Sex differences in the drivers of reproductive skew in a cooperative breeder.

Many cooperatively breeding societies are characterized by high reproductive skew, such that some socially dominant individuals breed, while socially subordinate individuals provide help. Inbreeding avoidance serves as a source of reproductive skew in many high-skew societies, but few empirical studies have examined sources of skew operating alongside inbreeding avoidance or compared individual attempts to reproduce (reproductive competition) with individual reproductive success. Here, we use long-term genetic and observational data to examine factors affecting reproductive skew in the high-skew cooperatively breeding southern pied babbler (Turdoides bicolor). When subordinates can breed, skew remains high, suggesting factors additional to inbreeding avoidance drive skew. Subordinate females are more likely to compete to breed when older or when ecological constraints on dispersal are high, but heavy subordinate females are more likely to successfully breed. Subordinate males are more likely to compete when they are older, during high ecological constraints, or when they are related to the dominant male, but only the presence of within-group unrelated subordinate females predicts subordinate male breeding success. Reproductive skew is not driven by reproductive effort, but by forces such as intrinsic physical limitations and intrasexual conflict (for females) or female mate choice, male mate-guarding and potentially reproductive restraint (for males). Ecological conditions or "outside options" affect the occurrence of reproductive conflict, supporting predictions of recent synthetic skew models. Inbreeding avoidance together with competition for access to reproduction may generate high skew in animal societies, and disparate processes may be operating to maintain male vs. female reproductive skew in the same species.

Individual dispersal delays in a cooperative breeder: Ecological constraints, the benefits of philopatry and the social queue for dominance.

Delayed dispersal is a key step in the evolution of familial animal societies and cooperative breeding. However, no consensus has been reached on the ecological and social circumstances driving delayed dispersal. Here, we test predictions from the ecological constraints and benefits of philopatry hypotheses as well as the recently proposed dual benefits hypothesis to better understand the evolution of group-living and cooperative breeding. Furthermore, we consider how individual social circumstances within groups affect dispersal decisions. We examine 11 years of life-history information on a wild population of cooperatively breeding southern pied babblers Turdoides bicolor. We investigate the effects of ecological conditions, natal-group membership and individual social context on male and female dispersal delays, disperser survival and acquisition of dominance. Female dispersal decisions are generally unconstrained by ecological or social circumstances. In contrast, males disperse in response to relaxed ecological constraints, decreases in nepotistic tolerance or when low social rank in the queue for dominance decreases their likelihood of gaining a dominant breeding position. Early dispersal by end-of-queue males often leads to a head-of-queue subordinate position in a non-natal group, thereby increasing access to dominant breeding positions. However, males and females remaining in natal groups gain benefits of philopatry via increased survival and, for head-of-queue males, very high likelihood of acquisition of a breeding position. Overall, predictions from the dual benefits hypothesis best describe these results, while some predictions from each of the ecological constraints and benefits of philopatry hypotheses were supported. The benefits of living and working together (collective action benefits) in large stable groups are of central importance in shaping dispersal delays in southern pied babbler societies. In addition, position in the subordinate social queue for dominance is the key in determining access to reproduction, particularly for males. This research highlights the importance of considering the costs and benefits of individual social circumstances in dispersal decisions and illustrates how the dual benefits hypothesis offers new perspectives in understanding delayed dispersal.

Exorcising Grice's ghost: an empirical approach to studying intentional communication in animals.

Language's intentional nature has been highlighted as a crucial feature distinguishing it from other communication systems. Specifically, language is often thought to depend on highly structured intentional action and mutual mindreading by a communicator and recipient. Whilst similar abilities in animals can shed light on the evolution of intentionality, they remain challenging to detect unambiguously. We revisit animal intentional communication and suggest that progress in identifying analogous capacities has been complicated by (i) the assumption that intentional (that is, voluntary) production of communicative acts requires mental-state attribution, and (ii) variation in approaches investigating communication across sensory modalities. To move forward, we argue that a framework fusing research across modalities and species is required. We structure intentional communication into a series of requirements, each of which can be operationalised, investigated empirically, and must be met for purposive, intentionally communicative acts to be demonstrated. Our unified approach helps elucidate the distribution of animal intentional communication and subsequently serves to clarify what is meant by attributions of intentional communication in animals and humans.

Interspecific signalling between mutualists: food-thieving drongos use a cooperative sentinel call to manipulate foraging partners.

Interspecific communication is common in nature, particularly between mutualists. However, whether signals evolved for communication with other species, or are in fact conspecific signals eavesdropped upon by partners, is often unclear. Fork-tailed drongos (Dicrurus adsimilis) associate with mixed-species groups and often produce true alarms at predators, whereupon associating species flee to cover, but also false alarms to steal associating species' food (kleptoparasitism). Despite such deception, associating species respond to drongo non-alarm calls by increasing their foraging and decreasing vigilance. Yet, whether these calls represent interspecific sentinel signals remains unknown. We show that drongos produced a specific sentinel call when foraging with a common associate, the sociable weaver (Philetairus socius), but not when alone. Weavers increased their foraging and decreased vigilance when naturally associating with drongos, and in response to sentinel call playback. Further, drongos sentinel-called more often when weavers were moving, and weavers approached sentinel calls, suggesting a recruitment function. Finally, drongos sentinel-called when weavers fled following false alarms, thereby reducing disruption to weaver foraging time. Results therefore provide evidence of an 'all clear' signal that mitigates the cost of inaccurate communication. Our results suggest that drongos enhance exploitation of a foraging mutualist through coevolution of interspecific sentinel signals.

Deception by flexible alarm mimicry in an African bird.

Deception is common in nature, but victims of deception discriminate against and ultimately ignore deceptive signals when they are produced too frequently. Flexible variation of signals could allow evasion of such constraints. Fork-tailed drongos (Dicrurus adsimilis) use false alarm calls to scare other species away from food that they then steal. We show that drongos mimic the alarms of targeted species. Further, target species reduce their response to false alarm calls when they are repeated. However, the fear response is maintained when the call is varied. Drongos exploit this propensity by changing their alarm-call type when making repeated theft attempts on a particular species. Our results show that drongos can evade the frequency-dependent constraints that typically limit deception payoffs through flexible variation of their alarm calls.

The ecological economics of kleptoparasitism: pay-offs from self-foraging versus kleptoparasitism.

Animals commonly steal food from other species, termed interspecific kleptoparasitism, but why animals engage in kleptoparasitism compared with alternate foraging tactics, and under what circumstances they do so, is not fully understood. Determining what specific benefits animals gain from kleptoparasitism could provide valuable insight into its evolution. Here, we investigate the benefits of kleptoparasitism for a population of individually recognizable and free-living fork-tailed drongos (Dicrurus adsimilis) in the southern Kalahari Desert. Drongos engaged in two foraging behaviours: self-foraging for small insects or following other species which they kleptoparasitized for larger terrestrial prey that they could not capture themselves. Kleptoparasitism consequently enabled drongos to exploit a new foraging niche. Kleptoparasitism benefitted drongos most in the morning and on colder days because at these times pay-offs from kleptoparasitism remained stable, while those from self-foraging declined. However, drongos engaged in kleptoparasitism less than expected given the overall high (but more variable) pay-offs from this behaviour, suggesting that kleptoparasitism is a risky foraging tactic and may incur additional foraging costs compared with self-foraging. This is the first study to comprehensively investigate the benefits of facultatively engaging in kleptoparasitism, demonstrating that animals may switch to kleptoparasitism to exploit a new foraging niche when pay-offs exceed those from alternate foraging behaviours.

Inbreeding and inbreeding depression of early life traits in a cooperative mammal.

Mating between relatives often results in negative fitness consequences or inbreeding depression. However, the expression of inbreeding in populations of wild cooperative mammals and the effects of environmental, maternal and social factors on inbreeding depression in these systems are currently not well understood. This study uses pedigree-based inbreeding coefficients from a long-term study of meerkats (Suricata suricatta) in South Africa to reveal that 44% of the population have detectably non-zero (F > 0) inbreeding coefficients. 15% of these inbred individuals were the result of moderate inbreeding (F ≥ 0.125), although such inbreeding events almost solely occurred when mating individuals had no prior experience of each other. Inbreeding depression was evident for a range of traits: pup mass at emergence from the natal burrow, hind-foot length, growth until independence and juvenile survival. However, we found no evidence of significant inbreeding depression for skull and forearm length or for pup survival. This research provides a rare investigation into inbreeding in a cooperative mammal, revealing high levels of inbreeding, considerable negative consequences and complex interactions with the social environment.

Fork-tailed drongos use deceptive mimicked alarm calls to steal food.

Despite the prevalence of vocal mimicry in animals, few functions for this behaviour have been shown. I propose a novel hypothesis that false mimicked alarm calls could be used deceptively to scare other species and steal their food. Studies have previously suggested that animals use their own species-specific alarm calls to steal food. However none have shown conclusively that these false alarms are deceptive, or that mimicked alarm calls are used in this manner. Here, I show that wild fork-tailed drongos (Dicrurus adsimilis) make both drongo-specific and mimicked false alarm calls when watching target species handling food, in response to which targets flee to cover abandoning their food. The drongo-specific and mimicked calls made in false alarms were structurally indistinguishable from calls made during true alarms at predators by drongos and other species. Furthermore, I demonstrate by playback experiments that two of these species, meerkats (Suricata suricatta) and pied babblers (Turdoides bicolor), are deceived by both drongo-specific and mimicked false alarm calls. These results provide the first conclusive evidence that false alarm calls are deceptive and demonstrate a novel function for vocal mimicry. This work also provides valuable insight into the benefits of deploying variable mimetic signals in deceptive communication.

Adaptive suppression of subordinate reproduction in cooperative mammals.

Attempts to account for observed variation in the degree of reproductive skew among cooperative breeders have usually assumed that subordinate breeding has fitness costs to dominant females. They argue that dominant females concede reproductive opportunities to subordinates to retain them in the group or to dissuade them from challenging for the dominant position or that subordinate females breed where dominants are incapable of controlling them. However, an alternative possibility is that suppressing subordinate reproduction has substantive costs to the fitness of dominant females and that variation in these costs generates differences in the net benefits of suppression to dominants which are responsible for variation in the frequency of subordinate breeding that is not a consequence of either reproductive concessions or limitations in dominant control. Here, we show that, in wild Kalahari meerkats (Suricata suricatta), the frequency with which dominants evict subordinates or kill their pups varies with the costs and benefits to dominants of suppressing subordinate breeding, including the dominants' reproductive status, the size of their group, and the relatedness of subordinates. We review evidence from other studies that the suppression of reproduction by subordinates varies with the likely costs of subordinate breeding to dominants.