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Evolution of Sociality in Marmots Author(S): Kenneth B American Society of Mammalogists Evolution of Sociality in Marmots Author(s): Kenneth B. Armitage Source: Journal of Mammalogy, Vol. 80, No. 1 (Feb., 1999), pp. 1-10 Published by: American Society of Mammalogists Stable URL: http://www.jstor.org/stable/1383202 Accessed: 05/10/2009 12:50 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=asm. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Mammalogists is collaborating with JSTOR to digitize, preserve and extend access to Journal of Mammalogy. http://www.jstor.org EVOLUTION OF SOCIALITY IN MARMOTS KENNETHB. ARMITAGE Department of Systematics and Ecology, The University of Kansas, Lawrence, KS 66045-2106 C. Hart Merriam Presentation Presented at the 78th Annual Meeting of the American Society of Mammalogists Virginia Polytechnic Institute and State University Blacksburg, VA 6-10 June 1998 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, subordinate 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 immergence 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 immer- gence 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 a 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 age 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 al- loparental care. Subordinate adults also may reproduce or succeed to territorial status. Al- loparental care occurs during hibernation when subordinate adults assist in social thermo- regulation of closely related young. Thus, marmots have the characteristics of cooperative breeding. The following sequence of events is hypothesized to have occurred in the evo- lution 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. Key words: Marmota, delayed dispersal, sociality, mass, delayed reproduction, mass: length ratios. Sociality may be viewed as a life-history gate to exploit a resource or to reduce pre- tactic, that, in the general sense, increases dation risk (Alexander, 1974). Once social the fitness of group members (Armitage, groups are formed, behaviors such as alarm 1981). The environmental factors that in- calling (Blumstein et al., 1997; Sherman, teract with other life-history traits to select 1997) and alloparental care (Arnold, 1993) for sociality are not always understood. can evolve. However, sociality also has Generally, animals are expected to aggre- costs; time to first reproduction is increased, Journal of Mammalogy, 80(1):1-10, 1999 1 2 JOURNAL OF MAMMALOGY Vol. 80, No. I a smaller proportion of females breed, and of a reproductive territorial pair, subordi- litter size is reduced as social complexity nate adults, yearlings, and young. The mat- increases (Blumstein and Armitage, 1998). ing system is monogamous, but polyandry Thus, sociality is associated with consider- may occur in some species; e.g., M. mar- able reproductive skew (Blumstein and Ar- mota (Arnold, 1990a), M. bobac, and M. mitage, in press; Keller and Reeve, 1994; baibacina (Rymalov, 1994). Sherman et al., 1995). Presumably the ben- This diversity of social systems raises the efits of sociality outweigh the costs, but the question of what factors led to their evo- benefits and costs differ widely among in- lution. Before we can frame hypotheses for dividuals (e.g., Armitage, 1986; Clutton- future testing, we must consider two critical Brock et al., 1988; Faulkes and Abbott, features of marmot biology. First, marmots 1997). The nature of the ecological con- are relatively large-they are the largest straints that lead some individuals to accept true hibernator. Thus, much of their biology reduced reproductive success as a conse- revolves around the requirement of suc- quence of living in a group and the behav- cessful hibernation. There is also consider- ioral, physiological, and ecological deter- able variation in body size; the body mass minants of reproductive success within at emergence from hibernation of the larg- groups remain major questions of social bi- est species, M. bobak, is 1.76 times that of ology (e.g., Emlen, 1997; Solomon and the smallest species, M. flaviventris. Sec- French, 1997). ond, they have a short active season ranging Marmots (Marmota) are an excellent tax- among species from about 3.5 to 7.5 on for studying the evolution of sociality. months (X = 4.8 months). During this lim- Fourteen recognized species are distributed ited time they must reproduce, grow, and across the northern hemisphere (Barash, prepare for hibernation. The interaction of 1989). Because they are diurnal and bur- these two factors establishes the basis for row-dwelling, they can be easily located for the evolution of sociality. trapping, marking, and observation. Most BODYMASS importantly, they have a variety of social systems. Current knowledge suggests that What accounts for the diversity of body marmot species may be placed in one of mass? Because marmots do not store food, four social systems (Armitage, 1996). Only one possibility is that body mass is related the woodchuck (Marmota monax) leads a to hibernation; e.g., those species that have solitary existence; the mating system is po- longer periods of hibernation lose more lygynous. Yellow-bellied marmots (M. fla- mass. Note that hibernation period refers to viventris) form female kin groups, with as- the time from disappearance above ground sociated young (less than one year old) and in the autumn to emergence from the bur- yearlings, that may persist through time as row the following spring. Hibernation re- matrilines. An adult male resides with one fers to the period when characteristic bouts or more matrilines to produce a harem-po- of torpor occur. This suggestion is not sup- lygynous mating system (Armitage, 1998). ported as there is no correlation among spe- The social system of the Olympic (M. olym- cies between body mass or mass loss during pus) and hoary (M. caligata) marmots con- hibernation and the length of the hiberna- sists of an adult male with two females and tion period (Armitage and Blumstein, in a litter of yearlings or young; the mating press). However, immergence mass (body system is bigamous, but some populations mass at entry into hibernation) is highly of the hoary marmot are monogamous correlated (r = 0.91) with mass loss during (Holmes, 1984). As far as is known, all oth- hibernation. These relationships suggest er species are characterized as living in that marmots are large because they use family groups (Bibikow, 1996) consisting considerable mass during the hibernation 1999 ARMITAGE--C.HART MERRIAM PRESENTATION 3 7400 6900 Y = 2064.4 + 14.87 X soly P= 0.008 R2 = 0.49 6400 sehim CO O ooal *bob E 5900 S 5400 vanba E mare E 4900 S*cam emon 4400 3900 - *sib ecau *men 3400 1- I fla 350 400 450 500 550 Head and body length (mm) FIG. 1.-The relationshipbetween immergencemass and head and body length for 13 species of marmots. bai = M. baibacina, bob = M. bobac, cal = M. caligata, cam = M. camtschatica, cau = M. caudata, fla = M. flaviventris, him = M. himalayana, mar = M. marmota, men = M. menzbieri, mon = M. monax, oly = M. olympus, sib = M. sibirica, van = M. vancouverensis. Body mass for M. bobak from Mashkin, 1997; for M. menzbierifrom Mashkin and Baturin, 1993; for M. marmota from Mann et al., 1993; for all other species from Armitageand Blumstein, in press. Head and body length measurementsfor M. flaviventris and M. monax from specimens in the Kansas University Natural History Museum; for M.
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