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The Round-Eared Sengi and the Evolution of Social Monogamy: Factors That Constrain Males to Live with a Single Female M

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The Round-Eared Sengi and the Evolution of Social Monogamy: Factors That Constrain Males to Live with a Single Female M Trinity University Digital Commons @ Trinity Biology Faculty Research Biology Department 2009 The Round-Eared Sengi and the Evolution of Social Monogamy: Factors That Constrain Males to Live with a Single Female M. Schubert N. Pillay David O. Ribble Trinity University, [email protected] C. Schradin Follow this and additional works at: https://digitalcommons.trinity.edu/bio_faculty Part of the Biology Commons Repository Citation Schubert, M., Pillay, N., Ribble, D. O., & Schradin, C. (2009). The oundr -eared sengi and the evolution of social monogamy: Factors that constrain males to live with a single female. Ethology, 115(10), 972-985. doi:10.1111/j.1439-0310.2009.01684.x This Article is brought to you for free and open access by the Biology Department at Digital Commons @ Trinity. It has been accepted for inclusion in Biology Faculty Research by an authorized administrator of Digital Commons @ Trinity. For more information, please contact [email protected]. Ethology The Round-Eared Sengi and the Evolution of Social Monogamy: Factors that Constrain Males to Live with a Single Female Melanie Schubert* , Neville Pillay , David O. Ribbleà & Carsten Schradin § * Department of Animal Physiology, University of Bayreuth, Bayreuth, Germany School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Wits, South Africa à Department of Biology, Trinity University, San Antonio, TX, USA § Department of Animal Behaviour, Zoological Institute, University of Zurich, Zurich, Switzerland Correspondence Abstract Melanie Schubert, Department of Animal Physiology, University of Bayreuth, Animal dispersion in space and time results from environmental pres- Universita¨ tsstraße 30, NWI, 95440 Bayreuth, sures, and affects the outcome of a species’ social organization. When Germany. E-mail: [email protected] females are solitary, males may either roam or be pair-living. We stud- ied possible environmental influences affecting the social organization of Received: January 27, 2009 the round-eared sengi (Macroscelides proboscideus) in a semi-desert in Initial acceptance: March 25, 2009 South Africa, using trapping and radio-tracking across 2.5 yr. Adult sex Final acceptance: June 11, 2009 (J. Wright) ratios did not deviate from 1:1 and we found no indication of sexual dimorphism in body mass. Females maintained exclusive areas, which doi: 10.1111/j.1439-0310.2009.01684.x had little overlap (<4%) with neighbouring females (NF), and males overlapped predominately only with the home range of single females. Generally, inter- and intra-sexual overlap with neighbouring individuals was low (3–6%) for both sexes, indicating territoriality and pair-living. Pairs were perennial and territories were maintained year-round. How- ever, males generally maintained much larger areas than females, which were sensitive to population density. Male space use appeared to be primarily limited by the presence of neighbouring males. Female home ranges were smaller-sized despite changes in population density, possi- bly for energetic efficiency. Some paired males attempted to take over widowed females, but shifted back to their original home range follow- ing the intrusion of an un-paired male. We conclude that social mono- gamy is the predominant social organization in round-eared sengis in a semi-desert that may have resulted from females living solitarily in small exclusive territories, balanced sex ratios, and from a low variation of body mass between males. Brotherton 1997; Brotherton & Komers 2003). How- Introduction ever, social systems are rarely attributable to a single Animal dispersion in space and time results from factor (Sandell & Liberg 1992), and female dispersion environmental pressures and affects the outcome of per se is insufficient to account for the evolution a species’ social organization (Brown & Orians of monogamy, since males could opt for other 1970). In species lacking paternal care, dispersed tactics, such as roaming (Kraus et al. 2003; Eberle & living females present an important prerequisite for Kappeler 2004; Martin & Martin 2007). Thus, other the evolution of social monogamy, i.e. pair-living, factors may constrain males into socially monoga- because for males, the chances to encounter other mous relationships. Low population densities have potential mating partners are reduced (Komers & been emphasized in some species, like prairie voles 972 Ethology 115 (2009) 972–985 ª 2009 Blackwell Verlag GmbH M. Schubert et al. Social Monogamy in the Round-Eared Sengi (Microtus ochrogaster, Getz et al. 2003) or Mentawai organization of the round-eared sengi in a semi- snub-nosed langurs (Simias concolor, Watanabe desert, by testing for ecological and physical corre- 1981), and balanced adult sex ratios have been lates of social organization. The following predictions acknowledged to drive the occurrence of social were made: monogamy in other mammalian species, such as Firstly, we investigated space use of female round- Townsend’s voles (M. townsendii, Lambin & Krebs eared sengis. Environmental factors that influence 1991) and oribis (Ourebia ourebi, Adamczak & Dun- female space use ultimately influence the social bar 2007). organization of a population, because male space use Social monogamy is rare in mammals (Kleiman is affected by female distribution in space and time 1977), but is believed to occur in all species of a (Emlen & Oring 1977). We predicted that females unique order, the sengis (Macroscelidea, Ribble & live independently of each other, thereby decreasing Perrin 2005; Rathbun & Rathbun 2006). Sengis (or male monopolization potential for several females. If elephant-shrews) represent an ancient monophyletic males employ a roaming strategy, we suggest that clade with an early radiation from the Eutheria they will maintain much larger home ranges than (Corbet & Hanks 1968), and comprise 17 species from females in order to search widely for fertile females four genera that are all endemic to the African conti- in breeding season (Michener & McLean 1996), and nent. The assumption that all sengi species are that male home ranges will be characterized by large monogamous is based on field studies of six sengis overlap with male competitors (Sandell 1989; species (Rathbun 1979; FitzGibbon 1995, 1997; Leirs Gliwicz 1997). Alternatively, males may monopolize et al. 1995; Neal 1995; Ribble & Perrin 2005; single females resulting in pair-living. In this situa- Rathbun & Rathbun 2006). The social organization tion, male space is predicted to be similar to that of of sengis in these studies has been determined by females, i.e. pairs use similar-sized areas that have investigating space use predominately, but to date, little intra- and inter-sexual overlap with neighbour- detailed studies regarding environmental parameters ing individuals of both sexes (Komers & Brotherton related to the social organization are absent. 1997). In the present study, we investigated potential Secondly, given that males may roam, we expected ecological and physical parameters affecting the that round-eared sengis lack sexual dimorphism, social system in the round-eared sengi (Macroscelides because male body mass is not necessarily related to proboscideus), a small-bodied (35 g) omnivorous the roaming ability for female mates (Schwagmeyer & mammal (Sauer 1973; Kerley 1995), which is found Woonter 1986). Alternatively, the lack of sexual in the more arid regions of South Africa, Namibia dimorphism may also be a characteristic of pair-living and Botswana (Skinner & Smithers 1990). In con- (Kleiman 1977). trast to other sengi species, individual round-eared Thirdly, we studied the male searching efficiency sengis occupy undefended home ranges, reaching for female mates by documenting adult sex ratios in over 100 ha and resulting in a solitary life style with round-eared sengis. Since sex ratios co-evolve with non-territorial females and roaming males (Sauer & social systems, we assume that sex ratios will be Sauer 1971, 1972; Sauer 1973). In Sauer’s studies, more female biased promoting a male roaming tactic, the habitat was characterized by low food abundance because of a high searching efficiency for female and few shelter sites for individuals, and associated mates (Sandell & Liberg 1992; but see Eberle & with an extremely low population density of one Kappeler 2004). In contrast, low searching efficiency individual per 100 ha and an irregular dispersion caused by balanced adult sex ratios may favour the of round-eared sengis, resulting in small isolated monopolization of single females (Sandell & Liberg populations. 1992). Generally, population density of round-eared sen- Fourthly, we determined the length of breeding gis is positively correlated with cover (Joubert & season and synchronization of female receptivity. Ryan 1999) and food availability (van Deventer & Generally, if females reproduce asynchronously it is Nel 2006). Since Sauer’s study was conducted in a more likely that males will adopt a roaming strategy, desert and the present study in a semi-desert, demo- since this provides the opportunity to obtain matings graphical differences between the two study sites, with multiple females (Ims 1987; Ostfeld 1990). which reflect the differential availability of key Asynchronous breeding may also intensify male– resources, may promote different social organizations male competition because it increases the costs of (Lott 1984; Schradin & Pillay 2005a). Thus the aim territorial defence and decreases the monopoliza- of the current study was to determine
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