A review of the influence of climate change on species distributions in CA:

montane and desert biomes

Brit Brown
Abstract
Our aim in this paper is to provide an overview of the potential effects of climate change on
various ecosystems present throughout California. Presented here are predictions for desert and montane
biomes gathered from case studies performed by experts in the field of ecology. According to studies,
mountain ecosystems are expected to experience decreased snowfall and elevated temperatures further
upslope. In addition, species shifts upslope or to higher latitudes to counteract the rise in temperature are
predicted in both plants and animals. The data further predicts that desert species are expected to
experience intense pressure to disperse to higher latitudes to maintain thermal optimums as global
temperatures increase. Finally, due to the low elevation of most deserts, experts say that many species are
projected to experience extreme range reduction and possible local extinction with increasing
Climate Change & the CA Flora

temperatures.

Introduction
A study by Loarie in 2008 showed that California
is considered a major biodiversity hotspot with 2387
endemic plant species found through out the state. The
traditional definition of a hotspot is an area that contains a
minimum of 1500 endemic plant species and has already
lost at least 70% of other plant species. The high

Figure 1. Study area and Maxent diversity projections of the best known 591 species. (A) The province divided into six floristic regions
(solid lines): Northwestern California (NW), Central Western California (CW), Southwestern California (SW), the Cascade Ranges (CaR), the Great Central
Valley (GV), and the Sierra Nevada (SN). The province includes most of California (dashed line) and portions of Oregon and Mexico. We include a
surrounding buffer of equal area (colored areas outside solid line). Colors represent elevation in meters. (B) Projected present diversity. (CJ)
Projected diversity 80 years from now modeled with increasing amounts of future climate change: (CF) Plants cannot disperse. (GJ) Plants can
disperse to all suitable areas. (C, F, G, H) Simulations based on the lower sensitivity PCM model. (E, F, I, J) Simulations based on the highersensitivity HadCM3 model. (C, E, G, I) Lower emissions scenario (B1). (D, F, H, J) Higher emissions scenario (A1FI).
doi:10.1371/journal.pone.0002502.g001

amount of biodiversity in hotspots means that

Figure 1. Plant diversity projections of the 591 best

known species
in present
California
overof 2068
thespecies
next change
80 years.
in range size. In high emission scenarios (A1FI) with
Figure 2B shows
diversity projections
dispersal, we project species range centroids to shift by an average
from the MLGLM generalized linear model. Diversity patterns in
(A) represents
the tomajor
floristic
provinces
into CA
with(see Fig. S1).
of up
151 kilometers
figure 2B are similar
those in figure
2A except that
the range
As one might expect, species tend to move to higher elevations
derived diversity is lower in Northwestern California and
divisionsmap
drawn
in solid
and
dashed
lines.
shows
and(B)
often northward
(see Fig. S2). Interestingly, these trends result
Southwestern
California.
The patterns
in figure
2B differ from
the
in divergent projections for elements of the flora. Given
projections in that diversity is lower in the Sierra Nevada
the bestMaxent
representation
present-day
among
Californias geography,
movement to higher elevations often
and
Southwestern California andof
higher
in coastal Northwesterndiversity
means taking a southward path. Figure 3A illustrates two
California.
plant species.
(C-J)
represent
varying
representativeof
species that presently have essentially adjacent
Changing patterns
of diversity
projected from
the multileveldegrees
ranges. In the future, we project their ranges to be widely separate,
model are very similar to the patterns of diversity projected from
with one
moving
south to higher elevation regions of the Sierra
projected
future
diversity
with
(Cthe&coast
G)andbeing
the
bestMaxent.
In general,
diversity shifts
towards
Nevada, and the other moving north and towards the coast.
northwards, and the degree depends on the dispersal assumptions,
Figure
3B illustrates the broader consequences, based on analysis
case scenarios
and
(Fsensitivity
& J)of climate
being
the The
worst
case
emission scenarios,
and the
simulations.
of the centroids of the species ranges. They are ecologically
following results on species movement and range size change are
dramatic.
Within
the six major regions, substantial numbers of
scenarios.
In all
cases,
there
isspecies.
an obvious
shift
north
from Maxent
projections
of the 591
best known
species move in diametrically opposite directions typically north
of northwest, and south of southeast. In the Cascade Ranges and
and toward
the
coast.
(Loarie
et
al.,
2008)
Species movement
the Sierra Nevada, species at high elevations tend to move south to

conservation of these areas would preserve a high

amount of the worlds species richness. According
to Ackerly et al. in 2010, the diverse terrain in
California is predicted to be a major factor in how

Changes in diversity reflect the overall consequences of local

extirpation and species dispersal. These patterns do not address
the potential fate of individual species. For that reason, we also
examined individual species fate in terms of projected geographic
shifts in species mean elevation, range centroid, and percent
PLoS ONE | www.plosone.org

higher elevations. Those at lower elevations, like those in other

regions, are a mix of species, some of which move south and others
that move north. (See Fig. S3 for scenarios not shown here).
The results shown here are for the largest projected changes in
temperature (HadCM3, A1FI), allowing dispersal. We obtain
3

June 2008 | Volume 3 | Issue 6 | e2502

NATURE | Vol 462 | 24/31 December 2009

13

A2
13
A1B
12
B1
11

A2

A1B

Temperature (C)

12 states that plant distribution and abundance

the state will be affected by the changing global climate. He
11

B1

will play a major role in maintaining species diversity. Multiple sources predict that plant species ranges
10

10

8
and community compositions will be drastically affected by1950
increasing
temperatures
(Loik
et al.,
2012,
1950
2000
2050
2100
2000
2050
2100
10

1.00

A1B speed (km yr1)

Dolanc et al., 2014, McLaughlin et al., 2012, Cole et al., 2011, & Concilio et al., 2013) . Dispersal

northward and upslope is commonly thought to alleviate stress caused by increasing temperatures, but not
all species are capable of this shift. Another interesting variable discovered by Rapacciuolo in 2014 0.1
is
c

0.01
that movement upslope in California generally corresponds with travel in a southern direction since the

Figure 2 | The velocity of temperature change globally. a, Temporal

highest elevations can be found further south near the Mojave

desert. from
This
meansacross
thatthree
as emissions
speciesscenarios
disperse
gradients calculated
20002100
(A2,
to either higher latitudes or higher elevations,

A1B and B1). b, Temporal gradients calculated from 20002050 and

20502100 across three emissions scenarios. Trends plotted here are the
populations
may
end
upland
splitting
small
refugia.
A global
map ofisolated
climate velocity
average
of the
global
surface. c,into
calculated using the 20502100 Special Report on Emissions Scenarios
(SRES) A1B emissions scenario temporal gradient.

According to Rapacciuolo, timing in phenology, dispersal style, and varied habitats will all affect whether
of protected areas vary greatly across biomes (see Supplementary
20). To explore the interaction between protected area sizes
and velocities required to keep pace with climate change, we calculated residence times, defined as the diameter of each protected area
divided by velocity (km/km yr21 5 yr). Assuming that protected

or not species can thrive in the predicted warmer climate.

Fig.

Montane/Alpine Systems
In order to survive in a changing climate,

areas are circular an

as the time for curre
dence times exceed
tected areas. Figure
decreasing residence
the inverse of velocit
with the slowest vel
times. There are also
size of protected are
and coniferous fores
biomes despite relat
increased from 4, 7 a
mean residence time
(Fig. 3). In contrast,
their rank despite
example, the deserts
To guide the interp
tions. First, climate c
perature, precipitatio
on mean annual tem
perature is a useful s
change. The directio
less uncertain than
examples document
tions responding as
repeated all analyses
Interestingly, precipi
tainous areas due to
effects. As a result th
overall patterns are s
Second, there is u
dients of climate cha
contributing factors
upper estimates of v
1. Tundra, 74.6

Loarie et al. (2009) have suggested that both plants

2. Montane grasslands and sh

and animals respond by shifting their habitat ranges

1. Tropical and subtropical coniferous forests, 0.08

8. Tropical and subtropical dry broadleaf

forests, 0.42

2. Temperate coniferous forests, 0.11

9. Boreal forests/taiga, 0.43

3. Montane grasslands and shrublands, 0.11

10. Temperate grasslands, savannas and

shrublands, 0.59

habitat, while flat grasslands and deserts require

4. Mediterranean forests, woodlands and scrub, 0.26

11. Tropical and subtropical grasslands, savannas,

shrublands, 0.67

species to travel greater distances to keep pace with

5. Tundra, 0.29

12. Deserts and xeric shrublands, 0.71

climate change.

6. Tropical and subtropical moist broadleaf

forests, 0.33

13. Mangroves, 0.95

In his 2009 study, Loarie demonstrated that

7. Temperate broadleaf and mixed forests, 0.35

14. Flooded grasslands and savannas, 1.26

to more favorable geographic locations. He further

states that mountainous biomes offer the most
dramatic change for the least amount of dispersal in

3. Tropical and subtropical co

4. Tropical and subtropical m

forests, 43.8

5. Tropical and subtropical dr

forests, 22.8

6. Deserts and xeric shrublan

mountains provide many species with an effective

0.001 0.01 0.1

1
Speed (km yr1)

10

0.001 0.01 0.1

1
Speed (km yr1)

10

Figure 3 | The velocity of temperature change by biome. A map of biomes

and histograms of the speed of temperature change within each biome.
Figure
2. Different
velocities
of geometric
Histograms
are orderedestimated
by increasingrates
velocityofaccording
to their
means. needed to keep pace with a future changing
change

shelter against increasing temperatures because of

their low velocity of climate change. It is widely

7. Tropical and subtropical gr

shrublands, 15.0

10

Residence

Figure 4 | Climate resi

represent the ratio of p
(km h21), and are orde
The vertical bar indica

climate. Mountainous and high elevation landscapes

Publishers Limited. All rights reserved
offer the lowest velocity while flatter,2009
moreMacmillan
homogenous
landscapes require a much higher velocity. (Loarie et al.,
2009)

known that minimal shifts upslope can provide

mountainous species with relief from rising

temperatures experienced at lower elevations. This particular strategy is likely to lead to an initial
increase in species richness at higher elevations as populations attempt to evade increasing temperatures.
Rapacciuolo et al. (2014) state that it may also lead to an upslope shift in the lower limits of mountainous
ecosystems. After reviewing the literature, it is apparent that the biggest factor affecting the survival of
montane-alpine biomes is potential habitat availability. As stated by Loarie (2009), when temperatures
rise, species will have to continue to shift upslope to maintain status quo with the environment. The
problem with this strategy is that movement upslope reduces both range size and resource availability, as
mountains are limited in potential available space. A northward shift at stable elevation might enable
mountainous species to survive increasing temperatures while reducing the potential for reduced habitat
range that occurs with simply moving further upslope.

Field data collected by McLaughlin and Zavaleta in 2012

2308 B . C . M C L A U G H L I N & E . S . Z A V A L E T A

show that California valley oak (Quercus lobata) saplings are

more likely to be affected by climate change than adult trees, and
contrary to bioclimate model projections, range contractions are
expected to produce a shift toward bodies of water as opposed to
the more commonly expected northward shift. According to the
authors, bioclimate models are widely used to predict different
species responses to climate change because of their ability to

Fig. 7 Valley oak bioclimate model area inputs based on the

area of the current potential adult distribution (black) (Kueppers et al., 2005), and areas inputs that would be excluded in a
model based on the sapling climate threshold (white). Map
credit: A. Cole.

assess regional-based patterns. In their study, McLaughlin and Zavaleta

point out that different species-environment interactions reduce the
reliability of bioclimate models by introducing complicating factors.
The authors found that the projected bioclimate model for Q. lobata
failed to take into account different life history traits of saplings vs

Figure 3. Bioclimate model

projections for adult Q. lobata
(in black) and estimated
Constriction to microrefugia
sapling projections based on
Evidence, mainly from Quaternary phylogeographic
collected fieldwork.
studies (Keppel et al., 2011), suggests that regional-scale
(McLaughlin
& Zavaleta,
climate refugia
were important
for species persistence
through rapid
climate fluctuations during the last gla2012).

cial maximum and later allowed for postglacial recolonization (reviewed in Dobrowski, 2010; Keppel et al.,
2011). In a current extension of this concept, Loarie
et al. (2008) map predictions of future regional-scale climate refugia for endemic species in California and suggest that areas that harbor species with shrinking
ranges, such as scattered mountainous areas, may provide good conservation targets. While much historical
work focused on large-scale climate refugia there is also
some evidence for species use of local scale or microrefugia (also referred to as cryptic refugia), microclimates that supported small populations of species
beyond the climatic limits of their main distributions
(Rull, 2009; Dobrowski, 2010). Recently, restriction to
favorable microclimates has been shown to occur near
the climatic edges of species distributions (Hennenberg
& Bruelheide, 2003) and it has been proposed, in
theory, that a spatial shift to particular topographic conditions may act as an early indicator of climate change
impacts on a species distribution (Korner, 1999; Thomas et al., 2001). Protecting microrefugia has been suggested as a potential in situ conservation option for

adults. The prediction that the species will be able to persist in its current region with the changing

climate was rejected when compared to actual field data. McLaughlin and Zavaleta believe that the shift
in sapling dispersal toward bodies of water may demonstrate the importance of ground water and drought-

species threatened b
2010), however, actu
related constriction in
demonstrated with cur
Work on microrefug
graphic microclimates (
2010), yet other forms
also may exist. Becau
groundwater (Jepson,
fin, 1973), we expected
might exist for this sp
ter availability, and th
tant for recruitment wi
areas. Our finding of
saplings around surfa
expanding/persisting
this hypothesis. In the
lings were more restri
that a climate-based sh
oaks may be already u
in the form of higher
play a role in mediatin
change. We saw evide
with streams and sprin
scale with rivers, lakes
ter basins (Fig. 4ac).
Our finding that va
water bodies in the pr
scales, likely indicates
water availability duri
seedling-to sapling tr
2003; Tyler et al., 2006)
and lower precipitatio
seedlings there likely e
stress than seedlings
expanding areas. With
tracting areas may be
stress thresholds that
access to the water
areas where the water
We also found that
the local scale) saplin
water bodies than adu
areas. Reasons for this
may indicate that sa
restricted to higher gr
peratures are lower a
tively reducing the ov
and increasing the im
stressors such as herbiv
lin and Zavaleta, in pre
ing areas, saplings ar
adults and our data d
of the distribution.

stress vulnerability in this species. Higher projected temperatures and lower annual precipitation are
known to be key factors involved in potential future climate change.

An assessment by McIntyre et al. (2015) of historical and current-day alpine forest composition in
the Sierra Nevada range shows a trend toward denser forests with smaller trees. Their study found that
tree density has increased 30% while overall tree biomass has decreased 19%. Large tree declines were
found to be most dramatic in regions that experienced climatic water deficits. A study by Dolanc et al. in
2012 demonstrated a shift from pine-dominated forests to oak-dominated forests is linked to the increased
water stress experienced in this region. Dolanc found that the higher density of small trees is positively
correlated to greater turn-over rates in Sierra Nevada forest communities. He believes that drought stress
due to climate change is the mechanism behind this process. The authors also found that a shift in
community structure to include more shade-tolerant species filling the inter-forest spaces was linked to
denser tree stands. In 2013, Concilio et al. predicted that increasing temperatures coupled with
precipitation in the form of rainfall would allow invasive species to expand previous ranges to include
high-elevation ecosystems. The authors point out that most future climate models predict that
precipitation will occur more often as rain rather than snowfall, due to increasing temperatures at higher
elevations. Their study focused on cheatgrass (Bromus tectorum), an annual invasive species. Until
recently, studies have shown that invasion by B. tectorum into high-elevation ecosystems has been
reduced due to an inability to cope with annual snowpack. Concilio believes that reduction of annual
snowpack will open up previously unavailable resources, allowing B. tectorum to expand its current
range into high elevation forests.

In 2013, Devitt et al. used species distribution models (SDM) from the present and two late
Quaternary time periods to examine the effects of climate change habitat range shifts on populations of
Large-blotched Ensatina (Ensatina eschscholtzii klauberi). His models predicted a split of populations
into two isolated refugia, one in southern California and the other in northern Baja California at the
southern limit of the species current range. Devitts team used genetic testing to support the predicted
split, and found evidence of gene flow and connectivity between populations in northern refugia, but not

Downloaded from http://rspb.royalsocietypublishing.org/ on March 10, 2015

4284

T. L. Morelli et al.

in the south. Surveys of populations of Beldings

Climate refugia ameliorate decline

(a)

(b)

(c)

ground-squirrel (Urocitellus beldingi) by Morelli et

al. in 2012 were conducted to study the effects of
changing climate on the trailing edge (the region of
habitat range closest to an inhospitable changing

environment) of the species current range. Morelli

25 50

300 km

probability of presence

0 0.2
0.2 0.4
0.4 0.6
0.6 0.8
0.8 1.0

observed a range reduction of 255m upslope, which

Figure 4. Urocitellus beldingi species distribution model (GAM) results projecting (a) current distribution from historical presences and absences, (b) future distribution from 2080 Hadley, A2 scenario and (c) future distribution from 2080
CCCMA, A2 scenario. Probability of persistence is marked by quantile (0 0.2 not shown, 0.20.4 in beige, 0.40.6 in
light blue, 0.6 0.8 in blue and 0.81 in dark blue).

Figure 4. (A) Current range projections of U.

beldingi. (B&C) Estimated range contractions
of U. beldingi under two different climate
edge
of U.
activity instances,
(i.e. A2 scenario)
[60] might help to disentangle
the direct
effects of climateIn both
change
scenarios.
Ushowed greater than 50 per
cent range retraction of preferred habitats for U. beldingi
change from other causes.
in California
from reduction.
current habitat suitability, which
beldingi undergoes a drastic
range
already has been reduced by 42 per cent compared with
(b) Anthropogenic refugia
buffer climate
(Morelli
et al.,change
2012) a century ago. The wetter future of CCCMA A2 pro-

he used to project future range contractions of up to

72-99% of historic ranges along the southwest
beldingis current habitat by 2080.

Desert Systems

The absolute elevation limits of the U. beldingi California

jected the greatest decline, with up to 99 per cent loss
range have not changed over the past century, despite
of a suitable habitat. Although these projections only contheir disappearance from more than 40 per cent of surveyed
sider climate and are probably not reliable numeric
sites and the strong negative correlations between elevation
predictions
[61,62], they corJanuary 2011 and extirpation. Anthropogenic
PAST, PRESENT,
AND
FUTURE
OF JOSHUA
TREE of future U. beldingi habitat
145
refugia
appear
to explain
roborate our other results that indicate that the species is
this discrepancy. The elevational range of U. beldingi
being, and will continue to be, negatively impacted by cliwould
have60retracted
by at255
m if sites with
migration over
the next
to 90 yearsupslope
are shown
2-km
mate change along the warmer, and potentially also the
human-mediated
increases
in food and
water availability
expansion outward
from current
populations,
correwetter, trailing edge of its distribution.
Human
also had a positive
sponding to were
a rateexcluded.
of 22 to 33
m/year,modification
in order to make
effect on
Likewise,
was less affected
these areas visible
onabundance.
Fig. 5, although
our occupancy
observations
climate
change
in the Sierra
Nevada
for avian species
for past and by
current
rates
of migration
were only
;2 m/
5. CONCLUSIONS
that frequently
use areas
human-modified
year. The models
project large
of potentialhabitats
future [26].
Long-term occupancy data are needed to examine how
Our
study
demonstrates
a
potential
protective
effect of
habitat well outside of this range of natural migration,
species ranges have shifted as a result of recent changes
anthropogenic
for Base
montane
species. Food and/or
especially across
the Nellis refugia
Air Force
of southern
in climate and land-cover. This study provides a rare
water
or other
aspects of human presNevada (where
it issupplementation,
likely already present),
northwestern
glimpse of the effect of a century of climate and landence,
could
modify
the
effects
of
warming
and
shifting
Arizona, and southwestern Utah.
use change along the trailing edge of the range of a
precipitation regimes. Although we found that specific
montane mammal that was facilitated by the detailed hisDISCUSSION had a positive effect, activities that
human modification
torical field notes of Joseph Grinnell and colleagues. Our
convert
meadow
habitat
into
more
urban
or
suburban
Climatic tolerances for species in disequilibrium
study shows an extreme population reduction related to
uses would likely not support U. beldingi populations.
Inferring Identifying
species climate
limits
through
geographic
warm and wet climate, with a 255 m upslope retraction
both artificial and natural refugia that are bufcorrelations fered
between
a current
mediated only by anthropogenic refugia, and points to
from
climatespecies
changerange
may and
allowrecent
the persistence of
the need to understand the exceptions as well as the
climates relies
on the assumption
thatface
theofspecies
at
vulnerable
species in the
futureisclimate
change.
patterns of shifts in the trailing edge of species ranges.
least somewhat in geographic equilibrium with the
baseline climate period used. That is, at least a portion
We thank the MVZ faculty, staff and students who helped
(c) Future
beldingi
of the species
current projections
range must ofbeUrocitellus
close to the
with data collection and analysis, especially Jeni Chan,
distribution
geographic limit of each important variable comprising
Chris Conroy, Michele Hershey, Michele Koo, John
We projected that the range of U. beldingi along its trailing
Perrine, Bill Monahan, Karen Rowe, Kevin Rowe, Emily
its suitable climate model. The greater the proportion of
edge would decrease severely in a warmer future using
Rubidge and Jim Patton. We are grateful to David Wright
the species range analyzed, the more likely this
historically validated models. Two projections based on
for sharing data. Early drafts of this manuscript were
assumption is to be true because our observations of
greenhouse gas emissions within the scope of current
greatly improved by members of the Beissinger and Moritz
its realized niche will extend across the greatest portion
of its potential
niche
Proc. R.
Soc. Bspace
(2012)(Jackson and Overpeck
2000).
But the Joshua tree example used here does have an
added complication in that its migrational capacity to
respond to changing climates seems to be extremely
limited. There are no historical records of Joshua tree
invasions into new habitat and even few documented
instances of recent seedling establishment. Although the
rapidly warming climate of the early Holocene
(Steffensen et al. 2008, Cole 2010) would seem to have
opened up vast new areas of potential range to the
north, the fossil record does not record any significant
northward expansion over the last 11 700 years. These
facts coalesce with morphological observations of the
plants indehiscent fruits and the abundance of fruits
FIG. 4. (A) Suitable climate model for Joshua tree created
and seeds in fossil ground sloth dung to support the
with 20th-century ANUSPLIN mean precipitation variables
concept that the species current mobility is constrained and extreme mean monthly maximum and minimum temperby the earlier extinction of the Shasta ground sloth and ature events (Cole et al. 2008b). (B) Future suitable climate
other possible seed vector(s) (Janzen and Martin 1982, model run from high-resolution (;75 km) late-21st-century
projections downscaled to a ;1-km grid (Cole et al. 2008b).
Lenz 2001).
Because of the constrained migrational capacity of
Joshua tree, it is possible that our climate-window model predicted mortality of current southerly populations due
underestimates its potential tolerance to colder temper- to warmer temperatures.
atures. Populations could have prospered further north
The early Holocene retreat of Joshua tree to the
than its current range might suggest. This may be the
northern periphery of its extensive Pleistocene range
case for other western tree species as well such as singleleaves
little doubt that it is strongly influenced by rising
needle pinyon (Pinus monophylla; Cole et al. 2008a),
especially because some may have not completely temperatures. And its future movements, as modeled by
equilibrated to the warming over the last 150 years these results, suggest a repeat of the pattern seen at the
after the end of the little ice age. Because of this, it is close of the Pleistocene, except starting from a much
possible that our model has underestimated the extent of more restricted distribution. The results predict the
potential areas for relocation throughout the Great survival of some natural Joshua tree populations
Basin Desert regions of California, Nevada, and Utah. throughout the next century, but most will be greatly
But this factor would be unlikely to mitigate the future reduced in area.

According to Lehart, a study performed at the University of

Arizona showed that annual precipitation in the southwestern region of

the USA is expected to drop 5-10% by the end of the century. The study
indicates that this is due to a predicted increase in the size of the arid

region where Hadley Cell circulation touches down. It was found that

expansion of this desert zone and increasing downward hot-air masses

could potentially push the winter jet stream north, further reducing the
amount of winter precipitation this region receives. Lehart also stated

that precipitation events in the southwest will most likely become more

destructive when they do occur due to the large build up of water vapor
in the warmer air masses.

Joshua trees (Yucca brevifolia) are widely known to be an

endemic and keystone species to the Mojave Desert in central
and southern California. The iconic tree provides a much

Figure 5. (A) Current distribution of

suitable habitats for Y. brevifolia within
JTNP. (B) Predicted distribution of
suitable habitats within JTNP as global
temperature increases. (Barrows et al.,
2012)

needed vertical habitat in an otherwise flat landscape, allowing

for niche partitioning and specialization among species. In 2012, Barrows et al. used a fine-scale

assessment of sensitivity to climate change gradients to show a possible 90% reduction in current Y.
brevifolia distribution throughout Joshua Tree National Park (JTNP), the southern-most point of the
species range. The assessment projected only a few isolated pockets as potential refugia within JTNP,
however the possibility of future dispersal and range expansion from these pockets was estimated to be
unlikely. In 2012, Smith et al. compared range expansions and contractions of Y. brevifolia and four
specialist pollinator moths by examining phylogenetics and paleodistribution coupled with Pleistocene
packrat midden records and found no significant evidence of mutualistic contraction at the end of the last
glacial movement (LGM). A distribution model developed by Cole et al. in 2011 predicted total
elimination of Y. brevifolia throughout the southern portion of its current range. According to Cole, a lack
of potential dispersers and an affinity for a very specific climate will prevent Y. brevifolia from expanding
to a more favorable northern latitude during periods of increasing global temperatures.

In 2014, Lovich et al. showed that Agassizs desert tortoise (Gopherus agassizii) populations at the
Mojave-Sonoran interface in JTNP experienced a large decline from 1996-2012. Lovich suggests that the
declining population numbers correspond to intense periods of drought which were also recorded within
that time frame. Field data collection in the 2014 study reports that postures of dead specimens were
consistent with evidence of dehydration and/or starvation .

It is commonly known that desert tortoises

are long-lived species that are very resistant to changes in habitat and do not do well in attempts at
relocation. In conjunction with the evidence presented by Lovich, it seems reasonable to predict that
increases in atmospheric temperature and decreases in reliable sources of water may lead to a drastic
reduction in G. agassizii populations due to their inability to relocate to more favorable habitats. Further
desert studies by Barrows in 2011 looked at climate change sensitivity in both G. agassizii and
chuckwallas (Sauromalus ater) at the Mojave-Sonoran interface in JTNP. Barrows determined that
tortoise habitats were estimated to undergo a 88% reduction in size in the Sonoran portion and a 66%
reduction in size within the Mojave portion of the park. He further stated that chuckwallas are expected
to experience a 92% habitat reduction within the Sonoran portion of JTNP and 120% increase in habitat
size within the Mojave portion of the park. These results would indicate that both species are expected to
transition almost entirely out of the Sonoran portion of JTNP in the coming years.

The Great Basin Desert (GBD) is a North American desert that lies within the Basin and Range
Province, spreading from California to Utah. Its
location and the unique topography of the area make
the GBD the highest elevation cold desert in North
America. Within the GBD lies Death Valley, an
area that has been suggested by some to be one of
the hottest locations on earth. A 2014 study by
FIG. 3.Variation in adult woodrat body size over various timescales. a) Box plot of body mass of female woodrats (Neotoma lepida) from
Figure 6.dataThe
average
of N.lepida
populations
over over the period of record.
Furnace Creek (20032008);
generally
represent amass
4-year average.
Also plotted is
the mean monthly temperature

Smith et al. centered on mammalian adaptations

Error bars represent 95% confidence intervals. b) Median mass of the Furnace Creek population from 2003 to 2008 (gray histogram). Also plotted
time compared to elevational changes. Larger body sizes
in this region. Smiths research on adaptive

are mean (closed circles) and maximum (open circles) temperature for each month. Note that mean maximum temperature exceeds 418C for 34
months every year.
Mean body length with
of museum
specimens
of N. lepida collected
at various low-elevation
sites on the valley floor from 1891
arec) correlated
higher
elevation
populations
and colder
to 2008. Each datum represents the average of 9 to . 40 individuals; males (open squares) and females (open circles with dots) are plotted
temperatures,
sizes
are
linkeddifferences
to lower
separately. Woodrat
body mass scales aswhile
the cube smaller
of length, so body
fluctuations
represent
considerable
in mass. Values are plotted with

elevation warm temperature populations. (Smith et al., 2014)

responses to temperature change in woodrats

(Neotoma spp.) demonstrates the ability of each species to adjust to a potentially changing environment.
Results from the study showed evidence of a correlation between body size and the ability to handle
increasing temperatures. Smith determined that the larger N. cinerea is more adapted to montane slopes
and not well-suited to hot climates, while the smaller N. lepida lives on the valley floor and can withstand
increasing temperatures. Phenotypic changes in body mass and upslope range shifts were predicted for
both N. cinerea, and N. lepida as a way to cope with the changing climate.

Conclusions

Overall, the major predicted trend in range shifts is a move to higher, cooler climates as a way to
deal with global change. Multiple sources confirm that increases in atmospheric temperature will cause
species to disperse into new habitats (Loik et al., 2012, Dolanc et al., 2014, McLaughlin et al., 2012, Cole
et al., 2011, & Concilio et al., 2013). Populations that can disperse to higher elevations or higher latitudes
appear to be at an advantage when it comes to surviving in a warmer climate. Species that cant keep up
with their local velocity of change will most likely experience drastic habitat reduction and possible local
extinction (Loarie et al., 2009).

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