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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London B. S. putting the focus on elevational gradients. 2005). E. R.B. Hannah. R. C. 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. 1878). Baker. 16). University of California. T. Glob. Miles. A. A. J. V. Press. 2. Colwell. Biogeogr. University of California. A. Using occupancy modeling to control for variation in detectability. (Yale Univ. 389 (2002). in Climate Change 2007: Fourth Assessment Report of the Intergovernmental Panel on Climate Change. Am. Rahbek.C. C. Hanna.2 Chris J. T. J. Christensen et al. high global extinction rates. 11. Kutzbach. enabling precise identification of both species and sampling sites. 46. 1795 (2008). Silman. L. B. S. Bush. Lenoir. D. Acad. Proc. M.A. Berkeley.1. M. P.K. L. B. 539 (2004). CA 94720. Acad. 34. Fox. Brehm.7°C over the past 100 years. D. UCLA (A. Hansen et al. L. Marquet.C. K. Tewksbury. K. 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CA 94720. permitting the unbiased estimation of species’ “absences” from elevational bands in both periods (23). P.C. D. 21. J. 5852 (2003). Clark. R. 32. 9. 5738 (2007). B. G. M. L.. we show substantial (~500 meters on average) upward changes in elevational limits for half of 28 species monitored. P. *To whom correspondence should be addressed. Acad. Bush. R. USA.A. contractions of species’ ranges are less well A 1 Museum of Vertebrate Zoology. Kassim. Press. J. 14. Materials and methods are available as supporting material on Science Online. W. A. Kuhn. Philos. 18. Sci. Though some high-elevation species are threatened. and allowed us to examine long-term responses to climate change without confounding effects of land-use change.sciencemag. 358 (2006). In contrast to most early– 20th century records. S. A. Colwell.org on May 5.1.

S3). of which 10 increased (mean = +475 m) and two. Range shifts were significant if Pfa ≤ 0. 2B).sciencemag. including Yosemite (25).Future warming is predicted to cause substantial turnover of species within North American National Parks. and to an averaged elevational profile (lower panel). For example. and this occurred more frequently for lower than upper limits (c2 = 4. Range collapse—increased lower limits and decreased upper limits—was observed in two high-elevation species: the bushy-tailed woodrat (Neotoma cinerea) and the shadow chipmunk (T. and elevation (fig. P. ornatus). Map of surveyed sites in Grinnell (Historic) and Current surveys relative to the Yosemite National Park boundary and life zones (upper panel). 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. and altered community composition within elevational bands (9). and the harvest mouse (Reithrodontomys megalotis)] or expanded lower limits (two shrews: Sorex monticola and S. Finally. the park alone. S3). boylii. High-elevation species contracting (Table 1 and Fig. Range contractions due to increases in lowerelevation limits were also observed for two species formerly at mid.. S3). both shrews. we applied occupancy modeling (22. with similar numbers of upward (n = 4.sciencemag. maniculatus). 4%). records from ad hoc collecting) were included in Pfa tests but not in occupancy models.to high elevations [the golden-mantled ground squirrel (Spermophilus lateralis) and the long-tailed vole (Microtus longicaudus)] (Table 1). table S2).8. and table S4) declined from the Grinnell era to the present (repeated measures analysis of variance. we excluded shifts that were statistically significant but biologically trivial (Fig.3 species. as expected given the upward expansions of formerly Sonoran zone taxa and the range shifts of high-elevation species (Table 1). californicus). To test for elevational shifts.7 df = 1. In contrast. Closely related species responded idiosyncratically to climate change (Table 1). Parallel trends were observed on the east slope of the Sierra for N.org on May 5. fig. upward shift of the entire range or expansion of the upper limit for low. 1). and most life zones. 3C. senex) (Fig. Exceptions occurred when occupancy models were weak (i. Species richness averaged across five estimators (26) that account for nonobserved species (Fig.to mid-elevation species. era.05.012).004). some species of Peromyscus mice showed elevation range shifts (P. Given marked regional warming over the past century. we trapped small mammals at 121 sites compared to 56 in Grinnell’s time (table S1). cinerea and S. P = 0. where a species was not observed in one sampling period but was in the other (Table 1). Community similarity between Grinnell’s period and the present was high (mean similarity. truei). 10 OCTOBER 2008 VOL 322 SCIENCE www. High-elevation species typically experienced range contractions. both upper and lower limits increased by ~500 m. P = 0. Species composition was least similar for the Transition and Hudsonian-Arctic zones. df = 2. Elevational range shifts resulted in modest changes in species richness and composition at varying spatial scales. On the west slope.to high-elevation species. For each species we evaluated eight hypothesized relationships of occupancy. Range expansions resulted from either expanded upper limits [the pocket mouse (Chaetodippus californicus).e. 2C). 10). There were fewer sites on the east side in both time periods (9 for Grinnell.4 species. 1). decreased (mean = −744 m). Elevational ranges of species and their habitats differed markedly between the gradual western and steep eastern slopes of the transect (19) (Fig. 2010 REPORTS . This should manifest as upward contraction of the lower range limit for mid. Elevation limits shifted mostly upward (Table 1 and Fig. Belding’s ground squirrel (Spermophilus beldingi). df = 1.039). 12 for resurveys) because of limited extent (Fig. insufficient data) or detected changes in occupancy at elevations other than range limits. 3). occupancy models agreed (Table 1 and fig. whereas low-elevation species expanded their ranges upward (c2 = 8. mean = −309 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. but not within YNP (+1. water shrew (Sorex palustris). we predicted that species ranges should have shifted upward (5. 1. F = 32. the California vole (M. but overall effort and elevational range (~50 to 3300 m) were comparable (22). 2 and fig. The same is Fig. Twelve of 28 (43%) west slope species showed significant shifts in lower limits.. and pika (Ochotona princeps). 2A) included the alpine chipmunk (Tamais alpinus). the pinyon mouse (Peromyscus truei) translocated upward (Fig. beldingi (fig. Species richness was reduced within each life zone.e. mean = +501 m) and downward shifts (n = 3. whereas others did not (P.26. P = 0.9) for the whole transect. Richness estimators suggest a slight decrease across the whole transect (currenthistoric mean estimates = −4. In most cases where the Pfa test indicated an elevation shift. 3A). a pattern expected with increased temperature. upper limits changed significantly in only seven instances. Only one lowland species contracted—the kangaroo rat (Dipodomys heermanni) showed a modest increase in lower limit and a larger decrease in upper limit since Grinnell’s time. Lower range limits contracted in 50% of the high-elevation species but in only 262 10% of low-elevation species. but why species vary in response is not clear. S > 0. Conservatively.org Downloaded from www. Our analyses of richness and turnover focused on species detectable by standardized trapping (37 species) or by observation (6 species. or when nonstandard data (i. S4. 3B). 23) to the 23 west slope taxa with sufficient trapping records to estimate detectability in both periods (tables S1 and S2 and Fig. whereas 50% of low-elevation species expanded their upper range compared to none of the high-elevation species (Fig. with the largest change in the Lower and Upper Sonoran zones west of YNP. S2) using the 14 best detection models (table S3) to create model-averaged occupancy-elevation profiles (Fig. −9%).

9). Values in bold are further supported by occupancy models. and shrews (Sorex).00 NA NA 0.74 0.32 0.97 0. to species with mostly low.32 0.40 0. 8.85 1. accords with the predicted impacts of climate warming (5. 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. Although vegetation dynamics have likely contributed to changes at low to mid-elevation. confounding effects of land-use change are minimized.19 0.91 0. S4 for elevation plots and models of individual species.93 0. The elevational replacements among congeners. original elevation range. changes in upper (U) and lower (L) range limit that are significant by the Pfa tests. where L and H refer. Analyses of elevation change for 28 west slope species. S6).81 0.58 57–1160 models with those terms (w). The ~500-m average increase in elevation for affected species is also consistent with estimated warming of +3°C.74 0.62 0. www.97 193–914 549–914 2212–3287 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. life history and ecological traits were weak predictors of which species exhibited upward shifts of their range limits (tables S5 and S6). –293 U Era*Elev +614 L Era + Elev + Elev2 +159 L.sciencemag. Best occupancy model Range limit change (m) Lower–Upper Sonoran (L) +112 U +589 U. Even so. However.00 NA NA Transition–Hudsonian (H) Transition–Canadian (H) Canadian–Hudsonian (H) Canadian–Hudsonian (H) Canadian–Arctic-Alpine (H) †These species were encountered by observation and/or specialized trapping and were not subject to occupancy analyses. and original Lifezone classification (18).71 0.71 0. less fecund counterparts (table S5 and fig.90 0.99 Lower–Upper Sonoran (L) Upper Sonoran (L) 0.32 0.36 Lower Sonoran–Arctic-Alpine (H) Lower Sonoran–Transition (L) Lower Sonoran–Canadian (L) Lower Sonoran–Transition (L) Upper Sonoran–Transition (L) Transition–Canadian (H) Transition–Hudsonian (H) 0.98 0.org SCIENCE VOL 322 10 OCTOBER 2008 263 . californicus and west slope P. −64 U Era + Elev + Elev2 +1007 L. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an Table 1. are now quite different. +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.98 0.83 0. Several small-mammal taxa that responded to changing temperature also showed large range fluctuations during late Quaternary climate fluctuations (28). ground squirrels (Spermophilus).99 18 19 20 21 22 23 24 w 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).23 0. maniculatus covered the entire transect.48 0. The preponderance of upward range shifts. habitat change at higher elevations is limited (15) (fig. the best supported form of the occupancy model (Elev. 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.70 0. documented so carefully in the early 20th century (19). vegetation has changed within YNP over this period.78 0. we provide an unbiased comparison of changes in species’ ranges at the centennial scale.90 0.88 0. the cumulative Akaike’s Information Criterion weight for all No.99 NA 0. 1 2 Species P(G) P(C) Original elevation range (m) 0.87 0.39 0.81 0. NA.48 0. 2010 true for chipmunks (Tamias). respectively.78 1.99 0. leading to contraction of high-elevation species and expansions of low-elevation taxa.56 NA Lower–Upper Sonoran (L) Transition–Hudsonian (H) Transition–Hudsonian (H) Canadian (H) Transition–Hudsonian (H) Canadian–Hudsonian (H) Canadian–Arctic-Alpine (H) Canadian–Arctic-Alpine (H) Hudsonian–Arctic-Alpine (H) Canadian–Arctic-Alpine (H) 0. As examples.98 0. This was especially true for high-elevation species with upward contraction of their lower range limit. +468 L 0.92 NA 0. elevation.16 0.99 0. Beyond original elevation range (high versus low).98 57–3287 57–1676 61–2734 183–1646 183–2469 1160–2286 1217–3155 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 1229–3185 No change NA 0. whereas the downward shift of S.99 NA 0.82 0.99 0.72 NA 0.REPORTS survey. S5).28 0.90 0. in part due to fire suppression (22). See fig.60 0.98 0. L) 0.to mid-elevation ranges (<2000 m) and mid.sciencemag. truei are associated with fire-related conversion of conifer to shrub habitats.00 NA NA 1494–2210 1768–3155 1905–3155 2469–3353 +50 U +128 L. The park was hardly pristine in the early 20th century. S4).50 0.37 Upper Sonoran (L) Upper Sonoran (L) Canadian–Hudsonian (H) 0.95 0.93 57–1160 183–1220 5 6 0. with ranching of introduced herbivores in Yosemite Valley and the high country recovering from historical overgrazing.36 0.39 0.71 0.98 0. Original life zone (H. assuming a change of temperature with elevation of ~6°C per km.50 0. monticola could reflect recovery of their preferred wet meadow habitats. Given are average detectability per site for Grinnell [P(G)] and current [P(C)] periods.89 0. and some have declined regionally (29). voles (Microtus).95 1. expansion by C. Downloaded from www. P. −334 U Elev +Elev2 +244 L Era*(Elev + Elev2) +512 L Era + Elev + Elev2 +609 L.95 NA 57–1025 623–3287 1291–3185 1402–2743 1646–3200 1658–3155 1798–3287 2286–3287 2307–3353 2377–3871 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† 0.89 0. not analyzed). −719 U Era*(Elev + Elev2) +355 L Elev +629 L Era + Elev +153 L NA No change NA 0.98 0. Because much of the transect spans a longprotected National Park.53 0.org on May 5.to high-elevation ranges (>2000 m) in Grinnell’s time.

Berger.. C. Burns. R. Wood. 6. and range collapse (N. 28. See supporting material on Science Online. 19. Syst. 242 (2007). 21. including some endemic to the high Sierra (e. 37 (2003). Rowe.S. 415 (2006). D. PLoS Biol. Chang. D. T. alpinus refer to ad hoc records. 22. C. Nature 391. 32. Hijmans. Mammal. 181 (2004). C. 13. Westfall. Nature 421. and L. Rowe. Monahan. A. 16. J. Inferring Patterns and Dynamics of Species Occurrence (Elsevier. 637 (2006).1126/science. 12. S. B. 100. S. N. L. J. Grayson. Manage. 125. Graumlich. J. Sci. Vidakovic. 23. L. 269. A. 3A). Shoo. in Encyclopedia of Statistical Sciences. Roy. D. historic. Grinnell. alpinus) (Fig. Quat. For. Annu.g. Nature 427. Species were classified as “No Change” if range shifts were biologically trivial (<10% of previous elevation range) or of small magnitude (<100 m). the National Parks Service. C. Alp. A. and NSF (DEB 0640859). Read. Colwell. R. J..to high-elevation ranges (Table 1) across the transect. CA. Storer. S1 to S6 Tables S1 to S6 18 July 2008. W. R. 9. (B) Comparison of changes in elevation-range limits for species that formerly had low. 389 (2002). Front. 3. J. Conserv. C. J. B. Lawton. A. F. Pop. Franco. Example elevation plots from the west slope transect of upward range expansion (T. 2006). 4. Switzerland. A. D. J. Thompson (Yosemite National Park) stimulated and participated in the project. E. Hill. Nevertheless. Thomas et al. D. W. 1924).. San Diego. Change Biol. and the entire transect. 2010 References and Notes 1. 10. 145 (2004). 450 (2006). M. J. Am. 1704 (2006). Climate Change 2007: Synthesis Report Contribution of Working Groups I. 37. C. H. E. 13.sciencemag. Pearson et al. 30. K.to mid-elevation versus mid. 475 (2007). D. truei colonized high elevations west of the Sierra crest from the eastern slope. 353 (2008). de Ruffray. L. Mon. Hamer. Yohe. Supporting Online Material www. 1873 (2007). 3. 26. (Wiley. 20. Hickling. B. . Soc. 2964 (2006). Grinnell. Delany. Biol. D. Biogeogr. 51. I. Chow (U. R. Nature 416.. C. D. Res. D. 1883 (2005). 115 (1917). Walther et al. 8. King. 24. Org. J. Shorrocks. Gégout. R. Nat. 33. J. 11474 (2003). and M. J. 84. Schmitz. S. J. and III to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change (IPCC. Chao. Hill et al. Ecol. alpinus and P. 12. Johnston. Assistance was received from A. Am. J. J. Science 320. Marquet.org/cgi/content/full/322/5899/261/DC1 Materials and Methods Figs. Berkeley. J. R. L. S4) and community similarity (points) for individual life zones. 15. Shofi. Lond. H. B. Sci. Evolution 21. D. Our results confirm that protecting large-scale elevation gradients retains diversity by allowing species to migrate in response to climate and vegetation change. D. 25. Gutierrez. C. Beever.05) shifts are colored green for range expansion and red for contraction (Table 1). Jenkinson. Evol. because range expansions compensated for retractions.. S2). C. 2005). K. 18.. 11.sciencemag. 17. MacKenzie et al. Supported by funding from the Yosemite Foundation. Glob. of California Press. 2007). J. C. Balakrishnan. R. species diversity within Yosemite has changed little. Davis. Williams. Jetz.to high-elevation species. (A) Summary of elevational range changes across all species in relation to life zones. J. Tsao. Recent trends do not bode well for several mid.org on May 5. Sci. Red marks for historical elevation profile of T. Biol. Fig. Change Biol. Sci. P. see also table S4 and fig.A. Comments from R. M. R. Natl. Evolution Int. 1768 (2008). Taylor. K.REPORTS Fig. Yosemite National Park. Shoo improved this manuscript.1163428 264 10 OCTOBER 2008 VOL 322 SCIENCE www. E. Parmesan. Rev. Fox. V. Thomas. Nature 453. 2.org Downloaded from www. Millar. 2163 (2002). Proc. Biogeogr. Hill. 36. Wilson. M. 27. Jackson. Environ 5. Guarin. P. Williams. probability of false absence (Pfa). 14. 29. G. H. 5. Parmesan. U. 77. 170. Brisse. 5. R. P. Yang. truei) (A and C). Shown are occupied (black) and unoccupied (gray) sites. G. accepted 9 September 2008 10. 229 (2005). Hero. Geneva. B.S. J. K. 335 (2005). Gutierrez. Antarct. cinerea) (B). S.sciencemag. J. Geological Survey Yosemite) and S. E. P. P. 783 (1998). W. e157 (2007). Monserrat. II. Intergovernmental Panel on Climate Change. T. T. Animal Life in the Yosemite (Univ. Dobson. Glob. 2. The long-recognized importance of protected landscapes has never been greater. G. 25. Arct.. Significant (Pfa < 0. 163 (1910). J. R. Lenoir. H. Ecol. Wilcove. A. Rosenzweig et al. and model-averaged occupancyelevation profiles (table S3 and fig. J. Nat. K. J. 37 (2003). A. New York. J. A. Ecol. A. S. Acad. J. Koo. O. present. P. P. (C) Mean (T SE) estimates of species richness by era (bars: H. Eds. K. C. Brussard. J. Thomas. Proc. I. Power. Grinnell. 218. L. 7. M. Rev.

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