Ground-dwelling Squirrels Research Articles

Niches of three sympatric montane ground-dwelling squirrels: Relative importance of climate, topography, and landcover.

Species with different ecological niches will likely exhibit distinct responses to a changing environment. Differences in the magnitude of niche specialization may also indicate which species may be more vulnerable to environmental change, as many life-history characteristics are known to affect climate change vulnerability. We characterized the niche space of three sympatric high-elevation ground-dwelling squirrels, yellow-bellied marmot (Marmota flaviventer), Belding's ground squirrel (Urocitellus beldingi), and golden-mantled ground squirrel (Callospermophilus lateralis), in the alpine and upper subalpine regions of the Sierra Nevada in California. We used 5879 observations of individual squirrels, collected from 4 years (2009-2012) of transect survey data, to quantify which ecogeographical variable types (climate, topography, or landcover) were most important in defining the niche of each species. We conducted Ecological Niche Factor Analysis to quantify the niche and generate indices of "marginality" (magnitude of selection) and "specialization" (narrowness of niche space). All three species demonstrated differential use of niche space when compared to the available niche space. Moreover, the relative importance of the variables shaping the niche differed among these species. For example, the presence of meadows was important in defining the niche for U. beldingi and M. flaviventer, but the presence of conifers was important to C. lateralis. Precipitation was important in defining the niche for all three species, positively so for U. beldingi, and negatively for the other two species. The niche breadth of these three species was also positively associated with geographic range size. Mammals in high-elevation mountain systems often are perceived as vulnerable to climate shifts, but our results underscore the importance of also including non-climate-based factors in defining the niche. The overall magnitude of niche selection for all three species was driven by a combination of topographic, climatic, and landcover factors; thus, efforts to forecast areas where these species can persist in the future need to evaluate from more than just a climatic perspective.

Social organization in a North African ground squirrel

AbstractResearch on sociality in temperate ground-dwelling squirrels has focused on female philopatry and other life history trade-offs, which are influenced by constraints in the duration of the active growing season. Temperate ground-dwelling squirrels that experience high predation pressure, are large in body size, and have a short active season, show a more complex social organization. In contrast, African ground squirrels are active year-round, suggesting that instead of a short active season, distinct selective pressures influence their social organization. We examined the social organization of Barbary ground squirrels, Atlantoxerus getulus, and compared the social organization of temperate and African ground-dwelling sciurids. Anecdotal accounts on Barbary ground squirrels’ social organization suggested that they were either solitary or gregarious, or live in small family groups. We recorded the group size, composition, cohesion, and genetic relatedness, of the population on the arid island of Fuerteventura, Spain. Our data indicate that females live in small (1–8) all-female kin groups separate from adult males, and that unrelated adult males share sleeping burrows with immature individuals of either sex. We observed sex-biased dispersal with males primarily the dispersing sex and females primarily philopatric. Females sleep solitarily during gestation and lactation and nest either communally or singly after juvenile emergence. During the day, males and females can be active in the same area. Barbary ground squirrels are social because the squirrels share sleeping burrows and show spatiotemporal overlap. Barbary ground squirrels’ social organization resembles that of the closely related Cape ground squirrel rather than that of the temperate ground-dwelling sciurids, although the former are more temperate, seasonal breeders. In addition to describing the social organization of a previously unstudied species, this paper sheds light on the ecological drivers of sociality, and the evolution of distinct social organizations in ground-dwelling sciurids.

Ice-Age Climate Adaptations Trap the Alpine Marmot in a State of Low Genetic Diversity

SummarySome species responded successfully to prehistoric changes in climate [1, 2], while others failed to adapt and became extinct [3]. The factors that determine successful climate adaptation remain poorly understood. We constructed a reference genome and studied physiological adaptations in the Alpine marmot (Marmota marmota), a large ground-dwelling squirrel exquisitely adapted to the “ice-age” climate of the Pleistocene steppe [4, 5]. Since the disappearance of this habitat, the rodent persists in large numbers in the high-altitude Alpine meadow [6, 7]. Genome and metabolome showed evidence of adaptation consistent with cold climate, affecting white adipose tissue. Conversely, however, we found that the Alpine marmot has levels of genetic variation that are among the lowest for mammals, such that deleterious mutations are less effectively purged. Our data rule out typical explanations for low diversity, such as high levels of consanguineous mating, or a very recent bottleneck. Instead, ancient demographic reconstruction revealed that genetic diversity was lost during the climate shifts of the Pleistocene and has not recovered, despite the current high population size. We attribute this slow recovery to the marmot’s adaptive life history. The case of the Alpine marmot reveals a complicated relationship between climatic changes, genetic diversity, and conservation status. It shows that species of extremely low genetic diversity can be very successful and persist over thousands of years, but also that climate-adapted life history can trap a species in a persistent state of low genetic diversity.

The importance of individual variation in the alarm calls of Gunnison's prairie dogs

Gunnison's prairie dogs, Cynomys gunnisoni, frequently utter a repetitive series (a ‘bout’) of alarm calls (called ‘barks’) in response to a variety of terrestrial predators. Previous work has suggested that the structure of barks is finely tuned in ways that provide highly specific information about the identity of the predator. We re-evaluated these findings by presenting individually marked adult females with taxidermy mounts of three terrestrial predators of prairie dogs: a coyote, Canis latrans, a bobcat, Lynx rufus, and an American badger, Taxidea taxus, as well as one control stimulus (a cardboard box). Significant variation among individuals was evident from analyses of 10 measures associated with the general pattern of barking (e.g. latency to begin barking, bout duration, average number of barks/bout, overall bark rate, interbout interval) and also from eight structural features of individual barks (e.g. frequency and duration measures of the fundamental frequency band). When controlling for the identity of alarm callers, we found little evidence for differences in these same variables in response to different stimuli. We conclude that, like several other species of ground-dwelling squirrels, Gunnison's prairie dogs have an individually distinctive alarm call that does not differ for different predators. Our results conflict with earlier claims of extreme specificity in the alarm calls of Gunnison's prairie dogs.

Both familiarity and kinship influence odour discrimination by females in a highly social African ground squirrel

Kin recognition can be important in species where inbreeding avoidance or nepotism (favouritism towards kin) rely on identifying kin, particularly in species with alloparental care. The mechanisms that facilitate kin discrimination, where recognition is determined through cues that correlate with relatedness, usually include either prior association (familiarity) or phenotype matching or both. Odour is an important cue used in a number of mammalian species to discern kin, particularly the ground-dwelling squirrels. Cape ground squirrels, Xerus inauris, are a cooperative-breeding species, living in tight-knit family groups. However, group fission, promiscuity and the large home ranges of breeding males result in a high variance in relatedness both within and among social groups, making them a good species to investigate kin discrimination. We examined whether females are capable of discriminating between the odours of familiar versus unfamiliar females that varied in their relatedness to the focal female using odour experiments. Overall, the average duration of sniffing of the odours of familiar and unfamiliar females did not differ. Similarly, females did not adjust their sniff duration relative to the degree of relatedness of familiar females. However, females appeared to discriminate by the degree of relatedness of unfamiliar (stranger) females, spending longer sniffing odours from females that were not related to them. Thus, females were capable of discriminating the degree of relatedness from odour but they did not do so within their family group. We conclude that Cape ground squirrels are able to discriminate kin. However, whether females in this facultative cooperative breeder use the degree of relatedness in direct social interactions and nepotistic behaviours remains to be investigated.

Impacts of Inference Method and Data set Filtering on Phylogenomic Resolution in a Rapid Radiation of Ground Squirrels (Xerinae: Marmotini).

Phylogenomic data sets are illuminating many areas of the Tree of Life. However, the large size of these data sets alone may be insufficient to resolve problematic nodes in the most rapid evolutionary radiations, because inferences in zones of extraordinarily low phylogenetic signal can be sensitive to the model and method of inference, as well as the information content of loci employed. We used a data set of $>$3950 ultraconserved element (UCE) loci from a classic mammalian radiation, ground-dwelling squirrels of the tribe Marmotini (Sciuridae: Xerinae), to assess sensitivity of phylogenetic estimates to varying per-locus information content across four different inference methods (RAxML, ASTRAL, NJst, and SVDquartets). Persistent discordance was found in topology and bootstrap support between concatenation- and coalescent-based inferences; among methods within the coalescent framework; and within all methods in response to different filtering scenarios. Contrary to some recent empirical UCE-based studies, filtering by information content did not promote complete among-method concordance. Nevertheless, filtering did improve concordance relative to randomly selected locus sets, largely via improved consistency of two-step summary methods (particularly NJst) under conditions of higher average per-locus variation (and thus increasing gene tree precision). The benefits of phylogenomic data set filtering are variable among classes of inference methods and across different evolutionary scenarios, reiterating the complexities of resolving rapid radiations, even with robust taxon and character sampling.

Spatial behaviour of an overlooked alien squirrel: The case of Siberian chipmunks Eutamias sibiricus

Alien species of concern within the European Union have been recently listed and their populations need to be monitored, to plan addressed eradication or control programs. Therefore, the assessment of their presence should be rapidly carried out, particularly for elusive species or for those living at low densities. The Siberian chipmunk Eutamias sibiricus is a ground-dwelling squirrel, naturally distributed in northern and eastern Asia. Many introduced populations occur in Europe and Italy too. This species has been listed within the invasive species concern within the European Union and, thus, monitoring is mandatory to manage its potential range expansion. We carried out a hair-tube survey on 31 wood patches in northern and central Italy, where reproductive populations of Siberian chipmunk have been recorded. Hair tubes provided reliable data in assessing the presence of the Siberian chipmunk, with only 1% pseudo-absence and a high detection probability. The occurrence of Siberian chipmunk was positively influenced by study site and by the distance from release site, confirming low dispersal abilities by this species. Dense understorey also affected the presence of chipmunks, preventing them to search for food on the ground and to dig burrows.

Trait‐specific processes of convergence and conservatism shape ecomorphological evolution in ground‐dwelling squirrels

Our understanding of mechanisms operating over deep timescales to shape phenotypic diversity often hinges on linking variation in one or few trait(s) to specific evolutionary processes. When distinct processes are capable of similar phenotypic signatures, however, identifying these drivers is difficult. We explored ecomorphological evolution across a radiation of ground-dwelling squirrels whose history includes convergence and constraint, two processes that can yield similar signatures of standing phenotypic diversity. Using four ecologically relevant trait datasets (body size, cranial, mandibular, and molariform tooth shape), we compared and contrasted variation, covariation, and disparity patterns in a new phylogenetic framework. Strong correlations existed between body size and two skull traits (allometry) and among skull traits themselves (integration). Inferred evolutionary modes were also concordant across traits (Ornstein-Uhlenbeck with two adaptive regimes). However, despite these broad similarities, we found divergent dynamics on the macroevolutionary landscape, with phenotypic disparity being differentially shaped by convergence and conservatism. Such among-trait heterogeneity in process (but not always pattern) reiterates the mosaic nature of morphological evolution, and suggests ground squirrel evolution is poorly captured by single process descriptors. Our results also highlight how use of single traits can bias macroevolutionary inference, affirming the importance of broader trait-bases in understanding phenotypic evolutionary dynamics.

Social security: social relationship strength and connectedness influence how marmots respond to alarm calls

Individuals vary in the number and types of social relationships they maintain. If beneficial, social relationships may reduce predation risk and thus increase an individual’s sense of security. We tested this hypothesis by studying the responses of female yellow-bellied marmots (Marmota flaviventer) to broadcast alarm calls from unfamiliar individuals. First, we quantified affiliative interactions of animals in the field to calculate a set of social network measures. Because attributes of sociality are often correlated, we used principal component analysis to reduce our social network metrics to two unrelated factors and used the social network measure that accounted for the most variance for each principal component in further analyses. We then quantified the change in time allocated to vigilance and foraging following alarm call playback from baseline levels to the first 30 s and the second 30 s period (i.e., 31–60 s) following playback. We expected that if marmots with strong affiliative relationships felt more secure, they would forage more and allocate less time to vigilance after their immediate vigilance response to the broadcast alarm calls. Using mixed effects models that controlled for variation explained by a number of biologically important covariates and permutation tests to test the significance of social network variables, we found that marmots with a higher incloseness allocated significantly more time to vigilance in both the first and second 30 s after hearing a novel alarm call. Additionally, and while not significant (the observed parameter estimate fell between the 90 and 95% CI), marmots with a higher outstrength increased foraging in the second 30 s after hearing a novel alarm call. If we assume that time allocated to foraging is a measure of security, then marmots with strong affiliative relationships reacted more to an alarm call from a novel individual and seemingly felt more secure than their counterparts. Our results, therefore, suggest that strong social relationships increase perceptions of security and illustrate an effect of social relationships on predation risk assessment. In many species, including humans, there are benefits from maintaining good social relationships. These benefits include better health and greater longevity. We studied yellow-bellied marmots, a ground-dwelling squirrel that is notable because it has variable social relationships. Capitalizing on this social variation, we found that marmots that maintain strong affiliative relationships with other marmots may indeed forage more after hearing an artificially broadcast alarm call than marmots with weaker affiliative relationships. This result suggests that marmots that are more socially connected with others in their group may feel relatively more secure, possibly because they have others that can help them assess risk, and that a benefit of maintaining strong friendly social ties is that it permits animals to forage more. Thus, we suggest a new way that social relationships can be beneficial: they increase perceptions of security and this ultimately may facilitate foraging.

Paternity and male mating strategies of a ground squirrel ( Ictidomys parvidens ) with an extended mating season

Abstract Animal mating systems are driven by the temporal and spatial distribution of sexually receptive females. In mammals, ground-dwelling squirrels represent an ideal clade for testing predictions regarding the effects of these parameters on male reproductive strategies. While the majority of ground squirrel species have a short, highly synchronous annual breeding season that occurs immediately after females emerge from hibernation, the Mexican or Rio Grande ground squirrel ( Ictidomys parvidens ) differs markedly in having an extended mating season (2 months) and a long delay between emergence from hibernation and female receptivity (1–2 months). Both traits are expected to favor polygyny by increasing the chances that a male can secure matings with multiple females (e.g., females that come into estrus on different days). To test this prediction, we used microsatellite markers to characterize the mating system of a population of Rio Grande ground squirrels from Carlsbad, New Mexico. Our analyses indicated a high frequency of multiple paternity of litters in this population. Paternity was not related to spatial overlap between known mothers and assigned fathers, suggesting that territory defense is unlikely to be an effective male reproductive strategy in the study population. Dominance interactions among males were frequent, with heavier males typically winning dyadic interactions. Surprisingly, however, males with lower dominance scores appeared to have higher reproductive success, as did males that were active over a greater extent of the study site. Collectively, these results suggest that the mating system of the Rio Grande ground squirrel is best described as scramble competition polygyny, with the primary male reproductive strategy consisting of searching for estrous females. Similar patterns of male–male competition have been reported for a few other ground squirrel species, providing potentially important opportunities for comparative studies of the factors favoring this form of male reproductive strategy.

Climate change and the conservation of marmots

Conservation of marmots, large ground-dwelling squirrels restricted to the northern hemisphere, was impacted by direct human activity through hunting or modifying ecosystem dynamics. Regulating human activities reduced the threat of extinction. Climate change, an indirect human impact, threatens marmot survival through global warming and extreme weather events. Most marmot species occupy a harsh environment characterized by a short growing season and a long, cold season without food. Marmots cope with seasonality by hibernating. Their large size increases the efficiency of fat accumulation and its use as the sole energy source during hibernation. Marmot physiology is highly adapted to coping with low environmental temperatures; they are stressed by high heat loads. Global warming since the last ice age reduced the geographic distribution of some of the 15 species of marmots. Recent warming resulted in a movement upslope of their lower elevation boundary. This process likely will continue because warming is associated with drier unpalatable vegetation. Drought reduces reproduction and increases mortality; thus decreased summer rainfall in the montane environments where marmots live may cause local extinction. Snow cover, a major environmental factor, is essential to insulate hibernation burrows from low, stressful temperatures. However, prolonged vernal snow cover reduces reproduction and increases mortality. Montane areas currently lacking marmot populations because vernal snow cover persists beyond the time that marmots must begin foraging may become colonized if warming causes earlier snow melt. This benefit will be short-lived because decreased precipitation likely will result in unpalatable vegetation. Although some marmot populations are physiologically adapted to a warmer climate, global warming will increase too rapidly for any significant evolutionary response to dryness. The species that live in high, alpine meadows where tree and shrub invasions occur are most threatened with extinction. Captive breeding can preserve marmot species in the shortrun, but is impractical over the long-term. Widespread species are unlikely to be endangered in the foreseeable future, but local, low elevation populations will be lost.

Factors influencing burrow length and depth of ground-dwelling squirrels

We reviewed the literature on burrows of North American ground-dwelling squirrels to assess factors influencing burrow length and depth. Prior research suggested that soil characteristics and burrow age can influence burrow dimensions; we extended these findings by evaluating the influence of body size, temperature stress, and degree of sociality. We found that body size was positively correlated with burrow length and depth, probably because larger species require more space for underground activities. This relationship did not hold for Marmota monax and M. flaviventris, for unknown reasons. Burrow length scaled allometrically with body mass (b = 0.72), possibly because the metabolic effects of limited ventilation in burrows placed a constraint on burrow length. Temperature stress was not related to burrow depth, perhaps because of the insulation provided by snow and soil. The more highly social species excavated longer burrow systems, probably because of the greater space needs of multiple residents per burrow.

Conflicting research on the demography, ecology, and social behavior of Gunnison's prairie dogs (Cynomys gunnisoni)

Abstract Gunnison's prairie dogs (Cynomys gunnisoni) are rare, diurnal, colonial, burrowing, ground-dwelling squirrels. Studies of marked individuals living under natural conditions in the 1970s, 1980s, and 1990s showed that males are heavier than females throughout the year; that adult females living in the same territory are consistently close kin; and that females usually mate with the sexually mature male(s) living in the home territory. Research from 2007 through 2010 challenges all 3 of these findings. Here we discuss how different methods might have led to the discrepancies.

Stochastic population dynamics of a montane ground-dwelling squirrel.

Understanding the causes and consequences of population fluctuations is a central goal of ecology. We used demographic data from a long-term (1990–2008) study and matrix population models to investigate factors and processes influencing the dynamics and persistence of a golden-mantled ground squirrel (Callospermophilus lateralis) population, inhabiting a dynamic subalpine habitat in Colorado, USA. The overall deterministic population growth rate λ was 0.94±SE 0.05 but it varied widely over time, ranging from 0.45±0.09 in 2006 to 1.50±0.12 in 2003, and was below replacement (λ<1) for 9 out of 18 years. The stochastic population growth rate λs was 0.92, suggesting a declining population; however, the 95% CI on λs included 1.0 (0.52–1.60). Stochastic elasticity analysis showed that survival of adult females, followed by survival of juvenile females and litter size, were potentially the most influential vital rates; analysis of life table response experiments revealed that the same three life history variables made the largest contributions to year-to year changes in λ. Population viability analysis revealed that, when the influences of density dependence and immigration were not considered, the population had a high (close to 1.0 in 50 years) probability of extinction. However, probability of extinction declined to as low as zero when density dependence and immigration were considered. Destabilizing effects of stochastic forces were counteracted by regulating effects of density dependence and rescue effects of immigration, which allowed our study population to bounce back from low densities and prevented extinction. These results suggest that dynamics and persistence of our study population are determined synergistically by density-dependence, stochastic forces, and immigration.

Sociality, Bateman’s gradients, and the polygynandrous genetic mating system of round-tailed ground squirrels (Xerospermophilus tereticaudus)

Historically, most mammals have been classified as polygynous; although recent molecular evidence suggests that many mammals may be polygynandrous, particularly the ground-dwelling sciurids. We genotyped 351 round-tailed ground squirrels (Xerospermophilus tereticaudus) using seven microsatellite loci to determine paternity in 31 litters from 2004 to 2007. Polygyny was evident in all years except in 2007, when the population size was reduced. Multiple paternity occurred in the majority of litters (55%) with 2.5 ± 0.26 sires/litter (n = 31). Forty-nine percent of resident males (n = 114) sired offspring, and of males that sired offspring (n = 56) 27% sired young in multiple litters in a single breeding season. Litter size was positively correlated with the number of sires. Through an indirect analysis of paternity, we found 21 litters (68%) with an average relatedness of 0.5 or less. Males had a greater opportunity for sexual selection (I s = 1.60) than females (I s = 0.40); Bateman’s gradient was also greater in males (1.07 ± 0.04, n = 56) than females (0.82 ± 0.08, n = 31). The mating system in round-tailed ground squirrels defined through genetic analyses and Bateman’s gradients is polygynandrous compared to the previously suggested polygynous mating system as established by behavioral observations and fits within the predictions of the ground squirrel sociality models. Upon evaluating the predictions of the sociality models among sciurid species, we found a negative relationship between the level of sociality with litter size and the average percentage of multiple paternity within a litter. Thus, recent genetic information and reclassification of mating systems support the predictions of the ground-dwelling squirrel sociality models.

Comparison of the ITS1 and ITS2 rDNA in Eimeria callospermophili (Apicomplexa: Eimeriidae) from Sciurid Rodents

The taxonomy of the coccidia has historically been morphologically based. The purpose of this study was to establish if conspecificity of isolates of Eimeria callospermophili from 4 ground-dwelling squirrel hosts (Rodentia: Sciuridae) is supported by comparison of rDNA sequence data and to examine how this species relates to eimerian species from other sciurid hosts. Eimeria callospermophili was isolated from 4 wild-caught hosts, i.e., Urocitellus elegans, Cynomys leucurus, Marmota flaviventris , and Cynomys ludovicianus . The ITS1 and ITS2 genomic rDNA sequences were PCR generated, sequenced, and analyzed. The highest intraspecific pairwise distance values of 6.0% in ITS1 and 7.1% in ITS2 were observed in C. leucurus. Interspecific pairwise distance values > 5% do not support E. callospermophili conspecificity. Generated E. callospermophili sequences were compared to Eimeria lancasterensis from Sciurus niger and Sciurus niger cinereus and to Eimeria ontarioensis from S. niger. A single, well-supported clade was formed by E. callospermophili amplicons in neighbor joining and maximum parsimony analyses. However, within the clade, there was little evidence of host or geographic structuring of the species.

Natal dispersal of introduced Siberian chipmunks, Tamias sibiricus, in a suburban forest

The Siberian chipmunk, Tamias sibiricus, was introduced into the suburban forest of Senart in the late 1970s where it naturalised. Juvenile dispersal of this solitary ground-dwelling squirrel is not known, while it is an important component of the invasion process. From 2003 to 2007, mark–recapture techniques were used to determine how dispersal movements differed between year, cohort, sex and body mass. Distances moved were calculated between the location of first capture during the year of birth and the arithmetic centre of the trapping locations obtained the next year. Excluding heaviest individuals, e.g. 80 g, that must have already dispersed and thus lowered the estimation of averaged distance moved, our results shows that the dispersal distance in juvenile males (168 ± 24 m, n = 39) was greater than in females (83 ± 11 m, n = 66), with no effect detected for either year or cohort. Juveniles are thus involved in the spread of this ground squirrel, bearing in mind that movements estimated may have been underestimated due to the use of capture–mark–recapture methods on a limited trapping area.

Odontometric Patterns in the Radiation of Extant Ground-dwelling Squirrels within Marmotini (Sciuridae: Xerini)

I document odontometric variation across ground-dwelling squirrels of the Holarctic tribe Marmotini. Dental size, which correlates well with published average body mass values across species, accounts for most odontometric variation across the clade. Dental shape variation primarily reflects relative size of P3 (upper cheek teeth) and relative width of p4–m1 + length of m3 (lower cheek teeth). Shape variables and relative tooth crown height covary significantly across species, suggesting a common functional complex or shared genetic control. When dental morphology is mapped on published DNA-based phylogenies, Sciurotamias (Chinese rock squirrels), Ammospermophilus (antelope squirrels), and basal subgenera within Spermophilus (ground squirrels [Callospermophilus and Otospermophilus]) are shown to retain inferred primitive dental morphology—small to moderate dental size, relatively small P3, relatively narrow p4–m1 + shortened m3, and relatively low toothcrown height. Other clades depart from this morphotype in size (very small in Tamias [chipmunks] and very large in Marmota [marmots]), tooth shape (especially 2 clades representing Eurasian and North American subgenus Spermophilus), or in both attributes (notably Cynomys [prairie dogs]), with frequent homoplasy. A plot of odontometric distance against published estimates of divergence time between sister clades suggests a roughly ‘‘clocklike’’ accumulation of odontometric change through time but highlights episodes of rapid odontometric evolution during the origins of Marmota, Cynomys, and Spermophilus parryii (arctic ground squirrel).

Effects of Tourists on Behavior and Demography of Olympic Marmots

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.

PARENTAGE, MULTIPLE PATERNITY, AND BREEDING SUCCESS IN GUNNISON'S AND UTAH PRAIRIE DOGS

Feasibility of assigning parentage using variable microsatellite loci was assessed for 2 species of prairie dogs. Parentage was determined from 7 microsatellite loci for 46% of juveniles born during 1994 in a colony of Gunnison's prairie dogs (Cynomys gunnisoni), and for 53% and for 45% of juveniles born during 1996 and 1997, respectively, in a colony of Utah prairie dogs (C. parvidens). Frequency of multiple paternity estimated for Gunnison's (77%) and Utah (71% and 90%) prairie dogs was greater than that detected previously for black-tailed prairie dogs (5%–10%) but within the range reported for other ground-dwelling squirrels. Of the 84 adult females and 33 adult males present during 1994 in the colony of Gunnison's prairie dogs, 75 (89%) and 22 (67%), respectively, produced weaned offspring. Breeding success for Utah prairie dogs was relatively low in 1996 (45% for females and 32% for males) but increased in 1997 (80% for females and 81% for males).