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

48 0.95 0. and shrews (Sorex). truei are associated with fire-related conversion of conifer to shrub habitats.89 0. californicus and west slope P. accords with the predicted impacts of climate warming (5.sciencemag. life history and ecological traits were weak predictors of which species exhibited upward shifts of their range limits (tables S5 and S6).36 0.sciencemag. assuming a change of temperature with elevation of ~6°C per km. we provide an unbiased comparison of changes in species’ ranges at the centennial scale. 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. −719 U Era*(Elev + Elev2) +355 L Elev +629 L Era + Elev +153 L NA No change NA 0.98 0. −334 U Elev +Elev2 +244 L Era*(Elev + Elev2) +512 L Era + Elev + Elev2 +609 L.93 57–1160 183–1220 5 6 0. expansion by C. See fig. respectively.78 0.16 0. The park was hardly pristine in the early 20th century. Best occupancy model Range limit change (m) Lower–Upper Sonoran (L) +112 U +589 U. Downloaded from www.98 0. Increased prevalence of mesic small mammals following cessation of grazing has also been reported for an Table 1. S6).28 0. Given are average detectability per site for Grinnell [P(G)] and current [P(C)] periods.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 0.98 0. S4).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. not analyzed). and some have declined regionally (29).78 1.93 0.REPORTS survey.50 0. Original life zone (H. whereas the downward shift of S.90 0. Beyond original elevation range (high versus low).99 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).53 0.32 0. 8. 1 2 Species P(G) P(C) Original elevation range (m) 0.90 0.70 0.39 0. The ~500-m average increase in elevation for affected species is also consistent with estimated warming of +3°C. 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. +468 L 0.90 0. the best supported form of the occupancy model (Elev.98 0. –293 U Era*Elev +614 L Era + Elev + Elev2 +159 L. and original Lifezone classification (18). Because much of the transect spans a longprotected National Park.71 0.83 0. in part due to fire suppression (22).74 0.95 1. leading to contraction of high-elevation species and expansions of low-elevation taxa.88 0.99 0. The elevational replacements among congeners.81 0. where L and H refer.40 0.99 NA mid-elevation ranges (<2000 m) and mid. Even so.89 0.00 NA NA 1494–2210 1768–3155 1905–3155 2469–3353 +50 U +128 L. www. less fecund counterparts (table S5 and fig.87 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. As examples.32 0. However. −64 U Era + Elev + Elev2 +1007 L. maniculatus covered the entire transect.39 0.98 0.98 0.48 0. confounding effects of land-use change are minimized.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. Several small-mammal taxa that responded to changing temperature also showed large range fluctuations during late Quaternary climate fluctuations (28).19 0. voles (Microtus). S5). S4 for elevation plots and models of individual species. L) 0. the cumulative Akaike’s Information Criterion weight for all No.92 NA 0. are now quite different.99 0. NA. This was especially true for high-elevation species with upward contraction of their lower range limit.00 NA NA 0.23 0.81 0.82 0. P. ground squirrels (Spermophilus).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. with ranching of introduced herbivores in Yosemite Valley and the high country recovering from historical overgrazing. Although vegetation dynamics have likely contributed to changes at low to mid-elevation.74 0. 9).org on May 5.99 Lower–Upper Sonoran (L) Upper Sonoran (L) 0. The preponderance of upward range shifts. 2010 true for chipmunks (Tamias). documented so carefully in the early 20th century (19).50 0.99 NA 0. monticola could reflect recovery of their preferred wet meadow habitats. habitat change at higher elevations is limited (15) (fig. vegetation has changed within YNP over this high-elevation ranges (>2000 m) in Grinnell’s time.60 0.62 0.85 1.95 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† 0.58 57–1160 models with those terms (w).72 NA 0. Analyses of elevation change for 28 west slope species.97 0.37 Upper Sonoran (L) Upper Sonoran (L) Canadian–Hudsonian (H) 0. to species with mostly low.32 SCIENCE VOL 322 10 OCTOBER 2008 263 .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. elevation. original elevation range.91 0.71 0. changes in upper (U) and lower (L) range limit that are significant by the Pfa tests. Values in bold are further supported by occupancy models.

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