Marginal habitats provide unexpected survival benefits to the alpine marmot

Age-specific survival trajectories can vary significantly among wild populations. Identifying the environmental conditions associated with such variability is of primary importance to understand the dynamics of free-ranging populations. In this study, we investigated survival variations among alpine marmot (Marmota marmota) families living in areas with opposite environmental characteristics: the typical habitat of the species (alpine meadow) and a marginal area bordering the forest. We used data collected during an 11-year study in the Gran Paradiso National Park (Italy) and performed a Bayesian survival trajectory analysis on marked individuals. Furthermore, we investigated, at a territorial level, the relationships among demographic parameters and habitat variables by using a path analysis approach. Contrary to our expectations, for most of the marmot's lifespan, survival rate was higher in the marginal site closer to the forest and with lower visibility than in the alpine meadow site. Path analysis indicated that the number of families living close to each other negatively affected the stability of the dominant couple, which in turn affected both juvenile survival and reproduction. Given the lower number of neighboring families which inhabited the marginal site and the potentially different predation pressure by the most effective predator in the area (Aquila chrysaetos), our results suggest that species adapted to live in open habitats may benefit from living in a marginal habitat. This study highlights the importance of habitats bordering the forest in the conservation of alpine marmots.     

The successful introduction of the alpine marmot Marmota marmota in the Pyrenees,Iberian Peninsula,Western Europe

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Genetic characterization of four wild species of Chinese marmots using microsatellite markers

Marmots are large ground squirrels, and 14 species have been reported in the world, including four species of marmots (Himalayan marmot, Tarbagan marmot, gray marmot and long-tailed marmot) living in China. Although these biological resources are abundant in China, information regarding their genetic features is lacking, hampering further study regarding them. The aims of this research were to evaluate genetic variations of four species of Chinese wild marmots, and analyzed kinship of these marmot populations. In the current study, we collected samples of four species of Chinese wild marmot and analyzed the effective allele number, gene diversity, the Shannon index, and polymorphism information to evaluate genetic variations using 13 microsatellite loci. Based on Nei’s genetic distance using the unweighted pair group method, we constructed a dendrogram to analyze the population kinship. We determined that all four Chinese marmot species had high genetic polymorphisms and departure from Hardy-Weinberg equilibrium. The Chinese marmots to be divided into two large groups: Himalayan marmot was independent group. Tarbagan marmot, gray marmot and long-tailed marmot were others; Tarbagan marmot and gray marmot showed a close kinship with each other, but long-tailed marmot did not have a close relationship with the other species. The high polymorphisms and the kinship of Chinese marmot populations were correlated with geographical terrain of their habitat. Himalayan marmot was characterized as living in unique alpine meadows in Qinghai-Tibet plateau and was affected by terrain; however, Tarbagan marmot, gray marmot and long-tailed marmot were characterized as living in grassland or alpine grassland and were not affected by terrain. Genetic features of Chinese wild marmots were investigated in this study. This may give using information regarding protection of Chinese wild marmot resource and further application of biomedical research.     

Spatial distribution of <Emphasis Type="Italic">Marmota baibacina</Emphasis> and <Emphasis Type="Italic">M. sibirica</Emphasis> (<Emphasis Type="Italic">Marmota</Emphasis>, Sciuridae,Rodentia) in a zone of sympatry in Mongolian Altai: Bioacoustic analysis

The spatial distribution of two marmot species Marmota baibacina and M. sibirica in a zone of coexistence was studied by using their alarm call as a diagnostic trait. It was found that M. baibacina prefers to inhabit bouldery screes, whereas M. sibirica inhabits all suitable biotopes. The difference in biotopic distribution of these species could be explained by M. sibirica forcing M. baibacina out of optimum habitats. Cases of coexistence of both species in one family group sites were registered, which might contribute to the appearance of hybrids.     

Phylogeny and systematics of marmots (<Emphasis Type="Italic">Marmota</Emphasis>, Sciuridae,Rodentia) inferred from inter-SINE PCR data

Phylogenetic and taxonomic relationships in the genus Marmota were examined using inter-SINE PCR. The primers used were complementary to the consensus sequences of two short retroposons, MIR and B1-dID. The results suggest long-term genetic isolation of Nearctic and Palearctic marmots, but do not support subgeneric subdivision because of relatively low genetic differences between the marmot groups. Confirmation was received for the isolation of bobak and camtschatica, but not the caudata intrageneric species groups. Based on comparison of the mitochondrial and nuclear genome differences, the possibility of ancient hybridization between M. menzbieri and M. caudata was recognized. Species independence of M. kastschenkoi within the suggested superspecies of M. baibacina was supported.     

Inter- and Intraspecific Variation in the Acoustic Habitats of Three Marmot Species

Closely related species often have remarkably different vocalizations. Some of the variation in acoustic structure may result from species adapting their calls to maximize transmission through their acoustic environments. We document the relative magnitude of inter- and intraspecific variation in acoustic transmission properties of the habitats of three closely related marmot species to study the relative importance that the acoustic environment may have played in selecting for species-specific marmot alarm calls. We used spectrogram correlation to quantify the degree to which pure tones and alarm calls changed as they were broadcast through marmot home ranges to describe the acoustic habitats of golden (M. candata aurea), yellow-bellied (M. flaviventris), and alpine (M. marmota L.) marmots. Species lived in quantifiably different acoustic habitats. One analysis partitioned variation between species and between marmot social groups (nested within species). We found significant interspecific variation in the acoustic transmission fidelity of the three species' habitats and insignificant intraspecific variation between social groups. Further analysis of a larger sample of alarm calls broadcast through golden marmot social groups found significant intraspecific variation. Interspecific variation greater than intraspecific variation suggests that variable acoustic habitats may be responsible for at least some of the interspecific variation in alarm call structure. This is the first study to use spectrogram correlation to describe habitat acoustics. We discuss aspects of the method that may be useful for others seeking to quantify habitat acoustics.     

Rewarming rates of two large hibernators: Comparison of a monotreme and a eutherian

We measured body temperatures in two large hibernating mammals, the eutherian alpine marmot (Marmota marmota) and the egg-laying echidna (Tachyglossus aculeatus) from unrestrained animals in their natural environment. In both species hibernation is broken every 13 days on average by rewarming to euthermic temperatures. We found that the time course of a rewarming could be closely fitted with a sigmoid curve, allowing calculation of peak rewarming rate and corresponding body temperature. Maximum rewarming rates were twice as high in marmots as in echidnas (12.1±1.3 °C h⁻¹, n=10 cf. 6.2±1.2 °C h⁻¹, n=10). Peak rewarming rates were positively correlated with body temperature in echidnas, but negatively correlated in marmots.     

Winter weather versus group thermoregulation: what determines survival in hibernating mammals?

For socially hibernating mammals, the effectiveness of huddling as a means of energy conservation should increase with group size. However, group size has only been linked to increased survival in a few hibernating species, and the relative importance of social structure versus winter conditions during hibernation remains uncertain. We studied the influence of winter weather conditions, social group composition, age-structure, and other environmental factors and individual attributes on the overwinter survival of hoary marmots (Marmota caligata) in the Yukon Territory, Canada. Juvenile hoary marmot survival was negatively correlated with the mean winter (November to May) Pacific Decadal Oscillation (PDO) index. Survival in older age-classes was negatively correlated with PDO lagged by 1 year. Social group size and structure were weakly correlated with survival in comparison to PDO. The relationship between winter PDO and survival was most likely due to the importance of snowpack as insulation during hibernation. The apparent response of hoary marmots to changing winter conditions contrasted sharply with those of other marmot species and other mammalian alpine herbivores. In conclusion, the severity of winter weather may constrain the effectiveness of group thermoregulation in socially hibernating mammals.     

Quantum Speciation of <Emphasis Type="Italic">Yersinia pestis</Emphasis> Plague Microbe in a Heteroimmune Environment: In the Populations of Hibernating Tarbagan Marmots (<Emphasis Type="Italic">Marmota sibirica</Emphasis>)

This review is devoted to the ecological mechanism for the transformation of the population of the psychrophilic saprozoobiont pseudotuberculosis microbe Yersinia pseudotuberculosis O:1b into the population of the obligate blood plague microbe Yersinia pestis in the host–parasite system of the marmot flea (Marmota sibirica–Oropsylla silantiewi). The correspondence of this mechanism to the evolutionary principle of quantum speciation has been shown. The most significant population–genetic microbial transformations took place in the populations of hibernating marmots. The main factor of rapid speciation was the heterothermic and, accordingly, heteroimmune state of host marmots during hibernation. During winter awakenings, the body temperature of marmots increases in an S-shaped manner from 5 to 37°C within a short period of time (from dozens of minutes to several hours). A drastic acceleration of metabolic and immune processes occurs in the temperature range of 20–30°C. Rapid adaptation to the “explosive” increase in the immune activity of the primary host (tarbagan marmot) during its regular winter awakenings was the essence of the process of Y. pestis speciation.     

Ecological,Evolutionary and Social Constraints on Reproductive Effort: Are Hoary Marmots Really Biennial Breeders?

Biennial breeding is a rare life-history trait observed in animal species living in harsh, unproductive environments. This reproductive pattern is thought to occur in 10 of 14 species in the genus Marmota, making marmots useful model organisms for studying its ecological and evolutionary implications. Biennial breeding in marmots has been described as an obligate pattern which evolved as a mechanism to mitigate the energetic costs of reproduction (Evolved Constraint hypothesis). However, recent anecdotal evidence suggests that it is a facultative pattern controlled by annual variation in climate and food availability (Environmental Constraint hypothesis). Finally, in social animals like marmots, biennial breeding could result from reproductive competition between females within social groups (Social Constraint hypothesis). We evaluated these three hypotheses using mark-recapture data from an 8-year study of hoary marmot (Marmota caligata) population dynamics in the Yukon. Annual variation in breeding probability was modeled using multi-state mark-recapture models, while other reproductive life-history traits were modeled with generalized linear mixed models. Hoary marmots were neither obligate nor facultative biennial breeders, and breeding probability was insensitive to evolved, environmental, or social factors. However, newly mature females were significantly less likely to breed than older individuals. Annual breeding did not result in increased mortality. Female survival and, to a lesser extent, average fecundity were correlated with winter climate, as indexed by the Pacific Decadal Oscillation. Hoary marmots are less conservative breeders than previously believed, and the evidence for biennial breeding throughout Marmota, and in other arctic/alpine/antarctic animals, should be re-examined. Prediction of future population dynamics requires an accurate understanding of life history strategies, and of how life history traits allow animals to cope with changes in weather and other demographic influences.     

Habitat requirements of the long-tailed ground squirrel (Spermophilus undulatus) in the southern Altai

The long-tailed ground squirrels Spermophilus undulatus represent the most abundant burrowing herbivorous species in the southern Altai grasslands and are suggested to play an important role in the maintenance of this mountain ecosystem. The aim of this study was to identify the key features that influence their habitat use in the southern part of the Altai Republic (south-west Siberia, Russia). The research area represents a complete sequence of altitudinal vegetation zones from steppe, forest-steppe, forest, sub-alpine and alpine tundra. Our results suggest that S. undulatus prefers short-grass steppes, near the water source and with a thin layer of a chernozem soil containing a large amount of coarse clastics. The species strictly avoids forests and tolerates only a low density of bush cover. Altitude and exposure to sun do not represent significant factors in the habitat choice of S. undulatus . Neither the presence of pikas nor the presence of marmots influences habitat selection of the ground squirrels. Strong preferences for habitats near a water source may limit the distribution of the species to mountain areas. Degree of human disturbance was not a significant factor affecting distribution and the species even displays slight preferences for heavily grazed habitat near human settlements and roads. Intensive grazing prevents shrubs and forest invasion, keeps vegetation low and thus provides appropriate conditions for the ground squirrels, favouring an open habitat where predators can be easily detected by sight. Our results suggest that the habitat selection of ground squirrels may be determined rather by a protection from predators and burrowing conditions than by food availability.     

Phylogenetic and functional structures of succession in plant communities on mounds of Marmota himalayana in alpine regions on the northeast edge of the Qinghai–Tibet Plateau

Few studies have examined the succession of plant communities in the alpine zone. Studying the succession of plant communities is helpful to understand how species diversity is formed and maintained. In this study, we used species inventories, a molecular phylogeny, and trait data to detect patterns of phylogenetic and functional community structure in successional plant communities growing on the mounds of Himalayan marmots (Marmota himalayana) on the southeast edge of the Qinghai-Tibet Plateau. We found that phylogenetic and functional diversities of plant communities on marmot mounds tended to cluster during the early to medium stages of succession, then trended toward overdispersion from medium to late stages. Alpine species in early and late stages of succession were phylogenetically and functionally overdispersed, suggesting that such communities were assembled mainly through species interactions, especially competition. At the medium and late stages of succession, alpine communities growing on marmot mounds were phylogenetically and functionally clustered, implying that the communities were primarily structured by environmental filtering. During the medium and late stages of succession the phylogenetic and functional structures of plant communities on marmot mounds differed significantly from those on neighboring sites. Our results indicate that environmental filtering and species interactions can change plant community composition at different successional stages. Assembly of plant communities on marmot mounds was promoted by a combination of traits that may provide advantages for survival and adaptation during periods of environmental change.     

Nutritional ecology of yellow-bellied marmots in the White Mountains of California

This study examined the basis of diet preference in a mammalian hibernator, the yellow-bellied marmot Marmota flaviventris , in the White Mountains of California. Preference indices calculated from dietary data and food plant availability in the habitat indicated that forb species were preferred over graminoids, particularly in the spring and early summer. Chemical analysis of food plant species suggested that marmots preferred forbs over graminoids because of their lower fiber (cellulose) content and higher concentrations of phosphorus, sodium and possibly protein. Inclusion of graminoids in the diets was probably due to their greater availability in the habitat coupled with the need to maximize the rate of nutrient intake during the short (4–5 month) active season. Seasonal changes in diet composition were noted which may reflect, in part, temporal changes in nutrient demands.     

Concepts of macro- and microevolution as related to the problem of origin and global expansion of the plague pathogen Yersinia pestis

The relationship between macro- and microevolutionary processes is considered with reference to the ecological scenario of the origin of the plague pathogen and its subsequent natural and anthropogenic global expansion. The macroevolutionary transformation of the ancestral pseudotuberculosis microbe clone into the initial plague microbe Yersinia pestis tarbagani occurred in Central Asia at the end of the Late Pleistocene by a “vertical” Darwinian way in an inadaptive heterothermal continual intermediate environment—the Mongolian marmot Marmota sibirica—flea Oropsylla silantiewi system—via a sequence of unstable and currently extinct intermediate forms. Its natural geographic expansion on the “oil spot” principle in the postglacial time led to the microevolutionary formation of 20–30 hostal subspecies circulating in populations of the background species of burrowing rodents and pikas in arid areas of Eurasia. The intercontinental spread of the “marmot” and “rat” pathogen subspecies in the past few centuries has been exclusively anthropogenic, with the involvement of synanthropic (ship) rats.     

Freezing in a warming climate: Marked declines of a subnivean hibernator after a snow drought

Recent snow droughts associated with unusually warm winters are predicted to increase in frequency and affect species dependent upon snowpack for winter survival. Changes in populations of some cold‐adapted species have been attributed to heat stress or indirect effects on habitat from unusually warm summers, but little is known about the importance of winter weather to population dynamics and how responses to snow drought vary among sympatric species. We evaluated changes in abundance of hoary marmots (Marmota caligata) over a period that included a year of record‐low snowpack to identify mechanisms associated with weather and snowpack. To consider interspecies comparisons, our analysis used the same a priori model set as a concurrent study that evaluated responses of American pikas (Ochotona princeps) to weather and snowpack in the same study area of North Cascades National Park, Washington, USA. We hypothesized that marmot abundance reflected mechanisms related to heat stress, cold stress, cold exposure without an insulating snowpack, snowpack duration, atmospheric moisture, growing‐season precipitation, or select combinations of these mechanisms. Changes in marmot abundances included a 74% decline from 2007 to 2016 and were best explained by an interaction of chronic dryness with exposure to acute cold without snowpack in winter. Physiological stress during hibernation from exposure to cold, dry air appeared to be the most likely mechanism of change in marmot abundance. Alternative mechanisms associated with changes to winter weather, including early emergence from hibernation or altered vegetation dynamics, had less support. A post hoc assessment of vegetative phenology and productivity did not support vegetation dynamics as a primary driver of marmot abundance across years. Although marmot and pika abundances were explained by strikingly similar models over periods of many years, details of the mechanisms involved likely differ between species because pika abundances increased in areas where marmots declined. Such differences may lead to diverging geographic distributions of these species as global change continues.     

The social behaviour of the hoary marmot (Marmota caligata)

The social behaviour of four colonies of hoary marmots (Marmota caligata) was studied in Glacier National Park, Montana, during the summer of 1970. Colony structure involved a dominant male with a few females (3 years or older), 2-year-olds, yearlings and pups. Patterns of burrow use, greetings, play, and aggressive chasing are described, indicating a closely-integrated social structure with reproductive patterns suggesting late dispersal and maturation. A close resemblance to the behaviour of the Olympic marmot (M. olympus) is proposed.     

Social behaviour and individual differences in free-living Alpine marmots (Marmota marmota)

I studied the behaviour of free-living Alpine marmots, Marmota marmota, from 1 June to 19 August 1973 in Vanoise National Park, France. Comparing animals of the same sex and age class, individual differences in behaviour generally were greater between colonies than within colonies. Individual differences among adults exceeded those of yearlings, and greater individual differences occurred with regard to social behaviour than with relatively asocial behaviour. General patterns of social biology resembled those of the North American marmots, exceptions being a tendency for adult males to remain somewhat away from and at higher elevation than their colonies, and a distinctly lower frequency of greeting behaviour. Implications of these findings for the evolution of marmot social behaviour are discussed.     

Detection,Prevalence and Phylogenetic Relationships of Demodex spp and further Skin Prostigmata Mites (Acari,Arachnida) in Wild and Domestic Mammals

This study was conceived to detect skin mites in social mammals through real-time qPCR, and to estimate taxonomic Demodex and further Prostigmata mite relationships in different host species by comparing sequences from two genes: mitochondrial 16S rRNA and nuclear 18S rRNA. We determined the mite prevalence in the hair follicles of marmots (13%) and bats (17%). The high prevalence found in marmots and bats by sampling only one site on the body may indicate that mites are common inhabitants of their skin. Since we found three different mites (Neuchelacheles sp, Myobia sp and Penthaleus sp) in three bat species (Miotis yumanensis, Miotis californicus and Corynorhinus townsendii) and two different mites (both inferred to be members of the Prostigmata order) in one marmot species (Marmota flaviventris), we tentatively concluded that these skin mites 1) cannot be assigned to the same genus based only on a common host, and 2) seem to evolve according to the specific habitat and/or specific hair and sebaceous gland of the mammalian host. Moreover, two M. yumanensis bats harbored identical Neuchelacheles mites, indicating the possibility of interspecific cross-infection within a colony. However, some skin mites species are less restricted by host species than previously thought. Specifically, Demodex canis seems to be more transmissible across species than other skin mites. D. canis have been found mostly in dogs but also in cats and captive bats. In addition, we report the first case of D. canis infestation in a domestic ferret (Mustela putorius). All these mammalian hosts are related to human activities, and D. canis evolution may be a consequence of this relationship. The monophyletic Demodex clade showing closely related dog and human Demodex sequences also supports this likely hypothesis.     

Habitat Effects on the Breeding Performance of Three Forest-Dwelling Hawks

Habitat loss causes population declines, but the mechanisms are rarely known. In the European Boreal Zone, loss of old forest due to intensive forestry is suspected to cause declines in forest-dwelling raptors by reducing their breeding performance. We studied the boreal breeding habitat and habitat-associated breeding performance of the northern goshawk (Accipiter gentilis), common buzzard (Buteo buteo) and European honey buzzard (Pernis apivorus). We combined long-term Finnish bird-of-prey data with multi-source national forest inventory data at various distances (100–4000 m) around the hawk nests. We found that breeding success of the goshawk was best explained by the habitat within a 2000-m radius around the nests; breeding was more successful with increasing proportions of old spruce forest and water, and decreasing proportions of young thinning forest. None of the habitat variables affected significantly the breeding success of the common buzzard or the honey buzzard, or the brood size of any of the species. The amount of old spruce forest decreased both around goshawk and common buzzard nests and throughout southern Finland in 1992–2010. In contrast, the area of young forest increased in southern Finland but not around hawk nests. We emphasize the importance of studying habitats at several spatial and temporal scales to determine the relevant species-specific scale and to detect environmental changes. Further effort is needed to reconcile the socioeconomic and ecological functions of forests and habitat requirements of old forest specialists.     

Sympatric speciation of the plague microbe <Emphasis Type="Italic">Yersinia pestis</Emphasis>: Monohostal specialization in the host–parasite marmot–flea (<Emphasis Type="Italic">Marmota sibirica–Oropsylla silantiewi</Emphasis>) system

An ecological scenario of the origin of the plague microbe that is interpreted in the light of modern Darwinism (synthetic theory of evolution) is presented. It is shown that the plague microbe emerged from a clone of the psychrophilic saprozoonotic pseudotuberculosis microbe Yersinia pseudotuberculosis O:1b in the mountain steppe landscapes of Central Asia in the Sartan time, 22000–15000 years ago, in the monohostal Mongolian marmot (Marmota sibirica)–flea (Oropsylla silantiewi) host–parasite system. It was noted that the evolutionary process described corresponds to the sympatric form of speciation by transition of the clone of migrant founders to a new, already-existing ecological niche. It was established that monohostal specialization of the plague microbe was made possible due to heterothermia (5–37°C) of marmots in the hibernation period. The factors of the speciation process—isolation, the struggle for existence, and natural selection—were analyzed.