Richard McElreath’s research while affiliated with Max Planck Institute for Evolutionary Anthropology and other places

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Publications (206)


Posterior means and 89% lower (Lo) and higher (Hi) posterior density intervals for each latent learning parameter, by treatment and model. ¯ θ baseline , ¯ θ t=1 , and ¯ θ t=25 denote the average of individual-specific parameter estimates from the EWA baseline model, and from material choice 1 and 25 in the EWA monotonic model, respectively.
Dynamic strategic social learning in nest-building zebra finches and its generalisability
  • Preprint
  • File available

June 2025

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8 Reads

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Tristan Eckersley

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Animals often balance asocial and social information strategically, adjusting when and from whom they copy based on context. Yet the cognition driving this dynamic—and its broader implications—remains poorly understood. Here, we show that nest-building zebra finches follow a copy-if-dissatisfied strategy, but only when social information aligns with recent experience: after observing a conspecific build a nest, males were more likely to choose the demonstrated 'social' material—particularly on their first choice—if they had previously used low-quality material. Using Bayesian cognitive modelling, we estimated how latent learning mechanisms underpinned males' material choices, identifying two asocial and two social governing parameters. These parameters provide the first formal evidence for the cognitive basis of birds' nest building. Forward simulations informed—but not prescripted—by these parameters approximated observed material-choice trajectories, supporting their causal role. Additional simulations targeting simplified real-world contexts showed that payoff structure—not (dis)satisfaction—drove social material choice, though higher rewards did not proportionally boost its use, offering preliminary insights into the mechanisms underlying material-use variation more broadly. Our study illustrates how computational modelling can robustly link behaviour to underlying learning mechanisms and probe the generalisability of animal cognition—a rarity in this field.

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Fig. 1 | Descriptive AF-MTG evidence. A High resolution colour FA image overlay on an anatomical image of one exemplary chimpanzee. Note the descending blue line representing white matter fibres running in top-down orientation, lateral to the optic radiation (green). The absence of this blue line has been reported in previous chimpanzee studies, but it is present in humans 15 . B Deterministic tractography (purple) overlay on an anatomical image demonstrating the descending arcuate fascicle (AF-)MTG connection at the location from the blue line in the colour FA image (see (A)). Also, this structural image shows a clear contrast between the optic radiation and the lateral structure representing the AF. OR Optic radiation. C Example individual in sagittal view depicting a long AF-MTG connection as assessed by virtual dissection deterministic tractography. STG superior temporal gyrus, MTG middle temporal gyrus.
Fig. 2 | Quantitative MTG connectivity of the arcuate fascicle (AF). Group average of the probabilistic AF-STG (blue) and AF-MTG (red) tractography results for the right and left hemispheres and for chimpanzees (A) and humans (B). Average tractography shows the pathway at more than 10 probabilistic streamlines per voxel for chimpanzees (A) and more than 100 streamlines per voxel for humans (B). For different thresholds see Supplementary Fig. S6. C Individual normalised connectivity strength for the AF-STG (blue) and AF-MTG (red) on a log scale for the left hemispheres (N = 20) and right hemispheres (N = 19). D Comparison of AF-STG and AF-MTG connection strength between humans and chimpanzees, highlighting an inverse pattern of strength. In chimpanzees, median AF-STG connection was found to be 14.3-(left, N = 20) and 45.3-(right, N = 19) times stronger than AF-MTG pathway for the left and right hemispheres, respectively. In humans median AF-MTG connection was found to be 6.32-(left, N = 20) and 2.5-(right, N = 20) times stronger than the AF-STG pathway. The boxplots show the median (black thick line), the 25% and 75% quartiles and the individual data points. The orange horizontal lines represent the model mean estimates and the 95% Credible Interval (CI). Comparison of AF-MTG lateralisation between chimpanzees and humans is shown in Supplementary Fig. S8. Individual tractography results are shown in Supplementary Fig. S5 and are provided as supplementary information files. STG superior temporal gyrus, MTG middle temporal gyrus.
Fig. 3 | Asymmetry quotient for individual chimpanzee brains for the AF-STG (right panel) and AF-MTG (left panel) connections. Left: For the AF-STG, all but one individual are lateralised (19 individuals meet the inclusion criteria) and 10 (53%) chimpanzees are left lateralised whereas 8 (42%) are right lateralised. For the AF-MTG connection, 11 (65%) chimpanzees are left lateralised, and 6 (35%) are right lateralised. Wild individuals (purple), zoo-housed individuals (orange), sanctuaryhoused individuals (grey). The corresponding individual connectivity values are shown in the Supplementary Fig. S9. AF arcuate fascicle, STG superior temporal gyrus, MTG middle temporal gyrus.
Fig. 4 | Schematic summary of the findings on the temporal endings of the AF (purple). The figure shows the AF-STG (blue) and AF-MTG (red) in the left hemisphere of macaques, chimpanzees, and humans (Supplementary Fig. S1). Our results replicate the AF-STG connection in chimpanzee 8,11,14,15,19 and macaques 7,9,19
Long arcuate fascicle in wild and captive chimpanzees as a potential structural precursor of the language network

May 2025

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185 Reads

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2 Citations

The arcuate fascicle (AF) is the main fibre tract in the brain for human language. It connects frontal and temporal language areas in the superior and middle temporal gyrus (MTG). The AF’s connection to the MTG was considered unique to humans and has influenced theories of the evolution of language. Here, using high-resolution diffusion MRI of post-mortem brains, we demonstrate that both wild and captive chimpanzees have a direct AF connection into the MTG, albeit weaker than in humans. This finding challenges the notion of a strictly human-specific AF morphology and suggests that language-related neural specialisation in humans likely evolved through gradual evolutionary strengthening of a pre-existing connection, rather than arising de novo. It is likely that this neural architecture supporting complex communication was already present in the last common ancestor of hominins and chimpanzees 7 million years ago, enabling the evolution of language processes in the human lineage.


Bayesian longitudinal social network models: An implementation in R using STRAND

May 2025

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4 Reads

Across the social and biological sciences, researchers study social networks that are often longitudinal in nature, featuring the same set of individuals interacting with one another over long periods of time. Such network data may be collected via panel studies in human communities, or via long-form behavioral observation programs in animal communities. Important research questions often hinge on how dyadic ties or flows in the network at one time-step are associated with dyadic ties or flows at subsequent time-steps. Similar questions focus on the relationship between nodal characteristics across time-steps: for example, is an individual with a high out-degree at one time-step more likely to have a high in-degree in the next? These questions are effectively addressed by longitudinal extensionsof generative social network models like the Social Relations Model (SRM). Here, we present a novel longitudinal extension of the SRM, provide an implementation of the model in the STRAND R package, and provide tutorials teaching end-users how to fit the model to their own data. We provide worked examples of data analysis, parameter visualization, and results interpretation, using data-sets from both social science and animal behavior. The software package allows end-users to deploy complex longitudinal network analysis models using only simple, base-R model syntax.


Spectrograms illustrating different types of utterances: single vocal units and vocal sequences
Spectrograms are represented with frequency along the y-axis and time in seconds along the x-axis (A) depicts an instance of a single vocal unit/call type twitter (TW) consisting of repetitions of twitter elements, (B) depicts an instance of the single vocal unit/call type grunt (GR) consisting of repetitions of grunt elements, and (C) depicts a vocal sequence combining the GR and TW vocal units to form a GR-TW-GR sequence. All three examples fall into the overarching category of “utterance” regardless of the number of different vocal units that compose them.
Proportion of sooty mangabey vocal utterances containing 1–11 different call types, categorised by age class
The total number of recordings and individuals making up each age class are listed on the x axis. The gradient of blue indicates sequences of increasing length from lighter to darker. ‘Iterated units’ refers to the repeated emission of a vocal unit that already occurs earlier in the vocal sequence, such that GR-TW-GR is treated as a different sequence to GR-TW. The ‘no iterated units’ condition would remove the repeated instance of GR and treat those two utterances as equivalent.
Sex and age effects on mangabey vocal repertoire size and maximum utterance length
A Depicts the model predictions presented alongside our raw data for repertoire size (i.e., the number of different utterance types, including both vocal units produced singly and vocal sequences) when iterated vocal units are included, categorised by age class and sex. B Presents the same information for the scenario where iterated vocal units have been removed from sequences. C Depicts the maximum utterance length with iterated vocal units included and (D) presents it with iterated units removed. The thick horizontal black lines indicate the mean posterior prediction derived from the model and the vertical black line the lower and upper bounds of the 89% credible interval. Circles indicate the individuals contributing to our sample, with the size of the circle indicating how many recordings were collected for that individual.
Vocal sequence diversity and length remain stable across ontogeny in a catarrhine monkey (Cercocebus atys)

March 2025

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118 Reads

Communications Biology

During childhood, human speech utterances increase steadily in complexity, length and diversity. In contrast, the vocal repertoire of non-human primates has long been considered fixed from birth. Recent studies showing the acquisition of vocal sequences during ontogeny in chimpanzees and marmosets challenge this view. Here we further explore the potential flexibility of non-human primate vocal production by comparing the vocal sequence repertoire across age groups in sooty mangabeys, a species with a rich sequence repertoire for a catarrhine monkey. We recorded 1844 utterances from 75 individuals from two wild groups in Taï National Park, Ivory Coast. We used custom-made Bayesian models specifically designed to estimate the individual repertoire size of vocal sequences while accounting for under-sampling of certain vocalisations in certain individuals. We hereby provide a tool to estimate vocal repertoire size applicable to other taxa. We found no relevant ontogenetic changes in vocal repertoire size and utterance length. Ontogenetic vocal sequence expansion is therefore not universal among primates that routinely use vocal sequences to communicate. Rather, this feature may have evolved independently in distantly-related taxa due to social features thought to promote vocal complexity, such as the complex social organisation of chimpanzees and the cooperative breeding systems of marmosets.


Social learning preserves both useful and useless theories by canalizing learners’ exploration

January 2025

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85 Reads

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2 Citations

In many domains, learning from others is crucial for leveraging cumulative cultural knowledge, which encapsulates the efforts of successive generations of innovators. However, anecdotal and experimental evidence suggests that reliance on social information can reduce the exploration of the problem space. Here, we experimentally investigate the extent to which cultural transmission fosters the persistence of arbitrary solutions in a context where participants are incentivized to improve a physical system across multiple trials. Participants were exposed to various theories about the system, ranging from accurate to misleading. Our findings indicate that even under conditions conducive to exploration, the transmission of cultural knowledge canalizes learners’ focus, limiting their consideration of alternative solutions. This effect was observed in both the theories produced and the solutions attempted by participants, irrespective of the accuracy of the provided theories. These results challenge the notion that arbitrary solutions persist only when they are efficient or intuitive and underscore the significant role of cultural transmission in shaping human knowledge and technologies.



Bridging theory and data: A computational workflow for cultural evolution

November 2024

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15 Reads

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4 Citations

Proceedings of the National Academy of Sciences

Cultural evolution applies evolutionary concepts and tools to explain the change of culture over time. Despite advances in both theoretical and empirical methods, the connections between cultural evolutionary theory and evidence are often vague, limiting progress. Theoretical models influence empirical research but rarely guide data collection and analysis in logical and transparent ways. Theoretical models themselves are often too abstract to apply to specific empirical contexts and guide statistical inference. To help bridge this gap, we outline a quality-assurance computational workflow that starts from generative models of empirical phenomena and logically connects statistical estimates to both theory and real-world explanatory goals. We emphasize and demonstrate validation of the workflow using synthetic data. Using the interplay between conformity, migration, and cultural diversity as a case study, we present coded and repeatable examples of directed acyclic graphs, tailored agent-based simulations, a probabilistic transmission model for longitudinal data, and an approximate Bayesian computation model for cross-sectional data. We discuss the assumptions, opportunities, and pitfalls of different approaches to generative modeling and show how each can be used to improve data analysis depending on the structure of available data and the depth of theoretical understanding. Throughout, we highlight the significance of ethnography and of collecting basic cultural and demographic information about study populations and call for more emphasis on logical and theory-driven workflows as part of science reform.


Strategic housing decisions and the evolution of urban settlements: optimality modelling and empirical application in Ulaanbaatar, Mongolia

October 2024

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60 Reads

Investments in housing influence migration and landscape construction, making them a key component of human–environment interactions. However, the strategic decision-making that builds residential landscapes is an underdeveloped area of research in evolutionary approaches to human behaviour. Our contribution to this literature is a theoretical model and an empirical test of this model using data from Ulaanbaatar, Mongolia. We develop a model of strategic housing decisions using stochastic dynamic programming (SDP) to explore the trade-offs between building, moving and saving over time, finding different trade-offs depending on optimization scenarios and housing costs. Household strategies are then estimated using data on 825 households that settled in the Ger districts of Ulaanbaatar between 1942 and 2020. The Ger districts are areas of self-built housing that feature both mobile dwellings (gers) and immobile houses (bashins). Using approximate Bayesian computation (ABC), we find the parameters of our dynamic programming model that best fit the empirical data. The model is able to capture the time horizon of housing changes and their bi-directionality, showing that moving from a fixed to mobile dwelling can also be an optimal strategy. However, the model underpredicts household persistence in dwelling types. We discuss deviations from model predictions and identify a more detailed exploration of risk and population mixes of strategies as key steps for future research.


Assumed relationship between refinement level and payoff increment, illustrated by refinements in hammer technology
The payoff to a behavior is given by its basic (unrefined) payoff plus an increment that is a function of refinement level. Refining resulted in payoff increments (illustrated by the black curve) surpassing the highest basic payoff (dashed blue line) after approximately 10 refine moves. The inset illustrates an example distribution for the basic payoffs–most payoffs are low, and few payoffs are high.
Scores and learning in Stage 1
Relationship between score and (A) the proportions of learning (INNOVATE+OBSERVE+REFINE) moves, (B) Score as a function of REFINE moves, averaged over each entry in Stage 1. (C) Distribution of the proportion of learning moves averaged by entry, for each extension over all entries and (D) for the top-ten best-performing entries.
Cultural diversity measured across extensions for Stages 2 and 3
(A-D) Cumulative culture led to plummeting diversity in both the behaviors performed and known about, as populations converged on heavily refined behaviors, that (E-H) persist for long periods of time. ‘Behavior’ refers to the acts that the population was using at each timepoint, and ‘Knowledge’ refers to the acts present in the repertoire of at least one individual, but not necessarily used. ‘Amount’ captures the proportion of behaviors or knowledge known about within the population (i.e., mean proportion of possible behaviors used or known by at least one agent in each round in the last quarter of the simulations), ‘evenness’ measures the flatness of the frequency distribution (using Pielou’s evenness index, see Materials and methods), and ‘persistence’ refers to the length of time the behavior or knowledge persisted in the population (i.e. mean or maximum number of rounds that a behavior within a population was exploited or that knowledge of it persisted within the population, without a break; see Materials and methods for more detail). Data for the ‘No extension’ case give a baseline comparison and are from 1,000 simulations using the top-ten tournament entries with randomly chosen parameter values.
The refinement paradox
(A) and (B). Predicted scores from a linear mixed model accounting for between-entry variation, using Stage 1 data, for (A) all entries, and (B) top-ten entries, that did and did not play REFINE, in refined compared to non-refined environments (Tables B and C in S1 Supporting Information). Top-ten entries used refinement strategically, achieving higher scores, and constructing maximally refined environments beneficial to all. (Circles indicate entry means and diamonds show group means, as predicted by the model. Environments were defined as refined when a population reached the highest refinement level). (C) Relative fitness of ‘clever’ REFINE entries over a blind copier (OBSERVE-once-then-EXPLOIT-forever; see Materials and methods for details). ‘Clever’ entries had the advantage at low refinement levels, but were vulnerable to invasion at higher refinement levels.
The refinement paradox and cumulative cultural evolution: Complex products of collective improvement favor conformist outcomes, blind copying, and hyper-credulity

September 2024

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126 Reads

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4 Citations

Social learning is common in nature, yet cumulative culture (where knowledge and technology increase in complexity and diversity over time) appears restricted to humans. To understand why, we organized a computer tournament in which programmed entries specified when to learn new knowledge and when to refine (i.e. improve) existing knowledge. The tournament revealed a ‘refinement paradox’: refined behavior afforded higher payoffs as individuals converged on a small number of successful behavioral variants, but refining did not generally pay. Paradoxically, entries that refined only in certain conditions did best during behavioral improvement, while simple copying entries thrived when refinement levels were high. Cumulative cultural evolution may be rare in part because sophisticated strategies for improving knowledge and technology are initially advantageous, yet complex culture, once achieved, favors conformity, blind imitation and hyper-credulity.


A causal framework for the drivers of animal social network structure

June 2024

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142 Reads

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1 Citation

A bstract A major goal of behavioural ecology is to explain how phenotypic and ecological factors shape the networks of social relationships that animals form with one another. This inferential task is notoriously challenging. The social networks of interest are generally not observed, but must be approximated from behavioural samples. Moreover, these data are highly dependent: the observed network edges correlate with one another, due to biological and sampling processes. Failing to account for the resulting uncertainty and biases can lead to dysfunctional statistical procedures, and thus to incorrect results. Here, we argue that these problems should be understood—and addressed—as problems of causal inference. For this purpose, we introduce a Bayesian causal modelling framework that explicitly defines the links between the target interaction network, its causes, and the data. We illustrate the mechanics of our framework with simulation studies and an empirical example. First, we encode causal effects of individual-, dyad-, and group-level features on social interactions using Directed Acyclic Graphs and Structural Causal Models. These quantities are the objects of inquiry, our estimands . Second, we develop estimators for these effects—namely, Bayesian multilevel extensions of the Social Relations Model. Third, we recover the structural parameters of interest, map statistical estimates to the underlying causal structures, and compute causal estimates from the joint posterior distribution. Throughout the manuscript, we develop models layer by layer, thereby illustrating an iterative workflow for causal inference in social networks. We conclude by summarising this workflow as a set of seven steps, and provide practical recommendations.


Citations (61)


... We understand this core assumption of our framework to be empirically justified in light of research on causally opaque cultural knowledge showing that humans depend on socially transmitted information without necessarily understanding the causal mechanisms that determine the effectiveness of any particular cultural element (e.g. Boyd et al. (2011); Derex et al. (2019Derex et al. ( , 2025; Harris et al. (2021); Henrich (2021)). If such causal opacity dominates even in the relatively simple domain of tasks necessary for survival as a subsistence forager, then it is surely also present in the domain of norm cognition that governs cooperation in complex modern societies 1 , see also (Jagiello et al., 2022;Whitehouse, 2021). ...

Reference:

Societal and technological progress as sewing an ever-growing, ever-changing, patchy, and polychrome quilt
Social learning preserves both useful and useless theories by canalizing learners’ exploration

... /2025 expertise into their models and a pooling of information from disparate sources while controlling for confounds in detailed ways. Thus, this form of modeling is useful for causal inference in fields such as anthropology in which experiments are not possible (Deffner et al., 2022(Deffner et al., , 2024Friederici et al., 2024). Primarily the field of neuroimaging has utilized innovations in machine learning algorithms that maximize prediction accuracy to "decode" the complexity of the brain (Grootswagers et al., 2017). ...

Bridging theory and data: A computational workflow for cultural evolution
  • Citing Article
  • November 2024

Proceedings of the National Academy of Sciences

... Archaeoriddle relies on 'research gamification', inspired by previous work in the social and behavioural sciences (Axelrod, 1980;Rendell et al., 2010; but see also Miu et al., 2024 for a similar, more recent approach), where participants searched for solutions to specific questions as they made their way through tournaments. Similar attempts to investigate the robustness of inferential techniques have been carried out in statistical science. ...

The refinement paradox and cumulative cultural evolution: Complex products of collective improvement favor conformist outcomes, blind copying, and hyper-credulity

... In some conditions, they can additionally choose whether or not-and how-to use the information provided by a third-party arbitrator to determine their transfer decisions. To classify individual-level strategy-use on the basis of a sequence of game-play decisions, we use a Bayesian finite mixture model [18][19][20]. This model evaluates the probability of the game-play sequence of individual i under each strategy s-e.g., ATFT-out of a set of S possible strategies, and then determines the relative probability weight that individual i deploys strategy s. ...

IPDToolkit: An R package for simulation and Bayesian analysis of iterated prisoner’s dilemma game-play under third-party arbitration
  • Citing Preprint
  • June 2024

... Dyadic non-cooperative games are those where every player has only two possible actions (pure strategies) [3], [4]. A classic example of the dyadic game is the prisoner's dilemma game [5], [6] and other similar games dealing with economic [5], [7], [8], ecologic [9], [10], jurisprudential [11], political [12], social [7], [13] dilemmas of choice and involving more than just two players [9], [10], [14]. ...

Evidence of direct and indirect reciprocity in network-structured economic games

Communications Psychology

... Our results contribute to the debate on humans' propensity to rely on social information and the implications for the persistence of arbitrary solutions. Some scholars argue that individuals are inclined to rely on social information, which can facilitate the adoption of hard-to-devise, unintuitive solutions [12,22,23]. Others contend that individuals should be cautious about social information to avoid the risk of being accidentally or intentionally misled [24,25]. ...

The refinement paradox and cumulative cultural evolution: collective improvement in knowledge favors conformity, blind copying and hyper-credulity

... We are op6mis6c, however, that these beginnings are evidence of a strong founda6on for expanding HBE scholarship in China. In order to do so, research teams must cram new projects with care, ensuring that ques6ons are appropriately framed and assump6ons made explicit, ideally with clear framing of sets of alterna6ves (PlaL 1964) that are rooted in op6mality frameworks (McElreath & Koster 2024), thereby avoiding common pixalls that create a sense of "just-so" story telling (e.g., Gould & Lewon6n 1979; but see Reeve & Sherman 1993). Without a clear set of alterna6ves being specified, any rela6onship between a behavior and some proxy of fer6lity may be spurious and claims for natural selec6on poten6ally untenable (McElreath & Koster 2024). ...

The End of Human Behavioral Ecology
  • Citing Chapter
  • March 2024

... However, subsequent studies using the same database of captive individuals have not observed these traits (AF-MTG connection 8,10-12 , left lateralisation 11,12,16 ), leading to the consensus that both traits are unique to humans. Here, we re-examine chimpanzee AF anatomy using independent high-resolution data from the Evolution of Brain Connectivity Project (EBC) 17,18 , which includes data from both wild and captive individuals. ...

Brain structure and function: A multidisciplinary pipeline to study hominoid brain evolution

Frontiers in Integrative Neuroscience

... As argued above, proxies are central to the scaffolding, amplification and diversification of human intelligence. This is consistent with ideas that proxies and culture coevolve (Brinkmann et al. 2023), and that the "individual" can extend to proxies (Krakauer et al. 2020) and the emergence of a new level of individuality (Rainey 2023). TIS has the ingredients to explain the generality of human intelligence, but would need to be further developed into explicit mathematical models in order to understand the varied contributions of proxies. ...

Machine culture
  • Citing Article
  • November 2023

Nature Human Behaviour

... Sampling effort is an exposure variable, which strongly-indeed, proportionally-influences the outcome counts. To account for variation in sampling effort across dyads in a way that fully propagates uncertainty (see Ross et al., 2023, for derivation), a Bayesian model of network data will generally take the form: ...

Modelling animal network data in R using STRAND