Impact of a Century of Climate Change on

Small-Mammal Communities in Yosemite National

Park, USA
Craig Moritz, et al.
Science 322, 261 (2008);
DOI: 10.1126/science.1163428
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REPORTS

References and Notes

1. C. Parmesan, Annu. Rev. Ecol. Evol. Syst. 37, 637
(2006).
2. R. Wilson, D. Gutierrez, J. Gutierrez, V. Monserrat, Glob.
Change Biol. 13, 1873 (2007).
3. J. Lenoir, J. Gegout, P. Marquet, P. de Ruffray, H. Brisse,
Science 320, 1768 (2008).
4. M. B. Bush, Glob. Ecol. Biogeogr. 11, 463 (2002).
5. Materials and methods are available as supporting
material on Science Online.
6. M. B. Bush, H. Hooghiemstra, in Climate Change and
Biodiversity, T. E. Lovejoy, L. Hannah, Eds. (Yale Univ.
Press, New Haven, CT, 2005), pp. 125137.
7. J. A. Pounds, M. P. L. Fogden, K. L. Masters, in Climate
Change and Biodiversity, T. E. Lovejoy, L. Hanna, Eds.
(Yale Univ. Press, New Haven, CT, 2005), pp. 7074.
8. C. L. Cardels, R. K. Colwell, J. E. Watkins, J. Ecol. 94,
144 (2006).
9. G. Brehm, R. K. Colwell, J. Kluge, Glob. Ecol. Biogeogr.
16, 205 (2007).
10. A. R. Wallace, Tropical Nature and Other Essays
(Macmillan, London, 1878).

11. T. Dobzhansky, Am. Sci. 38, 209 (1950).

12. L. Hannah, R. A. Betts, H. H. Shugart, in Tropical
Rainforest Responses to Climatic Change, M. B. Bush,
J. R. Flenley, Eds. (Springer Praxis, 2007), pp. 351366.
13. J. R. Flenley, Clim. Change 39, 177 (1998).
14. M. B. Bush, M. R. Silman, D. H. Urrego, Science 303, 827
(2004).
15. F. E. Mayle, D. J. Beerling, W. D. Gosling, M. B. Bush,
Philos. Trans. R. Soc. London B Biol. Sci. 359, 499
(2004).
16. M. Bush, M. Silman, J. Quat. Sci. 19, 677 (2004).
17. D. D. Ackerly et al., Bioscience 50, 979 (2000).
18. J. Hansen et al., Proc. Natl. Acad. Sci. U.S.A. 103, 14288
(2006).
19. C. A. Deutsch et al., Proc. Natl. Acad. Sci. U.S.A. 105,
6668 (2008).
20. D. A. Clark, S. C. Piper, C. D. Keeling, D. B. Clark, Proc.
Natl. Acad. Sci. U.S.A. 100, 5852 (2003).
21. K. J. Feeley, S. Joseph Wright, M. N. Nur Supardi,
A. R. Kassim, S. J. Davies, Ecol. Lett. 10, 461 (2007).
22. O. L. Phillips, S. L. Lewis, T. R. Baker, Y. Malhi, in Tropical
Rainforest Responses to Climatic Change, M. B. Bush,
J. R. Flenley, Eds. (Springer Praxis, 2007), pp. 317332.
23. L. Miles, A. Grainger, O. Phillips, Glob. Ecol. Biogeogr.
13, 553 (2004).
24. S. A. Cowling et al., Philos. Trans. R. Soc. London B Biol.
Sci. 359, 539 (2004).
25. J. W. Williams, S. T. Jackson, J. E. Kutzbach, Proc. Natl.
Acad. Sci. U.S.A. 104, 5738 (2007).
26. M. B. Bush, M. R. Silman, C. McMichael, S. Saatchi, Philos.
Trans. R. Soc. London B Biol. Sci. 363, 1795 (2008).
27. R. L. Peters, J. D. S. Darling, Bioscience 35, 707 (1985).
28. G.-R. Walther et al., Nature 416, 389 (2002).

Impact of a Century of Climate Change

on Small-Mammal Communities in
Yosemite National Park, USA
Craig Moritz,1,2* James L. Patton,1,2 Chris J. Conroy,1 Juan L. Parra,1,2
Gary C. White,3 Steven R. Beissinger1,4
We provide a century-scale view of small-mammal responses to global warming, without
confounding effects of land-use change, by repeating Grinnells early20th century survey across
a 3000-meter-elevation gradient that spans Yosemite National Park, California, USA. Using
occupancy modeling to control for variation in detectability, we show substantial (~500 meters on
average) upward changes in elevational limits for half of 28 species monitored, consistent with the
observed ~3C increase in minimum temperatures. Formerly low-elevation species expanded their
ranges and high-elevation species contracted theirs, leading to changed community composition at
mid- and high elevations. Elevational replacement among congeners changed because species
responses were idiosyncratic. Though some high-elevation species are threatened, protection
of elevation gradients allows other species to respond via migration.
lthough human-driven global warming
(1) has changed phenology of species
and contributed to range expansions
(26), contractions of species ranges are less well

A
1

Museum of Vertebrate Zoology, University of California,

Berkeley, CA 94720, USA. 2Department of Integrative
Biology, University of California, Berkeley, CA 94720, USA.
3
Department of Fish, Wildlife, and Conservation Biology,
Colorado State University, Fort Collins, CO 80523, USA.
4
Department of Environmental Science, Policy and Management, University of California, Berkeley, CA 94720,
USA.
*To whom correspondence should be addressed. E-mail:
craigm@berkeley.edu

documented (710). Models of future climatechange scenarios predict large range shifts, high
global extinction rates, and reorganized communities (11, 12), but model outcomes are also highly uncertain (13, 14). Most studies of species
responses span only a few decadestypically
from the 1960 or 1970s, which was a relatively
cool period, to the present. Such results can be
confounded by decadal-scale climate oscillations
(15) and landscape modification (8, 16). Furthermore, range shifts are uncertain when confounded
by false absences due to limited historic sampling
and inability to control for changes in detectability
between sampling periods (17, 18).

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VOL 322

29. C. Ghalambor, R. Huey, P. Martin, J. Tewksbury, G. Wang,

Integr. Comp. Biol. 46, 5 (2006).
30. R. G. Pearson, T. P. Dawson, Glob. Ecol. Biogeogr. 12,
361 (2003).
31. S. Williams, E. E. Bolitho, S. Fox, Proc. R. Soc. London B.
Biol. Sci. 270, 1887 (2003).
32. J. H. Christensen et al., in Climate Change 2007: Fourth
Assessment Report of the Intergovernmental Panel on
Climate Change, S. Solomon et al., Eds. (Cambridge Univ.
Press, Cambridge, 2007), pp. 847940.
33. J. Kluge, M. Kessler, R. R. Dunn, Glob. Ecol. Biogeogr.
15, 358 (2006).
34. Supported by the Organization for Tropical Studies; the
Tropical Ecology Assessment and Monitoring (TEAM) project
of Conservation International; University of Connecticut
(R.K.C. and C.L.C.); UCLA (A.C.G.); U.S. NSF (DEB-0072702:
R.K.C., G.B., and J.T.L.; DEB-0640015: J.T.L.;
DEB-0639979: R.K.C.; Research Fellowship and
Dissertation Improvement Grant: C.L.C.); Sigma Phi,
Explorers Club, and Steven Vavra Plant Systematics Fund
(A.C.G.); and the Deutsche Forschungsgemeinschaft
(BR 2280/1-1: G.B.). We thank M. B. Bush, R. L. Chazdon,
D. A. Clark, R. R. Dunn, K. M. Kuhn, C. Rahbek, T. F. L. V. B.
Rangel, M. R. Silman, and our peer reviewers for comments.

Downloaded from www.sciencemag.org on May 5, 2010

pear unlikely, putting the focus on elevational

gradients, where range-shift gaps will develop
early for the great numbers of narrow-ranged
species. The lowland tropics lack a source pool of
species adapted to higher temperatures to replace
those driven upslope by warming, raising the
possibility of substantial attrition in species richness in the tropical lowlands.

Supporting Online Material

www.sciencemag.org/cgi/content/full/322/5899/258/DC1
Materials and Methods
Figs. S1 and S2
References
30 June 2008; accepted 2 September 2008
10.1126/science.1162547

We quantified the impact of nearly a century

of climate change on the small-mammal community of Yosemite National Park (YNP) in California, USA, by resampling a broad elevational
transect (60 to 3300 m above sea level) that
Joseph Grinnell and colleagues surveyed from
1914 to 1920 (19) (Fig. 1). Their work documented the diversity and distribution of terrestrial
vertebrates in California to establish a benchmark
for future comparison (20), and led to the concept
of the ecological niche, the importance of temperature as determinant of range boundaries, and
the notion that species respond uniquely to environmental changes (21). In contrast to most early
20th century records, the Yosemite Transect
was densely sampled across elevations (Fig. 1)
and is amply documented by specimens (n =
4354), field notes (>3000 pages), and photographs
(~700) (22), enabling precise identification of both
species and sampling sites. From daily trapping
records, we estimated detectability of species in
historical as well as current surveys, permitting
the unbiased estimation of species absences
from elevational bands in both periods (23). The
transect spans YNP, a protected landscape since
1890, and allowed us to examine long-term responses to climate change without confounding
effects of land-use change, although at low to midelevations there has been localized vegetation
change relating to seral dynamics, climate change,
or both (24). Finally, analyses of regional weather
records pointed to substantial increase of the average minimum monthly temperature of 3.7C
over the past 100 years, with notable increases
from 1910 to 1945 and from 1970 to the present
(15, 22) (fig. S1).

10 OCTOBER 2008

261

Future warming is predicted to cause substantial turnover of species within North American
National Parks, including Yosemite (25). Given
marked regional warming over the past century,
we predicted that species ranges should have
shifted upward (5, 10). This should manifest as
upward contraction of the lower range limit for
mid- to high-elevation species, upward shift of
the entire range or expansion of the upper limit
for low- to mid-elevation species, and altered community composition within elevational bands (9).
Elevational ranges of species and their habitats differed markedly between the gradual western and steep eastern slopes of the transect (19)
(Fig. 1). On the west slope, we trapped small
mammals at 121 sites compared to 56 in Grinnells
time (table S1), but overall effort and elevational
range (~50 to 3300 m) were comparable (22).
There were fewer sites on the east side in both
time periods (9 for Grinnell, 12 for resurveys)
because of limited extent (Fig. 1). Our analyses
of richness and turnover focused on species detectable by standardized trapping (37 species) or
by observation (6 species; table S2). To test for
elevational shifts, we applied occupancy modeling (22, 23) to the 23 west slope taxa with sufficient trapping records to estimate detectability
in both periods (tables S1 and S2 and Fig. 2). The
best detection model in a set of 36 (table S3) was
used to calculate the probability of a false absence (Pfa) across trapping sites, where a species
was not observed in one sampling period but was
in the other (Table 1). Range shifts were significant if Pfa 0.05. For each species we evaluated
eight hypothesized relationships of occupancy,
era, and elevation (fig. S2) using the 14 best detection models (table S3) to create model-averaged
occupancy-elevation profiles (Fig. 2 and fig. S3).
Conservatively, we excluded shifts that were statistically significant but biologically trivial (Fig.
3). In most cases where the Pfa test indicated an
elevation shift, occupancy models agreed (Table
1 and fig. S3). Exceptions occurred when occupancy models were weak (i.e., insufficient data)
or detected changes in occupancy at elevations
other than range limits, or when nonstandard data
(i.e., records from ad hoc collecting) were included in Pfa tests but not in occupancy models.
Elevation limits shifted mostly upward (Table
1 and Fig. 3A), and this occurred more frequently
for lower than upper limits (c2 = 4.26, df = 1, P =
0.039). Twelve of 28 (43%) west slope species
showed significant shifts in lower limits, of which
10 increased (mean = +475 m) and two, both
shrews, decreased (mean = 744 m). In contrast,
upper limits changed significantly in only seven
instances, with similar numbers of upward (n = 4,
mean = +501 m) and downward shifts (n = 3,
mean = 309 m).
High-elevation species typically experienced
range contractions, whereas low-elevation species
expanded their ranges upward (c2 = 8.8, df = 2,
P = 0.012), a pattern expected with increased
temperature. Lower range limits contracted in
50% of the high-elevation species but in only

262

10% of low-elevation species, whereas 50% of

low-elevation species expanded their upper range
compared to none of the high-elevation species
(Fig. 3B). High-elevation species contracting
(Table 1 and Fig. 2A) included the alpine chipmunk (Tamais alpinus), Beldings ground squirrel (Spermophilus beldingi), water shrew (Sorex
palustris), and pika (Ochotona princeps). Range
collapseincreased lower limits and decreased
upper limitswas observed in two high-elevation
species: the bushy-tailed woodrat (Neotoma cinerea)
and the shadow chipmunk (T. senex) (Fig. 2B).
Parallel trends were observed on the east slope of
the Sierra for N. cinerea and S. beldingi (fig. S3).
Range contractions due to increases in lowerelevation limits were also observed for two
species formerly at mid- to high elevations [the
golden-mantled ground squirrel (Spermophilus
lateralis) and the long-tailed vole (Microtus
longicaudus)] (Table 1). Only one lowland
species contractedthe kangaroo rat (Dipodomys heermanni) showed a modest increase in
lower limit and a larger decrease in upper limit
since Grinnells time. Range expansions resulted
from either expanded upper limits [the pocket
mouse (Chaetodippus californicus), the California
vole (M. californicus), and the harvest mouse
(Reithrodontomys megalotis)] or expanded
lower limits (two shrews: Sorex monticola
and S. ornatus). Finally, the pinyon mouse
(Peromyscus truei) translocated upward (Fig.
2C); both upper and lower limits increased by

~500 m, but it now also occupies montane

conifer habitats on the west slope 800 to 1400
m higher after its east slope population expanded
upward by ~1000 m to cross the Sierra crest.
Elevational range shifts resulted in modest
changes in species richness and composition at
varying spatial scales. Species richness averaged
across five estimators (26) that account for nonobserved species (Fig. 3C, fig. S4, and table S4)
declined from the Grinnell era to the present
(repeated measures analysis of variance, F = 32.7
df = 1, P = 0.004). Richness estimators suggest a
slight decrease across the whole transect (currenthistoric mean estimates = 4.4 species, 9%), but
not within YNP (+1.3 species, 4%). Species richness was reduced within each life zone, with the
largest change in the Lower and Upper Sonoran
zones west of YNP. Community similarity between Grinnells period and the present was high
(mean similarity, S > 0.9) for the whole transect,
the park alone, and most life zones. Species composition was least similar for the Transition and
Hudsonian-Arctic zones, as expected given the
upward expansions of formerly Sonoran zone
taxa and the range shifts of high-elevation species
(Table 1).
Closely related species responded idiosyncratically to climate change (Table 1), but why species vary in response is not clear. For example,
some species of Peromyscus mice showed elevation range shifts (P. truei), whereas others
did not (P. boylii, P. maniculatus). The same is

Fig. 1. Map of surveyed sites in Grinnell (Historic) and Current surveys relative to the Yosemite
National Park boundary and life zones (upper panel), and to an averaged elevational profile (lower
panel).

10 OCTOBER 2008

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REPORTS

REPORTS
survey, we provide an unbiased comparison of
changes in species ranges at the centennial scale.
Because much of the transect spans a longprotected National Park, confounding effects of
land-use change are minimized. Even so, vegetation has changed within YNP over this period,
in part due to fire suppression (22). The park was
hardly pristine in the early 20th century, with
ranching of introduced herbivores in Yosemite
Valley and the high country recovering from historical overgrazing. As examples, expansion by
C. californicus and west slope P. truei are associated with fire-related conversion of conifer to
shrub habitats, whereas the downward shift of
S. monticola could reflect recovery of their
preferred wet meadow habitats. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an

Table 1. Analyses of elevation change for 28 west slope species. Given are
average detectability per site for Grinnell [P(G)] and current [P(C)] periods, original
elevation range, changes in upper (U) and lower (L) range limit that are significant
by the Pfa tests, the best supported form of the occupancy model (Elev, elevation;
NA, not analyzed), the cumulative Akaikes Information Criterion weight for all
No.

1
2

Species

P(G)

P(C)

Original
elevation
range (m)

0.81

0.58

571160

models with those terms (w), and original Lifezone classification (18), where L and H
refer, respectively, to species with mostly low- to mid-elevation ranges (<2000 m)
and mid- to high-elevation ranges (>2000 m) in Grinnells time; P. maniculatus
covered the entire transect. Values in bold are further supported by occupancy
models. See fig. S4 for elevation plots and models of individual species.
Best
occupancy
model

Range limit
change (m)

LowerUpper Sonoran (L)

+112 U
+589 U, +468 L

0.50
0.99

LowerUpper Sonoran (L)

Upper Sonoran (L)

0.32
0.74
0.37

Upper Sonoran (L)

Upper Sonoran (L)
CanadianHudsonian (H)

0.48
0.74
0.53
0.48
0.78
0.39
0.83
0.32
0.56
NA

LowerUpper Sonoran (L)

TransitionHudsonian (H)
TransitionHudsonian (H)
Canadian (H)
TransitionHudsonian (H)
CanadianHudsonian (H)
CanadianArctic-Alpine (H)
CanadianArctic-Alpine (H)
HudsonianArctic-Alpine (H)
CanadianArctic-Alpine (H)

0.72
NA
0.89
0.62
0.60
0.40
0.36

Lower SonoranArctic-Alpine (H)

Lower SonoranTransition (L)
Lower SonoranCanadian (L)
Lower SonoranTransition (L)
Upper SonoranTransition (L)
TransitionCanadian (H)
TransitionHudsonian (H)

0.87
0.93

571160
1831220

5
6

0.28
0.32
0.99

0.19
0.93
0.97

193914
549914
22123287

7
8
9
10
11
12
13
14
15
16

Dipodomys heermanni
Microtus longicaudus
Zapus princeps
Tamias senex
Spermophilus lateralis
Sorex palustris
Neotoma cinerea*
Spermophilus beldingi*
Tamias alpinus
Ochotona princeps

0.16
0.99
0.98
0.95
0.70
0.39
0.90
0.98
0.92
NA

0.98
0.98
0.90
0.71
0.89
0.23
0.71
0.98
0.95
NA

571025
6233287
12913185
14022743
16463200
16583155
17983287
22863287
23073353
23773871

17

Peromyscus
maniculatus*
Thomomys bottae
Spermophilus beecheyi
Neotoma macrotis
Peromyscus boylii
Sorex trowbridgii
Microtus montanus*
Tamiasciurus
douglasi*
Tamias
quadrimaculatus
Tamias speciosus*
Thomomys monticola
Marmota flaviventris

0.99
NA
0.50
0.90
0.98
0.71
0.81

0.99
NA
0.82
0.91
0.97
0.88
0.98

573287
571676
612734
1831646
1832469
11602286
12173155

No change
No change
250 U
+67 U
122 L
No change
No change

Era*(Elev + Elev2)
NA
Era*(Elev + Elev2)
Elev + Elev2
Elev + Elev2
Elev + Elev2
Elev + Elev2

NA

NA

12293185

No change

NA

0.95
1.00
NA
NA

0.85
1.00
NA
NA

14942210
17683155
19053155
24693353

+50 U
+128 L, +65 U
No change
No change

Era*(Elev + Elev2)
Era*(Elev + Elev2)
NA
NA

25
26
27
28

* Similar trends are observed for east-side populations (see fig. S4).

Original life zone (H, L)

0.36

0.99
0.99

18
19
20
21
22
23
24

Range expansions
+505 U
Elev

Microtus californicus
Reithrodontomys
megalotis
Peromyscus truei*
Chaetodippus
californicus
Sorex ornatus
Sorex monticolus

3
4

analogous community in the Rocky Mountains

(27).
The preponderance of upward range shifts,
leading to contraction of high-elevation species
and expansions of low-elevation taxa, accords with
the predicted impacts of climate warming (5, 8, 9).
Although vegetation dynamics have likely contributed to changes at low to mid-elevation, habitat change at higher elevations is limited (15)
(fig. S6). The ~500-m average increase in elevation for affected species is also consistent with
estimated warming of +3C, assuming a change
of temperature with elevation of ~6C per km.
Several small-mammal taxa that responded to
changing temperature also showed large range
fluctuations during late Quaternary climate fluctuations (28), and some have declined regionally (29).

Downloaded from www.sciencemag.org on May 5, 2010

true for chipmunks (Tamias), ground squirrels

(Spermophilus), voles (Microtus), and shrews
(Sorex). Beyond original elevation range (high
versus low), life history and ecological traits were
weak predictors of which species exhibited upward shifts of their range limits (tables S5 and
S6). This was especially true for high-elevation
species with upward contraction of their lower
range limit. However, lowland species that are
short-lived and lay more litters per year (so-called
fast life-style species) were more likely to expand
their range upward than were their long-lived,
less fecund counterparts (table S5 and fig. S5).
The elevational replacements among congeners,
documented so carefully in the early 20th century
(19), are now quite different.
By applying occupancy modeling to a thoroughly documented historical record and the re-

Elev
Era*(Elev + Elev2)

+800 U
Era*(Elev + Elev2)
485 L
Era *(Elev + Elev2)
1003 L
Era + Elev + Elev2
Range contractions
+63 L, 293 U
Era*Elev
+614 L
Era + Elev + Elev2
+159 L, 64 U
Era + Elev + Elev2
+1007 L, 334 U
Elev +Elev2
+244 L
Era*(Elev + Elev2)
+512 L
Era + Elev + Elev2
+609 L, 719 U
Era*(Elev + Elev2)
+355 L
Elev
+629 L
Era + Elev
+153 L
NA
No change

NA
0.78
1.00
NA
NA

TransitionHudsonian (H)
TransitionCanadian (H)
CanadianHudsonian (H)
CanadianHudsonian (H)
CanadianArctic-Alpine (H)

These species were encountered by observation and/or specialized trapping and were not subject to occupancy analyses.

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10 OCTOBER 2008

263

REPORTS
Fig. 2. Example elevation plots
from the west slope transect of upward range expansion (T. alpinus
and P. truei) (A and C), and
range collapse (N. cinerea) (B).
Shown are occupied (black) and
unoccupied (gray) sites, probability of false absence (Pfa),
and model-averaged occupancyelevation profiles (table S3 and
fig. S2). P. truei colonized high
elevations west of the Sierra crest
from the eastern slope. Red
marks for historical elevation
profile of T. alpinus refer to ad
hoc records.

Recent trends do not bode well for several

mid- to high-elevation species, including some endemic to the high Sierra (e.g., T. alpinus) (Fig. 3A).
Nevertheless, species diversity within Yosemite
has changed little, because range expansions compensated for retractions. Our results confirm that
protecting large-scale elevation gradients retains
diversity by allowing species to migrate in response to climate and vegetation change. The
long-recognized importance of protected landscapes has never been greater.

Fig. 3. (A) Summary of elevational

range changes across all species in
relation to life zones. Significant
(Pfa < 0.05) shifts are colored green
for range expansion and red for contraction (Table 1). Species were classified as No Change if range shifts
were biologically trivial (<10% of
previous elevation range) or of small
magnitude (<100 m). (B) Comparison of changes in elevation-range
limits for species that formerly had
low- to mid-elevation versus mid- to
high-elevation ranges (Table 1) across
the transect. (C) Mean (T SE) estimates of
species richness by era (bars: H, historic;
P, present; see also table S4 and fig. S4)
and community similarity (points) for individual life zones, Yosemite National Park,
and the entire transect.

Supporting Online Material

www.sciencemag.org/cgi/content/full/322/5899/261/DC1
Materials and Methods
Figs. S1 to S6
Tables S1 to S6
18 July 2008; accepted 9 September 2008
10.1126/science.1163428

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References and Notes

1. Intergovernmental Panel on Climate Change, Climate
Change 2007: Synthesis Report Contribution of Working
Groups I, II, and III to the Fourth Assessment Report of
the Intergovernmental Panel on Climate Change (IPCC,
Geneva, Switzerland, 2007).
2. C. Rosenzweig et al., Nature 453, 353 (2008).
3. C. Parmesan, G. Yohe, Nature 421, 37 (2003).
4. G. R. Walther et al., Nature 416, 389 (2002).
5. C. Parmesan, Annu. Rev. Ecol. Evol. Syst. 37, 637 (2006).
6. J. Lenoir, J. C. Ggout, P. A. Marquet, P. de Ruffray,
H. Brisse, Science 320, 1768 (2008).
7. R. J. Rowe, J. Biogeogr. 32, 1883 (2005).
8. R. Hickling, D. B. Roy, J. K. Hill, R. Fox, C. D. Thomas,
Glob. Change Biol. 12, 450 (2006).
9. R. J. Wilson, D. Gutierrez, J. Gutierrez, V. J. Monserrat,
Glob. Change Biol. 13, 1873 (2007).
10. C. D. Thomas, A. M. A. Franco, J. K. Hill, Evolution Int. J.
Org. Evolution 21, 415 (2006).
11. C. D. Thomas et al., Nature 427, 145 (2004).
12. J. W. Williams, S. T. Jackson, Front. Ecol. Environ 5, 475
(2007).
13. A. J. Davis, L. S. Jenkinson, J. H. Lawton, B. Shorrocks,
S. Wood, Nature 391, 783 (1998).
14. R. G. Pearson et al., J. Biogeogr. 33, 1704 (2006).
15. C. I. Millar, R. D. Westfall, D. L. Delany, J. C. King,
L. J. Graumlich, Arct. Antarct. Alp. Res. 36, 181 (2004).
16. W. Jetz, D. S. Wilcove, A. P. Dobson, PLoS Biol. 5, e157
(2007).
17. J. K. Hill et al., Proc. R. Soc. Lond. B. Biol. Sci. 269, 2163
(2002).
18. L. P. Shoo, S. E. Williams, J. M. Hero, Biol. Conserv. 125,
335 (2005).
19. J. Grinnell, T. Storer, Animal Life in the Yosemite (Univ. of
California Press, Berkeley, CA, 1924).
20. J. Grinnell, Pop. Sci. Mon. 77, 163 (1910).
21. J. Grinnell, Am. Nat. 51, 115 (1917).
22. See supporting material on Science Online.
23. D. I. MacKenzie et al., Inferring Patterns and Dynamics of
Species Occurrence (Elsevier, San Diego, 2006).
24. A. Guarin, A. H. Taylor, For. Ecol. Manage. 218, 229 (2005).
25. C. E. Burns, K. M. Johnston, O. J. Schmitz, Proc. Natl.
Acad. Sci. U.S.A. 100, 11474 (2003).
26. A. Chao, in Encyclopedia of Statistical Sciences,
N. Balakrishnan, C. B. Read, B. Vidakovic, Eds.
(Wiley, New York, 2005).
27. R. J. Rowe, Am. Nat. 170, 242 (2007).
28. D. K. Grayson, Quat. Sci. Rev. 25, 2964 (2006).
29. E. A. Beever, P. E. Brussard, J. Berger, J. Mammal. 84, 37
(2003).
30. Supported by funding from the Yosemite Foundation, the
National Parks Service, and NSF (DEB 0640859).
Assistance was received from A. Chang, H. Shofi, K. Tsao,
D. Yang, W. Monahan, R. Hijmans, and M. Koo. L. Chow
(U.S. Geological Survey Yosemite) and S. Thompson
(Yosemite National Park) stimulated and participated in
the project. Comments from R. Colwell, M. Power,
F. Hamer, and L. Shoo improved this manuscript.