Argens Jean-Baptiste de Boyer, marquis d' 1738. Lettres juives ou Correspondance philosophique, historique et critique entre un juif voyageur et ses correspondans en divers endroits. [Jewish letters or critical, historical, and philosophical correspondence between a Jewish traveller and his correspondents in varied places]. La Haye, P. Paupie. Num. BNF de l'éd. de, Paris : INALF, 1961- (Frantext ; N736Reprod. de l'éd. de, La Haye : P. Paupie, 1738.
En français, in French.
Littérature française, French literature, marmotte, marmot, Savoyard, Argens Jean-Baptiste de Boyer 1704-1771.
Extrait/Extract pdf

Ariagno D. 1984. La marmotte Marmota marmota [The marmot Marmota marmota]. In Atlas des mammifères sauvages de France, S.F.E.P.M., 148-149.
En français, in French.
Marmota marmota, marmotte alpine, alpine marmot, faunistique, fauna, France.
Pdf

Ariagno D. & Bussy J. 1971. Les mammifères de la Dombes [Mammals in the Dombes]. Bull. Soc. Nat. Archéol. Ain, 85 : 11-20.
Mammifères, mammals, France.

Арипджанов М.П. (Aripdjanov, Aripdzhanov M.P.) & Есипов А.В. (Esipov A.V.) 199. [Mammifères rares du S-O du Tien Chan. Rare Mammals in SW Tian Shan]. Ecological Problems of Wild Life Protection, Abstracts to All-Union Conf. Moscow, 80-81.
En russe, in Russian.
Mammifères, mammals, Marmota, Tien Chan, Tien Shan.

Buste Timbre

Aristote 1964-1969. Histoire des animaux [Animal history]. Livre VIII, Les Belles Lettres, Paris. En français, in French. Marmota marmota. Les anciens savaient que les Loirs s'endorment profondément l'hiver, et Aristote émet à ce sujet cette idée assez singulière : que le sommeil doit mieux les nourrir que ne sauraient le faire les aliments pour les autres animaux, car on les trouve toujours abondamment pourvus de graisse ; d'après le m ême auteur, le Loir et l'Ours brun se retireraient, en hiver, dans des tanières où ils engraisseraient considérablement sans prendre de nourriture. Appendice Dubois, 1896a.

Platon et Aristote dans l'Ecole d'Athènes de Raphaêl (Vatican)

Aristotle 350 B.C.E. The History of Animals [Histoire des animaux]. Translated by D'Arcy Wentworth Thompson, 1910, Book VIII, Part 17, The Internet Classics Archive. Retrieved August 10, 2005, from the World Wide Web: http://classics.mit.edu/Aristotle/history_anim.8.viii.html
Extrait Pdf Extract

Тарульян В.О. (Armanb A.D.) & Тарульян В.О. (Taroul'ïan, Tarul'yan V.O.) 1974. Nekotorye irintsipialnye ogranitcheniya eksperimenta i modelirovaniya v geografii [Quelques limites importantes d'expérimentation et de modélisation en géographie. Some of the main experimental and model limits in geography]. Izv. AN SSSR, geogr., 4.
En russe, in Russian
Géographie, geography, modèle, model..

Armitage K.B. 1961. Frequency of melanism in the golden-mantled marmot [Fréquence du mélanisme chez la marmotte doérée]. J. Mamm., 42:100-101.
En anglais, in English.
Marmota flaviventris, mélanisme, melanism.

Armitage K.B. 1962. Social behaviour of a colony of the yellow-bellied marmot (Marmota flaviventris) [Comportement social d'une colonie de marmottes à ventre jaune]. Anim. Behav., 10: 319-331.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, social, EUA, USA, Wyoming.
Étude en juin-août 1955, en juillet-août 1956, juin-août 1957, juin-aout 1961. La colonie est située sur une terrasse près de la rivière Snake à l'entrée du parc de Yellowstone. Le terrier principal est celui où les jeunes sont élevés, où les animaux passent la nuit, et où ils vont à la suite d'un signal d'alarme. Les terriers auxiliaires sont des refuges temporaires. Les terriers sont connect és par des sentiers. Les animaux préfèrent les terriers centraux à la colonie. L'organisation des domaines vitaux est de trois types : sans chevauchement, faibles chevauchements, chevauchements importants. Les animaux qui utilisent le même domaine vital fréquemment s'évitent les uns les autres. La forme du domaine vital dépend de la proximité des zones alimentaires. Les comportements agonistiques modifient à la fois la taille et la forme du domaine vital. Les patrons de domaines vitaux ont tendance à changer au cours des deux premières semaines d'août lorsque certains animaux hiberrnent. Les changements des domaines vitaux d'une année sur l'autre sont corrélés aux changements du nombre d'individus occupant ces domaines. Les animaux émergent des terriers au lever du soleil. Une possible interaction entre l'émergence matinale et la défécation est postulée. L'activité est bimodale avec un pic matinal et un pic vespéral. Tous les animaux sont rentrés dans les terriers 30 mn après le coucher du soleil. La communication entre les animaux utilise la vue, l'odorat, le toucher et surtout audition. Le cri d'alarme est produit en réponse à toute intrusion dans la colonie. Il n'y a pas d'animal sentinelle. La communication olfactive consiste en salutations par reniflement des joues. Un ou deux animaux sont dominants. Les femelles avec des jeunes peuvent être agressives à proximit é de leur terrier et soumises dans les autres parties de la colonie. La dominance est caractérisée par une indépendance d'action, battements de queue et toilettage. La soumission est caractérisée par l'évitement des autres, une posture furtive avec la queue basse et la soumission au toilettage. Le comportement territorial ne semble pas exister chez la marmotte à ventre jaune. L'hibernation commence la deuxième semaine d'août. Les conflits et les cris d'alarme sont moins nombreux et l'activité décroît. Il est fait l'hypothèse que le comportement agonistique est l'un des facteurs limitant le nombre d'individus d'une colonie.

Armitage K.B. 1965. Vernal behaviour of the yellow-bellied marmot [Comportement printanier de la marmotte à ventre jaune]. Anim. behav., 13: 59-68.
En anglais, in English.
Marmota flaviventris, éthologie, ethology.

Armitage K.B. 1973. Population changes and social behaviour following colonization by the yellow-bellied marmot [Changements de population et comportement social suivant la colonisation de la marmotte à ventre jaune]. J. Mammal., 54 : 842-854.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, population, social.
In most years, the population of a colony of yellow-bellied marmots was fluid because of dispersal of yearlings and addition of young: the adult population remained relatively stable within any year. Recruitement to the adult population occured solely from animals born in the colony. Total number of young produced was closely correlated with the number of females of reproductive age : the number of young per female decreased at higher population densities because of a greater percentage of nonbreeding 2-years-old females in the population. There was no density-dependant relationship between numbers and survival of young between number of yearlings and number of adults. The rate of social interactions was generally highest in June dand decreased thereafter. Year-to-year and seasonal variations in rates of social interactions are interpreted in relation to the social structure of the population. Rates of social interaction were not directly correlated with population density. The dispersion of the population increased as the population increased. Adult males attempted to obtain exclusive use of the area and directed most of their agonistic behavior toward other males, both adults and yearlings. Agonistic behavior among females exceeded amicable behavior in only one of the five years of study. The growth of the harem may be possible because of mutual tolerance among females. Play figthing occured only among yearlings. A complex interaction occurs between the behavorial characteristics of the individual animals and density effects related to the number, age, and sex of the animals.

Armitage K.B. 1974a. Demography of yellow-bellied marmot populations [Démographie des populations de marmotte à ventre jaune]. Ecology, 55: 1233-1245.
En anglais, in English.
Marmota flaviventris, dynamique population, population dynamics.
Marmot (Marmota flaviventris) populations are colonial or satellite. The number of adults of colonial populations is relatively stable; fluctuations occur primarily because of changes in numbers of young and yearling. Population trends among five colonial populations are dissimilar. Satellite populations are unstable and reproduce at a lower rate than do colonial populations. Satellite marmots are shorter resident than colonial marmots. Both colonial and satellite females usually are longer resident than males. All adult colonial males and 41% of adult colonial females are recruited from other places; all satellite adult are recruited from other places. Losses of colonial marmots are attributed primarily to mortality during hibernation and emigration. Predation appears to be a minor source of mortality of colonial marmots, but may be of greater significance to satellite populations. Demographic relationships of individual colonies appear to be density -independent. Dispersal of colonial animals occurs primarily among yearlings, which have a higher expectation of reaching sexual maturity than young have. The major cause of dispersal is social pressure, but social stress is not simply density-dependent. The colonial social organization is more adaptive than the more nearly solitary (=satellite).

Armitage K.B. 1974b. Male behaviour and territoriality in the yellow-bellied marmot [Comportement des mâles et territorialité chez la marmotte à ventre jaune]. J. Zool., London, 192 (2) : 233-265.
Marmota flaviventris, éthologie, ethology, mâle, male, social, territoire, territory, Colorado, EUA, USA.
Male behaviour and territoriality were studied in five populations of marmots in the East River Valley, near the Rocky Mountain Biological Laboratory, Gothic, Colorado, at an elevation of 2900 m from 1962 through 1972. Males may be classified according to age as yearlings, marmots in their second summer of life; two-year olds, in their third summer; and adults, animals three years old or older. Socially, marmots may be categorized as colonial, peripheral, or transient. A colonial male lives with a harem of females, a peripheral male lives near a colony, and a transient male moves through a locality, remaining only a few days near a colony. Social behaviour was classified broadly as amicable or agonistic. Both types of behaviour occurred between male yearlings and female yearlings, male adults, and female adults. Social behaviour among male yearlings was characterized as play-fighting. Play-fighting has elements of adult sexual and of adult fighting behaviour. However, play-fighting cannot be classified in traditional behavioural categories and may be a developmental pattern in establishing adult modes of behaviour. Social interactions between yearling males and yearling females were primarily amicable. When agonistic interactions occurred, the female usually had the subordinate role. Social interactions between yearling males and adult females were agonistic twice as many times as amicable. The yearling male was subordinate. Social behaviour between yearling males and adult males was nearly always agonistic. The yearling was always subordinate. Amicable behaviour between yearling males and adult males occurred only under unusual circumstances. Most male yearlings dispersed from their parent colonies during their yearling summer. Male yearlings disperse because of the continued presence of an adult male. Agonistic behaviour between an adult male and yearling males is not necessary to cause yearling male dispersal, but supplements the natural avoidance patterns of the yearling males. The behaviour of adult males is characterized by its conspicuousness. Flagging of the tail advertizes the males presence. Some males patrol their territory. Patrolling is evident at those localities where topography or vegetation prevent the male from viewing his territory from one or two central lookouts. Adult male social behaviour is predominantly agonistic. Only with adult females does the occurrence of amicable behaviour exceed that of agonistic behaviour. There was considerable variation among the colonies in the relative amounts of amicable and agonistic behaviour between adult males and adult females. Mean rates or amicable and of agonistic behaviour between adult males and adult females were greater for years of male turnover than for years of male returns. Rates of social behaviour between male adults or male yearlings and the four age-sex categories showed little relationship to measures of population density. A colonial male vigorously defends his territory from strange males. The removal of a territorial male results in nearby males increasing their home range to occupy all or part of the vacant territory, or in a new male occupying the territory. All males introduced into established territories dispersed. The mean size of 24 typical territories is 0.67 ha. Territories may be larger when the number of males is reduced, or may be smaller when the number of territories at a locality is increased. Territories are co-extensive with the habitat at smaller localities. At the larger localities, the number of territories depends on the vigour of the males and the availability of males to replace deceased territorial males. Reduction in the number of territories resulted in an increase in agonistic behaviour among adult males Fights are rare and occurred when the territorial system was unstable. There was no evidence of aggressive appetitive behaviour. Fighting occurs only when social stability is threatened. The major function of the territorial system are enhanced reproduction in a limited environment, enhanced out-breeding, and enhanced fitness of colonial males.

Armitage K.B. 1975. Social behavior and population dynamics of marmots [Comportement social et dynamique des populations des marmottes]. Oikos, 26 (3) : 341-354.
En anglais, in English.
Marmota, éthologie, ethology, social, dynamique population, population dynamics.
La dynamique des populations et le comportement social de cinq populations ont été étudiés pendant ans. Le comportement agonistique caractérise les interactions sociales entre les femelles adultes et les jeunes subordonnés. Les taux de comportement amical et agonistique varie fortement d'une année sur l'autre et d'un harem à l'autre au cours de la même année. Le nombre moyen d'interactions et le nombre moyen de résidents ne sont pas en corrélation. Les taux de comportements amicaux ne sont pas reliés à la densité de population dans aucune des colonies. Le taux de comportement agonistique est en corrélation avec la densité dans une seule des populations. Les taux de ces comportements entre les mâles adultes et les jeunes femelles, entre les femelles adultes et les jeunes femelles, et entre les mâles adultes sont indépendants des densité de population que les données soient regroupées entre toutes les colonies ou que les colonies soient traitées séparément. Aucune relation n'existe entre la taille moyenne du domaine vital et le nombre moyen de femelles résidentes. Par contre, le nombre moyen de femelles résidentes est en corrélation positive avec le pourcentage de recoupement des territoires vitaux. Les canevas de comportement social sont légèrement reliés à la taille du territoire vital. La probabilité qu'une jeune femelle devienne résidente augmente lorsque son territoire vital dépasse les 50% de lui d'une femelle adulte. Le recrutement de femelle était associé à un fort pourcentage de recoupement des territoires vitaux et une tolérance sociale. Le déclin du nombre de résidents ou l'absence de recrutement était associé avec une intolérance sociale. Un modèle faisant intervenir le comportement social et les changements de populations met l'accent sur la densité comportementale, fonction du nombre d'animaux de chaque type comportemental représenté dans la population, comme un facteur contrôlant le recrutement.

Armitage K.B. 1976. Scent marking by yellow-bellied marmots [Marquage olfactif chez les marmottes à ventre jaune]. J. Mammal., 57 (3) : 583-584.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, olfaction, sécrétion, secretion, marquage, marking.
Glandes anales intervenant dans les séquences comportementales lors de disputes territoriales. Marquage de la joue.

Armitage K.B. 1977. Social variety in the yellow-bellied marmot : a population-behavioral system [Diversité sociale chez la marmotte à ventre jaune : un système population-comportement]. Anim. Behav., 25 : 585-601.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, social, dynamique des populations, population dynamics.

Armitage K.B. 1979a. Cannibalism among yellow-bellied marmots [Cannibalisme parmi les marmottes à ventre jaune]. J. Mamm., 60: 205-207.
En anglais, in English.
Marmota flaviventris, marmotte à ventre jaune, yellow-bellied marmot, cannibalisme, cannibalism.

Armitage K.B. 1979b. Food selectivity by yellow-bellied marmots [Sélection de la nourriture par les marmottes à ventre jaune]. J. Mammal., 60 (3): 628-629.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, écologie, ecology, alimentation, feeding.
Test cafétaria. Heracleum lanatum (Panais) est la plus consommée ; Mertensia ciliata (Jacinthe), feuilles ; Potentilla gracilis (Potentille), feuilles. Les fleurs seules de Aquilegia caerula (Colombine), Lupinus floribundus (Lupin), Delphinium barneyi (Pied-d'alouette). Mais, ces trois espèces contiennent des alcaloïdes. Epilobium angustifolium n'est jamais consommé or il contient des tanins.

Armitage K.B. 1981. Sociality as a life-history tactic of ground squirrels [La socialité, tacique d'histoire de vie des écureuils terrestres]. Oecologia, 48 (1) : 36-49.
En anglais, in English.
Sciuridae, éthologie, ethology, écologie, ecology, social.
Multi-variate analysis of life-history traits of 18 species of burrowing sciurids indicates that reproductive effort is determined by body-size energetics. Other traits, such as age adult weight reached, age of dispersal, length of time of gestation, were significantly correlated with body size. A principal component analysis suggested that the complex of life-history traits could be reduced to four components : body size (=weight), seasonality, specific reproductive effort, and maturity. The variation in the sociality index was best explained by age of first reproduction and age adult weight reached. Generally, species are more social when large body size combined with a relatively short growing season is associated with delayed dispersal and occurs in those species typically breeding for the first time at age two or older. Sociality in these species may have evolved through retention of daughters within the maternal home range as a means of continuing reproductive investment beyond weaning.

Armitage K.B. 1982a. Yellow-bellied marmot [La marmotte à ventre jaune]. In Handbook of census method for terrestrial vertebrates, Davis D.E. ed, C.R.C. Press, Boca Raton : 148-149.
En anglais, in English.
Marmota flaviventris, méthodologie, methodolgy.

Armitage K.B. 1982b. Marmots and coyotes : behavior of prey-predator [Marmottes et coyottes : comportement proie-prédateur]. J. Mammal., 63 (3) : 503-505.
En anglais, in English.
Canidae, Marmota flaviventris, éthologie, ethology, prédation.
Deux cas de prédation de jeunes de l'année. Dans les deux cas, au moins une marmotte a vu le coyote sans émettre de cri d'alarme.

Armitage K.B. 1982c. Social dynamics of juvenile marmots : role of kinship and individual variability [Dynamique sociale des marmottes juvéniles : rôle de la parenté et de la variabilité individuelle]. Behav. ecol. sociobiol., 11 : 33-36.
En anglais, in English.
Marmota flaviventris, éthologie, parenté, kinship.
A population of eight juvenile female yellow-bellied marmots (Marmota flaviventris) was introduced into a marmot locality from which all other marmots were removed. Social interactions were monitored in the field and the individual behavioral profile of each animal was determined by mirror image stimulation. Social interactions were unequally distributed amomg the eight juveniles. Neither body size nor kinship were significantly related to frequencies of social interactions. Social interactions were significantly related to individual differences.

Armitage K.B. 1983. Hematological values for free-ranging yellow-bellied marmots [Valeurs hématologiques chez les marmottes à ventre jaune sauvages]. Comp. Biochem. Physiol., 74 (1) : 89-93.
En anglais, in English.
Marmota flaviventris, physiologie, physiology, sang, blood, saison, season, sexe, sex.
1. Hemoglobin, packed cell volume, erythrocyte, leucocytes. MCV, MCH and MCHC were determined for a population of Marmota flaviventris over a period of seven years. 2. There was no significant difference in hematology among years, between sexes or between seasons for adults and yearlings. 3. Early season juveniles had significantly lower PCV. Hb and erythrocyte counts than did late season juveniles. There were no significant differences in hematological value among adults, yearlings and late season juveniles. 4. Juveniles had significantly lower leucocycte counts than adults and yearlings. 5. PCV of marmots responds to acclimatization. 6. Hematological value of scuirids are adaptive to environmental factors such as hypoxia of burrows and high altitude, temperature and metabolic rate. 7. PCV of yellow-bellied marmots evidences an adaptive response to high altitude when compared to the closely-related woodchuck, M. monax.

Armitage K.B. 1984. Recruitment in yellow-bellied marmot populations : kinship, philopatry and individual variability [Le recrutement dans les populations de marmottes à ventre jaune : parenté, philopatrie et variabilité individuelle]. In Biology of ground-dwelling squirrels : Annual Cycles, Behavioral Ecology and Sociality, 377-403, J.O. Murie & G.R. Michener, eds., Lincoln : university of Nebraska Press.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, parenté, kinship.
Recruitment of females forms matrilines, most of which become extinct and are repaced by immigrants. Mother/daughter or sister/sister kin groups share space. Philopatry characterizes matrilines but is not essential for their persistence. Matrilineal bifurcation is associated with the partitioning of a single space into two or more spaces, each of which is shared by individuals who are related by 0,5. Although the number of residents may not change, the total space used increases when matrilines bifurcate. Recruiters differ from nonrecruiters primarily in the nunber of female yearlings produced. Social females recruit a greater proportion of their yearling daughters than do asocial animals. Male turnover and the presence of unrelated adult females do not significantly affect recruitment, whereas an immigrant adult female nearly always prevents recruitment Recruitment of yearlings is more likely if they are philopatric ; yearlings that wander widely disperse from their natal home ranges. lmmigration generally is associated with occupying an empty area. There is no evidence of sex ratio adjustment by successful recruiters.

Armitage K.B. 1986a. Individual differences in the behavior of juveniles yellow-bellied marmots [Différences individuelles dans le comportement des jeunes marmottes à ventre jaune]. Behav. ecol. sociobiol., 18 (6) : 419-424.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, jeunes, young.
Le comportement individuel est testé dans un labyrinthe, par MIS (mirror-image stimulation) et par le comportement social sur le terrain. Les résultats suggèrent que les marmottes ont des phénotypes comportementaux individuels qui s'expriment dans les interactions sociales avec leurs conspécifiques.
Yellow-bellied marmots express considerable individuality as measured by behavior in a maze, mirror-image stimulation (MIS), and social behavior in the field. Maze behavior discriminated between residents and dispersers; residents explored the maze more widely than did dispersers. Males could not be distinguished from females nor survivors from non-survivors by their maze behavior. A group of six yearling females was established to examine the relationship between individual behavioral phenotypes (as determined by MIS) and social behavior in the field. This experiment provided a situation in which social behavior was not influenced by age, sex, or reproduction (female yearlings are non-reproductive). The number of social interactions per individual ranged from 25 to 69. The number of observed interactions per individual differed significantly from the expected for greeting, allogrooming, total amicable, play, and total social interactions. Rankings of greeting, total amicable, and total interactions were directly correlated with rankings on the "avoidance" axis : play was inversely correlated with the "approach" axis. These results suggest that marmots have individual behavioral phenotypes that are expressed in their social interactions with their conspecifics.

Armitage K.B. 1986b. Marmot polygyny revisited : Determinants of male and female reproductive strategies [Révision de la polygynie : déterminants des stratégies de reproduction des mâles et des femelles. In Ecological aspects of social evolution, D.S. Rubenstein and R.W. Wrangham, eds., New-Jersey, Princeton University Press.
En anglais, in English.
Marmota, éthologie, ethology, reproduction, dynamique des populations, population dynamics.

Armitage K.B. 1986c. Individuality, social behavior and reproductive success in yellow-bellied marmots [Individualité, comportement social et succès reproducteur chez les marmottes à ventre jaune]. Ecology, 67 (5) : 1186-1193.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, écologie, reproduction, social, dynamique des populations, population dynamics.
Current theory suggests that population dynamics are the consequence of the reproductive strategies of individuals. Individual differences should be expressed in reproductive output, dispersal, social behavior, and recruitment. Mirror-image stimulation (MIS; i.e., exposure of the animal to a large mirror) was used as an independent measure of individuality, which could be distributed continuously or which could be grouped into two or more types. Three axes derived from a factor analysis of behavioral data obtained during MIS accounted for 85% of the variance among individual marmots. The rank order of 19 adult females on each of the three MIS axes was not correlated with the rank order of lifetime reproductive success measured as number of young weaned, number of yearlings produced, or number of young or yearlings produced per year of residency. This result suggests that individual differences are not continuous. Each female was assigned to one of three groups according to the MIS axis on which she had her highest factor score. Rankings for the number of female yearlings, number of recruits, and number of 2-year-old resident daughters varied significantly among the MIS groups. Mean values of these measures were highest for females in the "sociability" group. Although none of eight measures of lifetime social behavior for l8 females was significantly related to the three MIS groups. Several measures of lifetime amicable behavior were correlated with the production and recuitment of female yearlings. Behavior in the field is affected not only by individual behavioral phenotypes, but also by kinship and patterns of space use. Marmots may have a strategy of phenotypic plasticity. By producing young of varied phenotypes, a female increases the probability that over the long term some of her descendants will survive in varied and unpredictable social and ecoIogical environments.

Armitage K.B. 1987a. Do female yellow-bellied marmots adjust the sex-ratios of their offspring? [Les marmottes femelles à ventre jaune ajustent-elles les sex-ratios de leur descendance?]. Am. Nat., 129 (4): 501-519.
En anglais, in English.
Marmota flaviventris, jeunes, young, reproduction.
The overall sex ratio of weaned yellow-bellied marmots does not differ significantly from one. Litter size has no effect on the sex ratio of young. Stress, measured by eosinophil concentration and mirror-image stimulation, is not associated with biased sex ratios. Three-year-old females and females living in matrilines of two produce more daughters than expected. Females whose age is below the mean produce female-biased litters. Young females that are members of matrilineal social groups produce significantly more daughters than sons. By contrast, the lifetime sex ratios of young produced by subordinate females is male biased. These results are contrary to those predicted from the model of Trivers and Willard and the model of local resource competition. The patterns of sex-ratio variation in marmots, primates, red deer, Antechinus, and white-tailed deer suggest that females produce the sex that has a higher probability of future reproductive success, regardless of the costs of producing individuals of that sex.

Armitage K.B. 1987b. Social dynamics of mammals : reproductive success, kinship and individual fitness [Dynamique sociale des mammifères : succès reproducteur, parent é et valeur sélective individuelle]. Train. Evol., 2 : 279-284.
En anglais, in English.
Marmota, reproduction, parenté, kinship.
Apparent altruism, in which an individual seemingly decreases its evolutionary fitness by assisting others, can confer benefits if the individual assists kin. Thus, an animal can increase its total inclusive fitness by producing offspring (direct fitness) and/or helping kin to reproduce (indirect fitness). Although kin selection has been suggested as the mechanism underlying the formation of mammalian societies, many species act as if they attempt to maximize the direct fitness component of their inclusive fitness.

Armitage K.B. 1988. Resources and social organization of ground-dwelling squirrels [Ressources et organisation sociale des écureuils terrestres]. In The ecology of social behavior, C.N. Slobodchikoff Ed., Academic Press, New-York: 131-155.
En anglais, in English.
Sciuridae, éthologie, population, social.
Throughout I have argued that sociality is a mechanism whereby a female ground squirrel attempts to maximize her direct fitness by increasing the probability that one or more daughters will survive to maturity. The retention of daughters in the natal area implies that survivorship is lower in dispersers than in residents. Female recruits may lose at least one year of reproduction because their reproductive capacity is suppressed by older females, often their mothers. This apparent willingness to forego reproduction in order to be a resident further suggests that the costs of dispersing are considerable. No female should disperse from her birth site unless her fitness would be increased over what it would be if she attempted to become a resident. A juvenile female should "assess" the probability of achieving reproductive success as a resident and "decide" whether to emigrate or remain. The proximal mechanisms initiating this decision are incompletely known (Gaines and McClenaghan 1980; Holekamp 1984), but adult residents probably play a major role. When all adults were removed from a colony of yellow-bellied marmots, none of the yearling females dispersed (Brody and Armitage 1985). Probably the proximal mechanisms are subtle; the mere presence of an adult who expresses dominant behavior may be sufficient to initiate dispersal of some juveniles. The benefits and costs of dispersal in comparison to recruitment must be determined if we are to understand the function of sociality as an evolutionary stategy for increasing fitness. There is little evidence that resource manipulation changes the basic social structure of a species. Food supplementation has direct fitness consequences : the probability of producing reproductive offspring is increased by increasing litter size and decreasing the age of first reproduction. Both responses increase the lifetime reproductive output of an individual. This response has its physiological limitations ; every species has an upper limit to the biomass of offspring a female can produce. In ground-dwelling squirrels the age of first reproduction cannot be lower than one year of age because of the limitations imposed by hibernation or winter inactivity. If there are limits to maximizing fitness by increasing the probability of evolutionary success though the production of more offspring, then we would expect social mechanisms to evolve to increase that probability. If sociality is such a successful mechanism for increasing fitness, why are not all ground-dwelling squirrels social? The answer seems to lie in mother-offspring conflict, in other words, in the costs of sociality.
The costs are mediated though competition and expressed by loss of reproductive output or loss of space to distant kin (r<0.25) that otherwise could be used by closely related kin (r=0.5). Dispersal characteristically occurs in the summer before reproductive competition beging. This competition must make the costs of sociality too high for those species who are reproductively mature at age one. The establishment of separate home ranges in those species who do not form matrilineal groups is functionally similar to the subdivision of yellow-bellied marmot matrilines and the generally small size (x=1.47) of these matrilines. Females attempt to occupy a resource base and live in a social environment that will increase their direct fitness. Prairie dogs are an interesting example of this process. The Gunnison and black-tailed prairies dogs are highly social, but the white-tailed is much less so (Hoogland 1981b). White-tailed prairies dogs breed at age one ; most black-tailed prairie dogs breed at age two. A comparative study of the two species hypothesized that reduced predation may be the most important benefit of prairie-dog sociality (Hoogland 1981b). However, I suggest that the benefits of sociality must be explained in terms of reproductive success. Various selective pressures can modify behavior without inducing sociality. Vigilance and alarm calling are common behaviors among ground-dwelling squirrels, but only a few species have developed sociality. Sociality in prairie dogs can be explained in terms of reproductive benefits. In the not-too-distant past, black-tailed prairie dog populations extended for many miles and were (and still are) characterized by sparse vegetation (Koford 1958). The population is organized into female groups with one or two males called coteries (king 1955). Except for coteries on the periphery, all coteries are surrounded by competitors. Any disperser must enter hostile territory and the likelihood of finding a place to settle must be low. It does not matter whether a disperser would fail because of the inability to find a burrow site, or inability to obtain sufficient food, or by becoming prey ; the key issue is that the likelihood of reaching reproductive maturity is very low. The probability of failure would be greater the younger and smaller the disperser. Thus, the probability of reproductive success would increase by retaining offspring in their natal area until they approach reproductive maturity. At the time of approaching maturity, decisions could be made. If mortality reduced coterie membership, the offspring could remain. If the coterie could not accept an additional member, the offspring could disperse. Because it would be older and larger it would have some increased probability of survival. It is critical to coterie success that new members be recruited. Coterie members defend their area against intruding conspecifics (Kin 1955; Hoogland 1981a). Small cotteries may lose space (=resources) to larger, neighboring coteries. A female will maximize her fitness if all coterie members are her daughters. Demographic considerations suggest that situation would be difficult to achieve. In fact, coterie member-ships may include half-sisters, full first cousins, half-aunt-nieces, etc. (Hoogland 1986). Thus, females probably are forced to accept less closely related kin in order to have assistance in coterie defence. However, coterie space is limited and only a few of the young that are produced can expect to achieve residency. A female should initiate behaviors that will increase the likelihood that her daughters will become residents rather than nieces, granddaughters, cousins, sisters, etc. One strategy is to kill offspring who would compete with one's own offspring for the limited space. Although female prairie dogs defend their burrows against conspecifics, infanticide is common (Hoogland 1985). Competition among female black-tailed prairie dogs is greatest during late pregnancy when the danger of infanticide is high (Hoogland 1986). Infanticide in this context is readily understood as an evolutionary strategy. Furthermore, amicable behavior among females is not distributed according to relatedness but varies inversely with competition (Hoogland 1986). Thus animals do not aid the reproduction of kin ; they attempt to gain more by maximizing their direct fitness (see also Rubenstein and Wrangham 1980). Sociality has both cooperative and competitive behaviors. Because social groups are frequently kin groups too much attention has focused on cooperation under the assumption that individuals should assist kin. This assumption implies that indirect fitness is an important component of inclusive fitness. The blend of cooperation and competition suggest that ground-dwelling squirrels attempt to maximize direct fitness (see also Armitage 1986a). The indirect component to fitness is important only as it contributes to an animal's direct fitness. Thus, a female yellow-bellied marmot or a female black-tailed prairie dog tolerates a sister or a daughter, not because of indirect fitness benefits, but because these individuals contribute to her direct fitness. Black-tailed prairie dog are highly agonistic toward non-coterie members, and amicable behavior within the coterie is always greater than amicable behavior with non-coterie members (Hoogland 1986). Obviously, inclusive fitness is greater when sociality involves kin groups rather than unrelated individuals. That is not the issue. What is critical is whether an animal's behavior is directed toward maximizing direct fitness. I suggest that the behavior of ground-dwelling squirrels is directed toward maximizing direct fitness. Social behavior is both cooperative and competitive. In most species, "competitive decisions" are made in the first year of life ; dispersal occurs at this time and most species, including white-tailed prairie dogs, do not form matrilineal groups. When the competitive decisions come at age two or later, matrilineal groups are formed. But competition becomes a way of life as each individual plays its game of attempting to maximize direct fitness. I have sketched a scenario. This scenario can be verified or rejected only by focusing on the lifetime reproductive success of individuals of known relatedness. And these studies must include learning the fate of dispersers. Only by documenting the reproductive success of dispersers and resident can we construct a logical story of the relative importance of direct and indirect selection in animal sociality. I predict that direct selection will be by far the most important.

Armitage K.B. 1989a. The function of kin discrimination [La fonction de la discrimination de la parenté]. Ethol. Ecol. Evol., 1 : 111-121.
En anglais, in English.
Ethologie, ethology, parenté, kinship.

Armitage K.B. 1989b. Dynamics of emigration into yellow-bellied marmot colony [Dynamique de l'émigration dans une colonie de marmottes à ventre jaune]. 218p.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, écologie, ecology, dispersion.

Armitage K. B. 1991a. Social and population dynamics of yellow-bellied marmots : results from long-term research [Dynamique sociale et des populations chez les marmottes à ventre jaune : bilan d'une étude à long terme]. Annu. Rev. Ecol. Syst., 22 : 379-407.
En anglais, in English.
pdf disponible/available
Marmota flaviventris, éthologie, ethology, écologie, ecology, social, population.

Armitage K.B. 1991b. Factors affecting corticosteroid concentrations in yellow-bellied marmots [Facteurs affectant les concentration en corticostéroides chez les marmottes &agrave; ventre jaune]. Comp. Biochem. Physiol., 98A: 47-54.
En anglais, in English.
Marmota flaviventris, physiologie, physiology, endocrinologie, endocrinology.
1. Bound and total corticosteroid concentrations of yellow-bellied marmots (Marmota flaviventris) were lowest in May after emergence from hibernation and peaked in August prior to immergence.
2. Total corticosteroids were affected by age but not by sex or reproductive status.
3. There was no consistent relationship between measures of population density and concentrations of corticosteroids; when a significant relationship occurred, only 22-34% of the variation was explained.
4. Social status and social behavior were the major factors affecting corticosteroid concentrations.

Armitage K.B. 1992. Struttura sociale e strategia di fitness nelle marmotte. Social organization and fitness strategies of marmots [Organisation sociale et stratégies sélectives des marmottes]. Proc. 1st Inter. symp. on Alpine Marmot and gen. marmota, Bassano B., Durio P., Gallo Orsi U., Macchi E. eds., 89-94. En italien et anglais, in Italian and English.
Marmota, éthologie, ethology, écologie, ecology, social, évolution.
Marmots (Marmota spp.) occur in various social groupings : woodchucks (M. monax) are essentially solitary (Barash 1989: 167). Yellow-bellied marmots (M. flaviventris) form female kin groups to which a male is attached to form a harem (Armitage 199l) ; Olympic (M. olympus) and hoary (M. caligata) marmot groups consist of a male and usually two adult females that reproduce in alternate years (Barash 1973, 1974a) and the alpine marmot (M. marmota) groups contain one pair of breeding adults plus a variable number of non-breeding 3 and 4 year-old adults (Arnold 1990a). In all species, dispersal typically occurs the year before reproductive maturity is reached, but subordinate, non-breeding adults may remain for one or more additional years in alpine marmot groups (Arnold 1990b). Increasing sociality in marmot groups was correlated with the length of the growing season ; where the growing season is too short to permit animals to become reproductively mature at l year of age, offspring are retained in their natal area for one or more additional growing seasons (Barash, 1974b). Subsequently, delayed dispersal and the consequent retention of offspring in their natal area were viewed as a life-history tactic whereby females continue reproductive investment beyond weaning (Armitage 198l). Among North American ground-dwe1ling sciurids, variation in sociality was best explained by age of first reproduction and age adult body mass was reached ; sociality was not significantly correlated with several measures of the length of the active season (Armitage, 198l). The lack of a relationship between sociality and the length of the growing season suggests that a more fundamental factor underlies delayed dispersal. The widespread occurrence of female philopatry and the formation of female kin clusters and matrilines (Armitage, 1981, 1987, 1991; Michener 1983) indicate that sociality is a fitness strategy.

Armitage K.B. 1993. Unusual mortality in a yellow-bellied marmot population [Mortalité inhabituelle dans une population de marmottes à ventre jaune]. Abst. Intern. Conf. on marmots of the CIS-states, Gaidary, Ukraine, Moscow, 42-43.
En anglais, in English.
The winter of 1991-92 apparently reduced survivorship and reproduction of yellow-bellied marmots (Marmota flaviventris) in our study area in the East River Valley, Gunnison Country, Colorado, at 2900m. I divided the study area into two groups, lower valley where winter conditions are milder and upper valley where winter conditions are more severe. In the lower valley the rate of survivorship of adult females was 0.72 for the three previous winters and 0.75 for 1991-92. For the upper valley, the rate of survivorship of adult females was 0.76 for the thee previous winter and 0.42 for 1991-92. Likewise, survivorship of young to yearlings in the lower valley was 0.67 for the three previous winters and 0.73 for 1991-92. For the upper valley, juvenille survivorship was 0.61 for the three previous winters and 0.26 for 1991-92. the percentage of adult females reproducing in the lower valley was 47.4% for the three previous years and 91.5% for 1992. For the upper valley in average of 69% of the adult females produced litters in the three previous years whereas only 30% reproduced in 1992. In the lower valley of five adult females that reproduced in 1991 and survived to 1992, three successfully reproduced. Six females that did not reproduce in 1991, did so in 1992. In the upper valley, no female that reproduced in 1991 successfully reproduced in 1992 and five of the nine1991 reproductive females did not survive. The three females that weaned litters were all three years old and reproduced for the first time. Young in 1991 in the upper valley reached only 65.7% of the body mass on September 15 that young averaged the previous two years whereas young in the lower valley achieved 88% of the body mass. Similarly, reproductive females in the upper valley weighed less on September 15 in 1991 than they did in either of the two previous years. Conversely, reproductive females in the lower valley weighed more in 1991 than in either1989 or 1990. In 1991 the non-reproductive females (2-years-old) in the upper valley weighed 3.58 kg on September 15 whereas the average mass of the reproductive females was 3.29 kg. Thus, both survivorship in the Winter of 1991-92 and reproduction in the summer of 1992 are related to the mass of animals just prior to imergence into hibernation.

Armitage K.B. 1994a. Social dynamics, kinship, and population dynamics of Marmots [Dynamique sociale, parenté, et dynamique des populations chez les marmottes]. Abstracts 2d Conf. Intern. Marmots,,
En français et en anglais, in French and in English.
Probably the social systems of all marmot species, except M. monax are based on kinship and most species have a family with a dominant breeding pair and non-breeding subordinates. Social behavior of closely-related animals is primarily amicable whereas agonistic behavior characterizes the social interactions of more distantly-related or unrelated individuals. All social groups are characterized by reproductive suppression of younger/subordinate animals by older/dominant individuals. Only the solitary M. monax lacks reproductive suppression among adults. Population growth occurs when offspring are retained in their natal group. Recruitment of offspring occurs when they are treated cohesively; dispersal is associated with either a lack of cohesive behaviors or the presence of agonistic behaviors. Population saturation probably occurs in all species and the lack of nutritious food is proposed as the limiting factor. Major population decline usually is associated with unusual weather; e.g., drought or prolonged winter. The availability of wide-spread habitat provided ecological release that permitted M. monax and M. flaviventris to evolve different social systems as a mechanism for escaping reproductive suppression.

Armitage K.B. 1994b. Partage de ressource et parenté chez les marmottes à ventre jaune. Resource sharing and kinship in yellow-bellied marmots (Marmota flaviventris). Abstracts 2d Conf. Intern. Marmots, 20-21.
En français et en anglais, in French and in English.
Marmota flaviventris, parenté, kinship.
Resource sharing, measured as space-use overlap, is directed primarily toward close kin and does not support the proportional distribution of benefits hypothesis in which benefits are distributed in proportion to the degree of relatedness. Matrilineal formation is the social mechanism whereby female yellow-bellied marmots attempt to maximize inclusive fitness by emphasizing the direct fitness component.

Armitage K.B. 1994c. Gain de poids saisonnier chez la marmotte à ventre jaune (Marmota flaviventris). Seasonal mass gain in yellow-bellied marmots (Marmota flaviventris). Abstracts 2d Conf. Intern. Marmots, 22-23.
En français et en anglais, in French and in English.
Marmota flaviventris, masse corporelle, body mass, saison, season.
Les marmottes ont une saison d'activité relativement courte pendant laquelle elles doivent se reproduire, grandir et accumuler de la graisse pour l'hibernation. La saison de croissance courte fait que toutes les espèces de marmottes sauf M. monax ont besoin d'au moins une saison supplémentaire de croissance avant d'atteindre la maturité reproductrice. Toutes les classes d'âge de Marmottes à ventre jaune gagnent du poids pendant la saison active. Les femelles reproductrices commencent à gagner du poids à peu près 3 semaines plus tard que les mˆles adultes, les un-an et les femelles non-reproductrices. Ces classes commencent à gagner du poids vers la fin mai et peuvent atteindre la masse d'hibernation début août, même s'ils n'hibernent pas avant fin août ou début septembre. La durée de l'accroissement pondéral a été mesurée pour les différentes classes d'âge et de sexe :
Nombre de jours de croissance

Armitage K.B. 1994d. Unusual mortality in a yellow-bellied marmot population. Neobytchnaya smertnost' v popoulyatsii jeltobryukhogo sourka. In Actual problems of marmots investigation, Rumiantsev ed., ABF, Moscow, 5-13.
En anglais, in English.
Marmota flaviventris, mortalité, reproduction.
Survivorship and reproduction were reduced in a yellow-bellied marmot population following the winter of 1991-92. Survivorship of young was much lower in the Upper Valley than in the Lower Valley because young had a shorter growing season in the Upper Valley. Reduced survivorship of young and adults was associated with a body mass that was smaller than that of marmots in years when ovewinter survival was much greater. Reduced body mass was associated with a summer of low rainfall.

Armitage K.B. 1996a. Dynamique sociale, parentèle et dynamique des populations chez les marmottes. Social dynamics, kinship, and population dynamics of marmot. In Biodiversité chez les marmottes/Biodiversity in marmots, Le Berre M., Ramousse R. & L. Le Guelte eds., 113-128.
En français et en anglais, in French and in English.
Marmota flaviventris, social, parenté, kinship, dynamique des populations, population dynamics, reproduction.

Armitage K.B. 1996b. Partage des ressources et parentèle chez la marmotte à ventre jaune. Resource sharing and kinship in yellow-bellied marmots. In Biodiversité chez les marmottes/Biodiversity in marmots, Le Berre M., Ramousse R. & L. Le Guelte eds., 129-134.
En français et en anglais, in French and in English.
Marmota flaviventris, spatial, reproduction.

Armitage K.B. 1996c. Augmentation saisonnière de la masse chez la marmotte à ventre jaune. Seasonal mass gain in yellow-bellied marmots. In Biodiversité chez les marmottes/Biodiversity in marmots, Le Berre M., Ramousse R. & L. Le Guelte eds., 223-226.
En français et en anglais, in French and in English.
Marmota flaviventris, reproduction, masse corporelle, body mass.

Armitage K.B. 1997. Sotsial'naya dinamika ou jelto'ryukhikh sourkov : strategiya evolyutsionnogo ouspekha. Social dynamics of yellow-bellied marmots: Strategies for evolutionary success [Dynamique sociale des marmottes à ventre jaune : stratégies d'un succès évolutif]. In Sourki golarktiki kak faktor bioraznoobraziya, III Mejdounarodnaya Konferentsia po sourkam, Tezisy dokladov, 10-11 (Rousskie), 117-118 (Angliïskie) [In Holarctic marmot as a factor of biodiversity, III International Conference on Marmots, Abstracts, 10-11 (Russian), 117-118 (English).
Marmota flaviventris, social, stratégies évolutives.

Армитейдж К.Б. (Armitage K.B.) 1997. Социалъная динамика у желтоьрюхиф рурков : стратегия эволюционного успеха. In Сурки голарктикикак фактор биоразнобразия, 10-11.
En russe, in Russian.
Marmota flaviventris, social, stratégies évolutives.
Version pdf / Pdf version

Armitage K.B. 1998. Reproductive strategies of yellow-bellied marmots: energy conservation and differences between the sexes. J. Mammal.,79(2): 385-393.
En anglais, in English.
Marmota flaviventris, reproduction, énergie, energy, sexe, sex.
pdf disponible/available

Armitage K.B. 1999. Evolution of sociality in marmots [Évolution de la socialité chez les marmottes]. Journal of Mammalogy, 80(1): 1-10.
En anglais, in English.
Marmota, social, évolution.
pdf disponible/available
The 14 species of marmots (Marmota) can be placed in one of four social systems: solitary; female kin matriline; adult male with two females and immature offspring; and a family group consisting of a territorial pair, subordinates adults, and immature offspring. Body size and hibernation are two critical features of marmot biology from which sociality evolves. The high correlation (r=0.91) between emergence mass and mass loss suggests that species are heavier because they require more mass to survive the hibernation period. Marmots increase mass either by having a larger frame or by increasing the amount of mass per frame. Emergence and immergence mass are linearly related to body length; but change in mass is curvilinearly related to body length: large species use relatively more mass. An analysis of mass:length ratios reveals that species with a relatively high immergence mass use considerable mass during the hibernation period whereas species with a relatively high emergence mass use considerable mass after emergence. The consequence of large body size and short growing season is that young in all but one species require two or more growing seasons to reach reproductive maturity. All species reach a maturity index for dispersal by age one; however, only two species disperse by that age and several species delay dispersal beyond the age of first reproduction. All species are reproductively mature by two, but many species delay reproduction for one or more years. Delayed dispersal produces social groups of high relatedness. A cost of sociality is reproductive suppression; reproductive loss is partially compensated by increased survivorship and alloparental care. Subordinate adults also may reproduce or succeed to territorial status. Alloparental care occurs during hibernation when subordinates adults assist in social thermoregulation of closely related young. Thus, marmots have the characteristics of cooperative breeding. The following sequence of events is hypothesized to have occurred in the evolution of marmot sociality. Large size and a short growing season required the retention of offspring in their natal group for one or more additional years to reach maturity. Habitat saturation led to delayed dispersal, which in turn, increased survivorship. When subordinate adults remained in the social unit, they could participate in social thermoregulation and alloparental care.

Armitage Kenneth B. 2000. Energetics of hibernation in the wood-chuck (Marmota monax) [Energétique de l'hibernation chez la marmotte commune d'Amérique]. In Life in the Cold, 11th International Hibernation Symposium.
En anglais, in English.

Armitage K.B. 2002a. Social dynamics of yellow-bellied marmots: Strategies for evolutionary success. Социальная динамика желтобрюхих сурков : стратегия для достижения эволюционного успеха. [Tsocial'naya dinamika jeltobriukhikh sourkov : strategiya dlya dostijeniya evoliutsionnogo ouspekha. Dynamique sociale des marmottes à ventre jaune : stratégies de succès évolutif]. In Сурки голарктики как фактор биоразнообразия. [Sourki golarktiki kak faktor bioraznoobraziïa. Holarctic marmots as a factor of biodiversity], Armitage K.B. & Rumiantsev VY., Proceedings of the 3d International Conference on Marmots, Cheboksary, Russia, 9-21.
En russe et en anglais, in Russian and in English.
Marmota flaviventris, reproduction.
There is evidence for density-dependent population regulation acting through a density-dependent feedback loop. Neither amicable nor agonistic behavior is related to density but each is more closely-related to kinship and the age-sex structure of the population. Female yellow-bellied marmots form matrilines by retaining daughters in their natal area. Matrilines undergo fission, not all litters produce recruits, not all females reproduce each year, and most matrilines become extinct. Social dynamics includes cooperation and competition; cooperative behaviors are directed toward close kin and agonistic behaviors are directed toward more distant kin or unrelated animals. Population dynamics is a consequence of the reproductive strategies of individuals. Female behavior increases the probability of her daughters living to reproduce. Social dynamics is directed toward maximizing direct fitness and not toward population regulation. Key-words: population regulation, matrilines, direct fitness, cooperation, competition.
Les preuves d’une regulation densité-dépendante de la population grace à une boucle de rétroaction densité-dépendante sont rares. Ni le comportement amical ni le comportement agressif ne sont en liaison avec la densité, mais chacun d’eux est plus étroitement lié à la parenté et aux structures d’âge et de sexe de la population. Les marmottes femelles à ventrejaune constituent des lignées matrilinéaires en retenant leur filles dans leur air natale. Ces lignées subissent des éclatements. Tous les members des portées ne sont pas recrutés, toutes les femelles ne se reproduisent pas chaque année, et la plupart des lignées disparaissent. La dynamique sociale comprend cooperation et competition; les comportements coopératifs sont otientés vers les parents les plus proches alors que les comportements compétitifs le sont vers les parents plus éloignés ou les non-apparentés. La dynamique des populations résulte des strategies reproductives individuelles. Le comportement de la femelle accroît les chances de survie et de reproduction de leurs filles. La dynamique sociale tend à rendre maximum la valeur selective directe et non pas la regulation de la population. Mots-clés: regulation des populations, lignées matrilinéaires, valeur selective directe, cooperation, competition.
PDF

Armitage K.B. & Blumstein D.T. 1999. Body mass diversity in marmots [Variation de masse corporelle chez les marmottes]. In Holarctic marmots as a factor of biodiversity, Armitage K.B.& Rumiantsev V.Y. eds., ABF and International Marmot Network, Moscow.
En russe et en anglais, in Russian and in English.Marmota, masse corporelle, body mass.

Armitage K.B. & Blumstein D.T. (Армитейдж К.Б., Блюмштейн Д.Т. ) 2002. Body-mass diversity in marmots. Разнообразие массы тела у сурков. [Raznoobrazie massy tela ou sourkov. Diversité de la masse corporelle chez les marmottes]. In Сурки голарктики как фактор биоразнообразия. [Sourki golarktiki kak faktor bioraznoobraziïa. Holarctic marmots as a factor of biodiversity], Armitage K.B. & Rumiantsev VY., Proceedings of the 3d International Conference on Marmots, Cheboksary, Russia, 22-40.
En russe et en anglais, in Russian and in English.Marmota, masse corporelle, body mass.
pdf disponible/available
Emergence mass is highly correlated with immergence mass, but explains about 55% of the variation. There is a 2-fold difference in body mass among marmot species. Total mass loss during hibernation is correlated with immergence mass. All measures of mass loss are correlated with each other. Big species may be big because they require the use of considerable mass during hibernation; however, mass loss is not significantly correlated with the length of hibernation. Body mass is related to the age of first reproduction and length of the active season. Environmental harshness may provide a unifying theme that integrates life-history traits and patterns of mass loss. Evidence for environment harshness is derived from patterns of female reproduction, mass loss following emergence, variation in home-range size, time of mating, and reproductive skipping. The major sources of mass loss are maintenance during hibernation, reproduction, social thermoregulation, and post-emergence environmental stress. The importance of these four sources of mass loss vary among species and should determine the differences in body mass among species of marmots. Patterns of mass loss are related to social systems.
Key-words: mass, mass loss, environmental harshness, immergence, emergence, life-history traits.
La corrélation entre la masse corporelle à la sortie d’hibernation et celle à l’entrée en hibernation est importante, mais n’explique que 55% de la variation.La masse corporelle des espèces de marmottes varie du simple au double. La perte de masse corporelle au cours de l’hibernation est en corrélation avec la masse corporelle lors de l’entrée en hibernation.Toutes les mesures de perte de masse corporelles sont en corrélation entre elles. Les espèces pourraient être de grande taille car elles doivent dépenser une masse considérable pendant l’hibernation; cependant, la perte de masse n’est pas en corrélation avec la durée de l’hibernation. La masse corporelle est liée à l’âge de la première reproduction et à la durée de la saison d’activité. La dureté du milieu pourrait fournir le cadre unificateur, intégrant les traits d’histoire de vie et les canevas de perte de masse corporelle. L’importance de la sévérité du milieu découle des patrons de reproduction des femelles, des pertes de masse suivant la sortie d’hibernation, de la variation de la taille du domaine vital, du moment de l’accouplement et des arrêts de reproduction. Les principales sources de perte de poids sont dues à l’entretien pendant l’hibernation, la reproduction, la thermorégulation sociale et la pression du milieu après la sortie d’hibernation. L’importance de ces quatre sources de perte de poids varie suivant les espèces et pourrait entraîner les différences de masse corporelle des espèces. Les patrons de perte de poids sont liés aux systèmes sociaux.
Mots clés: masse, perte de poids, dureté du milieu, entrée en hibernation, sortie d’hibernation, sortie d’hibernation, traits d’histoire de vie.

Armitage K.B. & Downhower J.F. 1970. Interment behavior in the yellow-bellied marmot [Comportemet d'enterrement chez la marmotte à ventre jaune]. J. Mammal., 51 (1): 177-178.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, social.
The behavior did not occur in an attack-withdrawal conflict situation, but clearly occured in a dominant attacking animal. Interment behavior apparently is a special form of agonistic behavior by which the aggressive elements of the behavior are transferred to the burrow when the dominant animal is in high state of excitment and is unable to reach the subordinate animal.

Armitage K.B. & Downhower J.F. 1974. Demography of yellow-bellied marmot populations [Démographie des populations de marmottes à ventre jaune]. Ecology, 55: 1233-1245.
En anglais, in English.
Marmota flaviventris, écologie, ecology, reproduction, population, hibernation, Colorado, EUA, USA.
Marmot (Marmota flaviventris) populations are colonial or satellite. The number of adults of colonial populations is relatively stable : fluctuations occur primarily because of changes in numbers of young and yearlings. Population trends among five colonial populations are dissimilar. Satellite populations are unstable and reproduce at a lower rate than do colonial populations. Satellite marmots are shorter resident than colonial marmots. Both colonial and satellite females usually are longer resident than males. All adult colonial males and 4l&#37; of adult colonial females are recruited from other places : all satellite adults are recruited from other places. Losses of colonial marmots are attributed primarily to mortality during hibernation and emigration. Predation appears to be a minor source of mortality of colonial marmots, but may be of greater significance to satellite populations. Demographic relationships of individual colonies appear to be density-independent. Dispersal of colonial animals occurs primarily among yearlings, which have a higher expectation of reaching sexual maturity than young have. The major cause of dispersal is social pressure, but social stress is not simply density-dependent. The colonial social organization is more adaptive than the more nearly solitary (- satellite).

Armitage K.B., J.F. Downhower & G.E. Svendsen 1976. Seasonal changes in weights of marmots [Variations saisonnière du poids des marmottes]. Am. Midl. Nat., 96 (1): 36-51.
En anglais, in English.
Marmota flaviventris, saison, season, croissance, growth.
Les variations saisonnières de la masse corporele chez Marmota flaviventris sont étudiées pendant 8 années au Colorado. Les mâles ont, dans chaque classe de poids, un poids plus élevé que les femelles. Il n'y a pas d'une année sur l'autre de différence dans le taux croissance. Les jeunes de l'année ont moins de réserves graisseuses que les individus plus âgées. Chez les jeunes d'un an et les plus vieux, la masse corporelle est minimale en juin (saison de croissance). Le taux de croissance est plus élevé et rapide dans la période qui suit l'hibernation.
Seasonal changes in the body weights of yellow-belIied marmot (Marmota flaviventris) populations were studied for 8 years in western Colorado. Marmots may be classified by size into four age groups : juvenile, yearling, 2-year and 3-year-old or older, each group with significantly different mean body weights. For each age group, the mean body weight of males was significantly larger than that of females. Mean body weights in June of yearlings and older were smaller, the later the onset of the growing season. All age groups made significant weight gains each year; there was no significant difference in growth rates among years. When the rates of weight gain of juvenilles are corrected for time of appearance above ground, the regression coefficients are virtually identical. The earlier young marmots are weaned, the more they weigh at hibernation and the more likely they are to survive hihernation. The percentage of young surviving their first winter of hibernation is significantly greater the earlier spring begins. Marmots at 3400 m elevation had growth rates similar to those of marmots at 2900 m elevation. The adaptive strategy of marmots includes rapid growth rates, high tissue growth efficiencies and extension of the growing season by reproducing immediately following hibernation.

Armitage K.B. & G.E. Gurri Glass 1993. Communal nesting in yellow-bellied marmots [Elevage au nid communautaire chez les marmottes à ventre jaune]. Abst. Intern. Conf. on marmots of the CIS-states, Gaidary, Ukraine, Moscow, 43.
En anglais, in English.
Marmota flaviventris, reproduction.
In our study area in the East River Valley, Gunnison Country, Colorado, female yellow-bellied marmots (Marmota flaviventris) produced 233 litters of young in 178 colony-years (a colony-year is one colony syudied in one year) at 7 research sites (=colonies) from 1962 to 1992, inclusive. Fifty-six litters (24.0%) were weaned by females living in the same burrow system. Of 25 instances of communal nesting, six involved three females and the remaining nineteen involved two females. Kinship of the communally-nesting females was known in 21 instances ; eight involved litter mate sisters ; seven, mother : daughter pairs ; five, a mother : daughter : sister trio, and one, nonlittermate sisters. Young freely intermingled at emergence and no behavioral discrimination could be detected among the young or between the young and adult females. Space-use overlap between the adult females and young and among the young suggested that the adults and young formed one social group of closely-related kin. Communal nesting was not necessarily associated with communal nursing. Each adult female was injected with an unique antigen, such as Limulus hemocyanin, turkey egg albumin, and porcine thyroglobulin. Females formed large titers of antibodies. Blood samples were collected from the young and screened for antibodies. In one year, one young was detected with two antibodies, which indicated that she nursed more than one adult female. However, for three instances of sisters : sister communally nesting pairs, there was no evidence that a young nursed more thant one female. Thus, we conclude that communal nesting does not imply communal nursing.

Armitage K.B. & Gurri Glass G.E. 1994. Communal nesting in yellow-bellied marmots [Elevage au nid communautaire chez les marmottes à ventre jaune]. In Actual problems of marmots investigation, Rumiantsev ed., ABF, Moscow, 14-26.
En anglais, in English.
Marmota flaviventris, social.
This report of communal nesting is the first for a marmot (Marmota). Communal nesting occurred much less often than possible based on the number of adult females present in a colony and always involved closely-related females (r = 0.5). Space-use patterns of young and adults of a communally-nesting group suggest that the group is one social unit. Young cannot be assigned to their litters nor can maternity be assigned based on space-use overlap. Space-use overlap among members of different communal groups was much less than space-use overlap among members within a communal group. One instance of communal nursing was detected. We conclude that communal nesting does not imply communal nursing.

Armitage K.B. & Johns D.W. 1982. Kinship, reproductive strategies and social dynamics of yellow-bellied marmots [Parenté, stratégies reproductives et dynamiques sociales chez la marmotte à jaune]. Behav. ecol. sociobiol., 11: 55-63.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, écologie, reproduction, social, parenté, kinship.
Social behavior of yellow-bellied marmots was observed for three years in colonies where kinship was known and for one year in a high elevation colony where harems were contiguous. Social dynamics of yellow-bellied marmots is dependent on kinship, group composition, and age-sex classes. This pattern is a consequence of the reproductive strategies of males and females. Females behave cohesively toward sisters or daughters, but not with sons and agonistically toward other females. Males generally behave amicably toward females and agonistically toward males, including their sons. Thus, reproductive strategies limit nepotism. This behavior is consistent with a population process in which sons typically disperse as yearling. At least some of the variation in the expected patterns of social behaviors is attributable to individual differences. Because male and female reproductive strategies differ, a marmot population consists of two social subsystem. The female unit is the closely related kin group which may also be a burrow group. The male unit is a harem which usually consist of two or more competing female kin groups.

Armitage K.B., Johns D.W. & Andersen D.C. 1979. Cannibalism among yellow-bellied marmots [Le cannibalisme chez les marmottes à ventre jaune]. J. Mammal., 60 (1): 205-207.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, cannibalisme, cannibalism, EUA, USA, Colorado.
Description de trois incidents de cannibalisme de Marmota flaviventris observé au Colorado. Le cannibalisme ne fait pas partie d'une stratégie reproductrice mais doit être interprété comme un comportement de charognard.

Armitage K.B., Melcher J.C. & Ward J.M. Jr. 1990. Oxygen consumption and body temperature in yellow-bellied marmot populations from montane-mesic and lowland-xeric environments [Consommationd'oxygèet température corporelle chez les populations de marmottes à ventre jaune en milieux secs de montagne et de plaine]. J. Comp. Physiol., B 160: 491-502.
En anglais, in English.
Marmota flaviventris, physiologie, physiology, conductance, consommation d'Oxygène, oxygen consumption, thermorégulation.
Yellow-bellied marmots characteristically live in montane-mesic environments, but in several areas in western North America this species extended its range into lowland-xeric habitats. Body mass was significantly smaller in the lowland-xeric population from eastern Washington at 393 m than in the montane-mesic population from western Colorado at 2900 m. Oxygen consumption of marmots from montane-mesic and lowland-xeric environments was signiflcantly affected by ambient temperature (TA), water regimen, population, and a population x water regimen x temperature interaction. Lowland-xeric animals had a higher metabolic rate at low TAs, but a lower metabolic rate at higher TAs than the montane-mesic aminals. Oxygen consumption was lower on a restricted-water regimen than on ad libitum water in both populations. Coefficients relating oxygen consumption to body mass were affected by TA, water regimen, and population. These intraspecific coefficients are larger than the interspecific coefficients for all mammals. Body temperature (TB) was affected significantly by TA, water regimen, and population, TA, body mass, and a population x water regimen interaction significantly affected conductance. Conductance generally was higher in the lowland-xeric than in the montane-mesic marmots. Both populations increased conductance at high TA, but the lowland-xeric population dissipated a much higher proportion of the heat by evaporative water loss (EWL) than did the montane-mesic population. Metabolic water production exceeded or equaled EWL at 5-20 °C. Smaller body size, reduced metabolism at high TA, and increased EWL at high TA characterized the lowland-xeric population. Metabolic rates of yellow-bellied marmots were higher than predicted from body size during the reproductive season but decreased to 67% of that predicted from the KIeiber curve by late summer. Marmots minimize thermoregulatory costs by concentrating activity at times when the microclimate is favorable, by tolerating hyperthermia at high TA in the field, and by having a conductance lower than that predicted from body size.

Armitage K.B. & Rumiantsev V.Yu. 2002a. Holarctic marmots as a factor of biodiversity. Proceeedings of the 3rd International Conference on marmots, Cheboksary, Russia, 25-30 August 1997, Moscow ABF P.H., 411 p.
En anglais et en russe, in English and in Russian.
Marmota.

Armitage K.B. & Rumiantsev V.Yu. 2002b. Introduction. Vvedenie. In Holarctic marmots as a factor of biodiversity, Armitage K.B. & Rumiantsev V.Yu. eds., Moscow ABF P.H., 3-4.
En anglais et en russe et résumé en français, in English and in Russian with a French abstract.

Armitage K.B. & Salsbury C.M. 1992. Factors affecting oxygen consumption in wild-caught yellow-bellied marmotas (Marmota flaviventris) [Facteurs influençant la consommation d'oxygène des marmottes à ventre jaune]. Comp. Biochem. Physiol. A, 103: 729-737.
En anglais, in English.
Marmota flaviventris, physiologie, respiration.

Armitage K.B. & Salsbury C.M. 1993. The effect of molt on oxygen consumption of yellow-bellied marmots (Marmota flaviventris) [Effet de la mue sur la consommation d'oxygène des mamottes à ventre jaune]. Comp. Biochem. Physiol., 106A: 667-670.
En anglais, in English.
Marmota flaviventris, physiologie, mue, molt, respiration.

Armitage K.B., Salsbury C.M., Barthelmess E.M., Gray R.C. & Kovach A. 1996. Population time budget for the yellow-bellied marmot [Budget-temps populationnel de la marmotte à ventre jaune]. Ethol. Ecol. & Evol., 8: 67-95.
En anglais, in English.
Marmota flaviventris, éthologie, ethology, Etats-unis d'Amérique, Colorado.
Time budget for 17 behaviors were analyszed for cohort, day-period, season-period and interactions among in the Upper East River Valley in Western Colorado. These effects explained up to 79% of the variation in the behaviors. Marmots allocated more time (40-60%, 110-265 min daily) above-ground to sitting/lying than to any other activity. Foraging was the other major activity (12-23%, 37 to 94 min daily). Vigilance/alert varied from 1.1 to 14.5% and from 12.0 to 71.7 min daily. Social status affected the time budget, especially time allocated to vigilance/alert. All other behaviors averaged about 5% or less except for out-of-sight and enter-burrow. The adult male cohort spent significatively more time above-ground than all other cohorts and reproductive females allocated significantly more time foraging than the other cohorts. The amout of time spent above-ground decreased linearly from the down-river site to the up-river site. The proportion of time spent above-ground was significanti-vely less at mid day than in the morning or afternoon. Above-ground activity was lowest during gestation, increased during lactation and remined high during early post-lactation and declined during the final season-period. The following significant relationships common to the three colonies suggest species characterestics or common environmental influences: more time allocated to foraging and foraging-vigilance in the afternoon and more time allocated to foraging-alert, alert, and locomotion during gestation and lactation than during post-lactation. Marmots adjust their behaviors according to prevailing conditions. The remaining significant relationship can be attributed to specific age-sex cohorts or to habitat differences. Because marmots allocate so much tim to sitting/lying, we suggest that energy budgets are not constrained by foraging time but by time required to process ingested food. Similarly, time spent vigilant/alert does not seem to constrain energy intake. Social behavior is not limited by time, but could easily be expanded by spending less time inactive. In general, there do not seem to be tradeoffs among activities. Among other specis of ground-dwelling sciurids, social behavior occupies a small proportion of the time budget, but the amount of time allocated to foraging and sitting/lying varies videly.

Armitage K.B. & Schwartz O.A. 2000. Social enhancement of fitness in yellow-bellied marmots. Proceedings of the National Academy of Sciences of the United States of America, 97(22): 12149-12152.
En anglais, in English.
pdf disponible/available
Marmota flaviventris, écologie, ecology, comportement, behaviour, structure sociale, social structure, adaptation.
The yellow-bellied marmot (Marmota flaviventris) is a social, ground-dwelling squirrel that lives either individually or in kin groups of from two to five adult females. Philopatry and daughter recruitment lead to the formation and persistence of matrilines at habitat sites. By using 37 years of demographic data for 12 habitat sites, we could determine long-term trends in the effects of group size on two measures of fitness, survivorship and net reproductive rate, which otherwise are obscured by annual fluctuations in these measures. Both size and number of matrilines varied among sites and survivorship and net reproductive rate varied among sites and among matriline sizes. The role of social organization was explored further by examining the effect of matriline size, averaged over all years and sites, on fitness. For both survivorship and net reproductive rate the relationship with matriline size was curvilinear. Fitness increased with the increase in matriline size and then decreased in the largest groups. Decreased fitness in matrilines of four or five was associated with agonistic behavior, a large number of 2-year-old females in the social group, and reproductive suppression. There is no evidence that females acted to increase their fitness by increasing indirect fitness; i.e., by assisting relatives, but attempted to increase direct fitness. Direct fitness increased when mortality and fission of large matrilines reduced group size and the surviving females increased reproduction.

Armitage K.B. & Woods B.C. 2003. Group hibernation does not reduce energetic costs of young yellow-bellied marmots [L’hibernation en groupe ne réduit pas le coût énergétique des marmottes à ventre jaune]. Physiol. Biochem. Zool., 76(6): 888-898.
En anglais, in English.
Marmota flaviventris, hibernation.
We investigated mechanisms of energy conservation during hibernation. The amount of time torpid was significantly less for groups of three young marmots than for marmots hibernating singly. Mean daily mass loss (DML; as mg d(-1) g(-1) immergence mass) averaged 1.33 for single marmots and 1.46 for grouped young. Animals were active 17.3% of the time, which used 82.4% of the energy, and were torpid 82.7% of the time, which used 17.6% of the energy expenditure. During longer torpor bouts, more time was spent in deep torpor, which decreased the hourly cost of a complete bout. Bout oxygen consumption V dot o2, percent time in deep torpor, and body temperature (T(B)) during deep torpor changed seasonally and were curvilinearly related to when in the hibernation period the measurements were made and probably represent a stage in the circannual metabolic cycle. The decrease of environmental temperature (T(E)) to 2 degrees C significantly increased metabolism. Potential costs of low T(E) were reduced by allowing T(B) to decrease, thereby reducing the T(B) to T(E) gradient. Average monthly metabolic rate was high early and late in the hibernation period when time spent euthermic was greater and when VO2 was higher. Over the hibernation period, energy saved averaged 77.1% and 88.0% of the costs for winter and summer euthermic metabolism, respectively. Hibernation costs were reduced by the seasonal changes, the high percentage of time in torpor, the rapid decline in V dot o2 following arousal, and allowing T(B) to decline at lower T(E). Asynchrony in the torpor cycles increased energy expenditures in group hibernators, which negated possible beneficial effects of group hibernation.

Armitage K.B. & Wynne-Edwards K.E. (Армитейдж К.Б., Винн-Эдвардс К.Е. ) 1997. Концентрацтя прогестеронов у желтобрюхих сурков, отловенных в дикой природе. Progesterone concentrations in wild-caught yellow-bellied marmots [Concentrations de progesterone chez les marmottes à ventre jaune sauvages capturées]. In Сурки голарктикикак фактор биоразнобразия, Sourki golarktiki kak faktor bioraznoobraziya, Holarctic marmot as a factor of biodiversity,III Mejdounarodnaya Konferentsia po sourkam, Tezisy dokladov, III International Conference on Marmots, Abstracts, 12-13 (Rousskie, Russian), 119-120 (Angliïskie, English).
Marmota flaviventris, endocrinologie, endocrinology, progestérone, sang, blood.

Armitage K.B. & Wynne-Edwards K.E. (Армитейдж К.Б., Винн-Эдвардс К.Е. ) 2002. Progesterone concentrations in wild-caught yellow-bellied marmots. Концентрация прогестерона ц пойманых в природе желтобрюхих сцрков. [Kontsentratsiya progesterona ou poïmanykh v prirode jeltobrioukhikh sourkov. Concentrations de progesterone chez les marmottes à ventre jaune sauvages capturées]. In Сурки голарктики как фактор биоразнообразия. [Sourki golarktiki kak faktor bioraznoobraziïa. Holarctic marmots as a factor of biodiversity], Armitage K.B. & Rumiantsev V.Yu. eds., Moscow ABF P.H., 41-53.
En anglais et en russe et résumé en français, in English and in Russian with a French abstract.
Marmota flaviventris, endocrinologie, endocrinology, progestérone, sang, blood.
Unlike the majority of small animals and similar to other Marmotini, female reproduction in Marmota flaviventris was characterized by increased progesterone levels during lactation as well as gestation. Elevated levels of progesterone during lactation could be an important component of a feedback mechanism that prevents subsequent ovarian activity and ovulation in this obligate annual breeder. High levels of P4in non-breeding yearling females suggest that there may be critical priming events in a reproduction that occur the year before behavioral estrus. Nipple index is associated with the progesterone cycle and is a good index of breeding or attempted breeding by female marmots.
Key-words: progesterone, nipple index, gestation, lactation.
Contrairement à la majorité des petits mammifères, mais comme les autres Marmotini, la reproduction des femelles chez Marmota flaviventris est caractérisée par une augmentation des taux de progestérone pendant la lactation ainsi que pendant la gestation. Des taux élevés de progestérone au cours de la lactation pourraient constituer une composante importante d’un mécanisme de rétroaction qui prévient une activité ovarienne et une ovulation ultérieures chez cet animal à reproduction strictement annuelle. Des niveaux élevés de P4 chez des femelles d’un an non reproductrices suggèrent qu’il puisse y avoir des événements initiateurs de la reproduction qui se produisent l’année précédant l’oestrus comportemental. L’évolution des mamelons suit le cycle de la progestérone et constitue un bon indice de reproduction ou d’une tentative de reproduction chez les marmottes femelles.
Mots clés: progestérone, indice de reproduction, gestation, lactation.

Extrait pdf extract

Armitage Robert 1853. The penscellwood papers.Vol. I, Letter VII, Instinct of animals. Bentley Richard, London.
En anglais, in English.
Marmota bobac, fanage, haymaking, religion.
Extrait pdf extract

Armstrong D.M. 1972. Distribution of mammals in Colorado [Répartition des mammifères au Colorado]. Monograph, Museum of Natural History, University of Kansas, 3: 1–415.
En anglais, in English.
Mammifères, mammals, Colorado, EUA, USA.

Armstrong D.M. 1987. Rocky Mountain mammals: A handbook of mammals of Rocky Mountain National Park and vicinity [Les mammifères des Rocheuses : manuel des mammifères du Rocky Mountain National Park et de son voisinage]. Colorado Associate University Press, 223 pp.
En anglais, in English.
Marmota flaviventris.

Arnold W. 1985. Socioecology of alpine marmots [Socioécologie des marmottes alpines]. Abst. 19th International Ethological Conference, Université P. Sabatier, Toulouse (30 nov.-2d éc. 1985): 261.
En anglais, in English.
Marmota marmota, éthologie, ethology, écologie, social.
Alpine marmots have been studied since June 82 in Berchtesgaden National Park 230 animals have been marked. Every member of the 24 families in the main study area (l05 ha) is individually known. Marmots typically live in monogamous pairs with their offspring. Young usually disperse when sexually mature as 2-year-olds. Delayed dispersal or integration of strangers leads to groups with more than two adults. Mating occurs after hibernation; only the dominant female litters. Reproduction is not necessarily annual and, like litter size, depends on female body weight after hibernation. This weight is determined by both loss during hibernation and gain during the previous summer. Sex-ratio within litters is skewed in favour of males when hibernation weight losses are low. The most important 15 ecological variables were measured in each home-range and their influence on weight changes was tested. Factor analysis separated them into three categories : (l) abiotic conditions (e.g. radiation, thaw, soil moisture and mean soil temperature at the hibernacula during winter), (2) home-range size, (3) food quality within home-range. Mainly abiotic variables influence weight loss during winter and weight gain during summer. Thus, home-ranges with favourable hibernaculas are less suitable during summer, presumably due to thermal stress. Suboptimal wintering conditions might be counter-balanced by the fact that group members always hibernate together. In the lab body temperatures of all members of three groups were recorded during hibernation, using radiotelemetry. Torpor (12-16 days) alternates with short periods of activity (2-3 days), highly synchronized for all group members. Initial electrophoretic analyses of polymorphic allozymes to investigate the relationship within these groups showed that cuckoldry was present.

Arnold W. 1986a. Sozioëkologie des Alpenmurmeltieres [Socioécologie des marmottes]. Fachber. Biologie Diss., München, Ludwig-Maximilian-Univ.
En allemand, in German.
Marmota marmota, éthologie, ethology, écologie, ecology, social, Allemagne Germany.

Arnold W. 1986b. Die Murmeltier-gesellschaft [Organisation sociale des marmottes]. Mitteilungen aus der Wildforschung, 1-4.
En allemand, in German.
Marmota marmota, éthologie, ethology, écologie, ecology, social.

Arnold, W. 1986c.Ökosoziologie des Alpenmurmeltieres ( Marmota marmota marmota, Linne 1758). [Ecosociologie des marmottes alpines]. Dr. rer. nat. thesis, Ludwig-Maximilians-Universität München.
En allemand, in German.

Arnold W. 1987Die ökologischen Ursachen für das Familienleben des Alpenmurmeltieres (Marmota marmota). Max-Planck-Gesellschaft Jahrbuch , 439-441. 1987. Munich.
En allemand, in German.

Arnold W. 1988a. Social thermoregulation during hibernation in Alpine marmots (Marmota marmota) [Thermorégulation sociale pendant l'hibernation chez la marmotte alpine]. J. comp. Physiol., B 158: 151-156.
En anglais, in English.
Marmota marmota, éthologie, ethology, écologie, ecology, social, thermorégulation.
Au laboratoire, l'hibernation commence quelques jours apr ès avoir placé les animaux à 7-8°C. Les mâles adultes sortent plus tôt de l'hibernation et les juvéniles les derniers.
Body temperature (Tb) of socially hibernating alpine marmots, a pair and two family groups, was monitored continuously from October to March with implanted temperature-sensitive radiotransmitters. At the same time, the animals' behaviour was observed. The recurrent entrances: into and arousals from hibernation were highly synchronised within groups. Group members always lay huddled together when euthermic and also when torpid with a few exceptions at higher ambient temperatures (Ta). Body contact with euthermic nestmates warmed torpid marmots passively. The Tb of animals reentering hibernation did not fall to values close to Ta, as long as euthermic group members were present. Although animals presumably save energy through social thermoregulation, especially when euthermic, these benefits are not necessarily mutual among group members. Differences in thermoregulatory behaviour of individuals described in this study could be responsible for differential weight losses during winter as found in the natural habitat (Arnold 1986).

Arnold W. 1988b. Thermoregulation as a limit to habitat use in alpine marmots (Marmota marmota) [La thermorégulation, facteur limitant de l'utilisation de l'habitat chez les marmottes alpines]. Oecologia, 76 : 544-548.
En anglais, in English.
Marmota marmota, thermoregulation.

Arnold W. 1990a. The evolution of marmot sociality : I. Why disperse late? [Evolution de la socialité des marmottes : I. Pourquoise dispersent-elles tardivement?] Behav. Ecol. Sociobiolol., 27: 229-237.
En anglais, in English.
Marmota marmota; éthologie, ethology, écologie, ecology, dispersion.
Prolonged toleration of offspring in marmots was hypothesized to be (l) a means of preventing dispersal of undersized young (Barash 1974a) or more generaly (2) continued parental investment, increasing the probability of descendant survival and reproduction (Armitage 1981, 1987). These hypotheses are tested in this paper for one of the most social of marmot species, the alpine marmot. The animals studied lived in groups within territories defended by a dominant male and female, or as floaters lacking a well-defined home range. Offspring did not disperse before sexual maturity at age 2 (Fig. 1). Only territorial females bred, whereas territoral males were not able to monopolize reproduction likewise (Table 2). Dispersers had similar spring mass to nondispersers (Table 4). Hence, hypothesis 1 is not supported, at least not for adult-sized, (2 years old animals). During their residency, 19% of subordinates obtained their natal territory or a neighboring one (Fig. 2). Long distance dispersal bore a high mortality risk. Thus, toleration of mature offspring could well represent parentat investment. Other results, however, suggest additional influences on the timing of dispersal. (i) Males dispersed later than females (Fig. 3), possibly because of mate sharing by territorial males (see Emlen 1982). (ii) The higher mass loss of dispersers during the previous winter indicates that weak animals were forced to leave (Table 5) despite presumably lower chances of becoming territorial (Table 3). (iii) Subordinate animals which could not be the offspring of both territorials present were not more likely to disperse (Fig. 3). (iv) Lower dispersal rates when immatures lived in the group (Fig. 3) may indicate benefits from the subordinates presence for rearing young.

Arnold W. 1990b. The evolution of marmot sociality: II. Costs and benefits of joint hibernation [Evolution de la socialité des marmottes : II. Coûts et bénéfices de l'hibernation en groupe]. Behav. Ecol. Sociobiol., 27: 239-246.
En anglais, in English.
Marmota marmota, ecology, écologie, éthologie, ethology, hibernation.
Social groups of alpine marmots (Marmota marmota) were studied for 7 years. The groups consisted of a territorial pair and up to 18 lower ranking animals of various ages, mostly the pair's offspring (Tables 1, 2). Group members lived in a common home range and always hibernated together in one hibernaculum. Groups with older, subordinate animals experienced slightly higher summer mortality but significantly reduced winter mortality (Fig. 1). Infant winter mortality was further decreased if most older subordinates were potentially their full sibs (Fig. 2). Subordinate group members lost less mass during winter with increasing size of the hibernating group, but this trend was reversed when infants were present. Furthermore, augmented mass loss due to low hibernaculum quality became evident (Fig. 3). Apart from these effects, the presence of infants caused additional mass loss in potential full sibs. The opposite was found in subordinates certainly descending from other parents than those of the infants (Table 3). Winter mortality and mass loss data revealed (i) a general benefit of joint hibernation. (ii) an unavoidable cost of infants' presence to other group members, (iii) that only potential full sibs helped in warming infants, (iv) that helping was energetically expensive and increased infant survival. The evolution of postponed dispersal in ground-dwelling squirrels has been attributed to the direct fitness gained by enhanced reproductive chances of offspring when not expelled from the natal territory (Armitage 198l, 1987, 1988). This study shows that group living in alpine marmots has benefits during winter and indicates the additional importance of kin selection in marmot social evolution.

Arnold W. 1990c. Überlen in der Kälte. Sinn und Zweck des Familienlebens beim Alpenmurmeltier [A propos du froid. Sens et fonction de la vie en famille de la marmotte alpine]. Beilage zu Wildtiere, 7(1-7): 2.
En allemand, in German.
Marmota marmota, éthologie, ethology, écologie, ecology, hibernation.
Wer den harten Bergwinter uberleben will, muss sich den Bedingungen anpassen können. Fast 7 Monate verbringen Alpenmurmeltiere (Marmota marmota) eng aneinander-geschmiegt in ihrem Bau. Doch "Winterschlaf" allein reicht nicht aus. Die Frage ist : wie und mit wem? Je Frühjahr wieder erwachen. Das ausgeprägte soziale Verhalten der Alpenmurmeltiere ist ganz darauf ausgerichtet, der eigenen Familie über die lange Winterzeit zu helfen.

Arnold W. 1991a. Parental and alloparental care during social hibernation [Soins parentaux et alloparentaux pendant l'hibernation]. Abst. 1st Interrnational Symposium on Alpine Marmot (Marmota marmota ) and on Genus Marmota: 2-3.
En anglais, in English.
Marmota marmota, éthologie, ethology, soins, care, social, hibernation.
Metabolic heat production of hibernating alpine marmots increases again with decreasing ambient temperature (Ta) below 5°C. Such Ta exist in natural burrows for about 2/3 of the hibernation season and are couterbalanced by social hibernation. However, the presence of infants in socially hibernating groups inflitcts extra mass loss in parents and older full sibs. This indicate that they warm infants during hibernation (Arnold, 1990). The mode of parental and alloparental care was investigated during winter 1990/1991 in the natural habitat. The body temperature (Tb) of 37 alpine marmots of 7 hibernation groups as well as the Ta in their hibernacula were recorded continuously using temperature sensisitve radio transmitters. Fisrt results show that (i) infants have lower Tb during torpor, that (ii) group members huddle together in the nest and (iii) usually synchronize their regular changes from torpor to euthermia. Extra arousal of individuals from torpor occur only in groups with infants when Ta drops below 5°C. The presence of euthermic animals in the nest raises the Tb of still torpid group members. Typically, parents or helpers become euthermic earlier than expected. Hence, the cost of parental and alloparental care seems to result from an increased torpor metabolism due to warming infants as well as from additional arousal at low Ta in favor of infants. Final results will be presented at the conference.

Arnold W. 1991b. Fitness consequences of delayed dispersal in alpine marmots [Conséquences de la valeur sélective de la dispersion tardive chez les marmottes alpines]. Abst. 1st International Symposium on Alpine Marmot (Marmota marmota) and on Genus Marmota: 3.
En anglais, in English.
Marmota marmota, écologie, ecology, éthologie, ethology, dispersion.
Alpine marmot social groups usually consisted of a dominant territorial pair and their progeny which always remained at the natal site beyond sexual maturity. Among females, reproduction was restricted to dominant territorials, whereas subordinate males sometimes copulated. Subordinates had a fair chance of becoming territorial close to the natal site or to inherit the natal territory. On the other hand, long distance dispersal bore a high mortality risk. Benefits from group living arose during hibernation. The presence of adults additional to the territorials reduced winter mortality, particulary when infants were present. Subordinates were less likely to disperse in years when immatures lived in the group. They lost extra weight during winter when they hibernated with infants. Infant winter mortality was further decreased in such groups. Prolonged tolerance of offspring in ground squirrels has been hypothesized to be continued parental investment (Armitage, 1981; 1987). Delayed dispersal in alpine mamots demands further explanation. Subordinates accepted an avoidable cost during winter if closely related to infants, presumably because they warmed them. At least for females, this can be explained only by invoking kin selection. Subordinate males caring for infants may potentially be their fathers. Territorial males may yield part of their fitness to sons to prevent their dispersal and to secure their help in raising young.

Arnold W. 1992a. Evoluzione sociale ed obbligatorietà dell'ibernazione di gruppo nella marmotte. Social evolution and obligatory group hibernation in marmots [Evolution sociale et l'hibernation en groupe obligatoire chez les marmottes]. Proc. 1st Inter. symp. on Alpine Marmot and gen. marmota, Bassano B., Durio P., Gallo Orsi U., Macchi E. eds.: 41-54. En italien et en anglais, in Italian and English)
Marmota marmota, écologie, ecology, éthologie, hibernation.

Arnold W. 1992b. Adattamento al freddo. La fisiologia del letargo della marmotta. Adaptation to the cold. The physiology of marmot hibernation [Adaptation au froid. Physiologie de l'hibernation de la marmotte]. Proc. 1st Inter. symp. on Alpine Marmot and gen. marmota, Bassano B., Durio P., Gallo Orsi U., Macchi E. eds.: 31-39. En italien et anglais, in Italian and English)
Marmota marmota, physiologie, physiology, thermorégulation.

Arnold W. 1993a. Energetics of social hibernation [L'énergétique de l'hibernation sociale]. In Life in the cold. Ecological, Physiological and Molecular Mechanisms, Carey C., Florant G.L., B. Wunder & B. Orwitz eds., Westview Press, Boulder.
En anglais, in English.
Marmota, hibernation, thermorégulation, thermoregulation.

Arnold W. 1993b. Social evolution in marmots and the adaptative value of joint hibernation [L'évolution sociale chez les marmottes et la valeur adaptative de l'hibernation groupée]. Verh. Dtsch. Zool. Ges., 86: 79-93.
En anglais, in English.
Marmota, éthologie, ethology, social, hibernation.

Arnold W. 1993c. Das soziale leben der Murmeltiere. PdN-Biologie, 42: 13-16.
En allemand, in German.Marmota marmota, hibernation, social.

Arnold W. 1993d. Social evolution in marmots and the adaptative value of joint hibernation. Verhandlungen der Deutschen Zoologischen Gesellschaft, 86: 79-93.
En anglais, in English.
Marmota marmota, hibernation, social.

Arnold W. 1995a. Social behavior and telemetrical assessment of thermoregulation in hibernating marmots [Comportement social et évaluation télémétrique de la thermorégulation des marmottess hibernantes]. In Behavioural brain research in naturalistic and semi-naturalistic settings, Alleva, E. et al. (eds.), Kluwer, Dordrecht, pp 395-411.
En anglais, in English.
Marmota, hibernation, thermorégulation, thermoregulation.

Arnold W. 1995b. The study of behaviour in naturalistic, semi-naturalistic and laboratory environments [Etude comportementale en milieu naturel, semi-naturel et au laboratoire]. In Behavioural brain research in naturalistic and semi-naturalistic settings (Alleva, E., Fasolo, A., Lipp, H. P., Nadel, L., & Ricceri, L., eds.), pp. 435-437. Kluwer, Amsterdam.
En anglais, in English.
Ethologie, ethology.

Arnold W. 1996. Forschungsbeispiele: 1. Verzögerte Abwanderung und soziales Leben bei Murmeltieren: Strategien des Überlebens im alpinen Lebensraum. In Verhaltensbiologie, Frank, D. (eds.), 3rd edition, Thieme, Stuttgart, pp 156-160.
En allemand, in German

Arnold W. 1997. Wildtierökologie aktuell. In: Alles Jagd ... eine Kulturgeschichte (unknown, ed.), Ferlach.
En allemand, in German

Arnold W. 1999a. Allgemeine Biologie und Lebensweise des Alpenmurmeltieres (Marmota marmota). Stapfia, 63: 1-20.
En allemand, in German

Arnold W. 1999b. Wildtiere in Naturschutzgebieten - Probleme und Fragen von europäischer Dimension. In: Natura 2000 - eine Chance für den Naturschutz Europas (Schriftenreihe des Österreichischen Bundesministeriums für Umwelt, J. u. F., ed.), pp. 57-58.
En allemand, in German.

Arnold W. 1999c.Winterschlaf des Alpenmurmeltieres (Marmota marmota). In Murmeltiere (Preleuthner, M. & Aubrecht, G., eds.), Stapfia, 63: 43-56. OÖ Landesmuseums, Linz.
En allemand, in German.
Marmota marmota, hibernation.

Arnold W. 1999d. Allgemeine Biologie und Lebensweise des Alpenmurmeltieres (Marmota marmota). In Murmeltiere (Preleuthner, M. & Aubrecht, G., eds.), vol. 63, pp. 1-20. OÖ Landesmuseum, Linz.
En allemand, in German.

Arnold W. 2003. The role of polyunsaturated fatty acids in diet selection and hibernation in alpine marmots [Rôle des acides gras polyinsaturés dans le régime alimentaire et l'hibernation des marmottes alpines]. Hibernation workshop of the European Science Foundation, Marburg/Lahn, Deutchland, 6.10-8.10.
En anglais, in English.
Marmota marmota, marmotte alpine, alpine marmot, acides gras, fatty acides, alimentation, feeding, hibernation.

Arnold W., Bruns U., Frey-Roos F. & Ruf T. 2002. Dietary fatty acids and natural hibernation in alpine marmots. Régime alimentaire en acides gras et hibernation naturelle chez la marmotte alpine. Режтм питания жировыми кислотами и естественная спячка у альпийского сурка. In Abstracts-résumés IVth Marmot World Conference, Ramousse R., Allainé D. & Le Berre M, Eds; International Marmot Network, Lyon, 12-13.
En français et en anglais, in French and in English.
Marmota marmota, essential fatty acids (EFA), acides gras essentiels (AGE), white adipose tissue (WAT), tissu adipeux blanc (TABi), energy expenditure, dépense énergétique.
Nous avons étudié chez la marmotte alpine dans son milieu les effets des acides linoléiques (18:2n-6) et #\-linolénique (18:3n-3) sur la sélection du régime alimentaire, l'hibernation et la dépense d'énergie durant l'hiver. Le site d'étude était localisé dans les Grisons, Suisse, entre 2100-2300 m d'altitude. Les marmottes étaient capturées deux fois par an après la sortie d'hibernation et avant l'entrée en hibernation en automne. Les animaux étaient marqués individuellement, pesés et, en automne, un échantillon tissulaire du dépôt de graisse sous-cutanée inguinal était prélevé chirurgicalement. Pour enregistrer le comportement au cours de l'hibernation, des émetteurs radio de température ont été implantés intrapéritonéalement chez 57 marmottes de 13 groupes sociaux. Les températures corporelles de ces animaux comme les températures ambiantes dans les hibernacula ont été enregistrées en continue durant les hivers 1999 et 2001. La sélection alimentaire a été étudiée en comparant le contenu stomacal de marmottes tuées dans une population soumise à un programme de régulation au cours de trois périodes différentes dans l'année avec les plantes présentes dans le domaine vital d'un animal. Les contenus stomacaux et la nourriture disponible ont été analysés quantitativement pour connaître la composition des espèces végétales, la composition chimique (protéine, lipide, fibre), les acides gras essentiels (AGE) et leur contenu énergétique. Au cours de la saison d'activité, les marmottes se nourrissent sélectivement de plantes, et de parties de plantes aisément digestibles et riche en contenu énergétique. Au cours de la période d'engraissement, les marmottes ont montré, en plus, une préférence claire pour les plantes à fort contenu en 18:2n-6 et un évitement des plantes à forte concentration en 18:3n-3. Avant l'entrée en hibernation, le contenu du tissu adipeux blanc (TABl) en AGE variait considérablement suivant les individus (18:2n-6 : moyenne 8,8 %, étendue : 5,9-13,0 % ; 18:3n-3 : moyenne 24,3 %, étendue 15,1- 33,8 %) avec des conséquences marquées sur l'hibernation suivante. Plus le contenu en 18:2n-6 était élevé plus la température corporelle minimale au cours de la torpeur profonde était basse et plus les périodes de torpeur individuelle au cours de l'hiver étaient longues. La concentration en 18:3n-3 dans le TABl en automne avait, s'il y en avait, des effets contraires. La perte totale de masse durant l'hiver était influencée significativement par le 18:2n- 6 dans une grandeur écologiquement importante. La variation naturelle de la concentration en 18:2n-6 du TABl résultait en des différences dans la perte de masse au cours de l'hibernation jusqu'à 210 g.