Diane L. Delany

Response of Subalpine Conifers in the Sierra Nevada, California, U.S.A., to 20th-Century Warming and Decadal Climate Variability

Abstract Four independent studies of conifer growth between 1880 and 2002 in upper elevation forests of the central Sierra Nevada, California, U.S.A., showed correlated multidecadal and century-long responses associated with climate. Using tree-ring and ecological plot analysis, we studied annual branch growth of krummholz Pinus albicaulis; invasion by P. albicaulis and Pinus monticola into formerly persistent snowfields; dates of vertical branch emergence in krummholz P. albicaulis; and invasion by Pinus contorta into subalpine meadows. Mean annual branch growth at six treeline sites increased significantly over the 20th century (range 130–400%), with significant accelerations in rate from 1920 to 1945 and after 1980. Growth stabilized from 1945 to 1980. Similarly, invasion of six snowfield slopes began in the early 1900s and continued into snowfield centers throughout the 20th century, with significantly accelerated mean invasion from 1925 to 1940 and after 1980. Rate of snowfield invasion decreased between 1950 and 1975. Meadow invasion and vertical leader emergence showed synchronous, episodic responses. Pinus contorta invaded each of ten subalpine meadows in a distinct multidecadal pulse between 1945 and 1976 (87% of all trees) and vertical release in five krummholz P. albicaulis sites also occurred in one pulse between 1945 and 1976 (86% of all branches). These synchronies and lack of effect of local environments implicate regional climate control. Composite weather records indicated significant century-long increases in minimum monthly temperature and multidecadal variability in minimum temperature and precipitation. All ecological responses were significantly correlated with minimum temperature. Significant interactions among temperature, precipitation, Pacific Decadal Oscillation (PDO) indices, and multiyear variability in moisture availability further explained episodic ecological responses. Four multidecadal periods of the 20th century that are defined by ecological response (<1925; 1925–1944; 1945–1976; >1976) correlate with positive and negative PDO phases, as well as with steps in the rate of temperature increase. These diverse factors in spatially distributed upper-montane and treeline ecosystems respond directionally to century-long climate trends, and also exhibit abrupt and reversible effects as a consequence of interdecadal climate variability and complex interactions of temperature and moisture.

Response of high-elevation limber pine (Pinus flexilis) to multiyear droughts and 20th-century warming, Sierra Nevada, California, USA

Limber pine (Pinus flexilis James) stands along the eastern escarpment of the Sierra Nevada, California, experienced significant mortality from 1985 to 1995 during a period of sustained low precipitation and high temperature. The stands differ from old-growth limber pine forests in being dense, young, more even-aged, and located in warmer, drier microclimates. Tree growth showed high interannual variability. Relative to live trees, dead trees over their lifetimes had higher series sensitivity, grew more variably, and had lower growth. Although droughts recurred during the 20th century, tree mortality occurred only in the late 1980s. Significant correlations and interactions of growth and mortality dates with temperature and precipitation indicate that conditions of warmth plus sustained drought increased the likelihood of mortality in the 1985-1995 interval. This resembles a global-change-type drought, where warming combined with drought was an initial stress, trees were further weakened by dwarf mistletoe (Arceuthobium cyanocarpum (A. Nels. ex Rydb.) A. Nels.), and proximally killed by mountain pine beetle (Dendroctonus ponderosae Hopkins). However, the thinning effect of the drought-related mortality appears to have promoted resilience and improved near-term health of these stands, which suffered no additional mortality in the subsequent 1999-2004 drought.

New Records of Marginal Locations for American Pika (Ochotona princeps) in the Western Great Basin

Abstract. We describe 46 new site records documenting occupancy by American pika (lOchotona princeps) at 21 locations from 8 mountain regions in the western Great Basin, California, and Nevada. These locations comprise a sub- set of sites selected from regional surveys to represent marginal, isolated, or otherwise atypical pika locations, and to provide information for assessing environmental tolerance limits. Several locations are known from historic observations (Madeline Plain, Bodie Mtns., Wassuk Mtns., Mono Craters) and are included here to update current status. Site eleva- tions range from 1848 m to 3392 m; relative to the broad range of pika sites in the region, the new locations have cli- mates that are 2—4 °C warmer and receive approximately half the annual precipitation. Sites are located in lava flows and domes, inselbergs (isolated, rocky exposures on a small hill), eroding bedrock, rock-glacier till, talus slopes, and anthro- pogenic roadbed armaments and mining ore dumps. Several sites are situated in uncommon vegetation contexts, for example, montane desert scrub communities or locations where vegetation adjacent to taluses is sparse or lacking. Prox- imity to surrounding pika habitats (as a measure of marginality) was evaluated based on relative talus distribution pat- terns for 0.5-km, 2.5-km, and 5.0-km circular areas nested around each site. Seven idealized, schematic spatial patterns were used to assess potential connectivity among sites, ranging from “island” (no other talus within the respective areas) to “even” (many talus patches regularly distributed). Applying this approach to the 21 sites demonstrated a simple method for qualitatively assessing pika habitat relative to dispersal potential and metapopulation viability and also revealed complexities of biogeographic patterns related to marginality.

Thermal Regimes and Snowpack Relations of Periglacial Talus Slopes, Sierra Nevada, California, U.S.A.

Abstract Thermal regimes of eight periglacial talus slopes, at contrasting elevations, aspects, and substrates, in the Sierra Nevada, California, had complex microclimatic patterns partially decoupled from external conditions. Over three years, warm seasons showed mean talus matrix temperatures and daily variances lower than surfaces and cooler than free-air; talus surface and matrix positions low in the taluses were colder than higher positions, yielding highly positive altitudinal temperature differentials; ground surface temperatures had greater daily extremes than talus positions; and talus matrix temperatures lagged in response to surface temperature changes. Regulating processes in summer include evaporative cooling, cold-air drainage and Balch effect, and shading effects. In the cold season, talus matrices were warmer than surfaces; low talus positions were warmer than high; isothermal zero-curtain periods occurred before snow disappearance; and snow covered talus low positions more often and longer than higher in the taluses, which were often snow-free. Winter thermal processes likely include insulation from snow cover at talus bases, free exchange between talus matrix and external air in the upper talus, and latent heat from thaw-refreezing in late winter. Permanent ice may occur within high elevation talus slopes. Partially decoupled talus thermal regimes provide buffered habitats for mammals such as American pikas and are likely to be important refugia under future warming.

Radiocarbon dating of American pika fecal pellets provides insights into population extirpations and climate refugia

The American pika (Ochotona princeps) has become a species of concern for its sensitivity to warm temperatures and potential vulnerability to global warming. We explored the value of radiocarbon dating of fecal pellets to address questions of population persistence and timing of site extirpation. Carbon was extracted from pellets collected at 43 locations in the western Great Basin, USA, including three known occupied sites and 40 sites of uncertain status at range margins or where previous studies indicated the species is vulnerable. We resolved calibrated dates with high precision (within several years), most of which fell in the period of the mid-late 20th century bomb curve. The two-sided nature of the bomb curve renders far- and near-side dates of equal probability, which are separated by one to four decades. We document methods for narrowing resolution to one age range, including stratigraphic analysis of vegetation collected from pika haypiles. No evidence was found for biases in atmospheric 14C levels due to fossil-derived or industrial CO2 contamination. Radiocarbon dating indicated that pellets can persist for >59 years; known occupied sites resolved contemporary dates. Using combined evidence from field observations and radiocarbon dating, and the Bodie Mountains as an example, we propose a historical biogeographic scenario for pikas in minor Great Basin mountain ranges adjacent to major cordillera, wherein historical climate variability led to cycles of extirpation and recolonization during alternating cool and warm centuries. Using this model to inform future dynamics for small ranges in biogeographic settings similar to the Bodie Mountains in California, extirpation of pikas appears highly likely under directional warming trends projected for the next century, even while populations in extensive cordillera (e.g., Sierra Nevada, Rocky Mountains, Cascade Range) are likely to remain viable due to extensive, diverse habitat and high connectivity.

Thermal Components of American Pika Habitat—How does a Small Lagomorph Encounter Climate?

ABSTRACT Anticipating the response of small mammals to climate change requires knowledge of thermal conditions of their habitat during times of the day and year when individuals use them. We measured diurnal and seasonal temperatures of free air and of six habitat components for American pikas (Ochotona princeps) over five years at 37 sites in seven mountain ranges in the western Great Basin, United States. Talus matrices (subsurfaces) had low daily variances and, in the warm season, remained cool during the hottest times of the day relative to surfaces and free air. During winter, matrices were warmer than free air. Talus surfaces were warmer than free air in the warm and cold seasons, and had large daily variances. Summer forefield and dispersal environments were warmest of all habitat components. Talus surfaces in summer were highly responsive to solar radiation over the course of the day, warming quickly to high midday temperatures, and cooling rapidly in the evening. By contrast, matrices lagged the daily warm-up and remained warmer than free air at night. These differences afford diurnal and seasonal opportunities for pikas to adapt behaviorally to unfavorable temperatures and suggest that animals can accommodate a wider range of future climates than has been assumed, although warming of the dispersal environment may become limiting. Climate envelope models that use or model only surface air measures and do not include information on individual thermal components of pika habitat may lead to errant conclusions about the vulnerability of species under changing climates.

Distribution, climatic relationships, and status of American pikas (Ochotona princeps) in the Great Basin, USA

ABSTRACT To advance understanding of the distribution, climatic relationships, and status of American pikas (Ochotona princeps) in the Great Basin, United States, we compiled 2,387 records of extant pika sites surveyed since 2005, 89 records of documented extirpated sites (resurvey of historic sites), and 774 records of sites with old sign only. Extant sites extended across five degrees latitude and ten degrees longitude, encompassed six subregions, traversed forty mountain ranges, spanned 2,378 m in elevation (1,631–4,009 m), and comprised three of five currently described pika subspecies. A climate envelope for extant sites using the PRISM climate model expands the range of temperature and precipitation values that have been previously described. Extirpated and old-sign sites were mostly found within the geographic and climatic space of extant sites, but often in warmer and drier portions. Considerable overlap of extirpated, old, and extant groups within the same climate space suggests that nonclimatic factors have also contributed to population losses. The broad distribution and enlarged climate envelope of extant pika sites indicate that despite some localized extirpations, pika populations are persisting across Great Basin mountains, and appear to be able to tolerate a broader set of habitat conditions than previously understood.