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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S3). Range contractions due to increases in lowerelevation limits were also observed for two species formerly at mid. Elevational range shifts resulted in modest changes in species richness and composition at varying spatial scales. −9%). both upper and lower limits increased by ~500 m.3 species. df = 1. 3). 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. ornatus).. 4%). 1.to mid-elevation species.8. Lower range limits contracted in 50% of the high-elevation species but in only 262 10% of low-elevation species. including Yosemite (25). 10). High-elevation species typically experienced range contractions. water shrew (Sorex palustris). 23) to the 23 west slope taxa with sufficient trapping records to estimate detectability in both periods (tables S1 and S2 and Fig. with similar numbers of upward (n = 4. Finally. Belding’s ground squirrel (Spermophilus beldingi). whereas low-elevation species expanded their ranges upward (c2 = 8. both shrews. the California vole (M. where a species was not observed in one sampling period but was in the other (Table 1). S2) using the 14 best detection models (table S3) to create model-averaged occupancy-elevation profiles (Fig. 3B). mean = −309 m). the pinyon mouse (Peromyscus truei) translocated upward (Fig. we trapped small mammals at 121 sites compared to 56 in Grinnell’s time (table S1). df = 2. P = 0. Species richness averaged across five estimators (26) that account for nonobserved species (Fig.004). Elevational ranges of species and their habitats differed markedly between the gradual western and steep eastern slopes of the transect (19) (Fig. 2B).012). In most cases where the Pfa test indicated an elevation shift. upper limits changed significantly in only seven instances. the park alone. Range shifts were significant if Pfa ≤ 0. or when nonstandard data (i. To test for elevational shifts.26. maniculatus). Conservatively. a pattern expected with increased temperature. some species of Peromyscus mice showed elevation range shifts (P. upward shift of the entire range or expansion of the upper limit for low.to high-elevation species. P = 0. occupancy models agreed (Table 1 and fig. S4. 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. 2 and fig. table S2). 3C.039). 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. whereas others did not (P.. boylii. 2010 REPORTS . P = 0.7 df = 1. High-elevation species contracting (Table 1 and Fig. as expected given the upward expansions of formerly Sonoran zone taxa and the range shifts of high-elevation species (Table 1). fig. but why species vary in response is not clear.sciencemag.9) for the whole transect. decreased (mean = −744 m). For each species we evaluated eight hypothesized relationships of occupancy. For example. we excluded shifts that were statistically significant but biologically trivial (Fig. and most life zones. Richness estimators suggest a slight decrease across the whole transect (currenthistoric mean estimates = −4. 10 OCTOBER 2008 VOL 322 SCIENCE www. 1). and to an averaged elevational profile (lower panel). beldingi (fig. The same is Fig. and this occurred more frequently for lower than upper limits (c2 = 4. Parallel trends were observed on the east slope of the Sierra for N.org Downloaded from www. Closely related species responded idiosyncratically to climate change (Table 1). Elevation limits shifted mostly upward (Table 1 and Fig. 2C).e. Species composition was least similar for the Transition and Hudsonian-Arctic zones. era. P. truei).org on May 5. insufficient data) or detected changes in occupancy at elevations other than range limits. and pika (Ochotona princeps). S > 0. and table S4) declined from the Grinnell era to the present (repeated measures analysis of variance. and the harvest mouse (Reithrodontomys megalotis)] or expanded lower limits (two shrews: Sorex monticola and S.to high elevations [the golden-mantled ground squirrel (Spermophilus lateralis) and the long-tailed vole (Microtus longicaudus)] (Table 1). but not within YNP (+1.sciencemag. 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. of which 10 increased (mean = +475 m) and two. cinerea and S. mean = +501 m) and downward shifts (n = 3. records from ad hoc collecting) were included in Pfa tests but not in occupancy models.e. S3). This should manifest as upward contraction of the lower range limit for mid. 1).4 species. Map of surveyed sites in Grinnell (Historic) and Current surveys relative to the Yosemite National Park boundary and life zones (upper panel). with the largest change in the Lower and Upper Sonoran zones west of YNP. senex) (Fig. On the west slope. Twelve of 28 (43%) west slope species showed significant shifts in lower limits. 3A). Community similarity between Grinnell’s period and the present was high (mean similarity. whereas 50% of low-elevation species expanded their upper range compared to none of the high-elevation species (Fig. In contrast.05. 2). F = 32. we predicted that species ranges should have shifted upward (5. Range expansions resulted from either expanded upper limits [the pocket mouse (Chaetodippus californicus). There were fewer sites on the east side in both time periods (9 for Grinnell. Our analyses of richness and turnover focused on species detectable by standardized trapping (37 species) or by observation (6 species. Species richness was reduced within each life zone. Given marked regional warming over the past century. 12 for resurveys) because of limited extent (Fig. and altered community composition within elevational bands (9). but overall effort and elevational range (~50 to 3300 m) were comparable (22).Future warming is predicted to cause substantial turnover of species within North American National Parks. S3). we applied occupancy modeling (22. and elevation (fig. 2A) included the alpine chipmunk (Tamais alpinus). Exceptions occurred when occupancy models were weak (i. californicus).

95 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.98 0. and original Lifezone classification (18).36 0. changes in upper (U) and lower (L) range limit that are significant by the Pfa tests. 9). elevation.39 0.91 0.81 0.00 NA NA 0. Downloaded from www.95 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. 1 2 Species P(G) P(C) Original elevation range (m) 0. where L and H refer. documented so carefully in the early 20th century (19). 2010 true for chipmunks (Tamias). S6). less fecund counterparts (table S5 and fig. ground squirrels (Spermophilus).50 0. L) 0. to species with mostly low.74 0. Because much of the transect spans a longprotected National Park.00 NA NA 1494–2210 1768–3155 1905–3155 2469–3353 +50 U +128 L. are now quite different.to high-elevation ranges (>2000 m) in Grinnell’s time.93 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.60 0. assuming a change of temperature with elevation of ~6°C per km. expansion by C. This was especially true for high-elevation species with upward contraction of their lower range limit.50 0.97 193–914 549–914 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. californicus and west slope P. Analyses of elevation change for 28 west slope species. 8. habitat change at higher elevations is limited (15) (fig. truei are associated with fire-related conversion of conifer to shrub habitats. whereas the downward shift of S.48 0. S4).78 1.32 0.90 0.99 NA 0.40 0. As examples. monticola could reflect recovery of their preferred wet meadow habitats. 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. vegetation has changed within YNP over this period. −64 U Era + Elev + Elev2 +1007 L.REPORTS survey.32 0.82 0. Given are average detectability per site for Grinnell [P(G)] and current [P(C)] periods. the best supported form of the occupancy model (Elev.71 0. Best occupancy model Range limit change (m) Lower–Upper Sonoran (L) +112 U +589 U. Several small-mammal taxa that responded to changing temperature also showed large range fluctuations during late Quaternary climate fluctuations (28).19 0.62 0.78 0. See fig.97 0.99 0. S4 for elevation plots and models of individual species.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 0.37 Upper Sonoran (L) Upper Sonoran (L) Canadian–Hudsonian (H) 0.sciencemag.83 0. confounding effects of land-use change are minimized.89 0. original elevation range.81 0. S5). with ranching of introduced herbivores in Yosemite Valley and the high country recovering from historical overgrazing. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an Table 1.32 0.53 0.16 0.39 0.sciencemag.71 0. −719 U Era*(Elev + Elev2) +355 L Elev +629 L Era + Elev +153 L NA No change NA 0.98 0.to mid-elevation ranges (<2000 m) and mid. in part due to fire suppression (22).org SCIENCE VOL 322 10 OCTOBER 2008 263 .74 0. P.48 0. maniculatus covered the entire transect.99 0. The park was hardly pristine in the early 20th century.98 0. life history and ecological traits were weak predictors of which species exhibited upward shifts of their range limits (tables S5 and S6). the cumulative Akaike’s Information Criterion weight for all No. respectively. www.23 0. we provide an unbiased comparison of changes in species’ ranges at the centennial scale.99 Lower–Upper Sonoran (L) Upper Sonoran (L) 0. –293 U Era*Elev +614 L Era + Elev + Elev2 +159 L. Original life zone (H. NA. However. The elevational replacements among congeners.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) 0.58 57–1160 models with those terms (w). Even so.98 0.98 0.90 0.88 0. 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. Values in bold are further supported by occupancy models.70 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† 0.92 NA 0.90 0.99 NA 0. accords with the predicted impacts of climate warming (5. The ~500-m average increase in elevation for affected species is also consistent with estimated warming of +3°C. Although vegetation dynamics have likely contributed to changes at low to mid-elevation.28 0.93 57–1160 183–1220 5 6 0.89 0.71 0.87 0. not analyzed).org on May 5.85 1. and some have declined regionally (29).99 0. and shrews (Sorex). −334 U Elev +Elev2 +244 L Era*(Elev + Elev2) +512 L Era + Elev + Elev2 +609 L. Beyond original elevation range (high versus low). voles (Microtus). The preponderance of upward range shifts. +468 L 0.98 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). leading to contraction of high-elevation species and expansions of low-elevation taxa.72 NA 0.

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