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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Their work documented the diversity and distribution of terrestrial vertebrates in California to establish a benchmark for future comparison (20). 28. 19. and led to the concept of the ecological niche. S. J. Fox. 539 (2004). Piper. E-mail: craigm@berkeley. D. R.1. Brehm. C. Natl. 2007). 32. and our peer reviewers for comments. M. 461 (2007). T. permitting the unbiased estimation of species’ “absences” from elevational bands in both periods (23). J.S. pp. 38. Flenley. 13. and the notion that species respond uniquely to environmental changes (21). A. Sci. Acad. Glob. Philos.. DEB-0639979: R. M. and reorganized communities (11. R. 677 (2004). Sci. A. Ecol. H. the Tropical Ecology Assessment and Monitoring (TEAM) project of Conservation International. 13. Huey. R. and photographs (~700) (22). climate change. Beissinger1. S. J. field notes (>3000 pages). R. 8. although at low to midelevations there has been localized vegetation change relating to seral dynamics. 205 (2007). Hanna. (Cambridge Univ. Miles.C. R. G. Trans. Shugart. A. R. Acad. USA. Models of future climatechange scenarios predict large range shifts. 2007). J. B. P. D. D. M. E. L. Lenoir. University of Connecticut (R. 70–74. From daily trapping records. C. J. 27. J. 10. T. Policy and Management. Flenley. L. Colwell. 1795 (2008). CO 80523. Martin. UCLA (A.. (Springer Praxis. L. Hannah. protection of elevation gradients allows other species to respond via migration. Bush. 16. 499 (2004). Hansen et al. R. we show substantial (~500 meters on average) upward changes in elevational limits for half of 28 species monitored. Biogeogr. Bush. J. Kluge. K. enabling precise identification of both species and sampling sites.2 Gary C. Hannah. Bush. E. 103.C. V. S1 and S2 References 30 June 2008. Syst. CA 94720. and Steven Vavra Plant Systematics Fund (A. Annu. Ecol.1. F. 351–366. M. 18). R.1126/science. and allowed us to examine long-term responses to climate change without confounding effects of land-use change. Lovejoy. Sci. Ecol. Walther et al. 7. J. F.G. Williams. R.. B. Press. 31. Natl.B. by repeating Grinnell’s early–20th century survey across a 3000-meter-elevation gradient that spans Yosemite National Park. 463 (2002). P. L. and C.). S. Rahbek. the “Yosemite Transect” was densely sampled across elevations (Fig. Finally. J. where range-shift gaps will develop early for the great numbers of narrow-ranged species. 5852 (2003). USA. U. 12). K.C. E. P. without confounding effects of land-use change. Sci.1. 9. 2.org on May 5. Mayle. Parmesan. Using occupancy modeling to control for variation in detectability. Monserrat. 37. 20. Eds. Dobzhansky. Dunn. 2005). U. P. B. Fort Collins.and high elevations. B. University of California. www. Natl. Eds. B. Darling. 18. 637 (2006). A. T. 979 (2000). U. K. Gosling. Patton. H. B. a protected landscape since 1890. H. Evol. Lewis.. CA 94720. 19.REPORTS References and Notes 1. Trans. J. R. G. M. Ghalambor. R. raising the possibility of substantial attrition in species richness in the tropical lowlands. D. 1878). E. Dunn. Phillips.B. Science 303. A. 5738 (2007). Bush. Such results can be confounded by decadal-scale climate oscillations (15) and landscape modification (8. Silman. M. S1). Berkeley. Feeley. R. L.A. Wang. Parra. Change Biol.sciencemag. M. B. 15.1162547 We quantified the impact of nearly a century of climate change on the small-mammal community of Yosemite National Park (YNP) in California. Kuhn. T. R. R.. R. M. Cambridge. Joseph Wright. Research Fellowship and Dissertation Improvement Grant: C. W. Ecol. which was a relatively cool period. R. Bioscience 50.2 Chris J. Gegout. M.

004). maniculatus). but overall effort and elevational range (~50 to 3300 m) were comparable (22). S3).to high-elevation species. 4%). −9%). P = 0. S > 0. Species richness averaged across five estimators (26) that account for nonobserved species (Fig. 1). Exceptions occurred when occupancy models were weak (i. On the west slope. the California vole (M. There were fewer sites on the east side in both time periods (9 for Grinnell. For each species we evaluated eight hypothesized relationships of occupancy. we applied occupancy modeling (22. and altered community composition within elevational bands (9). Given marked regional warming over the past century. a pattern expected with increased temperature.org on May 5.Future warming is predicted to cause substantial turnover of species within North American National Parks. and this occurred more frequently for lower than upper limits (c2 = 4. and table S4) declined from the Grinnell era to the present (repeated measures analysis of variance. 3B). the pinyon mouse (Peromyscus truei) translocated upward (Fig. both upper and lower limits increased by ~500 m. 2). High-elevation species contracting (Table 1 and Fig.. P. occupancy models agreed (Table 1 and fig. 2B). 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. the park alone. Range contractions due to increases in lowerelevation limits were also observed for two species formerly at mid. but why species vary in response is not clear. S3). In contrast. beldingi (fig. This should manifest as upward contraction of the lower range limit for mid. or when nonstandard data (i. Our analyses of richness and turnover focused on species detectable by standardized trapping (37 species) or by observation (6 species. Community similarity between Grinnell’s period and the present was high (mean similarity. fig. and most life zones. P = 0. Closely related species responded idiosyncratically to climate change (Table 1). we excluded shifts that were statistically significant but biologically trivial (Fig.9) for the whole transect. Conservatively. upward shift of the entire range or expansion of the upper limit for low. Twelve of 28 (43%) west slope species showed significant shifts in lower limits. 10). we trapped small mammals at 121 sites compared to 56 in Grinnell’s time (table S1). 3C. S4. and elevation (fig. Richness estimators suggest a slight decrease across the whole transect (currenthistoric mean estimates = −4. 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. table S2). Species composition was least similar for the Transition and Hudsonian-Arctic zones. Range shifts were significant if Pfa ≤ 0. Finally. 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. S3).05. whereas 50% of low-elevation species expanded their upper range compared to none of the high-elevation species (Fig. To test for elevational shifts. Elevation limits shifted mostly upward (Table 1 and Fig. with similar numbers of upward (n = 4. 12 for resurveys) because of limited extent (Fig. Range expansions resulted from either expanded upper limits [the pocket mouse (Chaetodippus californicus). mean = +501 m) and downward shifts (n = 3. 3A). Parallel trends were observed on the east slope of the Sierra for N. Belding’s ground squirrel (Spermophilus beldingi). truei). 2 and fig. P = 0.to high elevations [the golden-mantled ground squirrel (Spermophilus lateralis) and the long-tailed vole (Microtus longicaudus)] (Table 1). as expected given the upward expansions of formerly Sonoran zone taxa and the range shifts of high-elevation species (Table 1). insufficient data) or detected changes in occupancy at elevations other than range limits. and pika (Ochotona princeps).sciencemag. The same is Fig. senex) (Fig. 2A) included the alpine chipmunk (Tamais alpinus).e.039). 2010 REPORTS .012). and to an averaged elevational profile (lower panel). 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. df = 1. 1).sciencemag. upper limits changed significantly in only seven instances.8. Elevational ranges of species and their habitats differed markedly between the gradual western and steep eastern slopes of the transect (19) (Fig. 3). water shrew (Sorex palustris). whereas others did not (P.4 species. 1.26. High-elevation species typically experienced range contractions. including Yosemite (25). In most cases where the Pfa test indicated an elevation shift.7 df = 1. df = 2.e. era. decreased (mean = −744 m). 23) to the 23 west slope taxa with sufficient trapping records to estimate detectability in both periods (tables S1 and S2 and Fig. californicus). some species of Peromyscus mice showed elevation range shifts (P. Species richness was reduced within each life zone. boylii. and the harvest mouse (Reithrodontomys megalotis)] or expanded lower limits (two shrews: Sorex monticola and S. but not within YNP (+1. S2) using the 14 best detection models (table S3) to create model-averaged occupancy-elevation profiles (Fig. of which 10 increased (mean = +475 m) and two.3 species. Map of surveyed sites in Grinnell (Historic) and Current surveys relative to the Yosemite National Park boundary and life zones (upper panel).to mid-elevation species. mean = −309 m). F = 32. records from ad hoc collecting) were included in Pfa tests but not in occupancy models.org Downloaded from www. 2C). ornatus). both shrews. cinerea and S. where a species was not observed in one sampling period but was in the other (Table 1).. whereas low-elevation species expanded their ranges upward (c2 = 8. with the largest change in the Lower and Upper Sonoran zones west of YNP. For example. 10 OCTOBER 2008 VOL 322 SCIENCE www. we predicted that species ranges should have shifted upward (5. Lower range limits contracted in 50% of the high-elevation species but in only 262 10% of low-elevation species.

S6). L) 0. 8.90 0.93 57–1160 183–1220 5 6 0.87 0.99 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. Analyses of elevation change for 28 west slope species.32 0. Even so.74 0. 1 2 Species P(G) P(C) Original elevation range (m) 0.00 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.19 0. habitat change at higher elevations is limited (15) (fig.98 0. −64 U Era + Elev + Elev2 +1007 L. 9).95 0.62 0. Beyond original elevation range (high versus low). voles (Microtus). 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. leading to contraction of high-elevation species and expansions of low-elevation taxa. +468 L 0. ground squirrels (Spermophilus). However.72 NA 0. Original life zone (H.99 Lower–Upper Sonoran (L) Upper Sonoran (L) 0. −719 U Era*(Elev + Elev2) +355 L Elev +629 L Era + Elev +153 L NA No change NA 0. we provide an unbiased comparison of changes in species’ ranges at the centennial scale. with ranching of introduced herbivores in Yosemite Valley and the high country recovering from historical overgrazing. monticola could reflect recovery of their preferred wet meadow habitats.org SCIENCE VOL 322 10 OCTOBER 2008 263 .99 0. and shrews (Sorex).39 0. −334 U Elev +Elev2 +244 L Era*(Elev + Elev2) +512 L Era + Elev + Elev2 +609 L. in part due to fire suppression (22). This was especially true for high-elevation species with upward contraction of their lower range limit.89 0.58 57–1160 models with those terms (w). elevation.98 0.78 1.60 0.to mid-elevation ranges (<2000 m) and mid. Values in bold are further supported by occupancy models.88 0. Best occupancy model Range limit change (m) Lower–Upper Sonoran (L) +112 U +589 U. –293 U Era*Elev +614 L Era + Elev + Elev2 +159 L. 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. life history and ecological traits were weak predictors of which species exhibited upward shifts of their range limits (tables S5 and S6).90 0. vegetation has changed within YNP over this period.71 0.99 NA 0.95 1.53 0.50 0. where L and H refer.48 0.98 0.83 0. not analyzed).00 NA NA 1494–2210 1768–3155 1905–3155 2469–3353 +50 U +128 L. Several small-mammal taxa that responded to changing temperature also showed large range fluctuations during late Quaternary climate fluctuations (28).org on May 5.99 0.97 0.00 NA NA 0. Although vegetation dynamics have likely contributed to changes at low to mid-elevation. Downloaded from www.81 0.93 0.sciencemag. See fig.37 Upper Sonoran (L) Upper Sonoran (L) Canadian–Hudsonian (H) 0.90 0. truei are associated with fire-related conversion of conifer to shrub habitats.28 0.36 0. The park was hardly pristine in the early 20th century. The elevational replacements among congeners.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. changes in upper (U) and lower (L) range limit that are significant by the Pfa tests.48 0.56 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) 0.95 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† 0.16 0. S5).98 0.to high-elevation ranges (>2000 m) in Grinnell’s time. respectively.91 0.78 0. and original Lifezone classification (18).sciencemag.82 0.99 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). The preponderance of upward range shifts. S4).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. less fecund counterparts (table S5 and fig.98 0.32 0. californicus and west slope P. The ~500-m average increase in elevation for affected species is also consistent with estimated warming of +3°C. accords with the predicted impacts of climate warming (5.89 0.92 NA 0. original elevation range.40 0.85 1.39 0.23 0. the cumulative Akaike’s Information Criterion weight for all No. Given are average detectability per site for Grinnell [P(G)] and current [P(C)] periods. documented so carefully in the early 20th century (19). www.32 0. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an Table 1. the best supported form of the occupancy model (Elev. are now quite different.98 0.71 0. maniculatus covered the entire transect.71 0. 2010 true for chipmunks (Tamias). Because much of the transect spans a longprotected National Park. +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.50 0. and some have declined regionally (29).REPORTS survey. P. S4 for elevation plots and models of individual species. confounding effects of land-use change are minimized.70 0. As examples. whereas the downward shift of S.81 0.74 0. expansion by C. assuming a change of temperature with elevation of ~6°C per km.99 NA 0. to species with mostly low. NA.

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