Cooperative Sentinel Behaviour in the Arabian Babbler
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ANIMAL BEHAVIOUR, 2001, 62, 973–979 doi:10.1006/anbe.2001.1838, available online at http://www.idealibrary.com on Cooperative sentinel behaviour in the Arabian babbler JONATHAN WRIGHT*, ELENA BERG†, SELVINO R. DE KORT‡, VLADIMIR KHAZIN§ & ALEXEI A. MAKLAKOV§ *School of Biological Sciences, University of Wales, Bangor †Department of Wildlife, Fisheries & Conservation Biology, University of California, Davis ‡Institute of Evolutionary and Ecological Sciences, Leiden University §Department of Zoology, Tel Aviv University (Received 25 September 2000; initial acceptance 2 November 2000; final acceptance 15 July 2001; MS. number: 6698) Cooperative sentinel behaviour in Arabian babblers, Turdoides squamiceps, appears generally consistent with state-dependent models of individually selfish antipredator behaviour. We examined further detailed aspects of this cooperative behaviour, including the suggestion that by engaging in this behaviour sentinels advertise their status and gain social prestige. Chosen sentinel locations were higher, but no more exposed, than the best alternative locations within 25 m. Sentinels started off closer to the centre of the foraging group than when sentinel bouts were terminated. Change-overs between sentinels were nearly always due to previous sentinels terminating their own bout. On the rare occasions when bouts were interrupted by upcoming sentinels, physical contact or aggression was extremely rare. Dominant males tended to terminate sentinel bouts of other birds, interrupting especially the dominant females. Other than this, there was no effect of an individual’s sex or dominance rank on any aspect of sentinel change-overs or location choice when acting as a sentinel. There were also no differences in any sentinel activity between simple family groups, and those more complex nonfamily groups within which individuals compete for reproduction. Rates of alarm calling did not differ between group members, although dominant males did make more territorial calls to neighbouring groups while acting as a sentinel. Therefore, we found relatively little evidence that individuals compete for the chance to act as a sentinel as a means of showing off within groups of Arabian babblers. Instead, our study confirms the primary function of sentinels as a system of cooperative vigilance. 2001 The Association for the Study of Animal Behaviour Sentinel behaviour usually involves one member from a likely that cooperative sentinel effort would need to be cooperative group standing guard in a prominent posi- maintained via mutualistic benefits from investing in tion, while the rest of the group forages in comparative future partners and allies, and from maintaining group safety (Gaston 1977; Rasa 1986, 1989; McGowan & size (i.e. group augmentation: Woolfenden & Fitzpatrick Woolfenden 1989; Zahavi 1990; Clutton-Brock et al. 1978, 1984; Ligon 1981; Connor 1995; Wright 1998). A 1999; Wright et al., in press). A variety of evolutionary recent model has shown that such mutualistic benefits explanations exist for this apparently altruistic behav- and kin selection may not be necessary, however, because iour. The high relatedness within cooperative groups cooperative sentinel behaviour can be evolutionarily suggests kin selection for the purpose of protecting rela- stable when based solely upon short-term individually tives (Hamilton 1964; Maynard Smith 1964; McGowan & selfish decisions by group members depending upon their Woolfenden 1987). Alternatively, regular rotation of internal state (Bednekoff 1997). unrelated sentinels might be maintained via reciprocity For Arabian babblers, Turdoides squamiceps, Zahavi with score keeping (Trivers 1971), although it seems more (1989, 1990; Zahavi & Zahavi 1997) suggested that senti- nel behaviour has evolved as a signal, with individuals Correspondence: J. Wright, School of Biological Sciences, University benefiting by being seen to act as a sentinel and thereby of Wales, Bangor, Gwynedd LL57 2UW, U.K. (email: j.wright@ gaining social prestige. According to this hypothesis, bangor.ac.uk.). E. Berg is at the Department of Wildlife, Fisheries & individuals perform sentinel duties and alarm call in Conservation Biology, University of California, Davis CA 95616, order to show off and obtain social returns in terms of U.S.A. S. De Kort is at the Institute of Evolutionary and Ecological Sciences, Leiden University, NL-2300 RA Leiden, The Netherlands. access to advantageous collaborations and reproductive A. A. Maklakov is at the Mitriani Department of Desert Ecology, Ben opportunities (Zahavi 1995; Wright 1999). This hypoth- Gurion University of the Negev, Midreshet Ben-Gurion, Sede Boqer esis assumes that sentinel behaviour is costly in terms 84990, Israel. of lost opportunities to forage and the predation risk 0003–3472/01/110973+07 $35.00/0 973 2001 The Association for the Study of Animal Behaviour 974 ANIMAL BEHAVIOUR, 62, 5 inherent in taking up an exposed position. Sentinels groups are usually newly formed or recently restructured therefore have to be effective look-outs and spot preda- after the death or replacement of a breeding individual or tors first, because they are the most exposed members of pair and subsequent departure of subordinate group the group. To defend their high-status positions, domi- members of one sex (usually the females). Complex nant group members should thus interfere with the groups contain more than one potential breeder of either competitive sentinel efforts of subordinates, and use allo- sex, these being any bird in a group containing at least feeding and aggression to displace them and take over as one unrelated adult of the opposite sex (close inbreeding sentinels in their place (Zahavi & Zahavi 1997; but see is avoided in this species, with subordinate males com- Wright 1999). peting and obtaining reproduction only in complex State-dependent models of cooperative selfish sentinel groups, see Lundy et al. 1998; Wright et al. 1999). The behaviour (Bednekoff 1997) have recently been sup- remaining two-thirds of groups are ‘simple family ported by observational and experimental evidence from groups’, containing a single breeding pair and their non- meerkats, Surricatta surricata (Clutton-Brock et al. 1999). breeding offspring. Simple family groups tend on average Wright et al. (in press) suggested similar conclusions for to be larger than complex nonfamily groups (Wright et al. Arabian babblers, showing that sentinel effort within 1999), but in the present study there was no significant groups was positively related to dominance rank within difference in size between the two types of group (Wright each sex, and males acted as sentinels more than females et al., in press). of similar rank. Larger groups showed greater overall The study site at Hazeva is a 25-km2 area of desert, sentinel effort, but with less effort per individual as only located 30 km south of the Dead Sea in the Arava rift one sentinel was on duty at any one time. All of these valley in southern Israel. Twenty groups of Arabian effects on sentinel behaviour in Arabian babblers were babblers have been studied continuously since 1971 at irrespective of relatedness and group social structure, and Hazeva by Amotz Zahavi and students from Tel Aviv could be explained by differences in body mass (i.e. University. Between 1992 and 1997, up to 40 groups were ‘state’) within and between individual birds (see Wright monitored on a weekly basis and habituated to human et al., in press). In addition, experimental food supple- observers via occasional hand feeding. All birds were mentation of individual babblers created similar increases individually colour ringed and their family histories in sentinel effort, again related simply to differences in known. Dominance rank within each sex was assigned body mass and not social dominance (Wright et al. 2001). over many observations, involving the direction of sup- Although generally supportive of the more straight- plants and/or outcomes of aggressive interactions over forward antipredator function of sentinel behaviour, provisioned food (e.g. pieces of bread). These behavioural these studies do not necessarily discount an additional observations regarding dominance were confirmed and more complex social role for it. Along with social through genetic analysis of group structures and access to prestige, sentinel behaviour might be used in competitive reproductive opportunities (see Lundy et al. 1998; Wright interactions within and between groups. By keeping et al. 1999). a look out, dominants might be more likely to spot a Hazeva is a very open habitat with sparse vegetation neighbouring group, or be more able to monitor the rest lining the bottom of dry river beds, comprising well- of the group (e.g. for the purposes of mate guarding). spaced trees (Acacia spp.) and a few low bushes and Sentinels may also gain foraging advantages as sit-and- annual plants. However, thicker Tamarix nilotica scrub wait predators (Rasa 1983; Munn 1986; McGowan & and reed beds (Phragmytes spp.) exist in one area receiving Woolfenden 1989). regular outflows of water from human settlements and In this study we examined additional aspects of senti- agriculture. nel behaviour. These involved rates of alarm calling and other vocalizations by sentinels, the nature of change- Data Collection overs between successive sentinels and the types of sen- tinel locations used relative to alternative locations and We collected data throughout the year from the foraging group. Our aim was