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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On the west slope. S2) using the 14 best detection models (table S3) to create model-averaged occupancy-elevation profiles (Fig. we excluded shifts that were statistically significant but biologically trivial (Fig. we predicted that species ranges should have shifted upward (5. era.sciencemag. and this occurred more frequently for lower than upper limits (c2 = 4. The same is Fig. df = 1. S3). water shrew (Sorex palustris). For each species we evaluated eight hypothesized relationships of occupancy. Exceptions occurred when occupancy models were weak (i. Our analyses of richness and turnover focused on species detectable by standardized trapping (37 species) or by observation (6 species. 3B). ornatus). There were fewer sites on the east side in both time periods (9 for Grinnell. 3C.05. we trapped small mammals at 121 sites compared to 56 in Grinnell’s time (table S1). with the largest change in the Lower and Upper Sonoran zones west of YNP. 2). Community similarity between Grinnell’s period and the present was high (mean similarity. whereas others did not (P. insufficient data) or detected changes in occupancy at elevations other than range limits. 1. but overall effort and elevational range (~50 to 3300 m) were comparable (22).7 df = 1. S > 0. Finally. but why species vary in response is not clear. S4. table S2). Belding’s ground squirrel (Spermophilus beldingi). P = 0. and the harvest mouse (Reithrodontomys megalotis)] or expanded lower limits (two shrews: Sorex monticola and mid-elevation species. whereas 50% of low-elevation species expanded their upper range compared to none of the high-elevation species (Fig. records from ad hoc collecting) were included in Pfa tests but not in occupancy models. some species of Peromyscus mice showed elevation range shifts (P. 10 OCTOBER 2008 VOL 322 SCIENCE www.26. S3). fig. P = 0. Map of surveyed sites in Grinnell (Historic) and Current surveys relative to the Yosemite National Park boundary and life zones (upper panel). F = 32. 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. High-elevation species typically experienced range contractions. truei).9) for the whole transect. 1).3 species.e. californicus). 2C). S3).to high-elevation species. P. 10). and altered community composition within elevational bands (9). and most life zones. df = 2. Species richness averaged across five estimators (26) that account for nonobserved species (Fig. upper limits changed significantly in only seven high elevations [the golden-mantled ground squirrel (Spermophilus lateralis) and the long-tailed vole (Microtus longicaudus)] (Table 1). To test for elevational shifts. cinerea and S. as expected given the upward expansions of formerly Sonoran zone taxa and the range shifts of high-elevation species (Table 1). and to an averaged elevational profile (lower panel). 1).004). Twelve of 28 (43%) west slope species showed significant shifts in lower limits.e. but not within YNP (+1. Elevational range shifts resulted in modest changes in species richness and composition at varying spatial scales. Elevational ranges of species and their habitats differed markedly between the gradual western and steep eastern slopes of the transect (19) (Fig. of which 10 increased (mean = +475 m) and two. and pika (Ochotona princeps). and table S4) declined from the Grinnell era to the present (repeated measures analysis of variance. mean = +501 m) and downward shifts (n = 3. 2A) included the alpine chipmunk (Tamais alpinus). with similar numbers of upward (n = 4. In contrast. 3A). both upper and lower limits increased by ~500 m. whereas low-elevation species expanded their ranges upward (c2 = Downloaded from www. Species richness was reduced within each life zone. −9%). mean = −309 m). 3). 12 for resurveys) because of limited extent (Fig. 23) to the 23 west slope taxa with sufficient trapping records to estimate detectability in both periods (tables S1 and S2 and Fig.. High-elevation species contracting (Table 1 and Fig. Conservatively. 2010 REPORTS . both shrews. and elevation (fig.012). we applied occupancy modeling (22. 4%). 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. decreased (mean = −744 m).sciencemag. the California vole (M. maniculatus). upward shift of the entire range or expansion of the upper limit for on May 5. Parallel trends were observed on the east slope of the Sierra for N. including Yosemite (25). P = 0. Range expansions resulted from either expanded upper limits [the pocket mouse (Chaetodippus californicus). Elevation limits shifted mostly upward (Table 1 and Fig. a pattern expected with increased temperature. boylii. where a species was not observed in one sampling period but was in the other (Table 1). In most cases where the Pfa test indicated an elevation shift. Given marked regional warming over the past century.. 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. beldingi (fig. This should manifest as upward contraction of the lower range limit for mid. 2 and fig. 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. Species composition was least similar for the Transition and Hudsonian-Arctic zones. Range contractions due to increases in lowerelevation limits were also observed for two species formerly at mid.039).Future warming is predicted to cause substantial turnover of species within North American National Parks. senex) (Fig. or when nonstandard data (i. Lower range limits contracted in 50% of the high-elevation species but in only 262 10% of low-elevation species. Closely related species responded idiosyncratically to climate change (Table 1).4 species. the pinyon mouse (Peromyscus truei) translocated upward (Fig. the park alone. Richness estimators suggest a slight decrease across the whole transect (currenthistoric mean estimates = −4.8. For example. 2B). occupancy models agreed (Table 1 and fig. Range shifts were significant if Pfa ≤ 0.

See SCIENCE VOL 322 10 OCTOBER 2008 263 .23 0.72 NA 0. accords with the predicted impacts of climate warming (5.48 0.81 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. original elevation range.50 0. 8.98 0.98 0. Several small-mammal taxa that responded to changing temperature also showed large range fluctuations during late Quaternary climate fluctuations (28).60 0.83 0.74 0.89 0. life history and ecological traits were weak predictors of which species exhibited upward shifts of their range limits (tables S5 and S6). and shrews (Sorex). P. Downloaded from www.32 0.16 0. As examples. The preponderance of upward range shifts. 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. S4 for elevation plots and models of individual species. Given are average detectability per site for Grinnell [P(G)] and current [P(C)] periods. −64 U Era + Elev + Elev2 +1007 L.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. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an Table 1.53 0. Original life zone (H.93 57–1160 183–1220 5 6 0. the cumulative Akaike’s Information Criterion weight for all No. 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. –293 U Era*Elev +614 L Era + Elev + Elev2 +159 L.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 0.99 Lower–Upper Sonoran (L) Upper Sonoran (L) 0. Because much of the transect spans a longprotected National Park.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) 0.87 0. californicus and west slope P. 1 2 Species P(G) P(C) Original elevation range (m) 0.95 1. in part due to fire suppression (22).85 1.sciencemag. Even so.91 0.88 0. −334 U Elev +Elev2 +244 L Era*(Elev + Elev2) +512 L Era + Elev + Elev2 +609 L. and original Lifezone classification (18). Analyses of elevation change for 28 west slope species. we provide an unbiased comparison of changes in species’ ranges at the centennial scale. vegetation has changed within YNP over this period.32 0.70 0.71 0. +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. expansion by C. The park was hardly pristine in the early 20th century.19 0. changes in upper (U) and lower (L) range limit that are significant by the Pfa tests.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. leading to contraction of high-elevation species and expansions of low-elevation taxa.90 0.00 NA NA 0.62 0.97 0.71 0.98 0.93 0. the best supported form of the occupancy model (Elev.37 Upper Sonoran (L) Upper Sonoran (L) Canadian–Hudsonian (H) 0.89 0.95 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. maniculatus covered the entire transect.98 0. truei are associated with fire-related conversion of conifer to shrub habitats. monticola could reflect recovery of their preferred wet meadow habitats. to species with mostly low.99 NA 0. not analyzed). ground squirrels (Spermophilus).00 NA NA 1494–2210 1768–3155 1905–3155 2469–3353 +50 U +128 L. where L and H on May 5.28 0. elevation. −719 U Era*(Elev + Elev2) +355 L Elev +629 L Era + Elev +153 L NA No change NA 0.74 0. Beyond original elevation range (high versus low). This was especially true for high-elevation species with upward contraction of their lower range limit.99 0. and some have declined regionally (29).32 0. S5). However.REPORTS survey.39 0.90 mid-elevation ranges (<2000 m) and mid. www. 9). Best occupancy model Range limit change (m) Lower–Upper Sonoran (L) +112 U +589 U.82 0. The elevational replacements among congeners.98 0. are now quite different.99 NA 0.58 57–1160 models with those terms (w). assuming a change of temperature with elevation of ~6°C per high-elevation ranges (>2000 m) in Grinnell’s time. confounding effects of land-use change are minimized.50 0. whereas the downward shift of S.48 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). documented so carefully in the early 20th century (19).99 0. voles (Microtus). habitat change at higher elevations is limited (15) (fig.78 1.sciencemag. S4).90 0.71 0.98 0. less fecund counterparts (table S5 and fig.36 0.39 0. NA.81 0.92 NA 0. S6). +468 L 0.78 0. 2010 true for chipmunks (Tamias).40 0. respectively.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. with ranching of introduced herbivores in Yosemite Valley and the high country recovering from historical overgrazing. L) 0.99 0. Values in bold are further supported by occupancy models.

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