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Walia Special Edition on the Bale Mountains (2011) - Zoologische ...

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No new packs were formed between 1998 and 1992 (Sillero-Zubiri and Gottelli 1995b)<br />

but an apparent attempt by a subordinate female to split a pack suggested that fissi<strong>on</strong> could be a<br />

mechanism for pack formati<strong>on</strong> (this ended when <strong>the</strong> subordinate’s litter succumbed, probably killed<br />

by <strong>the</strong> dominant female) (Sillero-Zubiri et al. 1996a). During this period female ascendancy to<br />

breeding status, ei<strong>the</strong>r by immigrati<strong>on</strong> or inheritance, <strong>on</strong>ly occurred after <strong>the</strong> death of a dominant<br />

and so <strong>the</strong> chances of a female ever securing a breeding positi<strong>on</strong> were low. Five out of 10 dominant<br />

females retained <strong>the</strong>ir breeding positi<strong>on</strong> throughout four years of observati<strong>on</strong>, whereas <strong>the</strong> remainder<br />

maintained that role until <strong>the</strong>y died. Breeding openings occurred at an average of 0.12 ± 0.09<br />

opportunities for a subordinate female per year per pack. During c<strong>on</strong>tests for a breeding positi<strong>on</strong>,<br />

resident females appeared to have an advantage over floaters (three breeding females were replaced<br />

by <strong>the</strong>ir daughters after <strong>the</strong>ir deaths).<br />

In late 1991 a rabies epidemic decimated <strong>the</strong> populati<strong>on</strong> in <strong>Bale</strong> and resulted in <strong>the</strong><br />

disintegrati<strong>on</strong> of three out of five packs in Web, causing <strong>the</strong> sudden opening of potential breeding<br />

opportunities (Sillero-Zubiri et al. 1996b; similar epidemics took place in 2003 and 2008, see<br />

Randall et al. 2004 and Johns<strong>on</strong> et al. 2009). Ra<strong>the</strong>r than forming smaller, new breeding units,<br />

<strong>the</strong> surviving packs maintained <strong>the</strong>ir social cohesi<strong>on</strong> and readjusted <strong>the</strong>ir territorial boundaries to<br />

occupy <strong>the</strong> habitat available (Marino 2003b). After a new epidemic in 2003, seven out of nine packs<br />

survived, and again packs remained cohesive and territory boundaries shifted (Tallents 2007). It<br />

took five years after <strong>the</strong> first die-off for new packs to start forming, in <strong>on</strong>e case by <strong>the</strong> fissi<strong>on</strong> of a<br />

large pack, and twice by <strong>the</strong> grouping of dispersing individuals, mostly from neighbouring packs.<br />

This suggested that <strong>the</strong> establishment of a new group depends not <strong>on</strong>ly <strong>on</strong> <strong>the</strong> availability of highquality<br />

habitat but also <strong>on</strong> <strong>the</strong> presence of sufficient number of helpers to defence <strong>the</strong> new territory<br />

from <strong>the</strong> ‘expansi<strong>on</strong>ist’ packs that survived (Marino 2003b). The populati<strong>on</strong> reducti<strong>on</strong> also opened<br />

breeding vacancies to subordinate females, but social factors delaying formati<strong>on</strong> of new breeding<br />

units maintained populati<strong>on</strong> growth low at low densities (Marino 2003b; Marino et al. 2006).<br />

Am<strong>on</strong>g Ethiopian wolves, inbreeding avoidance may be an additi<strong>on</strong>al adaptive advantage<br />

of female-biased dispersal and may underlie <strong>the</strong> observed mating behaviour. Male philopatry and<br />

<strong>the</strong> l<strong>on</strong>g tenure of breeding females increase <strong>the</strong> potential for incest within groups, which may be<br />

countered in part by female dispersal (Sillero-Zubiri et al. 2004). A detailed genetic study of Ethiopian<br />

wolves also found that although mean pairwise relatedness within packs (R = 0.39) was significantly<br />

greater than that estimated from random assignment of individuals to packs, breeding pairs were<br />

most often unrelated (Randall et al. 2007). This str<strong>on</strong>gly suggests that female-biased dispersal also<br />

reduces <strong>the</strong> number of incestuous matings in <strong>the</strong> populati<strong>on</strong>. However inbreeding is not entirely<br />

absent in <strong>the</strong> populati<strong>on</strong>. Kinship analysis showed that 11 of 15 (73%) of successful matings were<br />

between unrelated individuals (R = -0.26 to 0.22, P < 0.05). Four apparently incestuous matings<br />

resulted in 9 of 47 (19%) pups being potentially inbred (Randall et al. 2007). Given <strong>the</strong> potential<br />

negative c<strong>on</strong>sequences of inbreeding for <strong>the</strong> l<strong>on</strong>g-term genetic and demographic viability of <strong>the</strong><br />

populati<strong>on</strong>, inbreeding should be m<strong>on</strong>itored in this populati<strong>on</strong>, particularly in <strong>the</strong> face of fur<strong>the</strong>r<br />

habitat loss and/or disease outbreaks.<br />

<str<strong>on</strong>g>Walia</str<strong>on</strong>g>-<str<strong>on</strong>g>Special</str<strong>on</strong>g> <str<strong>on</strong>g>Editi<strong>on</strong></str<strong>on</strong>g> <strong>on</strong> <strong>the</strong> <strong>Bale</strong> <strong>Mountains</strong> 69

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