Michael N. Weiss’s research while affiliated with University of Exeter and other places

In many aquatic taxa, formation traveling can reduce the energetic expenditure of locomotion by exploiting the vorticity trails shed by neighbors or through drafting. Cetaceans, especially odontocetes, often swim in groups; nevertheless, the possibility that whales gain energetic benefits from swimming in formation remains poorly studied, apart from mother-calf pairs. Between June and September in 2019 and 2021, we recorded aerial videos of Southern Resident killer whales (Orcinus orca) in the Salish Sea (USA) travelling in groups. We estimated whale tailbeat and breathing frequencies as proxies of the relative energetic costs of swimming, and tested the effect of swimming speed, relative positioning (e.g., leaders, whales in the middle of groups, or followers), sex and estimated size on these observed proxies. Our results reveal a complex relationship between physical characteristics, relative positioning, and energetic proxies. Intervals between respiration lasted longer in large-sized trailing individuals, but the overall breathing frequency was similar for all whales regardless of their position. The tailbeat frequency was mainly associated to whale sex, size, and swimming speed; in addition, tailbeat frequency showed a decreasing trend as the number of individuals in the formation increased. We found moderate evidence that position-based energetic effects may be present in the formation swimming of killer whales, and it is likely that additional factors such as social ties and hierarchies, play a key role in determining individual positioning in travelling groups. Significance Swimming in formation has been extensively studied in fish and other aquatic animals and has been documented to provide energetic advantages. Our understanding of the potential energetic benefits of wild cetacean formation swimming has been constrained by the difficulties of studying the movement of whale groups from traditional observation platforms. In recent years, non-invasive observations of cetaceans using unoccupied aerial systems have significantly improved the observation of these species in the wild, providing an exciting opportunity to better understand their behaviors and habits. Our results show a tendency for formation swimming to affect two energetic proxies (tailbeat frequency and the duration of underwater intervals between surfacing events). The results of this study set the stage for further research to identify the multiple determinants affecting killer whale formation swimming which go beyond purely energetic advantages, e.g. social relationships.

Understanding 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.

The mean relatedness of an individual to others in its social group may change with age. Such kinship dynamics, which are expected to influence sex- and age-specific social behaviours, are sensitive to demography. Existing models of kinship dynamics, however, have focused chiefly on the influences of population-level demographic parameters (e.g., sex-specific dispersal rate and mating pattern). Here, we extend these models to explore the effects of local group size on kinship dynamics and the resulting patterns of selection for sex- and age-specific helping and harming. We show that individuals' average relatedness to others is higher in smaller than larger groups, and consequently, selective pressure for helping or harming is generally stronger in smaller than larger groups. Moreover, relatedness changes faster with age in smaller groups, particularly at younger ages. These patterns favour more extreme social traits in smaller groups and especially so at earlier life history stages. In particular we highlight that, while social systems characterized by bi-sexual philopatry with non-local mating (e.g., whales) are known to favour a shift from more helpful to more harmful behaviour as females age, a trend that has been invoked to explain the evolution of menopause and post-reproductive helping, the timing of this shift is predicted to be sensitive to local group size, potentially favouring earlier female reproductive cessation in smaller groups. Our study generates new insights into the effects of local group properties on kinship dynamics, and suggests that such effects might also help to explain widespread variation in age-linked social traits, such as the timing of menopause in social mammals.

Endangered Southern Resident killer whales (Orcinus orca) are fish-eaters that preferentially prey on adult Chinook salmon (Oncorhynchus tshawytscha). Despite being salmon specialists, individuals from all three killer whale pods (J, K, L) have been observed harassing and killing porpoises (family Phocoenidae) without consuming them. Retrospectively , we identified and analyzed 78 episodes of Southern Resident killer whales harassing porpoises between 1962 and 2020, of which 28 resulted in the porpoise's death ("phocoenacide"). Fifty-six episodes involved harbor porpoise (Phocoena phocoena), 13 involved Dall's porpoise (Phocoenoides dalli), and the porpoise species was unreported for nine episodes. Southern Resident killer whales often targeted young porpoises that were similar in size to adult Chinook salmon. † These authors contributed equally to the manuscript.

Animal social networks are often constructed from point estimates of edge weights. In many contexts, edge weights are inferred from observational data, and the uncertainty around estimates can be affected by various factors. Though this has been acknowledged in previous work, methods that explicitly quantify uncertainty in edge weights have not yet been widely adopted and remain undeveloped for many common types of data. Furthermore, existing methods are unable to cope with some of the complexities often found in observational data, and do not propagate uncertainty in edge weights to subsequent statistical analyses. We introduce a unified Bayesian framework for modelling social networks based on observational data. This framework, which we call BISoN , can accommodate many common types of observational social data, can capture confounds and model effects at the level of observations and is fully compatible with popular methods used in social network analysis. We show how the framework can be applied to common types of data and how various types of downstream statistical analyses can be performed, including non‐random association tests and regressions on network properties. Our framework opens up the opportunity to test new types of hypotheses, make full use of observational datasets, and increase the reliability of scientific inferences. We have made both an R package and example R scripts available to enable adoption of the framework.

Understanding the evolution of menopause presents a long-standing scientific challenge1,2,3-why should females cease ovulation prior to the end of their natural lifespan? In human societies, intergenerational resource transfers, for example, food sharing and caregiving, are thought to have played a key role in the evolution of menopause, providing a pathway by which postreproductive females can boost the fitness of their kin.4,5,6 To date however, other late-life contributions that postreproductive females may provide their kin have not been well studied. Here, we test the hypothesis that postreproductive female resident killer whales (Orcinus orca) provide social support to their offspring by reducing the socially inflicted injuries they experience. We found that socially inflicted injuries, as quantified by tooth rake marks, are lower for male offspring in the presence of their postreproductive mother. In contrast, we find no evidence that postreproductive mothers reduce rake marking in their daughters. Similarly, we find no evidence that either reproductive mothers or grandmothers (reproductive or postreproductive) reduce socially inflicted injuries in their offspring and grandoffspring, respectively. Moreover, we find that postreproductive females have no effect on reducing the rake marks for whales in their social unit who are not their offspring. Taken together, our results highlight that directing late-life support may be a key pathway by which postreproductive females transfer social benefits to their male offspring.

Sexual signals are often structured in bouts, which can be adjusted in response to changes in the signaller's physical and social environment. For example, we might expect individuals to adjust their own signalling behaviour in response to changes in the signalling behaviour of rivals, because this can affect their relative attractiveness to potential mates. In this study, we used a biomimetic robot to experimentally manipulate rival waving behaviour in a wild population of fiddler crabs ( Afruca tangeri ), and investigated whether this leads to changes in the activity and waving behaviour of a focal male. Analysing the focal male's behaviour using hidden Markov models and linear hurdle models, we found no evidence that the focal male's waving rate changed in response to changes in the behaviour of the robotic rival. However, bouts of waving lasted longer when the robotic rival was waving at a fast rate. Focal males were also less likely to enter their burrow when the robotic rival was waving, and spent less time in their burrow if they did enter. These results reveal tactical adjustment of behaviour by competing signallers, and highlight the flexible nature of bout-structured sexual displays.

Age-related changes in the patterns of local relatedness (kinship dynamics) can be a significant selective force shaping the evolution of life history and social behaviour. In humans and some species of toothed whales, average female relatedness increases with age, which can select for a prolonged post-reproductive lifespan in older females due to both costs of reproductive conflict and benefits of late-life helping of kin. Killer whales (Orcinus orca) provide a valuable system for exploring social dynamics related to such costs and benefits in a mammal with an extended post-reproductive female lifespan. We use more than 40 years of demographic and association data on the mammal-eating Bigg's killer whale to quantify how mother-offspring social relationships change with offspring age and identify opportunities for late-life helping and the potential for an intergenerational reproductive conflict. Our results suggest a high degree of male philopatry and female-biased budding dispersal in Bigg's killer whales, with some variability in the dispersal rate for both sexes. These patterns of dispersal provide opportunities for late-life helping particularly between mothers and their adult sons, while partly mitigating the costs of mother-daughter reproductive conflict. Our results provide an important step towards understanding why and how menopause has evolved in Bigg's killer whales.

Michael Weiss and Darren Croft introduce Orcas (Orcinus orca) also known as killer whales.

Accurate measures of lifespan and age-specific mortality are important both for understanding life-history evolution and informing conservation and population management strategies. The most accurate data to estimate lifespan are from longitudinal studies, but for many species - especially those such as toothed whales that are wide-ranging and live in difficult-to-access environments - these longitudinal data are not available. However, other forms of age-structured data - such as from mass-strandings - are available for many toothed species, and using these data to infer patterns of age-specific mortality and lifespan remains an important outstanding challenge. Here we develop and test a Bayesian mortality model to derive parameters of a mortality function from age-structured data while accounting for potential error introduced to these data by mistakes in age estimation, sampling biases and population growth. We then searched the literature to assemble a database of 269 published age-structured toothed whale datasets. We applied our mortality model to derive lifespan estimates for 32 species of female and 33 species of male toothed whale. We also use our model to characterise sex differences in lifespan in toothed whales. Our mortality model allows us to curate the most complete and accurate collection of toothed whale lifespans to date.