Benefits of Pair-Bond Duration on Reproduction in a Lifelong Monogamous Cooperative Passerine.

AbstractLong-term social and genetic monogamy is rare in animals except birds, but even in birds it is infrequent and poorly understood. We investigated possible advantages of monogamy in a colonial, facultative cooperatively breeding bird from an arid, unpredictable environment, the sociable weaver (Philetairus socius). We documented divorce and extrapair paternity of 703 pairs over 10 years and separated effects of pair duration from breeding experience by analyzing longitudinal and cross-sectional datasets. Parts of the colonies were protected from nest predation, thereby limiting its stochastic and thus confounding effect on fitness measures. We found that 6.4% of sociable weaver pairs divorced and 2.2% of young were extrapair. Longer pair-bonds were associated with more clutches and fledglings per season and with reproducing earlier and later in the season, when snake predation is lower, but not with increased egg or fledgling mass or with nestling survival. Finally, the number of helpers at the nest increased with pair-bond duration. Results were similar for protected and unprotected nests. We suggest that long-term monogamy is associated with a better capacity for exploiting a temporally unpredictable environment and helps to form larger groups. These results can contribute to our understanding of why long-term monogamy is frequently associated with unpredictable environments and cooperation.

Interplay of cooperative breeding and predation risk on egg allocation and reproductive output.

Predation risk can influence behavior, reproductive investment, and, ultimately, individuals' fitness. In high-risk environments, females often reduce allocation to reproduction, which can affect offspring phenotype and breeding success. In cooperative breeders, helpers contribute to feed the offspring, and groups often live and forage together. Helpers can, therefore, improve reproductive success, but also influence breeders' condition, stress levels and predation risk. Yet, whether helper presence can buffer the effects of predation risk on maternal reproductive allocation remains unstudied. Here, we used the cooperatively breeding sociable weaver Philetairus socius to test the interactive effects of predation risk and breeding group size on maternal allocation to clutch size, egg mass, yolk mass, and yolk corticosterone. We increased perceived predation risk before egg laying using playbacks of the adults' main predator, gabar goshawk (Micronisus gabar). We also tested the interactive effects of group size and prenatal predator playbacks on offspring hatching and fledging probability. Predator-exposed females laid eggs with 4% lighter yolks, but predator-calls' exposure did not clearly affect clutch size, egg mass, or egg corticosterone levels. Playback-treatment effects on yolk mass were independent of group size, suggesting that helpers' presence did not mitigate predation risk effects on maternal allocation. Although predator-induced reductions in yolk mass may decrease nutrient availability to offspring, potentially affecting their survival, playback-treatment effects on hatching and fledging success were not evident. The interplay between helper presence and predator effects on maternal reproductive investment is still an overlooked area of life history and physiological evolutionary trade-offs that requires further studies.

Egg components and offspring survival vary with group size and laying order in a cooperative breeder.

Prenatal resource allocation to offspring can be influenced by maternal environment and offspring value, and affect offspring survival. An important pathway for flexible maternal allocation is via egg components such as nutrients and hormones. In cooperative breeders, females with helpers may increase resource allocation to eggs-'differential allocation'-or reduce it-'load-lightening'. Yet, helper effects on egg composition have been poorly studied. Moreover, it is unknown how helpers' presence modulates laying order effects on egg content and survival. Here, we investigated how maternal allocation varied with group size and laying order in the cooperatively breeding sociable weaver (Philetairus socius). We estimated interactive effects of helpers and laying order on allocation to egg mass, yolk nutrients-yolk mass, proteins, lipids, carotenoids, vitamin A and vitamin E-and hormones-testosterone, androstenedione, and corticosterone. Results concurred with the 'differential allocation' predictions. Females with more helpers produced later-laid eggs with heavier yolks and more lipids, and laid eggs overall richer in lipids. Proteins, antioxidants, and hormones were not found to vary with helper number. We then analyzed how helper number modulated laying order effects on survival. Females with more helpers did not specifically produce later-laid eggs with higher survival, but eggs laid by females with more helpers were overall more likely to fledge. These findings show that some egg components (yolk mass, lipids) can positively vary according to females' breeding group size, which may improve offspring fitness.

Disentangling climatic and nest predator impact on reproductive output reveals adverse high-temperature effects regardless of helper number in an arid-region cooperative bird.

Climate exerts a major influence on reproductive processes, and an understanding of the mechanisms involved and which factors might mitigate adverse weather is fundamental under the ongoing climate change. Here, we study how weather and nest predation influence reproductive output in a social species, and examine whether larger group sizes can mitigate the adverse effects of these factors. We used a 7-year nest predator-exclusion experiment on an arid-region cooperatively breeding bird, the sociable weaver. We found that dry and, especially, hot weather were major drivers of nestling mortality through their influence on nest predation. However, when we experimentally excluded nest predators, these conditions were still strongly associated with nestling mortality. Group size was unimportant against nest predation and, although positively associated with reproductive success, it did not mitigate the effects of adverse weather. Hence, cooperative breeding might have a limited capacity to mitigate extreme weather effects.

Maternal allocation in relation to weather, predation and social factors in a colonial cooperative bird.

Females may adjust prenatal allocation in relation to ecological conditions that affect reproductive success, such as weather conditions or predation risk. In cooperative breeders, helpers might also influence reproductive success, and previous studies suggest that females can lay smaller eggs or larger clutches when breeding with more helpers. Although recent work suggests that helper effects can vary according to climatic variables, how social and ecological factors interact to shape prenatal allocation is poorly understood. Here, we examine how ecological and social components of the breeding environment covary with egg mass and clutch size, using as a model the sociable weaver Philetairus socius, a colonial, cooperatively breeding passerine. The study spanned 9 years and included over 1,900 eggs from over 550 clutches. Our analyses combined natural variation in weather conditions (rainfall before each reproductive event) with a nest predator-exclusion experiment and continuous monitoring of the mother's social environment, allowing us to estimate how individual females adjust allocation to reproduction as their number of helpers varies. We found that egg mass varied consistently within females and did not clearly differ in relation to rainfall or predation risk. Contrary to previous studies, there was no evidence for plastic adjustments as females gained and lost helpers, and egg mass was instead better predicted by mother size and identity. Females laid larger clutches when breeding in environments where predation risk was experimentally reduced and after higher rainfall levels. Yet, there was no evidence for increasing clutch size as the number of helpers increased, nor for an interaction between helper effects and ecological factors. We conclude that while sociable weaver females can vary their clutch size, they show high individual consistency in egg mass. In addition, we found no evidence that females may maximize fitness through plastic prenatal allocation in relation to the number of helpers, or that the presence/absence of helper effects is modulated by rainfall levels or predation risk. These results challenge our current knowledge on some of the possible benefits of breeding with helpers and call for more long-term analyses on reproductive allocation adjustments in other cooperative systems.

How to make methodological decisions when inferring social networks.

Social network analyses allow studying the processes underlying the associations between individuals and the consequences of those associations. Constructing and analyzing social networks can be challenging, especially when designing new studies as researchers are confronted with decisions about how to collect data and construct networks, and the answers are not always straightforward. The current lack of guidance on building a social network for a new study system might lead researchers to try several different methods and risk generating false results arising from multiple hypotheses testing. Here, we suggest an approach for making decisions when starting social network research in a new study system that avoids the pitfall of multiple hypotheses testing. We argue that best edge definition for a network is a decision that can be made using a priori knowledge about the species and that is independent from the hypotheses that the network will ultimately be used to evaluate. We illustrate this approach with a study conducted on a colonial cooperatively breeding bird, the sociable weaver. We first identified two ways of collecting data using different numbers of feeders and three ways to define associations among birds. We then evaluated which combination of data collection and association definition maximized (a) the assortment of individuals into previously known "breeding groups" (birds that contribute toward the same nest and maintain cohesion when foraging) and (b) socially differentiated relationships (more strong and weak relationships than expected by chance). This evaluation of different methods based on a priori knowledge of the study species can be implemented in a diverse array of study systems and makes the case for using existing, biologically meaningful knowledge about a system to help navigate the myriad of methodological decisions about data collection and network inference.