Marmota Vancouverensis Research Articles

Molecular data resolve placement of the Olympic marmot and estimate dates of trans-Beringian interchange

Abstract 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...

Nasal Osteosarcoma and Interstitial Cell Tumor in a Vancouver Island Marmot (Marmota vancouverensis)

A 6-yr-old male Vancouver Island marmot (Marmota vancouverensis) presented for poor hibernation, weight loss, and symmetric trunk alopecia. An abdominal interstitial cell tumor was identified and surgically removed. Serum levels of estrogen were markedly elevated before surgery and decreased after tumor removal, indicating that the tumor had been functionally secretory. Nine months later, the marmot presented with respiratory stridor. A large boney nasal mass was identified radiographically and evaluated by computed tomography (CT) prior to surgical debulking. The marmot did not recover from anesthesia. Pathologic findings included a nasal osteosarcoma with lysis of the cribriform plate, and endocardial fibrosis with degenerative changes within the adjoining myocardium. This is the first known report of nasal osteosarcoma and interstitial tumor in a Vancouver Island marmot.

The reproductive endocrinology and behavior of Vancouver Island marmot (Marmota vancouverensis)

The Vancouver Island marmot (Marmota vancouverensis; VIM) is one of North America's most endangered species with fewer than 150 individuals remaining in the wild. A captive breeding program was established across four facilities in Canada as an insurance population and source of animals for reintroduction to the wild. The purpose of this study was to gather information about the basic reproductive biology and behavior of this species, which is essential to improve captive breeding programs. Regular fecal samples were obtained from adult female (n = 14) and male (n = 10) marmots, 2 years of age and older, over 1-3 breeding seasons (2-3 months duration posthibernation) for steroid hormone analysis. Enzyme immunoassays were validated for quantifying fecal testosterone metabolite concentrations for males, and fecal estrogen and progesterone metabolite concentrations for females. Results indicated that fecal progesterone metabolite concentrations can be used to monitor ovulation and pregnancy. Behavioral monitoring through infrared video surveillance was conducted in four breeding pairs over a 2-year period (n = 7 behavioral profiles). Breeding behaviors correlated strongly with changes in reproductive endocrine profiles. A high frequency of play behavior or "wrestling" was observed in conjunction with breeding activity before an elevation in progesterone metabolite concentrations. Impending parturition was associated with increased aggression and exclusion of the male from the maternal nestbox as well as an increase in nesting activity. Observational data combined with hormonal analysis suggest that female VIMs are induced ovulators and that multiple breeding attempts may be required for ovulation and conception. Gestation appears to be approximately 34 days from peak breeding activity (32 days from estimated ovulation). Fecal testosterone concentrations suggest that testicular activity is seasonal with the reproductive activity occurring immediately posthibernation. Monitoring breeding behavior is a useful means of indicating estrus, conception and pregnancy, which can also be supported by the hormonal analysis of daily fecal samples of individual animals.

Social ‘meltdown’ in the demise of an island endemic: Allee effects and the Vancouver Island marmot

1. More than 75 years have passed since W.C. Allee proposed that breakdowns in sociality may shift animal populations to inverse density dependence at small sizes and thereby hasten spirals to extinction. Despite decades of attention, empirical evidence of this 'Allee effect' in wild populations remains scarce. 2. Here, we report on findings from a multi-year study of the population ecology and behaviour of the critically endangered Vancouver Island marmot (Marmota vancouverensis) and present quantitative evidence of an Allee effect and highlight the mechanisms that drive it. 3. The V.I. marmot is a large, social rodent endemic to Vancouver Island, Canada, and its population has declined by 80-90% since the 1980s. The species currently is represented in the wild by roughly 200 individuals. 4. This study compared characteristics of contemporary V.I. marmots (2002-2005) with (i) animals in the same population at an earlier time period (1973-1975) and (ii) congeners. Specifically, data on time allocation, social activity and ranging behaviour of animals in colonies in the late stages of decline were compared with historical data collected from colonies under more stable demographic conditions. 5. We found that contemporary V.I. marmots had home ranges that were 10-60x larger than historic animals and congeners, interacted with conspecifics at 10% of the historic rate, devoted 10x more time to anti-predator vigilance, and abandoned the bi-modal activity patterns previously described for this and other marmot species. Contemporary marmots also showed an 86% decline in feeding rate, and entered hibernation on average 20 days later than animals in historic populations. 6. Combined with results showing reduced per capita survival and reproduction in contemporary marmots, these findings suggest a strong role for Allee effects in the current plight of the Vancouver Island marmot. A positive link between aspects of fitness and population size emphasizes the need to identify threshold colony sizes and densities necessary to promote recovery. We discuss this and other implications of this species' social 'meltdown'.

Detecting biological distinctiveness using geometric morphometrics: an example case from the Vancouver Island marmot

The Vancouver Island marmot (M. vancouverensis) consists of a small insular population with an uncommon dark-brown pelage colour. This population provides an excellent example of how morphology and molecules can tell very different stories about the evolution of a group. Mitochondrial DNA indicates that the degree of divergence of M. vancouverensis is minimal and actually comparable or even smaller than among populations of continental species of marmots. Accurate geometric morphometric analyses on crania and mandibles from modern and subfossil samples, in contrast, convincingly demonstrate that the Vancouver Island marmot is one of the most distinctive populations among all living marmots. Thus, M. vancouverensis is not simply a dark variant of its sister species M. caligata but represents a highly morphologically derived population whose evolutionary significance is overlooked by molecular analyses. In this study, we show how geometric morphometrics can complement studies of genetic divergence by measuring the phenotypic dis- tinctiveness of a population. This was done by performing cross-validated discriminant analyses on mandibular shape in the Vancouver Island marmot and its closest Nearctic relatives. Thus, we found that using mandibles we can achieve in M. vancouverensis a classification accuracy as high as for ‘good species’ (i.e., species which are also significantly divergent in molecular analyses) and much higher than in populations within those same species. The approach used in this study is simple, effective and, at least in this specific case, fairly robust to sampling error in small samples. Thus, such morphological analysis seems promising as a tool complementary to molecular studies to aid detection of biological distinctiveness in natural populations. We suggest that its potential should be explored and its usefulness examined also in Palearctic marmots and other mammals. If successful, the ‘geometric morphometric approach to the study of biological diversity‘ would add a phenotypic dimension to the characterization of the evolutionary significance of populations. In turn, this would allow a more inclusive understanding of the complexity of adaptive variation with potential implications for conservation biology.

Reintroducing endangered Vancouver Island marmots: Survival and cause-specific mortality rates of captive-born versus wild-born individuals

Recovery of the endangered Vancouver Island marmot ( Marmota vancouverensis) is contingent upon releases of captive-born marmots into natural habitats. Success of such re-introduction programs largely depends on the ability of released animals to survive in the wild. However, whether and to what extent survival and cause-specific mortality rates of captive-born marmots differ from those of their wild-born counterparts remains unknown. We used radio-telemetry (1992–2007) and mark-resighting (1987–2007) data to estimate seasonal and annual survival rates of the Vancouver Island marmot, to compare survival and cause-specific mortality rates of captive-born marmots that have been released into the natural habitat with those of wild-born marmots, and to test for the effect of age-at-release on survival of the released marmots. Analysis of radio-telemetry data suggested no difference in survival of males versus females. However, annual survival of captive-born marmots released into the wild was low ( S = 0.605; 95% CI = 0.507–0.696) compared to wild-born marmots ( S = 0.854; 95% CI = 0.760–0.915). Marmots released as 2-year-old or older survived more successfully than those released as yearlings. Additional forensic evidence reinforced the idea that predation was the most important cause of mortality. Causes of death differed significantly between captive-born and wild-born marmots. Predation by golden eagles ( Aquila chrysaetus) was the most important cause of mortality for captive-born marmots, whereas predation by wolves ( Canis lupus) and cougars ( Felis concolor) was more important for wild-born marmots. Age-specific apparent annual survival rate, estimated using the combined mark-resighting and radio-telemetry data, was lowest for pups ( S = 0.500; 95% CI = 0.375–0.616) and highest for yearlings and adults ( S = 0.656; 95% CI = 0.604–0.705); and apparent survival of 2-year-old was similar to that of yearlings and adults ( S = 0.649; 95% CI = 0.527–0.754). Our results, based on the analysis of radio-telemetry data, suggest that delaying release of captive-born marmots until 2 years of age may enhance their probability of survival in the wild, and will likely improve the success of the release program.

Tempo and mode of evolutionary divergence in modern and Holocene Vancouver Island marmots (Marmota vancouverensis) (Mammalia, Rodentia)

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.

Patterns of within and between-colony microsatellite variation in the endangered Vancouver Island marmot (Marmota vancouverensis): implications for conservation

Among the 14 extant species of the genus Marmota the Vancouver Island marmot (Marmota vancouverensis) is the most endangered. In 2007 as few as 85 individuals were left in the wild, with an additional 162 individuals maintained in captivity. To facilitate genetic monitoring of both wild and captive populations, polymorphic genetic markers were identified. Thirty-three different microsatellite loci were tested for amplification and variability in ≥30 wild-born individuals. Only 11 of these loci proved to be polymorphic and were subsequently analysed in 105 samples collected from wild Vancouver Island marmots. The average number of alleles (A) at those 11 loci was only 2.1, and the intraspecific variation (HE between 8 and 23% within colonies) was low compared to other marmot species. Variation within the small and geographically isolated Mt. Washington colony was particularly low (A = 1.3, HE = 0.08). Genetic distances between the Mt. Washington colony (11 individuals) and those of the Nanaimo Lakes region (94 individuals) on southern Vancouver Island were large (D values ranging from 0.42 to 0.50), while genetic distances among colonies within the latter area were much smaller (D values from 0.01 to 0.13). Given the low within-population genetic variation, and the resulting risk of inbreeding depression at Mt. Washington, we support the decision to maximize overall genetic variation of the species by crossbreeding marmots from the two different areas despite the possibility of local adaptation.

The case of the missing marmots: Are metapopulation dynamics or range-wide declines responsible?

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.

Factors Influencing Reproduction in Captive Vancouver Island Marmots: Implications for Captive Breeding and Reintroduction Programs

Many factors important to reproduction are difficult to quantify for wild mammals, yet an understanding of them is often critical to species' recovery programs, particularly those involving captive breeding and reintroduction. We examined management variables employed by the Vancouver Island marmot captive breeding program during 1998–2005 to determine how such variables influenced production of young and litter sizes. We then tested the ability of factors that were identified as important to predict production of young in the breeding program in 2006. For previously paired animals, production of young was significantly greater when they had been paired with the same mate for ≥1 year before the breeding season. When these females had been with a mate for < 1 year, production of young was more than 2 times greater among 2–4 year olds compared to older age classes. Among previously paired females, 5–7 year olds and those that had produced young previously tended to produce young more often than other categories. Litter size was positively influenced by visual contact between a female and other pairs. For previously paired females, age, the amount of time paired with their mate, and previous production of young accurately predicted 75% of production of young in 2006. We propose that results from our study be combined with genetic considerations to plan future pairings in the breeding program and to assist in the selection of individuals for reintroduction to the wild. Methods reported herein may be applicable to recovery programs for other imperiled mammals.

Evolutionary acceleration in the most endangered mammal of Canada: speciation and divergence in the Vancouver Island marmot (Rodentia, Sciuridae)

The Vancouver Island marmot is the most endangered mammal of Canada. Factors which have brought this population to the verge of extinction have not yet been fully elucidated, but the effects of deforestation and habitat fragmentation on survival rates, as well as those of variation in rainfall, temperature, snowpack depth and snowmelt strongly suggest that marmots on the island are struggling to keep pace with environmental changes. Genetic analyses, however, seem to indicate that the Vancouver Island marmot may merely represent a melanistic population of its parental species on the mainland. Were it not for its black pelage colour, it is unlikely that it would have attracted much attention as a conservation priority. Our study uses three-dimensional coordinates of cranial landmarks to further assess phenotypic differentiation of the Vancouver Island marmot. A pattern of strong interspecific divergence and low intraspecific variation was found which is consistent with aspects of drift-driven models of speciation. However, the magnitude of shape differences relative to the putatively neutral substitutions in synonymous sites of cytochrome b is too large for being compatible with a simple neutral model. A combination of bottlenecks and selective pressures due to natural and human-induced changes in the environment may offer a parsimonious explanation for the large phenotypic differentiation observed in the species. Our study exemplifies the usefulness of a multidisciplinary approach to the study of biological diversity for a better understanding of evolutionary models and to discover aspects of diversity that may be undetected by using only a few genetic markers to characterize population divergence and uniqueness.

Predator discrimination and ‘personality’ in captive Vancouver Island marmots (Marmota vancouverensis)

AbstractA major impediment to recovering declining populations successfully is the mortality of reintroduced or translocated animals. We generally assume that captive‐born animals may lose their antipredator behaviour abilities in captivity, but studies rarely compare predator recognition abilities of captive‐born and wild‐captured animals to test this. To identify whether predator discrimination abilities of the critically endangered Vancouver Island marmots Marmota vancouverensis were lost in captivity, we presented wild‐captured and captive‐born marmots with taxidermic mounts of predators (a cougar Felis concolor and wolf Canis lupus) together with control stimuli (marmot, domestic goat Capra aegagrus, the cart on which all stimuli were presented and a ‘blank’ no‐stimulus control). Regardless of specific predator discrimination abilities, for some species overall ‘personality’ may be associated with response to predators and subsequent survival. Thus, to quantify overall reactivity in the presence of a predator, we also conducted a mirror‐image stimulus (MIS) presentation experiment where marmots were video‐recorded with or without the presence of a wolf. Marmots discriminated among these stimuli, responding the most to the wolf and cougar. The MIS results suggest that marmots varied along a continuum of reactivity. The amount of reactivity was unaffected by the presence of a wolf, and was correlated with our highest level of responsiveness (vigilance at the burrow and time within the burrow) to the wolf. Taken together, we conclude that marmots differentiate predators from non‐predators and that this ability has not been lost under the conditions in which they have been reared.

Reproductive rates of wild and captive Vancouver Island marmots (Marmota vancouverensis)

I evaluated reproductive rates of the critically endangered Vancouver Island marmot (Marmota vancouverensis Swarth, 1911) using data from captive and wild populations over the 1980–2004 period. Results were similar to those reported for other alpine-dwelling marmots, including the closely related Marmota caligata (Eschscholtz, 1829) and Marmota olympus (Merriam, 1898). Most females in captivity first bred at age 3 or 4 (Tbreed = 4.3 years, SD = 1.15, n = 9), an age not significantly different from that observed in the wild (Tbreed = 3.6 years, SD = 1.2, n = 16). Numbers of pups weaned per litter were similar in captivity (Npups = 3.0, SD = 1.4, n = 25) and in the wild (Npups = 3.4, SD = 1.1, n = 58). Females were capable of weaning pups in consecutive years (46.4%, n = 13) but often skipped 1 year (39.3%, n = 11) or 2 years (14.3%, n = 4) between litters. Two-year-old females weaned pups infrequently (Pbreed = 0.09, n = 43) and older females were far more likely to breed (Pbreed = 0.40, n = 200); in neither case were significant captive-wild differences found. The oldest breeding female was 10 years old, but sample sizes for marmots older than 8 years were small and maximum breeding age may be underestimated. Between-litter intervals in captivity (Tbetween = 1.4 years, SD = 0.7, n = 11) were significantly shorter than in the wild (Tbetween = 1.9 years, SD = 0.7, n = 17). Sex ratios of weaned pups did not differ from 1:1 in the wild (female/male = 1.04) but were significantly skewed towards males in captivity (female/male = 0.56). I conclude that reproductive performance in Vancouver Island marmots is limited both by body condition and social constraints.

Timing and causes of mortality in the endangered Vancouver Island marmot (Marmota vancouverensis)

We used radiotelemetry to evaluate seasonal survival rates and mortality factors for a critically endangered island endemic, the Vancouver Island marmot (Marmota vancouverensis Swarth, 1911). Recovery of radio transmitters and marmot remains suggested that predation was the major cause of mortality, accounting for at least 24 of 29 (83%) known-fate deaths recorded since radiotelemetry efforts began in 1992. Wolves (Canis lupus L., 1758) and cougars (Puma concolor (L., 1771)) apparently accounted for 17 deaths (59%). Three marmots (10%) were killed by golden eagles (Aquila chrysaetos (L., 1758)), four (14%) were killed by unknown predators that probably included all of the above species, two (7%) died from unknown causes, and three (10%) died during hibernation in a single burrow. Mortality rates varied seasonally. The daily probability of death during hibernation was very low (Pdeath = 0.016). The probability of death was also low from spring emergence through 31 July (Pdeath = 0.051), but was eight times higher in August (Pdeath = 0.395) and four times higher in September (Pdeath = 0.175). We concluded that predation was the proximate cause of recent declines in wild Vancouver Island marmot populations, that losses were highly concentrated in late summer, and that previous studies exaggerated the importance of winter mortality. We suggest that high predation rates were associated with forestry and altered predator abundance and hunting patterns.

Patterns of morphological evolution in Marmota (Rodentia, Sciuridae): geometric morphometrics of the cranium in the context of marmot phylogeny, ecology and conservation

Marmots are of great interest for both sociobiologists studying the evolution of mammal societies and conservationists trying to protect them from extinction. In contrast, their phylogeny and morphological evolution are poorly understood and studied. Recently, a phylogenetic analysis using cytochrome b provided the first reconstruction of marmot evolutionary history and suggested that a high level of sociality evolved at least twice independently in the two proposed marmot subgenera. A morphological analysis of the marmot mandible supported this subgeneric classification and showed interesting, and unexpected, patterns in the evolution of marmot skeletal characters. In the present study we investigated a more complex, and potentially informative structure, the ventral cranium. Geometric morphometric techniques were applied in the first analysis of cranial morphology including all marmot species. Three main phenetic groups were found, which reflect phylogeny (subgenus Petromarmota, and Palaearctic subgenus Marmota) or geographical distribution (Palaearctic vs. Nearctic subgenus Marmota). Convergence in skeletal characters due to size similarities, a common finding in the sciurid skeleton according to traditional morphological analyses, did not occur in the marmot ventral cranium. Despite a genetic distance between Marmota vancouverensis and Marmota caligata similar to that among different populations of the latter species, the Vancouver Island marmot had the most atypical ventral cranium in the subgenus Petromarmota. This finding confirmed results obtained with the mandible, and emphasized the uniqueness of M. vancouverensis and the usefulness of complementing molecular analyses with morphological studies for a thorough characterization of population divergence, and a careful planning of conservation strategies.

Comparison of discriminant function and classification tree analyses for age classification of marmots

We evaluated the predictive power of two classification techniques, one parametric – discriminant function analysis (DFA) and the other non‐parametric – classification and regression tree analysis (CART), in order to provide a non‐subjective quantitative method of determining age class in Vancouver Island marmots (Marmota vancouverensis) and hoary marmots (Marmota caligata). For both techniques we used morphological measurements of known‐age male and female marmots from two independent population studies to build and test predictive models of age class. Both techniques had high predictive power (69–86%) for both sexes and both species. Overall, the two methods performed identically with 81% correct classification. DFA was marginally better at discriminating among older more challenging age classes compared to CART. However, in our test samples, cases with missing values in any of the discriminant variables were deleted and hence unclassified by DFA, whereas CART used values from closely correlated variables to substitute for the missing values. Therefore, overall, CART performed better (CART 81% vs DFA 76%) because of its ability to classify incomplete cases. Correct classification rates were approximately 10% higher for hoary marmots than for Vancouver Island marmots, a result that could be attributed to different sets of morphological measurements. Zygomatic arch breadth measured in hoary marmots was the most important predictor of age class in both sexes using both classification techniques. We recommend that CART analysis be performed on data‐sets with incomplete records and used as a variable screening tool prior to DFA on more complete data‐sets.

Anti‐Predator Behavior of Vancouver Island Marmots: Using Congeners to Evaluate Abilities of a Critically Endangered Mammal

Behavioral comparisons between endangered species and their congeners may provide valuable data with which to test ideas about declining populations or the future direction of recovery efforts. We considered the case of the highly endangered Vancouver Island marmot (Marmota vancouverensis). Predation is a current source of mortality, and inadequate anti‐predator behavior could have profound ramifications for the future success of re‐introductions. We tested whether M. vancouverensis anti‐predator behavior was unusual or ‘deficient’ by quantifying it and comparing it to 13 other marmot species. We found no evidence that Vancouver Island marmots were unwary. If anything, the converse was true. Vancouver Island marmots were responsive and vigilant towards real and simulated predatory threats. They dug numerous escape burrows that reduced the likelihood of predation. Our results have several implications for future recovery efforts, one of which was to establish ‘baseline’ flight‐response targets that captive‐bred Vancouver Island marmots will have to meet or exceed prior to release into predator‐rich environments.

Serum fatty acid analysis and digestibility study in the Vancouver Island marmot (Marmota vancouverensis) fed a captive diet supplemented with α‐linolenic acid

AbstractMammals that hibernate must rely on endogenous lipid reserves to survive over winter. This study was conducted to compare the difference in serum fatty acid composition, dietary intake, and apparent digestibility in the Vancouver Island marmot (N = 6) fed the Metro Zoo lagomorph diet supplemented with α‐linolenic acid [C(18:3) n‐3 (α‐LA)]. The experiment was designed as a 3 × 3 Latin square with three 17‐day collection periods. The test diets contained 12.16, 14.85, and 17.05% α‐LA as a percentage of fatty acids in the diet supplied through the addition of flaxseed oil (∼53% α‐LA). Across treatments, dry matter intake (g/d), dry matter digestibility, apparent fat digestibility, and apparent neutral detergent fiber digestibility did not differ significantly (P &gt; 0.05). There were no significant differences in serum α‐LA concentration between the three levels of α‐LA supplementation. However, this supplementation did elevate serum α‐LA, eicosapentaenoic acid [C20:5 (n‐3)], and docosahexaenoic acid [C22:6 (n‐3)] levels compared with feeding the basal zoo lagomorph diet (P &lt; 0.05). Thus, supplementation of the basal zoo lagomorph diet with α‐LA elevated the serum levels of essential fatty acids in the Vancouver Island marmot. Zoo Biol 20:251–259, 2001. © 2001 Wiley‐Liss, Inc.

ALARM CALLING IN THREE SPECIES OF MARMOTS

AbstractMany 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.

Distribution and abundance of Vancouver Island marmots (Marmota vancouverensis)

We analysed historic records and annual counts to assess the population status of Vancouver Island marmots (Marmota vancouverensis). Since 1972, marmots have been found at 47 sites on 15 mountains. All but 2 colonies were located within 5 adjacent watersheds on south-central Vancouver Island. Counts underestimated actual marmot abundance. For most site–year combinations, observers probably counted 66–78% of adults and 75–89% of juveniles. Reproductive colonies typically contained fewer than 5 adults ([Formula: see text], SE = 0.61, n = 34). Most animals were found at elevations above 1000 m (81%), on south- to west-facing slopes (74%). After 1981, marmots colonized 11 habitats created by logging of forests above 700 m. Numbers of adults were above average (134–147%) during the early 1980s and have been near or below average since 1990 (58–99%). The current (1995) population contains 100–200 animals, including 50–100 animals in logged habitats. Marmota vancouverensis is rare primarily because of the small size and patchy distribution of natural subalpine meadows on Vancouver Island. The species is apparently adapted to a metapopulation life-style, in which a network of small colonies exhibit population fluctuations, local extinctions and recolonizations over time.