to extinction by contemporary climate

change (Parmesan, 2006). Endemic species restricted to low

and isolated mountains in the temperate and tropical zones

are especially threatened with extinction as a consequence of

range-shift responses to climate warming (Krajick, 2004), for

example, alpine endemics restricted to summits of low

mountains lacking nival belts or subalpine endemics restricted

to summits of low mountains lacking alpine and nival belts.

In the long run, these low mountains may act as climatic traps

for range-restricted species facing climate warming (Forero-

Medina et al., 2011). However, contemporary climate change at

these high latitudes (poles) and elevations (mountaintops)

also allow the immigration of new species from lower lati-

tudes and elevations, which generally host a higher diversity,

with the net result of these contemporary range shifts being a

gradual increase in species richness toward the poles and

mountain summits (Grabherr et al., 1994; Walther et al.,

2005a), though time lags in range-shift responses to climate

change are expected and may lead to both extinction debt and

immigration credit (Jackson and Sax, 2010). Not only gradual

increase in species richness is expected toward the poles and

mountain summits as a net result of contemporary range

shifts, but also lowland biotic attrition may occur in the Tro-

pics as a net result of range contractions and extinctions

without any possibility for compensation, contrary to higher

latitudes, where immigrants can come from lower latitudes

(Colwell et al., 2008). Similar attrition may also occur in extra-

tropical lowland areas due to migrational lags if source pools

are distantly located (Svenning and Condit, 2008).

Communities may also be strongly affected by con-

temporary climate-driven range shifts, mainly throughout

changes in species composition. Indeed, shifting species might

be forced to interact with a new set of resident species from the

recipient community, thus leading to community reorganiza-

tion (Walther, 2010). Such new species assemblages may result

in the formation of nonanalog communities, that is, com-

munities that are different in species composition from any

communities present at a selected reference point in time

(Keith et al., 2009). Range-shift-induced community reorgan-

ization may have important ecological consequences. For ex-

ample, plant community homogenization among alpine

summits has been reported in the Alps (Jurasinski and

Kreyling, 2007) as a consequence of the upward shift of alpine

plant species in the same region (Walther et al., 2005a).

However, such community reorganization occurs not

only ‘‘horizontally,’’ that is, within the same taxonomic group,

but also ‘‘vertically,’’ that is, across taxonomic groups and

trophic levels, and, thus, could also strongly affect ecological

networks.

Climate change effects on ecological networks could also

have consequences for ecosystem functioning (Walther, 2010).

For instance, species range shifts within a given trophic level

may induce trophic cascades through bottom-up or top-down

effects. An empirical example is the impact of sea surface

warming on the poleward shift of phytoplankton in the

Northeast Atlantic, which has been reported to propagate up

the food web throughout bottom-up control on copepod

herbivores and zooplankton carnivores (Richardson and

Schoeman, 2004). In this case, the tight trophic coupling be-

tween phytoplankton, copepods, and zooplankton carnivores

allows a preservation of the main functions ensured by this

marine food web. However, such ecosystem functioning can

be disrupted if the coupling between interacting species is too

loose or if interacting species respond differently to climate

change. The most striking evidence of such disruptions can be

found in reports of pest outbreaks after range shifts of pest

species released from the top-down control of their predators.

As a typical example, recent range shifts of the pine pro-

cessionary moth, Thaumetopoea pityocampa, has caused major

outbreaks in Europe by switching hosts (Battisti et al., 2005).

Ecological consequences of these large-scale herbivore out-

breaks may involve changes in the carbon sequestration

budgets. Other important implications of species range shifts

for ecological networks and ecosystem function concern the

loss or gain of new functional types. Indeed, some ecosystems

are likely to lose or gain functions as a result of species range

shifts and community reorganization. In northern Arizona,

recent vegetation shifts resulting from extreme mortality of

pinyon, Pinus edulis, have affected the structure of the pinyon-

juniper woodland, causing loss of avian seed dispersers, the

disappearance of ectomycorrhizas specific to pinyon, and

above- and below-ground competitive release of juvenile trees

(Mueller et al., 2005). In parallel, the recent expansion of

shrubs in the Arctic (Sturm et al., 2001) is changing the

physiognomy of the arctic vegetation by allowing shrubs, a

new functional type, to establish, with likely cascading eco-

logical consequences. Indeed, it is suggested that increased

vegetation cover in the Arctic may not only provide food re-

sources for invasions of new animal species such as moths,

already now causing outbreaks in the Arctic, but also cause

ecological changes that may affect the Earth’s climate itself,

notably via changes in regional land–atmosphere greenhouse

gas balances and carbon budgets (Post et al., 2009).

Conservation Implications

Latitudinal and elevational range shifts and their resulting

ecological consequences constitute a major challenge for con-

servation biology. Many recommendations for climate change

adaptation strategies for biodiversity conservation have been

recently listed in a literature review (Heller and Zavaleta,

2009). Here the focus is on two heavily discussed management

recommendations: the traditional habitat-management rec-

ommendations and the controversial species-management

recommendations. Most management recommendations so far

have focused on conserving or restoring habitat quality to

enhance species survival. The top-most-cited recommendation

for biodiversity conservation under climate change is to in-

crease habitat connectivity by establishing corridors, removing

barriers for dispersal, and planning reserves close to each other

so as to re-create series of stepping stones of high-quality

habitats and increase possibilities for species range shifts

(Heller and Zavaleta, 2009). Other habitat-management rec-

ommendations include creating and managing buffer zones

within and around nature reserves (Heller and Zavaleta, 2009).

Latitudinal and Elevational Range Shifts under Contemporary Climate Change 607

Author's personal copy

For instance, topographic heterogeneity is associated with

low climate change velocity (Loarie et al., 2009) and should

enhance the long-term conservation capacity of nature reserves

(Ackerly et al., 2010) by providing short-distance escapes for

species facing climate change (Scherrer and Korner, 2011).

However, depending on the magnitude of climate change

and the degree of topographic heterogeneity within species

ranges, topographic heterogeneity may just temporarily buffer

biodiversity before starting to act as a climatic trap by in-

creasing spatial isolation and extinction risks (Forero-Medina

et al., 2011).

Nevertheless, it is also increasingly clear that the traditional

habitat-management recommendations may be insufficient for

safeguarding biodiversity from climate-driven extinctions, and

therefore species-management recommendations are being

proposed, notably assisted migration (also called assisted col-

onization or managed translocation) (Thomas, 2011). Among

others, dispersal-limited or small-range species might not be

able to be saved by more traditional habitat-management rec-

ommendations, such as increasing landscape connectivity to

enhance species migration (Heller and Zavaleta, 2009; Thomas,

2011). Assisted migration is controversial, especially given the

fear that translocated species could increase the risk of native

species extinctions in their new recipient habitats (Ricciardi and

Simberloff, 2009). However, for climate-threatened, dispersal-

limited, or small-range species, it has been suggested that

translocation might be the only successful management strat-

egy, and therefore should be implemented provided that risks

of extinction to native species in recipient habitats are small

(Morueta-Holme et al., 2011; Thomas, 2011). To help managers

in making decisions between habitat- and species-management

recommendations, a decision framework has already been

proposed for assessing when species translocations are possible

(Hoegh-Guldberg et al., 2008).

Future Challenges

Despite the increasing number of studies documenting lati-

tudinal and elevational range shifts under contemporary cli-

mate change, a better understanding of current range shifts is

still needed for developing well-founded predictive models of

future range shifts. More research efforts are needed to better

understand climate change effects on species ranges at low

latitudes and especially within the Tropics, the most species-

rich part of the world and the area hosting most threatened

taxa (IUCN, 2004). So far, evidence of recent species range

shifts in the Tropics have been mostly documented along

elevational gradients (Pounds et al., 1997; Raxworthy et al.,

2008; Chen et al., 2009; Feeley et al., 2011; Juvik et al., 2011)

with at most a handful of reports for plants (Feeley et al., 2011;

Juvik et al., 2011). Although the magnitude of climate warming

varies geographically, most tropical areas have warmed just as

much as temperate areas (IPCC, 2007a). Furthermore, small

changes in temperature conditions in the Tropics may have

much stronger biotic impact than similar changes in tempe-

rate areas. Notably, recent research into the thermal tolerance

of terrestrial insects shows that tropical insects are closer to

their maximum thermal tolerance than temperate insects

(Deutsch et al., 2008), suggesting that tropical species and

their ranges may be particularly sensitive to climate warming.

Importantly, crash-range shifts are therefore more likely in the

Tropics (Pounds et al., 1997) than elsewhere. Even more

alarming, there do not exist biodiversity source pools now

living in hotter places (as such areas do not exist) that

are available to replace climatically displaced or declining

species in tropical lowland ecosystems, potentially leading to

lowland biotic attrition in the lowland tropical areas (Colwell

et al., 2008), with unknown consequences for ecosystem

functioning.

Another future challenge is to achieve a better under-

standing of the mechanisms involved in species range shifts.

Notably, it is still unresolved how the different population

dynamics involved at both the range edges is driving species

range shifts. Are colonization and establishment processes

occurring more rapidly at the front edge of a species’ range

than mortality and extinction processes at the rear edge? For

instance, the rear edge of a species’ range is likely to host a

relatively high genetic diversity due to selection pressure over

time (Hampe and Petit, 2005), thus having a high potential

for delaying range contractions under contemporary climate

change. Therefore, rear edges may be disproportionately im-

portant for the survival and evolution of species (Hampe and

Petit, 2005). Surprisingly, fewer studies have focused on con-

temporary range shifts at the trailing edge of species’ ranges in

comparison to the leading edge. This lack of focus on range

shifts at the trailing edge of species’ ranges is even more im-

portant along the latitudinal gradient for long-lived plants

such as woody species (Jump et al., 2009). In short, there is

also an urgent need to improve our understanding of species

range shifts at the trailing edge of species’ ranges. Additionally,

the strong variation in the magnitude of contemporary range

shifts among species raises questions about the underlying

reasons and whether the magnitude of shifts is linked to

species traits. A recent study focusing on the relationship be-

tween species traits and the magnitude of recent shifts at the

leading edge found low explanatory power, suggesting that

trait-based range shift forecasts face several challenges (Angert

et al., 2011). Finally, the occurrence of unexpected range shifts

(such as stasis or even downslope shifts) is an understudied

phenomenon of high importance for predicting range re-

sponses to future climate change (Lenoir et al., 2010a). This

is linked to our current lack of knowledge regarding the effect

of biotic interactions and dispersal constraints on species

distributions.

Addressing these challenges will allow the development of

improved predictive models of future range shifts. Indeed,

with exactly this aim, recent discussions have raised the need

for improving the field of species distribution modeling

(SDM) by developing more mechanistic, hybrid approaches

such as coupling stochastic population models with dynamic

bioclimatic habitat models (Keith et al., 2008) or by better

accounting for nonclimatic factors and ecological complexity,

notably biotic interactions and dispersal constraints (Morin

and Lechowicz, 2008).

See also: Climate Change and Extinctions. Climate, Effects of.

Evolution in Response to Climate Change. Introduced Species,

608 Latitudinal and Elevational Range Shifts under Contemporary Climate Change

Author's personal copy

Impacts and Distribution of. Migration. Phenological Shifts in

Animals Under Contemporary Climate Change. Species Distribution

Modeling. Translocation as a conservation strategy. Trophic Cascades

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