Robert D. Magrath

HATCHING ASYNCHRONY IN ALTRICIAL BIRDS

1. The review aims to provide a simple conceptual framework on which to place recent studies of hatching asynchrony in altricial birds and to assess the evidence used in support of specific hypotheses.

Seasonal changes in clutch size in British birds

In many species of birds the mean clutch size is not constant through the breeding season: Some species show a seasonal pattern of declining clutch size, while others can show a mid-season peak. We suggest an evolutionary interpretation of the different patterns of seasonal change in birds, and test predictions by analysis of nest-record data held by the British Trust for Ornithology (BTO). We assume that conditions for breeding will show a seasonal pattern of improvement and then decline, and that the optimal clutch size (Lack 1947) is larger for each individual in the population when the conditions for breeding are better

Interspecific information transfer influences animal community structure

Acquiring information from the cues and signals of other species of the same trophic level is widespread among animals, and can help individuals exploit resources and avoid predators. But can such interspecific information transfer also influence the spatial structure of species within communities? Whereas some species use heterospecific information without changing their position, we review research that indicates that heterospecific information is a driving factor in the formation or maintenance of temporary or stable mixed-species groups. Heterospecific information can also influence the organization of such groups, including leadership. Further, animals sometimes select habitats using heterospecific information. We survey interspecific information transfer, and evaluate the morphological, ecological and behavioral factors that make some species information sources and others information seekers.

Communicating about danger: urgency alarm calling in a bird

Vertebrate flee alarm calls can provide information about the type of predator, and some mammalian alarm calls also appear to communicate the degree of danger and therefore urgency of escape. However, because predators are usually rare, it has proved difficult to obtain observations differing only in the degree of danger, or to record sufficient naturally provoked alarm calls for fully replicated playback experiments. In this study, we took advantage of a system in which the major aerial predator was common, allowing repeated, matched observations of natural interactions between predator and prey, combined with a fully replicated playback experiment. We found that the aerial trill alarm call of the white-browed scrubwren, Sericornis frontalis, varied according to the distance from the suddenly appearing predator: the closer the predator, the greater the number of elements in the call and the higher their minimum frequency (pitch). Playback experiments showed that multi-element alarm calls prompted a more urgent response, including immediate fleeing to cover. Furthermore, the response was graded, such that an alarm with more elements provoked a more urgent response. Our study therefore isolated the effect of predator distance on alarm call design, and showed that individuals respond appropriately to calls in the absence of any other cues. To our knowledge, this is the most explicit demonstration that avian flee alarm calls can convey urgency.

Relatedness, polyandry and extra-group paternity in the cooperatively-breeding white-browed scrubwren (Sericornis frontalis)

Abstract We used DNA fingerprinting to examine the genetic parentage and mating system of the cooperatively breeding white-browed scrubwren, Sericornis frontalis, in Canberra, Australia. Our analyses revealed a remarkable variety of mating tactics and social organization. Scrubwrens bred in pairs or multi-male groups that consisted of a female and two or more males. Females were always unrelated to the pair male or alpha (dominant) male. Among multi-male groups we found three different mating tactics. Firstly, when alpha and beta (subordinate) males were unrelated, they usually shared paternity in the brood. This resulted in both males gaining reproductive benefits directly. Secondly, when beta males were not related to the female but were related to the alpha males, beta males sired offspring in some broods. In this situation, beta males gained reproductive benefits both directly and potentially indirectly (through the related alpha male). Thirdly, when beta males were related to the female or both the female and alpha male, they remained on their natal territory and did not sire any offspring. Thus beta males gained only indirect reproductive benefits. Overall, when group members were related closely, the dominant male monopolized reproductive success, whereas when the members were not related closely the two males shared paternity equally. This positive association between monopolization of reproduction and relatedness is predicted by models of reproductive skew, but has not been reported previously within a single population of birds. Other cooperatively breeding birds with both closely related and unrelated helpers may show a similar variety of mating tactics. Finally, we found that extra-group paternity was more common in pairs (24% of young) than in multi-male groups (6%), and we discuss three possible reasons for this difference.

Subordinate males are more likely to help if unrelated to the breeding female in cooperatively breeding white-browed scrubwrens

Abstract Subordinates in communally breeding groups of birds usually help to provision nestlings, but in some species helping-at-the-nest is facultative. In species in which groups usually contain relatives, subordinates either always feed young or are more likely to do so when breeding dominants are close relatives, suggesting that benefits of helping collateral kin are important. In other species, adult group members are unrelated to each other and males may only feed young if they have gained paternity, showing that cooperation is related to the mating system. The white-browed scrubwren, Sericornis frontalis, is a communally breeding species in which most groups consist of a simple pair or a dominant pair with a subordinate male. Subordinate males either fed nestlings in a given nest at a rate comparable to the dominants, or did not feed them at all. Breeding groups usually formed through natal philopatry of males, so that about 80% of subordinates were closely related to one or both members of the dominant pair. However, because of death and dispersal, 54% of subordinates were unrelated to the resident female. Although subordinates with their mother fed nestlings in 48% of cases, they fed offspring in 75% of cases if their mother had been replaced by an unrelated female, suggesting that their decision to help is influenced by the opportunity to mate with the female. Supporting this conclusion, relatedness to the dominant male did not affect subordinate behaviour, and genetic studies showed that subordinates often gained paternity if unrelated to the female. Thus, paradoxically, provisioning nestlings is related to the opportunity for mating in a society in which there is natal philopatry and subordinates are usually related to one or both members of the dominant pair.

Parental alarm calls suppress nestling vocalization

Evolutionary models suggest that the cost of a signal can ensure its honesty. Empirical studies of nestling begging imply that predator attraction can impose such a cost. However, parents might reduce or abolish this cost by warning young of the presence of danger. We tested, in a controlled field playback experiment, whether alarm calls cause 5–, 8– and 11–day–old nestlings of the white–browed scrubwren, Sericornis frontalis, to suppress vocalization. In this species, nestlings vocalize when parents visit the nest (‘begging’) and when they are absent (‘non–begging’), so we measured effects on both types of vocalization. Playback of parental alarm calls suppressed non–begging vocalization almost completely but only slightly reduced begging calls during a playback of parental feeding calls that followed. The reaction of nestlings was largely independent of age. Our results suggest two reasons why experiments ignoring the role of parents probably overestimate the real cost of nestling vocalizations. Parents can warn young from a distance about the presence of danger and so suppress non–begging vocalizations that might otherwise be overheard, and a parents presence at the nest presumably indicates when it is safe to beg.

Recognition of other species' aerial alarm calls: speaking the same language or learning another?

Alarm calls given by other species potentially provide a network of information about danger, but little is known about the role of acoustic similarity compared with learning in recognition of heterospecific calls. In particular, the aerial ‘hawk’ alarm calls of passerines provide a textbook example of signal design because many species have converged on a design that thwarts eavesdropping by hawks, and call similarity might therefore allow recognition. We measured the response of fairy-wrens (Malurus cyaneus) to playback of acoustically similar scrubwren (Sericornis frontalis) aerial alarm calls. First, if call similarity prompts escape independent of learning, then fairy-wrens should flee to playback of scrubwren calls outside their geographical range. However, fairy-wrens fled only in sympatry. Second, if call similarity is necessary for learning heterospecific calls, then fairy-wrens should not respond to sympatric species with different calls. We found, on the contrary, that fairy-wrens fled to the very different aerial alarm calls of a honeyeater (Phylidonyris novaehollandiae). Furthermore, response to the honeyeater depended on the specific structure of the call, not acoustic similarity. Overall, call similarity was neither sufficient nor necessary for interspecific recognition, implying learning is essential in the complex task of sifting the acoustic world for cues about danger.

Calling at a cost: elevated nestling calling attracts predators to active nests

Begging by nestling birds has been used to test evolutionary models of signalling but theory has outstripped evidence. Eavesdropping predators potentially impose a cost on begging that ensures signal honesty, yet little experimental evidence exists for such a cost at active nests because the use of artificial nests, long playback bouts and absence of parents may have exaggerated costs. We broadcast short periods (1 h) of either nestling vocalizations or background noise at active white-browed scrubwren, Sericornis frontalis, nests. Nestlings called naturally during both treatments, allowing us to test whether elevated calling increases risk, a key but rarely tested assumption of evolutionary models. Predators visited nests exclusively during periods of elevated calling. Furthermore, playbacks affected neither adult visits nor nestling activity, suggesting that calling alone attracted predators. Adults gave alarm calls and nestlings usually called less when predators approached nests. Predation risk to broods is, therefore, likely to fluctuate substantially over short periods of time, depending on nestling hunger and whether adults or young have detected predators. This study confirms a present-day cost of nestling begging, demonstrates that this cost can be incurred over short periods and supports the importance of parent–offspring antipredator strategies in reducing predation risk.

Temporal coordination signals coalition quality.

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