Abstract: In many cooperatively breeding species, group members form a dominance hierarchy or queue to inherit the position of breeder. Models aimed at understanding individual variation in helping behavior, however, rarely take into account the effect of dominance rank on expected future reproductive success and thus the potential direct fitness costs of helping. Here we develop a kin-selection model of helping behavior in multimember groups in which only the highest ranking individual breeds. Each group member can invest in the dominant's offspring at a cost to its own survivorship. The model predicts that lower ranked subordinates, who have a smaller probability of inheriting the group, should work harder than higher ranked subordinates. This prediction holds regardless of whether the intrinsic mortality rate of subordinates increases or decreases with rank. The prediction does not necessarily hold, however, where the costs of helping are higher for lower ranked individuals: a situation that may be common in vertebrates. The model makes two further testable predictions: that the helping effort of an individual of given rank should be lower in larger groups, and the reproductive success of dominants should be greater where group members are more closely related. Empirical evidence for these predictions is discussed. We argue that the effects of rank on stable helping effort may explain why attempts to correlate individual helping effort with relatedness in cooperatively breeding species have met with limited success.

Abstract: Mobbing signals advertise the location of a stalking predator to all prey in an area and recruit them into the inspection aggregation. Such behavior usually causes the predator to move to another area. However, mobbing calls could be eavesdropped by other predators. Because the predation cost of mobbing calls is poorly known, we investigated whether the vocalizations of the mobbing pied flycatcher, Ficedula hypoleuca, a small hole nesting passerine, increase the risk of nest predation. We used mobbing calls of pied flycatchers to examine if they could lure predators such as the marten, Martes martes. This predator usually hunts by night and may locate its mobbing prey while resting nearby during the day. Within each of 56 experimental plots, from the top of one nest-box we played back mobbing sounds of pied flycatchers, whereas blank tapes were played from the top of another nest-box. The trials with mobbing calls were carried out before sunset. We put pieces of recently abandoned nests of pied flycatchers and a quail, Coturnix coturnix, egg into each of the nest-boxes. Nest-boxes with playbacks of mobbing calls were depredated by martens significantly more than were nest-boxes with blank tapes. The results of the present study indicate that repeated conspicuous mobbing calls may carry a significant cost for birds during the breeding season.

Abstract: Neurobiological and behavioral research indicates that place learning and response learning occur simultaneously, in parallel. Such findings seem to conflict with theories of associative learning in which different cues compete for learning. The authors conducted place+response training on a radial maze and then tested place learning and response learning separately by reconfiguring the maze in various ways. Consistent with the effects of manipulating place and response systems in the brain (M. G. Packard & J. L. McGaugh, 1996), well-trained rats showed strong place learning and strong response learning. Three experiments using associative blocking paradigms indicated that prior response learning interferes with place learning. Blocking and related tests can be used to better understand how memory systems interact during learning.

Abstract: (Accepted: 23 June 2005)
Summary
Much research in animal communication is aimed at understanding the functional design
features of animal vocal signals. Our detailed analyses of the vocalizations and behavioural
responses elicited in white-faced capuchins by predators and other disturbances point to two
call variants that differ modestly in their acoustic structure and that are accompanied by
functionally distinct behavioural responses. The first variant is given exclusively to avian
predators and is almost invariably accompanied by the monkeys immediate descent from
the treetops where it is most vulnerable; therefore, we label this call variant the aerial
predator alarm?. The second variant, that differs only slightly but noticeably from the first,
is given to a wide range of snakes and mammals, including a range of species that represent
no predatory threat to the monkeys. This second call is also associated with more variable
responses from calling monkeys, from delayed retreat from the source of disturbance, to
active approach, inspection, and sometimes mobbing of the animal involved. We therefore
label this variant more generally as an “alerting call”. Although some other primate species
show a more diverse system of anti-predator calls, and the capuchins themselves may yet
be found to produce a greater variety of calls, a system of two call variants with varying
degrees of predator specificity and behavioural response is not uncommon among primates
and appears functionally appropriate for capuchins. The basic structure of the alerting call
allows conspecific listeners to localize the caller and the source of disturbance readily, thereby
allowing listeners to approach and assist in mobbing in cases where the disturbance warrants
it, or to avoid the area in cases where the disturbance is identified as a predatory threat.
Conversely, the aerial predator alarm is inherently less localizable and therefore conveys the

Abstract: Biological market models explain variability in reciprocity and interchange between groups. In groups with a shallow dominance gradient, grooming will be mostly exchanged for itself (i.e. exchange will occur). In groups with steep dominance hierarchies, interchange is expected: individuals will groom higher ranking individuals to get access to limited resources or commodities such as support in conflicts, and grooming will be traded for these commodities. We examine patterns of reciprocity in grooming and support, and of interchange of grooming for support or for tolerance in six captive groups of bonobos. We test whether differences between groups in patterns of reciprocity and interchange can be attributed to differences in a measure of steepness of dominance hierarchies, which is based on dyadic agonistic interactions. We found that grooming was reciprocal in some, but not all groups. Support was highly reciprocal, but this was a side effect of dominance in most groups. Interchange between grooming and support was observed in some groups. Corroborating earlier findings, this was a side effect of individuals preferring high ranking individuals as grooming and support partners, possibly because these high-ranking individuals provide more efficient support in conflicts. There was no evidence for interchange of grooming for tolerance. Variability in grooming reciprocity was explained by differences in steepness of dominance hierarchies, as predicted by the biological market models. In groups with a shallow dominance hierarchy, grooming was more reciprocal. This was not true for reciprocity in support. There was some evidence that individuals groomed dominants more frequently in groups with a steep dominance hierarchy. The variation in interchange relations between grooming and support did not depend on the steepness of dominance hierarchies. We suggest that grooming in itself is a valuable commodity in bonobos, especially under captive conditions, which can be exchanged reciprocally. Bonobos may interchange grooming for another value equivalent, with food sharing as a very likely candidate. This interchange effects seem more dependent on potential to monopolise food than on steepness of dominance hierarchies per se.