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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Impact of a Century of Climate Change on Small-Mammal Communities in Yosemite National Park. A. enabling precise identification of both species and sampling sites. *To whom correspondence should be addressed. 209 (1950). Trans.K. Philos. E. consistent with the observed ~3°C increase in minimum temperatures. Proc. B.L. 553 (2004). Eds. London. London B Biol. A. analyses of regional weather records pointed to substantial increase of the average minimum monthly temperature of 3. P.C. 10. Change Biol.S. 46. pp. The lowland tropics lack a source pool of species adapted to higher temperatures to replace those driven upslope by warming. Soc. R. Sci. Brisse.. Davies. Kuhn. J.2 Gary C. (Springer Praxis. 2007).1. M. J. Grainger. USA Craig Moritz. J. S. Eds. 3.. and photographs (~700) (22). (Cambridge Univ. 14. Kessler. B. DEB-0639979: R. 677 (2004). Chazdon. Science 320. G. B. 12). Lovejoy. Clark. S. but model outcomes are also highly uncertain (13. 144 (2006). T. Hansen et al. New Haven. 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boylii. Lower range limits contracted in 50% of the high-elevation species but in only 262 10% of low-elevation species. −9%). Species richness was reduced within each life zone. 2C). There were fewer sites on the east side in both time periods (9 for Grinnell. both shrews. In contrast. with similar numbers of upward (n = 4.9) for the whole transect.org on May 5. a pattern expected with increased temperature. 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.. Species richness averaged across five estimators (26) that account for nonobserved species (Fig. ornatus).4 species.8. 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. Given marked regional warming over the past century. S3). and this occurred more frequently for lower than upper limits (c2 = 4. and altered community composition within elevational bands (9).to mid-elevation species. upward shift of the entire range or expansion of the upper limit for low. 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. High-elevation species contracting (Table 1 and Fig. 2B). table S2). upper limits changed significantly in only seven instances. both upper and lower limits increased by ~500 m. beldingi (fig. 10 OCTOBER 2008 VOL 322 SCIENCE www. but overall effort and elevational range (~50 to 3300 m) were comparable (22). 10). S3). mean = −309 m). and pika (Ochotona princeps). df = 1.to high-elevation species. 4%). Elevation limits shifted mostly upward (Table 1 and Fig. and table S4) declined from the Grinnell era to the present (repeated measures analysis of variance.sciencemag. P = 0. For example. In most cases where the Pfa test indicated an elevation shift. Closely related species responded idiosyncratically to climate change (Table 1).004). To test for elevational shifts. truei). 2010 REPORTS . df = 2. where a species was not observed in one sampling period but was in the other (Table 1). mean = +501 m) and downward shifts (n = 3. californicus). 1). we trapped small mammals at 121 sites compared to 56 in Grinnell’s time (table S1). senex) (Fig. Conservatively. whereas 50% of low-elevation species expanded their upper range compared to none of the high-elevation species (Fig. Range shifts were significant if Pfa ≤ 0. This should manifest as upward contraction of the lower range limit for mid. 3). Belding’s ground squirrel (Spermophilus beldingi). S3). For each species we evaluated eight hypothesized relationships of occupancy. the pinyon mouse (Peromyscus truei) translocated upward (Fig.sciencemag. some species of Peromyscus mice showed elevation range shifts (P. F = 32.7 df = 1. 2A) included the alpine chipmunk (Tamais alpinus). Richness estimators suggest a slight decrease across the whole transect (currenthistoric mean estimates = −4. Elevational range shifts resulted in modest changes in species richness and composition at varying spatial scales. Community similarity between Grinnell’s period and the present was high (mean similarity. fig. Elevational ranges of species and their habitats differed markedly between the gradual western and steep eastern slopes of the transect (19) (Fig. Map of surveyed sites in Grinnell (Historic) and Current surveys relative to the Yosemite National Park boundary and life zones (upper panel).3 species. of which 10 increased (mean = +475 m) and two. and elevation (fig.039). 3A). whereas low-elevation species expanded their ranges upward (c2 = 8. 3C. and to an averaged elevational profile (lower panel). and most life zones. High-elevation species typically experienced range contractions.26. 1. S2) using the 14 best detection models (table S3) to create model-averaged occupancy-elevation profiles (Fig. P = 0. Our analyses of richness and turnover focused on species detectable by standardized trapping (37 species) or by observation (6 species. we applied occupancy modeling (22. The same is Fig.Future warming is predicted to cause substantial turnover of species within North American National Parks. water shrew (Sorex palustris). records from ad hoc collecting) were included in Pfa tests but not in occupancy models. but why species vary in response is not clear. S4. 3B). maniculatus). we predicted that species ranges should have shifted upward (5. P = 0. and the harvest mouse (Reithrodontomys megalotis)] or expanded lower limits (two shrews: Sorex monticola and S. 12 for resurveys) because of limited extent (Fig. cinerea and S. 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.. era. insufficient data) or detected changes in occupancy at elevations other than range limits.org Downloaded from www. P. decreased (mean = −744 m). 2 and fig. including Yosemite (25). occupancy models agreed (Table 1 and fig. Finally. the park alone. whereas others did not (P. we excluded shifts that were statistically significant but biologically trivial (Fig. S > 0.e. with the largest change in the Lower and Upper Sonoran zones west of YNP.e. or when nonstandard data (i. Exceptions occurred when occupancy models were weak (i. 1). On the west slope. Range contractions due to increases in lowerelevation limits were also observed for two species formerly at mid. Species composition was least similar for the Transition and Hudsonian-Arctic zones.012). the California vole (M. but not within YNP (+1. as expected given the upward expansions of formerly Sonoran zone taxa and the range shifts of high-elevation species (Table 1). 2). Twelve of 28 (43%) west slope species showed significant shifts in lower limits.to high elevations [the golden-mantled ground squirrel (Spermophilus lateralis) and the long-tailed vole (Microtus longicaudus)] (Table 1). Range expansions resulted from either expanded upper limits [the pocket mouse (Chaetodippus californicus). 23) to the 23 west slope taxa with sufficient trapping records to estimate detectability in both periods (tables S1 and S2 and Fig.05. Parallel trends were observed on the east slope of the Sierra for N.

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. 2010 true for chipmunks (Tamias).99 0.78 1. +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.36 0.sciencemag. expansion by C. Given are average detectability per site for Grinnell [P(G)] and current [P(C)] periods. –293 U Era*Elev +614 L Era + Elev + Elev2 +159 L. As examples. to species with mostly low. Even so.89 0.90 0.to high-elevation ranges (>2000 m) in Grinnell’s time.83 0.19 0.78 0. Values in bold are further supported by occupancy models. The ~500-m average increase in elevation for affected species is also consistent with estimated warming of +3°C.98 0.95 1.16 0. less fecund counterparts (table S5 and fig. accords with the predicted impacts of climate warming (5. habitat change at higher elevations is limited (15) (fig. ground squirrels (Spermophilus). S4 for elevation plots and models of individual species.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. S4).to mid-elevation ranges (<2000 m) and mid. maniculatus covered the entire transect.58 57–1160 models with those terms (w).39 0.92 NA 0. See fig. Original life zone (H.99 0. Beyond original elevation range (high versus low). 9).87 0.81 0. Because much of the transect spans a longprotected National Park.40 0. californicus and west slope P. confounding effects of land-use change are minimized.53 0.32 0.28 0.81 0.98 0. we provide an unbiased comparison of changes in species’ ranges at the centennial scale.88 0.48 0.98 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† 0.89 0. the cumulative Akaike’s Information Criterion weight for all No.91 0. P. 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.99 0. The preponderance of upward range shifts.97 0. S6).74 0. www. This was especially true for high-elevation species with upward contraction of their lower range limit. life history and ecological traits were weak predictors of which species exhibited upward shifts of their range limits (tables S5 and S6).90 0. assuming a change of temperature with elevation of ~6°C per km. and shrews (Sorex). and some have declined regionally (29). with ranching of introduced herbivores in Yosemite Valley and the high country recovering from historical overgrazing.99 NA 0. in part due to fire suppression (22). 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.23 0. monticola could reflect recovery of their preferred wet meadow habitats.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.REPORTS survey.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) 0.99 NA 0.org on May 5. respectively.93 0. Analyses of elevation change for 28 west slope species. truei are associated with fire-related conversion of conifer to shrub habitats. elevation.70 0.32 0.93 57–1160 183–1220 5 6 0. 1 2 Species P(G) P(C) Original elevation range (m) 0.62 0. the best supported form of the occupancy model (Elev. L) 0. whereas the downward shift of S.72 NA 0.org SCIENCE VOL 322 10 OCTOBER 2008 263 .98 0. Downloaded from www.50 0. S5).98 0. NA.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). However.37 Upper Sonoran (L) Upper Sonoran (L) Canadian–Hudsonian (H) 0.00 NA NA 1494–2210 1768–3155 1905–3155 2469–3353 +50 U +128 L. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an Table 1. where L and H refer.00 NA NA 0. not analyzed). −64 U Era + Elev + Elev2 +1007 L.60 0.99 Lower–Upper Sonoran (L) Upper Sonoran (L) 0. voles (Microtus).74 0. −334 U Elev +Elev2 +244 L Era*(Elev + Elev2) +512 L Era + Elev + Elev2 +609 L.85 1.82 0. Although vegetation dynamics have likely contributed to changes at low to mid-elevation.32 0. The elevational replacements among congeners.39 0.50 0.48 0. documented so carefully in the early 20th century (19). are now quite different. and original Lifezone classification (18).sciencemag.95 0. leading to contraction of high-elevation species and expansions of low-elevation taxa. +468 L 0. Best occupancy model Range limit change (m) Lower–Upper Sonoran (L) +112 U +589 U.71 0. changes in upper (U) and lower (L) range limit that are significant by the Pfa tests.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. original elevation range.71 0. Several small-mammal taxa that responded to changing temperature also showed large range fluctuations during late Quaternary climate fluctuations (28). 8. vegetation has changed within YNP over this period. −719 U Era*(Elev + Elev2) +355 L Elev +629 L Era + Elev +153 L NA No change NA 0.98 0.90 0. The park was hardly pristine in the early 20th century.71 0.

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