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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.

Although much research has examined the characteristics of Olympic marmot behavior and habitat, most studies mention the direct bioturbation effects of these marmots incidentally in an effort to examine other phenomena. A better understanding of marmot impacts on soil properties can provide a baseline for examining additional phenomena that are influenced by marmot activity. Soil surface and subsurface temperature, compaction, and moisture were measured in situ on abandoned marmot burrows in northeastern Olympic National Park, Washington. Soil samples were collected to analyze texture, pH, salinity, P, K, and organic matter content. Results indicated that soil compaction and organic matter were significantly reduced on abandoned marmot burrows when compared to undisturbed sites. Surface temperatures were typically warmer on marmot-disturbed soils, and subsurface temperatures were cooler. Soil moisture, texture, pH, salinity, and macronutrients were surprisingly similar on both disturbed and undisturbed so...

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.

256. Olympic Marmot Marmota olympus French: Marmotte des Olympic / German: Olympic-Mountains-Murmeltier / Spanish: Marmota de las Olimpicas Taxonomy. Marmota olympus Merriam, 1898, “From Timberline at head of Sole- duc River, Olympic Mountains, [Olympic Nat. Park] Washington [USA].” This species is monotypic. Distribution. Endemic to the Olympic Mts of W Washington (NW USA). Descriptive notes. Head-body 720-750 mm (males) and 670-690 mm (females), tail 210-237 mm (males) and 180-192 mm (females); weight 8.5-11 kg (males), 4.1-4.3 kg after hibernation and 6.7-7.5 kg (females), 2.7-3.5 kg after hibernation. The Olympic Marmot has dorsal pelage of straw yellow-brown to light brown that fades to yellow during summer. Snout is white often, extending to eyes. Venter is usually more rufous or gray than dorsal pelage. Tail is relatively long and densely haired, straw yellow to tan. Chromosomal complement is 2n = 44 and FN = 66. Habitat. Open grass, sedge, and forb subalpine and alpine meadows and talus slopes near and above timberline (most habitat is in the Olympic National Park, USA). Food and Feeding. The Olympic Marmotis an herbivore that feeds on sedges, herbs, roots, grasses and sometimes conifer bark and cambium. Dry grasses may be collected underground for either food or bedding. Foraging by the Olympic Marmot can remove ¢.30% of the biomass and promotes a high level of plant diversity. They will eat animal material when available; cannibalism has been observed. Breeding. The Olympic Marmotlives in multi-burrow colonies where most breeding occurs. Females typically reproduce in alternating years. Litters are 3-5 young, and young remain in natal areas until at least their third year when they disperse. Activity patterns. The Olympic Marmot is diurnal and is only active for 4-5 months each summer and hibernates within burrows for the remaining 7-8 months of the year; spring emergence is in April-May. Movements, Home range and Social organization. Burrows, often with small mounds at the entrance, are common in colonies and are used for maternal nests, sleeping dens, and escape. Sleeping chambers are often lined with dry plant material. Polygynous colonial groups commonly consisting of a single dominate adult male, less than three adult females and numerous two-year olds, one-year-olds, and young-of-the-year. The Olympic Marmot is among the most social of marmots. Colonial territories are 2-8-7 ha. Subordinate males may live on peripheries of colonies but are treated as non-residents. Mortality occurs mostly in winter, especially in years with little insulating snowpack. Scent marking of rocks and plants with oral and cheek glands is common. Four distinct alarm calls are produced in response to potential threats and number of whistles, tooth chatters, growls, “yips,” and “chucks” are used in a variety of contexts. Status and Conservation. Classified as Least Concern on The IUCN Red List. Population trend of the Olympic Marmot is declining. Habitat degradation due to climate change and unusually heavy predation by Coyotes (Canis latrans) and other predators in their highly isolated habitat are major challenges. Bibliography. Armitage (2013a, 2014), Barash (1973, 1989), Blumstein et al. (2001), Edelman (2003), Griffin et al. (2009), Thorington et al. (2012), Witczuk et al. (2013)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Sciuridae, pp. 648-837 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 822, DOI: 10.5281/zenodo.6840226

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.

Olympic marmots (Marmota olympus) are reported to skip at least one year between reproductive efforts. We observed several female marmots weaning infants in consecutive years. There was no evidence that reproductive skipping was more common than annual reproduction. High spring food availability resulting from climate change may allow females to wean consecutive litters regularly.

We established the phylogeny of 11 species of the genus Marmota based on the entire sequence of the mitochondrial cytochrome b (cyt-b) gene (1.1 kb) and a partial sequence of the NADH dehydrogenase subunit 4 (ND4) gene (1.2 kb). In three species (Marmota caligata, Marmota olympus, and Marmota bobac) full-sized nuclear pseudogenes of the mitochondrial cyt-b were identified. The mitochondrial cyt-b genes and the three pseudogenes form separate clusters in the maximum parsimony dendrogram. This finding suggests that the pseudogenes originated from a single transfer to the nucleus that may have occurred prior to the radiation of the genus Marmota. Notably, compared with their functional mitochondrial equivalents the pseudogenes show a much lower substitution rate. In the dendrograms deduced from the mitochondrial sequences two distinct clusters become apparent: one cluster consists of the North-west American species, the other contains the Eurasian species together with the North American species Marmota monax. The position of M. monax as a member of the Eurasian clade is in accordance with the evolution of chromosome numbers. The results are of special interest with respect to the evolution of social systems in the genus that vary from solitary species (M. monax) to highly social species living in family groups (e.g. Marmota marmota). The molecular phylogeny suggests a diphyletic origin of high sociality in the genus Marmota.

Many species produce alarm calls that vary according to situation. Theoretically, alarm call structure could covary with predator type and could communicate potentially ''referential information, or calls could covary with the degree of risk a caller experienced when it emitted a call. Using similar methods, I studied the ways in which Olympic (Marmota olympus), hoary (M. caligata), and Vancouver Island marmots (M. vancouverensis) communicated situational variation. I observed both natural alarm calling, and I artificially elicited alarm calls with simulated terrestrial and aerial predators. I used playback experiments to study marmots' responses to different alarm call variants. All three species produced four roughly similar but distinctive loud alarm vocalizations that could be categorized by their relative shape, duration, and whether calls were quickly repeated to create multi-note vocalizations. In addition, the Vancouver Island marmot produced a fifth loud alarm call-the kee-aw. Call micro-structure varied as a function of the distance the caller was from an alarming stimulus and the type of alarming stimulus. Two lines of evidence suggest that all three species had alarm calls associated with the caller's risk (i.e. they were not referential). First, marmots often changed call types within a calling bout: there were no unique stimulus-class specific vocalizations. Second, marmot responses to alarm calls were graded: marmots did not have unique responses to different call types. These three close taxonomic relatives with superficially similar calls, communicated risk differently.

Marmota vancouverensis is the only insular species among the 14 species of marmots. The evolutionary history of this species is unresolved. Although M. vancouverensis is strongly differentiated in osteological and other morphological characters, its low genetic divergence suggests recent evolution from an ancestral continental species. We used geometric morphometric techniques to assess the morphology of hemimandibles from 239 modern M. vancouverensis, Marmota caligata, Marmota flaviventris, Marmota olympus and 30 Holocene (9435–735 cal. yr bp) subfossil M. vancouverensis. Our results confirm that the mandible of M. vancouverensis is strongly differentiated in shape from continental marmot species, but similar in size to its mainland sister species M. caligata. Temporal variation in size and shape over the past 2500 years among allochronic samples of M. vancouverensis was minimal suggesting that the morphological divergence of this species occurred in a period of rapid change following its isolation from mainland populations in the late Pleistocene. Selection pressures associated with environmental changes and founder effects and genetic drift resulting from population bottlenecks created by population declines and habitat fragmentation are hypothesized as factors contributing to the morphological divergence of this species.

Marmota olympus (Merriam, 1898). Proc. Acad. Nat. Sci. Philadelphia, 50:352. TYPE LOCALITY: "From Timberline at head of Soleduc River, Olympic Mountains, [Olympic Nat. Park] Washington [USA]." DISTRIBUTION: Olympic Mtns of W Washington (USA). COMMENTS: Considered a subspecies of marmota (Rausch, 1953). Reviewed by Hoffmann et al. (1979) who confirmed its specific status., Published as part of Robert S. Hoffmann & Charles G. Anderson, 1993, Order Rodentia - Family Sciuridae, pp. 419-465 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 432, DOI: 10.5281/zenodo.7353130, {"references":["Rausch, R. L. 1953. On the status of some Arctic mammals. Arctic (Journal of the Arctic Institute of North America), 6: 91 - 148.","Hoffmann, R. S., J. W. Koeppl, and C. F. Nadler. 1979. The relationships of the amphiberingian marmots (Mammalia: Sciuridae). Occasional Papers of the Museum of Natural History, University of Kansas, 83: 1 - 56."]}

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.

This report presents the results of a 3-year study of the Olympic marmot in Olympic National Park. The animals inhabited sub-alpine and alpine meadows and formed closelyorganized colonies with extensive burrow systems. Southern exposures were preferred and no correlation of seasonal emergence with snow depth was apparent. Marmots ate nearly all meadow plants plus occasional animal matter early in the season. A variety of natural predators were present, and instances of predation by a cougar and coyote are reported. Visual and accoustic communication is described and sonograms of different vocalizations are presented. Activity patterns paralleled those reported for other marmot species, with early-morning and late afternoon peaks. Detailed activity budgets are presented, indicating behavioural differences among sex and age classes, and with changes in season. Records of weights are presented for all classes, indicating steady seasonal increases and sharp declines during hibernation. A black moult began in mid-summer and is apparently unique to the Olympic marmot. Most colonies were composed of one adult male, two adult females, a litter of yearlings and a litter of infants. Occasionally one or more 2-year-olds and one or two ‘satellite’ adult males were present. Colony members generally hibernated together but parous females and satellite males moved away to separate burrows shortly after emergence. Varying combinations of living arrangements are described. Greeting behaviour was analysed and quantified as a major component of marmot sociality. Greeting frequency varied significantly with age and sex class, season and time of day: it was especially high in the morning, among adult males in the spring, and among infants. Dominance relations were generally diffuse, except those between resident and satellite males (resident dominated) and between pregnant or lactating females and any other animal (female dominated). Individual territories and home ranges did not exist within colonies. Quantification of inter-individual distances revealed a tendency for greater aggregation early in the season among most animals but greater earlyseason repulsion between resident and satellite males. The ontogeny of infant activity is quantitatively described, demonstrating progressively more time spent above ground and increased distance travelled from the home burrow. The decline in association between females and their litters is also described. Upright, ‘play-fighting’ was common and showed age, sex and seasonal differences with high levels among adult males early in the season and among infants. ‘Playfulness’ was prominent in marmot social behaviour. Burrow visiting, greeting, social grouping and playfighting all reflected intense social interaction early in the season with gradual declines later. Flow charts of sexual activity are used to compare the behaviour patterns of parous and nonparous females and to analyse changes in pattern with time following emergence. Distinct oestrus behaviour was apparent for the parous females, peaking during the second week post-emergence. Interactions between colonies are also described: they generally evidenced lower frequencies and greater intensities of agonism than did withincolony behaviours. The social system of M. olympus was shown to be more closely integrated and less aggresively-organized than that of the yellow-bellied marmot, M. flaviventris, which in turn is more highly social than the woodchuck, M. monax. Survivorship data indicate that mortality rates were highest among infants and during the winter, especially during winters with scanty snowfall. Two-year-olds dispersed upon seasonal emergence in colonies where winter mortality was low; however, following high winter mortality and consequent small colony size at emergence, dispersal did not occur. Thus, colony population size was maintained essentially constant and relatively independent of yearly fluctuations in mortality or recruitment. Dispersal from large colonies was correlated with behavioural responses of the 2-year-olds to a relatively high level of social interaction in these colonies. Olympic marmot females mature at 3 years and bear young in alternate years only. The evolutionary significance of biennial breeding is considered, and this population system is interpreted as providing maximum effective reproduction under the conditions of extremely short growing season prevailing in the marmot meadows. Comparison with other North American marmots reveals a progressive decrease in length of growing season experienced by the wood-chuck, yellow-bellied and Olympic marmots. This in turn correlates with decreases in proportional growth rates and breeding frequency, and increases in age at dispersal, age at sexual maturity and in the degree of ‘sociality’ of these marmot species. Implications for the evolution of marmot behaviour are discussed, including the suggestion that exposure to progressively shortened growing seasons favours the development of less aggressively-organized social systems.

Marmota olympus (Merriam, 1898). Proc. Acad. Nat. Sci. Phila., 50:352. TYPE LOCALITY: U.S.A., Washington, timberline at head of Soleduc River, Olympic Mtns. DISTRIBUTION: Olympic Mtns, of W. Washington (U.S.A.). COMMENT: Reviewed by Hoffmann et al., 1979, Occas. Pap. Mus. Nat. Hist. Univ. Kans., 83:1 -56. ISIS NUMBER: 5301410002016007001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 1), pp. 345-382 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 355, DOI: 10.5281/zenodo.7353023, {"references":["Hoffmann, R. S., J. W. Koeppl, and C. F. Nadler. 1979. The relationships of the amphiberingian marmots (Mammalia: Sciuridae). Occasional Papers of the Museum of Natural History, University of Kansas, 83: 1 - 56."]}