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Although climate has been shown to have a considerable influence on alpine forest dynamics, many fine-scale processes contribute to observable forest patterns. Disturbances created by burrowing mammals may provide conifer seedlings with refuge in previously inhospitable environments and may result in conifer invasion that is not directly associated with climate change. In Olympic National Park, Washington, USA, Olympic marmot (Marmota olympus) populations have declined. Abandoned marmot burrows, which may provide ideal sites for conifer establishment, now dot the alpine landscape. Similarly, in Glacier National Park, Montana, USA, pocket gophers (Thomomys talpoides) have disturbed large areas of alpine meadows at treeline. Field and laboratory analysis of gopher ‘eskers’ and spoil mounds of abandoned marmot burrows illustrated that such sites are less compacted than soils in adjacent, undisturbed locations. Additionally, disturbed soils contained higher amounts of P and lower amounts of K and organic matter. Soils disturbed by gophers were cooler at the surface and warmer at depth than undisturbed soils, whereas marmot-disturbed soils were warmer at the surface and cooler at depth. Experimental seed germination of subalpine fir (Abies lasiocarpa) was higher on mammal-disturbed soils, and this was statistically significant for marmot-disturbed soils. These findings suggest that fossorial mammal disturbances have the potential to affect conifer germination and establishment. Furthermore, this research initiates the dialogue for future research on the distinct morphological and behavioral influences of different fossorial mammals as zoogeomorphic agents and ecosystem engineers. This article is protected by copyright. All rights reserved.

We investigated predation by nonnative coyotes (Canis latrans) on endemic Olympic marmots (Marmota olympus) in Olympic National Park, Washington, in 2005 and 2006. Although nearly the entire marmot range is protected within the park, declines and local extirpations of the species have been documented. Through analyses of carnivore scat across the range of the Olympic marmot we determined the distribution and relative density of coyotes and characterized the extent to which coyotes and native carnivores preyed on marmots. We used mitochondrial DNA analysis of scats to determine carnivore species, and microsatellite markers for individual coyote identification. Scat analysis indicated that invasive coyotes are widespread and the numerically dominant carnivore on sampled trails within the Olympic highlands—71% (301 of 426) of all scats verified to species arose from coyote. Out of all carnivore scats collected, 11.6% (111 of 958) contained marmot remains. For 85% of the samples with marmots, coyotes were confirmed as the predator. The remainder arose from bobcat (13%) and cougar (2%). Coyotes were the predominant marmot predator across all months and in most regions of the park. Twelve out of 13 coyote individuals identified with genetic markers preyed on marmots. Marmots ranked 5th in frequency of coyote diet items, after snowshoe hares (Lepus americanus), mountain beavers (Aplodontia rufa), voles, and cervids. Scat analysis indicated that in the Olympic Mountains, the coyote as an invasive generalist predator is subsidized by abundant multiple prey, and appears to be the primary terrestrial predator on the endemic Olympic marmot. We conclude that predation by coyotes on marmots is widespread and substantial across the marmot's species range, and therefore likely driving observed marmot declines and extinctions.

We reinvestigated the phylogeny of all 15 species of Marmota to resolve a conflict between 2 published analyses, one by Kruckenhauser et al. and another by Steppan et al., regarding the Olympic marmot (M. olympus) and to improve resolution in the genus. We acquired fresh samples of M. olympus, combined all available data on mitochondrial DNA (cytochrome b [Cytb] and ND3/ND4), new sequences for ND3/ND4, and 2,000 base pairs (bp) of the nuclear RAG1 gene. All analyses indicate that M. olympus is a much older, rather than more recent, offshoot of the widespread hoary marmot (M. caligata) or Vancouver Island marmot (M. vancouverensis). The mitochondrial data and some RAG1 results are largely congruent, but RAG1 differs on several points, including: the subgenus Marmota appears paraphyletic to Petromarmota, with reciprocally monophyletic Palearctic and Nearctic clades; and the long-tailed marmot (M. caudata) and Menzbier's marmot (M. menzbieri) are not sister species, suggesting mitochondrial introgre...

The Mahalanobis distance statistic (D 2 ) has emerged as an effective tool to identify suitable habitat from presence data alone, but there has been no mechanism to select among potential habitat covariates. We propose that the best combination of explanatory variables for a D 2 model can be identified by ranking potential models based on the proportion of the entire study area that is classified as potentially suitable habitat given that a predetermined proportion of occupied locations are correctly classified. In effect, our approach seeks to minimize errors of commission, or maximize specificity, while holding the omission error rate constant. We used this approach to identify potentially suitable habitat for the Olympic marmot (Marmota olympus), a declining species endemic to Olympic National Park, Washington, USA. We compared models built with all combinations of 11 habitat variables. A 7-variable model identified 21,143 ha within the park as potentially suitable for marmots, correctly classifying 80% of occupied locations. Additional refinements to the 7-variable model (e.g., eliminating small patches) further reduced the predicted area to 18,579 ha with little reduction in predictive power. Although we sought a model that would allow field workers to find 80% of Olympic marmot locations, in fact, ,3% of 376 occupied locations and ,9% of abandoned locations were .100 m from habitat predicted by the final model, suggesting that .90% of occupied marmot habitat could be found by observant workers surveying predicted habitat. The model comparison procedure allowed us to identify the suite of covariates that maximized specificity of our model and, thus, limited the amount of less favorable habitat included in the final prediction area. We expect that by maximizing specificity of models built from presence-only data, our model comparison procedure will be useful to conservation practitioners planning reintroductions, searching for rare species, or identifying habitat for protection.

range in the spring of a given year, 16% of 2-year-old males, 50% of 3-year-old males, 17% of 2-year-old females, and 29% of 3-year-old females subsequently dispersed. Dispersal rates for 3-year-olds were slightly lower when all animals were included in the analysis regardless of whether their dispersal history was known. Males dispersed farther than females (median 5 984 m, n 5 14 versus median 5 267 m, n 513) and 69% of females settled within 500 m of their original home range. If the observed dispersal patterns are representative of range-wide patterns and if Olympic marmot densities remain low, successful dispersal may be too infrequent to sustain reliable recolonization of vacant habitats or even genetic or demographic rescue of isolated marmot groups.

In the mid-1990s, anecdotal reports of Olympic marmot ( Marmota olympus ) disappearances from historically occupied locations suggested that the species might be declining. Concern was heightened by the precipitous decline of the Vancouver Island marmot ( Marmota vancouverensis ), coupled with reports that climate change was affecting other high-elevation species. However, it was unclear whether the Olympic marmot was declining or undergoing natural extinctions and recolonizations; distinguishing between normal metapopulation processes and population declines in naturally fragmented species can be difficult. From 2002–2006, we used multiple approaches to evaluate the population status of the Olympic marmot. We surveyed sites for which there were records indicating regular occupancy in the later half of the 20th century and we conducted range-wide surveys of open high-elevation habitat to establish current and recent distribution. We used these targeted and general habitat surveys to identify locations and regions that have undergone extinctions or colonizations in the past 1–4 decades. Simultaneously, we conducted detailed demographic studies, using marked and radio-tagged marmots, to estimate the observed and projected current population growth rate at nine locations. The habitat surveys indicate that local extinctions have been wide-spread, while no recolonizations were detected. Abundance at most intensive study sites declined from 2002–2006 and the demographic data indicate that these local declines are ongoing. Adult female survival in particular is considerably lower than it was historically. The spatial pattern of the extinctions is inconsistent with observed metapopulation dynamics in other marmot species and, together with very low observed dispersal rates, indicates that population is not at equilibrium.

If changes in animal behavior resulting from direct human disturbance negatively affect the persistence of a given species or population, then these behavioral changes must necessarily lead to reduced demographic performance. We tested for the effects of human disturbance on Olympic marmots (Marmota olympus), a large ground-dwelling squirrel that has disappeared from several areas where recreation levels are high. We assessed the degree to which antipredator and foraging behavior and demographic rates (survival and reproduction) differed between sites with high recreation levels (high use) and those with little or no recreation (low use). Compared with the marmots at low-use sites, marmots at high-use sites displayed significantly reduced responses to human approach, which could be construed as successful accommodation of disturbance or as a decrease in predator awareness. The marmots at high-use sites also looked up more often while foraging, which suggests an increased wariness. Marmots at both types of sites had comparable reproductive and survival rates and were in similar body condition. Until now, the supposition that marmots can adjust their behavior to avoid negative demographic consequences when confronted with heavy tourism has been based on potentially ambiguous behavioral data. Our results support this hypothesis in the case of Olympic marmots and demonstrate the importance of considering demographic data when evaluating the impacts of recreation on animal populations.

Monitoring of rare and declining species is one of the most important tasks of wildlife managers. Here we present a large-scale, long-term monitoring program for Olympic marmot (Marmota olympus) throughout its range across a logistically challenging mountainous park. Our multiple-stage process of survey design accounts for the difficulty imposed by access to remote habitats and funding constraints. The Olympic marmot is endemic to the Olympic Mountains, Washington State, USA. Although nearly all of its range is enclosed within Olympic National Park, declines and local extirpations of the species have been documented. We considered several possible alternative survey approaches, and propose a monitoring program designed to reflect extinction–recolonisation dynamics using presence–absence data. The sampling design is based on annual surveys of a set of at least 25 randomly selected clusters (closely located groups of sites with record of current or historical occupancy by marmots), and supplemented by sampling 15 never-occupied sites to test for new colonisations. The monitoring plan provides a framework that park managers can use for assessing changes over time in Olympic marmot distribution across the range of the species. Our sampling design may serve as a useful case study for establishing monitoring programs for other species with clumped distributions.