Measuring the complexity of social associations using mixture models

This is final version. Available on open access from Springer via the DOI in this record.We propose a method for examining and measuring the complexity of animal social networks that are characterized using association indices. The method focusses on the diversity of types of dyadic relationship within the social network. Binomial mixture models cluster dyadic relationships into relationship types, and variation in the preponderance and strength of these relationship types can be used to estimate association complexity using Shannon’s information index. We use simulated data to test the method, and find that models chosen using integrated complete likelihood give estimates of complexity that closely reflect the true complexity of social systems, but these estimates can be downwardly biased by low intensity sampling and upwardly biased by extreme overdispersion within components. We also illustrate the use of the method on two real data sets. The method could be extended for use on interaction rate data using Poisson mixture models, or on multidimensional relationship data using multivariate mixture models

Ecological knowledge, leadership, and the evolution of menopause in killer whales

This is the final version of the article. Available from Elsevier via the DOI in this record.There is another record in ORE for this publication: http://hdl.handle.net/10871/16488Classic life-history theory predicts that menopause should not occur because there should be no selection for survival after the cessation of reproduction [1]. Yet, human females routinely live 30 years after they have stopped reproducing [2]. Only two other species-killer whales (Orcinus orca) and short-finned pilot whales (Globicephala macrorhynchus) [3, 4]-have comparable postreproductive lifespans. In theory, menopause can evolve via inclusive fitness benefits [5, 6], but the mechanisms by which postreproductive females help their kin remain enigmatic. One hypothesis is that postreproductive females act as repositories of ecological knowledge and thereby buffer kin against environmental hardships [7, 8]. We provide the first test of this hypothesis using a unique long-term dataset on wild resident killer whales. We show three key results. First, postreproductively aged females lead groups during collective movement in salmon foraging grounds. Second, leadership by postreproductively aged females is especially prominent in difficult years when salmon abundance is low. This finding is critical because salmon abundance drives both mortality and reproductive success in resident killer whales [9, 10]. Third, females are more likely to lead their sons than they are to lead their daughters, supporting predictions of recent models [5] of the evolution of menopause based on kinship dynamics. Our results show that postreproductive females may boost the fitness of kin through the transfer of ecological knowledge. The value gained from the wisdom of elders can help explain why female resident killer whales and humans continue to live long after they have stopped reproducing.This research was funded by a Natural Environment Research Council grant (NE/K01286X/1) to D.P.C., D.W.F., and M.A.C

Common permutation methods in animal social network analysis do not control for non-independence

This is the final version. Available on open access from Springer via the DOI in this recordData availability: The R code required to repeat the simulations has been deposited at: https://doi.org/10.5281/zenodo.4903396).The non-independence of social network data is a cause for concern among behavioural ecologists conducting social network analysis. This has led to the adoption of several permutation-based methods for testing common hypotheses. One of the most common types of analysis is nodal regression, where the relationships between node-level network metrics and nodal covariates are analysed using a permutation technique known as node-label permutations. We show that, contrary to accepted wisdom, node-label permutations do not automatically account for the non-independences assumed to exist in network data, because regression-based permutation tests still assume exchangeability of residuals. The same assumption also applies to the quadratic assignment procedure (QAP), a permutation-based method often used for conducting dyadic regression. We highlight that node-label permutations produce the same p-values as equivalent parametric regression models, but that in the presence of non-independence, parametric regression models can also produce accurate effect size estimates. We also note that QAP only controls for a specific type of non-independence between edges that are connected to the same nodes, and that appropriate parametric regression models are also able to account for this type of non-independence. Based on this, we suggest that standard parametric models could be used in the place of permutation-based methods. Moving away from permutation-based methods could have several benefits, including reducing over-reliance on p-values, generating more reliable effect size estimates, and facilitating the adoption of causal inference methods and alternative types of statistical analysis.Engineering and Physical Sciences Research Council (EPSRC)European Research Council (ERC)National Institutes of Health (NIH)Natural Environment Research Council (NERC

The evolution of menopause in toothed whales

This is the final version. Available on open access from Nature Research via the DOI in this recordData availability: All data used in this analysis are available as a database at: github.com/samellisq/marinelifehistdataCode availability: All R and stan code used for this analysis are available at osf.io/26s7m/. In addition, the mortality model is implemented as an R package available from: github.com/samellisq/marinesurvivalUnderstanding how and why menopause has evolved is a long-standing challenge across disciplines. Females can typically maximize their reproductive success by reproducing for the whole of their adult life. In humans, however, women cease reproduction several decades before the end of their natural lifespan1,2. Although progress has been made in understanding the adaptive value of menopause in humans3,4, the generality of these findings remains unclear. Toothed whales are the only mammal taxon in which menopause has evolved several times5, providing a unique opportunity to test the theories of how and why menopause evolves in a comparative context. Here, we assemble and analyse a comparative database to test competing evolutionary hypotheses. We find that menopause evolved in toothed whales by females extending their lifespan without increasing their reproductive lifespan, as predicted by the 'live-long' hypotheses. We further show that menopause results in females increasing their opportunity for intergenerational help by increasing their lifespan overlap with their grandoffspring and offspring without increasing their reproductive overlap with their daughters. Our results provide an informative comparison for the evolution of human life history and demonstrate that the same pathway that led to menopause in humans can also explain the evolution of menopause in toothed whales.Leverhulme TrustNatural Environment Research Council (NERC

Calculating effect sizes in animal social network analysis

This is the final version. Available on open access from Wiley via the DOI in this record1. Because of the nature of social interaction or association data, when testing hypotheses using social network data it is common for network studies to rely on permutations to control for confounding variables, and to not also control for them in the fitted statistical model. This can be a problem because it does not adjust for any bias in effect sizes generated by these confounding effects, and thus the effect sizes are not informative in the presence of counfouding variables. 2. We implemented two network simulation examples and analysed an empirical data set to demonstrate how relying solely on permutations to control for confounding variables can result in highly biased effect size estimates of animal social preferences that are uninformative when quantifying differences in behaviour. 3. Using these simulations, we show that this can sometimes even lead to effect sizes that have the wrong sign and are thus the effect size is not biologically interpretable. We demonstrate how this problem can be addressed by controlling for confounding variables in the statistical dyadic or nodal model. 4. We recommend this approach should be adopted as standard practice in the statistical analysis of animal social network dataNatural Environment Research Council (NERC

Common datastream permutations of animal social network data are not appropriate for hypothesis testing using regression models

This is the final version. Available on open access from Wiley via the DOI in this record1. Social network methods have become a key tool for describing, modelling, and testing hypotheses about the social structures of animals. However, due to the non-independence of network data and the presence of confounds, specialized statistical techniques are often needed to test hypotheses in these networks. Datastream permutations, originally developed to test the null hypothesis of random social structure, have become a popular tool for testing a wide array of null hypotheses in animal social networks. In particular, they have been used to test whether exogenous factors are related to network structure by interfacing these permutations with regression models. 2. Here, we show that these datastream permutations typically do not represent the null hypothesis of interest to researchers interfacing animal social network analysis with regression modelling, and use simulations to demonstrate the potential pitfalls of using this methodology. 3. Our simulations show that, if used to indicate whether a relationship exists between network structure and a covariate, datastream permutations can result in extremely high type I error rates, in some cases approaching 50%. In the same set of simulations, traditional node-label permutations produced appropriate type I error rates (~ 5%). 4. Our analysis shows that datastream permutations do not represent the appropriate null hypothesis for these analyses. We suggest that potential alternatives to this procedure may be found in regarding the problems of non-independence of network data and unreliability of observations separately. If biases introduced during data collection can be corrected, either prior to model fitting or within the model itself, node-label permutations then serve as a useful test for interfacing animal social network analysis with regression modellingNatural Environment Research Council (NERC)NI

Kinship dynamics: patterns and consequences of changes in local relatedness

This is the author accepted manuscript. The final version is available from the Royal Society via the DOI in this recordMounting evidence suggests that patterns of local relatedness can change over time in predictable ways, a process termed kinship dynamics. Kinship dynamics may occur at the level of the population or social group, where the mean relatedness across all members of the population or group changes over time, or at the level of the individual where an individual’s relatedness to its local group changes with age. Kinship dynamics are likely to have fundamental consequences for the evolution of social behaviour and life history because they alter the inclusive fitness payoffs to actions taken at different points in time. For instance, growing evidence suggests that individual kinship dynamics have shaped the evolution of menopause and age-specific patterns of helping and harming. To date, however, the consequences of kinship dynamics for social evolution have not been widely explored. Here we review the patterns of kinship dynamics that can occur in natural populations and highlight how taking a kinship dynamics approach has yielded new insights into behaviour and life history evolution. We discuss areas where analysing kinship dynamics could provide new insight into social evolution and we outline some of the challenges in predicting and quantifying kinship dynamics in natural populations.Natural Environment Research Council (NERC)Leverhulme Trus

Midday measurements of leaf water potential and stomatal conductance are highly correlated with daily water use of Thompson Seedless grapevines

A study was conducted to determine the relationship between midday measurements of vine water status and daily water use of grapevines measured with a weighing lysimeter. Water applications to the vines were terminated on August 24th for 9 days and again on September 14th for 22 days. Daily water use of the vines in the lysimeter (ETLYS) was approximately 40 L vine−1 (5.3 mm) prior to turning the pump off, and it decreased to 22.3 L vine−1 by September 2nd. Pre-dawn leaf water potential (ΨPD) and midday Ψl on August 24th were −0.075 and −0.76 MPa, respectively, with midday Ψl decreasing to −1.28 MPa on September 2nd. Leaf g s decreased from ~500 to ~200 mmol m−2 s−1 during the two dry-down periods. Midday measurements of g s and Ψl were significantly correlated with one another (r = 0.96) and both with ETLYS/ETo (r = ~0.9). The decreases in Ψl, g s, and ETLYS/ETo in this study were also a linear function of the decrease in volumetric soil water content. The results indicate that even modest water stress can greatly reduce grapevine water use and that short-term measures of vine water status taken at midday are a reflection of daily grapevine water us

A long post-reproductive lifespan is a shared trait among genetically distinct killer whale populations

This is the final version. Available on open access from Wiley via the DOI in this recordData availability: Data to replicate the analyses are available from the online repository: https://doi.org/10.5061/dryad.6t1g1jwxx. Requests for access to raw data can be directed to the authors, the Center for Whale Research (www.whaleresearch.com) or Fisheries and Oceans Canada (www.dfo-mpo.gc.ca)The extended female post-reproductive lifespan found in humans and some toothed whales remains an evolutionary puzzle. Theory predicts demographic patterns resulting in increased female relatedness with age (kinship dynamics) can select for a prolonged post reproductive lifespan due to the combined costs of inter-generational reproductive conflict and benefits of late-life helping. Here we test this prediction using >40 years of longitudinal demographic data from the sympatric yet genetically distinct killer whale ecotypes: resident and Bigg’s killer whales. The female relatedness with age is predicted to increase in both ecotypes, but with a less steep increase in Bigg’s due to their different social structure. Here, we show that there is a significant post-reproductive lifespan in both ecotypes with >30% of adult female years being lived as post-reproductive, supporting the general prediction that an increase in local relatedness with age predisposes the evolution of a post reproductive lifespan. Differences in the magnitude of kinship dynamics however, did not influence the timing or duration of the post-reproductive lifespan with females in both ecotypes terminating reproduction before their mid-40s followed by an expected post reproductive period of ~20 years. Our results highlight the important role of kinship dynamics in the evolution of a long post-reproductive lifespan in long-lived mammals, while further implying that the timing of menopause may be a robust trait that is persistent despite substantial variation in demographic patterns among populations.Nuffield FoundationNatural Environment Research Council (NERC)Leverhulme TrustEarthwatch InstituteNOAA FisheriesFisheries and Oceans Canada Species At Risk Progra