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The Effects of Population Density and Sociality on Scent Marking in the Yellow Mongoose

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The Effects of Population Density and Sociality on Scent Marking in the Yellow Mongoose Le Roux, A; Cherry, M I; Manser, M B (2008). The effects of population density and sociality on scent marking in the yellow mongoose. Journal of Zoology, 275(1):33-40. Postprint available at: http://www.zora.uzh.ch University of Zurich Posted at the Zurich Open Repository and Archive, University of Zurich. Zurich Open Repository and Archive http://www.zora.uzh.ch Originally published at: Journal of Zoology 2008, 275(1):33-40. Winterthurerstr. 190 CH-8057 Zurich http://www.zora.uzh.ch Year: 2008 The effects of population density and sociality on scent marking in the yellow mongoose Le Roux, A; Cherry, M I; Manser, M B Le Roux, A; Cherry, M I; Manser, M B (2008). The effects of population density and sociality on scent marking in the yellow mongoose. Journal of Zoology, 275(1):33-40. Postprint available at: http://www.zora.uzh.ch Posted at the Zurich Open Repository and Archive, University of Zurich. http://www.zora.uzh.ch Originally published at: Journal of Zoology 2008, 275(1):33-40. The effects of population density and sociality on scent marking in the yellow mongoose Abstract We investigated scent marking behaviour in the yellow mongoose Cynictis penicillata, focusing on a low-density population where all offspring dispersed upon reaching sexual maturity. Dominant males appeared to be the main territory defenders and demarcators, with offspring foraging and marking only near the territory cores. The cheek-marking rates of dominant males increased during the breeding season and may have been involved in olfactory mate guarding. We compared our low-density population with a high-density population displaying natal philopatry. The two populations differed markedly in terms of individual contributions to territorial marking, as subordinate group members in the low-density population performed almost no territorial marking or defence, but were the primary scent-markers and territory defenders in the high-density population. We discuss scent-marking distinctions between populations in the context of ecological and social differences. 1 Journal of Zoology (2008), 275: 33-40 2 doi: 10.1111/j.1469-7998.2007.00404.x 3 4 5 The effects of population density and sociality on scent marking in the yellow 6 mongoose 7 8 Aliza Le Roux1, Michael I. Cherry1, Marta B. Manser2 9 10 1. Department of Botany and Zoology, University of Stellenbosch, South Africa 11 2. Animal Behaviour, Department of Zoology, University of Zurich 12 13 14 We investigated scent marking behaviour in the yellow mongoose Cynictis penicillata, 15 focusing on a low-density population where all offspring dispersed upon reaching sexual 16 maturity. Dominant males appeared to be the main territory defenders and demarcators, 17 with offspring foraging and marking only near the territory cores. The cheek-marking 18 rates of dominant males increased during the breeding season and may have been 19 involved in olfactory mate guarding. We compared our low-density population with a 20 high-density population displaying natal philopatry. The two populations differed 21 markedly in terms of individual contributions to territorial marking, as subordinate group 22 members in the low-density population performed almost no territorial marking or 23 defence, but were the primary scent-markers and territory defenders in the high-density 24 population. We discuss scent-marking distinctions between populations in the context of 25 ecological and social differences. 26 27 Keywords: mate guarding, population density, scent marking, territory, yellow 28 mongoose 1 1 Introduction 2 3 Scent-marking forms an integral part of the communicative repertoire of many 4 mammals (Eisenberg & Kleiman, 1974; Gorman & Trowbridge, 1989). Marking rates 5 typically increase around the onset of puberty (e.g. Woodmansee et al., 1991), and many 6 mammals are able to discriminate between individuals' scents (Swaisgood, Lindburg & 7 Zhou, 1999; Mendl, Randle & Pope, 2002; Mateo, 2006). In solitary animals, age, 8 gender and territory ownership appear to be the main determinants of scent-marking rates 9 (e.g. in honey badgers Mellivora capensis Begg, du Toit & Mills, 2003), whereas social 10 mammals are, additionally, affected by position in the dominance hierarchy, relative 11 contributions to territory defence, and the frequency of aggressive interactions (e.g. 12 coyotes Canis latrans Gese & Ruff, 1997). Although the dominant male in a group is 13 usually the primary scent marker, subordinate adult group members often contribute 14 substantially to marking and defence (Gese & Ruff 1997; Jordan 2007). Additionally, 15 males may overmark females’ scent, as a form of olfactory mate guarding (e.g. Brashares 16 & Arcese, 1999) or other males’ scents, indicating intrasexual dominance (e.g. Rich & 17 Hurst, 1999). Within a social group, allomarking – marking other individuals – often 18 occurs, probably to maintain a ‘familiar’ group smell and tolerance between group 19 members (e.g. in European badgers Meles meles Buesching, Stopka & MacDonald, 20 2003). 21 Olfactory communication is involved in a variety of functions as diverse as dominance 22 advertisement (e.g. in guinea pigs, Carla porcellus, Drickamer & Martan, 1992) and the 23 marking of food caches (e.g. in in the short-tailed shrew, Blarina brevicauda, Robinson 2 1 & Brodie, 1982). However, in solitary as well as social mammals, scent marks are usually 2 implicated in two primary functions, i.e. territory demarcation and sexual advertisement. 3 Territory holders typically mark and actively defend their home ranges (Ralls, 1971). 4 Scent marks serve as information exchange points (Eisenberg & Kleiman, 1974), 5 containing, inter alia, information on the ability of an owner to defend its territory (Rich 6 & Hurst, 1999). During the breeding season, this information may include indications of 7 oestrus in females (e.g. in kangaroo rats, Dipodomys spectabilis, Randall, 1986), and 8 males may also increase signaling during this season as a form of sexual advertisement 9 (Kappeler, 1998). In some species, the male may attempt to mask the female’s sexual 10 advertisement by overmarking her scent marks, thereby performing mate guarding (e.g. 11 Roberts & Dunbar, 2000; Lewis, 2005). 12 13 As there may be great intraspecific variation in territory size, group composition 14 and degree of sociality in vertebrates (Lott, 1991), scent marking behaviour within a 15 species is potentially variable. For example, the scent marking strategies of hyenas can be 16 strongly affected by territory size. Whereas marks were concentrated in the territory core 17 of large territories (the ‘hinterland’ marking strategy), they were concentrated along the 18 borders (border marking strategy) of smaller territories (Gorman & Mills, 1984). These 19 distinct strategies are seen as the most economical means of marking a territory, ensuring 20 that intruders will come across a scent mark before reaching the protected territory core. 21 However, relatively few studies have performed intraspecific comparisons of scent 22 marking behaviour between populations with different territory sizes, and in particular we 3 1 found no intraspecific comparisons between populations that show different social 2 structures. 3 4 The yellow mongoose Cynictis penicillata dens together with conspecifics in 5 groups ranging between two and 13 individuals in size, but individuals typically forage 6 alone or in pairs (Rasa et al., 1992; Cavallini, 1993). Females are polyestrous, 7 occasionally giving birth to two litters per season, and young typically disperse during 8 spring (Rasa et al., 1992). Similar to many mammals (Ralls, 1971; Gorman & 9 Trowbridge, 1989), the yellow mongoose is territorial and uses various forms of marking 10 in its territory (Earlé, 1981; Wenhold & Rasa, 1994). Yellow mongoose mark vegetation 11 and other prominent objects using anal marks, cheek marks (‘cheek wipes’) and body 12 rubs (‘sidewipes’) (after Earlé, 1981; Wenhold & Rasa, 1994). Body rubbing appears to 13 be a form of self-anointing with odours (including own scent marks) from the 14 environment, rather than the actual deposition of scent, as yellow mongooses lack scent 15 glands on their flanks (Pocock, 1916). Urination and defecation are considered to be 16 secondary forms of marking (e.g. Wenhold & Rasa, 1994), and we have not included the 17 analysis of defecation and urination in this study as there was no evidence of a primary 18 communicative function (le Roux 2007). 19 20 Scent marking in the yellow mongoose has been described in varying degrees of 21 detail in populations of intermediate to high densities [23 – 26 individuals per km2 22 (Balmforth, 2004) to 133 – 200 individuals per km2 (Earlé, 1981; Wenhold & Rasa, 23 1994)]. Of these studies, only Wenhold and Rasa’s (1994) quantified the marking 4 1 behaviour of a group of mongooses (n = 13 group members) and tested specific 2 hypotheses. They presented individuals' marking rates as averages over nine months 3 (April – December) (Wenhold & Rasa, 1994). Whereas Wenhold and Rasa (1994) 4 showed that subordinate adults were the primary territory defenders and markers, Earlé 5 (1981) found in the same Big Island (BI) population in the Vaal Dam (26º52’S, 28 6 º11’E), South Africa, that the dominant males played the main role in this respect. 7 8 We focused our research on a low-density population of yellow mongooses at the 9 Kuruman River Reserve (KRR) in South Africa. The mated pair constituted the only 10 adult members of each group and offspring were not involved in raising new litters but 11 dispersed on reaching sexual maturity. Similar to other low-density populations 12 (Cavallini, 1993), aggression between family members and neighbours was low. In 13 contrast, groups in the BI population consisted of the mated pair, recent offspring and 14 related adults that cooperated in the rearing of young and aggressive territory defence 15 (Earlé, 1981; Wenhold, 1990; Wenhold & Rasa, 1994). 16 17 Here we describe the scent marking behaviour of the KRR population and 18 compare it with the scent marking behaviour of the BI population (Wenhold & Rasa, 19 1994).
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