Delayed dispersal as a potential route to cooperative breeding in ambrosia beetles

Xyleborini are a species-rich tribe of ambrosia beetles, which are haplodiploid and typically mate among siblings within their natal brood chamber. Several characteristics of this tribe would predict the evolution of higher levels of sociality: high genetic relatedness within galleries due to inbreeding, high costs of dispersal and the potential benefit of cooperation in brood care within the natal gallery (e.g. by fungus gardening, gallery extension, offspring feeding and cleaning). However, information on the social system of these beetles is very limited. We examined the potential for cooperative breeding in Xyleborinus saxeseni by monitoring dispersal in relation to brood size and composition. Results show that adult female offspring delay dispersal despite dispersal opportunities, and apparently some females never disperse. The females' decision to stay seems to depend on the presence of eggs and dependent siblings. We found no indication that female offspring reproduce in their natal gallery, as colonies with many mature daughters do not contain more eggs than those with few or no daughters. There is a significant positive relationship between the number of females present and the number of dependent siblings (but not eggs), which suggests that cooperative brood care of female offspring raises colony productivity by improving survival rates of immatures. Our results suggest that cooperative breeding is likely to occur in X. saxeseni and possibly other xyleborine species. We argue that a closer look at sociality within this tribe may yield important information on the factors determining the evolution of cooperative breeding and advanced social organizatio

The influence of social experience on cooperative behaviour of rats ( Rattus norvegicus ): direct vs generalised reciprocity

Cooperation among non-kin has been attributed sometimes to reciprocal altruism: Two or more individuals exchange behaviour that benefits the respective partner. According to direct reciprocity, cooperation is based on past behaviour of a known partner. In contrast, in generalised reciprocity, cooperation is based on anonymous social experience where the identity of the partner is irrelevant. In a previous study, female Norway rats (Rattus norvegicus) were found to cooperate according to a generalised reciprocity mechanism. In this study, we tested whether Norway rats would also cooperate as predicted by a direct reciprocity mechanism and whether direct reciprocity would cause a higher propensity to cooperate than generalised reciprocity. Focal animals were experimentally manipulated to receive social experience from known or unknown, helpful or defecting partners in an instrumental cooperative task. Our first experiment shows that rats are more helpful towards a partner from which they had received help before than towards a partner that had not helped (i.e. direct reciprocity). Our second experiment revealed that after receiving help by others, rats were more helpful towards a partner from which they had received help before than towards a new partner (i.e. direct reciprocity generated a higher cooperation propensity than generalised reciprocity). We conclude that in female Norway rats, the tendency to cooperate is influenced by partner-specific information. To our knowledge, this is the first study to demonstrate direct reciprocity in rodents, and it is the first study testing direct vs generalised reciprocity in animal

Female choice of a non-bodily ornament: an experimental study of cichlid sand craters in Cyathopharynx furcifer

In many species of East African cichlids, males build sand craters or "bowers” to attract females for spawning. It is commonly assumed that these exaggerated sexual traits have a function similar to bodily ornaments. In non-bodily ornaments, however, the behaviour creating the structure may serve as an additional source of information to potential partners, providing multiple signals for mate choice. We tested whether and to what extent females use these signals for choosing males by observing the individual sampling behaviour of female Cyathopharynx furcifer in the field. In addition, we experimentally manipulated crater characteristics in the field and laboratory. We found that females spawn preferably with owners of large and well-maintained craters, but when crater size was enlarged or reduced, the resulting building activity of crater owners affected female choice more strongly than the manipulated crater size per se. We discuss the importance of multiple signals in species constructing extended phenotype

Sequential polyandry affords post-mating sexual selection in the mouths of cichlid females

Females mating with multiple males may obtain direct benefits such as nuptial gifts or paternal care or indirect (i.e. genetic) benefits resulting in higher-quality offspring. While direct benefits are easily identified, it is difficult to determine indirect benefits, and it is hence largely unclear how they are obtained. This is particularly true in species with external fertilisation, where females seem to have little control over fertilisation. In cichlids, most maternal mouthbrooders show sequential multiple mating, where females visit several males for egg deposition. Genetic data revealed that multiple paternity of eggs and young in the mouth of females is common, but behavioural data of female spawning decisions are missing. Here, we test four hypotheses to explain female multiple mating in the maternally mouthbrooding cichlid, Ophthalmotilapia ventralis: (1) fertilisation insurance, (2) genetic bet-hedging, (3) female choice and (4) ‘sperm shopping' (i.e. induction of sperm competition resulting in sexually selected sperm). Detailed observations of spawning behaviour in the field combined with histological analyses of the male reproductive organs suggest that fertilisation insurance, genetic bet-hedging and pre-mating female choice are unlikely to explain the sequential female multiple mating in O. ventralis. Instead, cryptic female choice by sperm shopping, i.e. post-mating sexual selection, is most compatible with our data and might be the major ultimate cause of multiple mating in females of this species and of mouthbrooding cichlids with maternal care in general. Our study provides new insight into ultimate causes of sequential polyandry in species with external fertilisation, as hitherto post-mating sexual selection by cryptic female choice has been assumed to be incompatible with external fertilisation mechanisms except by components of the ovarian flui

Commodity-specific punishment for experimentally induced defection in cooperatively breeding fish.

Coercion is an important but underrated component in the evolution of cooperative behaviour. According to the pay-to-stay hypothesis of cooperative breeding, subordinates trade alloparental care for the concession to stay in the group. Punishment of idle subordinates is a key prediction of this hypothesis, which has received some experimental scrutiny. However, previous studies neither allowed separating between punishment and effects of disruption of social dynamics, nor did they differentiate between different helping behaviours that may reflect either mutualistic or reciprocal interaction dynamics. In the cooperative breeder Neolamprologus pulcher, we experimentally engineered the ability of subordinates to contribute to alloparental care by manipulating two different helping behaviours independently from one another in a full factorial design. We recorded the treatment effects on breeder aggression, subordinate helping efforts and submissive displays. We found two divergent regulatory mechanisms of cooperation, dependent on behavioural function. Experimental impediment of territory maintenance of subordinates triggered punishment by dominants, whereas prevented defence against egg predators released a compensatory response of subordinates without any enforcement, suggesting pre-emptive appeasement. These effects occurred independently of one another. Apparently, in the complex negotiation process among members of cooperative groups, behaviours fulfilling different functions may be regulated by divergent interaction mechanisms

Adaptive behavioural syndromes due to strategic niche specialization

<p>Abstract</p> <p>Background</p> <p>Behavioural syndromes, i.e. consistent individual differences in behaviours that are correlated across different functional contexts, are a challenge to evolutionary reasoning because individuals should adapt their behaviour to the requirements of each situation. Behavioural syndromes are often interpreted as a result of constraints resulting in limited plasticity and inflexible behaviour. Alternatively, they may be adaptive if correlated ecological or social challenges functionally integrate apparently independent behaviours. To test the latter hypothesis we repeatedly tested helpers in the cooperative breeder <it>Neolamprologus pulcher </it>for exploration and two types of helping behaviour. In case of adaptive behavioural syndromes we predicted a positive relationship between exploration and aggressive helping (territory defence) and a negative relationship between these behaviours and non-aggressive helping (territory maintenance).</p> <p>Results</p> <p>As expected, helpers engaging more in territory defence were consistently more explorative and engaged less in territory maintenance, the latter only when dominant breeders were present. Contrary to our prediction, there was no negative relationship between exploration and territory maintenance.</p> <p>Conclusion</p> <p>Our results suggest that the three behaviours we measured are part of behavioural syndromes. These may be adaptive, in that they reflect strategic specialization of helpers into one of two different life history strategies, namely (a) to stay and help in the home territory in order to inherit the breeding position or (b) to disperse early in order to breed independently.</p

Working dogs cooperate among one another by generalised reciprocity.

Cooperation by generalised reciprocity implies that individuals apply the decision rule "help anyone if helped by someone". This mechanism has been shown to generate evolutionarily stable levels of cooperation, but as yet it is unclear how widely this cooperation mechanism is applied among animals. Dogs (Canis familiaris) are highly social animals with considerable cognitive potential and the ability to differentiate between individual social partners. But although dogs can solve complex problems, they may use simple rules for behavioural decisions. Here we show that dogs trained in an instrumental cooperative task to provide food to a social partner help conspecifics more often after receiving help from a dog before. Remarkably, in so doing they show no distinction between partners that had helped them before and completely unfamiliar conspecifics. Apparently, dogs use the simple decision rule characterizing generalised reciprocity, although they are probably capable of using the more complex decision rule of direct reciprocity: "help someone who has helped you". However, generalized reciprocity involves lower information processing costs and is therefore a cheaper cooperation strategy. Our results imply that generalised reciprocity might be applied more commonly than direct reciprocity also in other mutually cooperating animals

Sexual conflict over breeding substrate causes female expulsion and offspring loss in a cichlid fish

Females of the Lake Tanganyika cichlid Lamprologus callipterus exclusively breed in empty snail shells that males collect in their territories. Male-male competition for shells is severe, leading to frequent shell stealing and territory takeover. As a consequence, males have breeding females in their shells that spawned with competitors. In this field study, we investigated both naturally occurring and experimentally induced encounters of territorial males with females that had spawned with other males. We found that the breeding success of females that were taken over by a different male was significantly reduced. Behavioral observations after experimental shell relocation further showed that males recognized females that they had not spawned with: males directed more exploration and manipulation behavior toward such shells compared with controls. Reoccupation rate of emptied experimental shells was significantly higher than that of unmanipulated empty shells. This indicates that shell stealing and nest takeover, followed by female expulsion, contribute to the reproductive success of L. callipterus males. We also found that female mate choice reduces expulsion risk: females preferred to mate with large males, and male size correlated with dominance. We conclude that the limited availability of breeding substrate is a key determinant of both intrasexual competition and intersexual conflict in this specie

Rats play tit-for-tat instead of integrating social experience over multiple interactions

Funding was provided by SNF-grant 31003A_156152 to M.T. and P2BEP3 175269 to M.K.S.Theoretical models of cooperation typically assume that agents use simple rules based on last encounters, such as “tit-for-tat”, to reciprocate help. In contrast, empiricists generally suppose that animals integrate multiple experiences over longer timespans. Here we compared these two alternative hypotheses by exposing Norway rats to partners that cooperated on three consecutive days but failed to cooperate on the fourth day, and to partners that did the exact opposite. In additional controls, focal rats experienced cooperating and defecting partners only once. In a bar-pulling setup, focal rats based their decision to provide partners with food on last encounters instead of overall cooperation levels. To check whether this might be due to a lack of memory capacity, we tested whether rats remember the outcome of encounters that had happened three days before. Cooperation was not diminished by the intermediate time interval. We conclude that rats reciprocate help mainly based on most recent encounters instead of integrating social experience over longer timespans.PostprintPeer reviewe