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i personally think that social behavior can provide many benefits for living things specifically those that like in like groups for example social animals that work together to locate food defend against predators and care for their offspring and honestly i just think it increases the chance of survival

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at least one source to support your ideas.

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at least one source to support your ideas.

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a personal statement about what you think the benefits of social behavior as a whole are for living things.

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a personal statement about what you think the benefits of social behavior as a whole are for living things.

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 how is this behavior has become an adaptation

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 how is this behavior has become an adaptation

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Description of the main attributes of increased survival

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Description of the main attributes of increased survival

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health improvement

environment a safe and supportive environment can increase you chances of survival

adaptability organisms that can adapt to environmental change and to learn and apply new skills will help with survival to

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increased survival can become an adaption through the process of natural selection natural selection is the process where specific organisms are better suited to their environment and they are more likely to survive and then passing the trait down to their offspring

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Sociality and Living in Groups

There are a number of potential ecological and animal welfare benefits from living in groups which include: predator defense, cooperative foraging, mating opportunities, and reduced vulnerability to infanticide (14).

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Adaptive Rationality Group, Centre of Ecology and Conservation, University of Exeter, Penryn, United Kingdom

Emergent evidence of aspects of sociality, such as social structure and social learning, across many vertebrate taxa, warrant more detailed consideration of their influence on welfare outcomes for wildlife. Sociality can be dynamic across organismal development, it can: provide protection through safety in numbers; may influence breeding outcomes via mate choice and alloparental care; can influence foraging success through transmission of social information and co-operation; and it can provide opportunities for the spread of novel behavior. Social learning itself provides an important mechanism for resilience in changing environments, but also has the potential to increase vulnerability or facilitate the spread of maladaptive behaviors. The welfare consequences of vertebrates living in social groups are explored using Wilson's 10 qualities of sociality as a framework, and the implications of human activities are discussed. Focus to date has been on the importance of social networks for the welfare of farmed or captive animals. Here I consider the importance of social networks and sociality more generally for the welfare of wildlife and explore Mellor's five domain model for animal welfare within the context of wildlife sociality.

Introduction

Sociality is a measure of the degree to which animals interact or form long-term or transient associations and is prevalent across a broad range of vertebrate and invertebrate taxa. Animal welfare, on the other hand, is a multidimensional field, measured through a range of criteria from health and comfort, to the ability to express natural behaviors ( 1). While some natural behaviors may be considered, many aspects of sociality are often overlooked in animal welfare assessment and this is particularly the case for wildlife, where social settings may be complex.

Wild animal conservation and individual and group welfare are deeply intertwined ( 2). While the social environment is undoubtedly of significance to both the conservation and welfare of many species, quantifying the welfare implications of aspects of sociality can be challenging in the wild. The social environment has been described as being comprised of non-random and heterogeneous social interactions ( 3, 4). The social environment of vertebrate taxa is highly diverse, ranging from: the complex third order alliances of male bottlenose dolphins ( Tursiops sp.) ( 5); to matriarchal African elephant societies ( Loxodonta sp.) ( 6); to the hierarchical group dynamics of flocking birds ( 7). For long-lived, wide ranging species, such as blue whales ( Balaenoptera musculus), there are also important spatio-temporal considerations for interpretation of behavior and associations ( 8).

Aspects of sociality, such as social structure and social transmission of information, across many vertebrate taxa, warrant a more detailed exploration of their influence on individual and group welfare outcomes for wildlife. Quantifying the welfare of wild animals presents a number of unique challenges. Wildlife welfare is often framed in terms of physiology or individual behavior. Nevertheless, better understanding of the processes of social behavior may provide important insights for the welfare of wildlife. While the focus of this review is vertebrates, there are some notable invertebrate taxa, such as cephalopods ( 9), which might also benefit from similar consideration.

Wilson's 10 qualities of sociality ( 10) have previously been suggested as a framework for evaluating the importance of sociality for individual and group welfare for cetaceans (whales and dolphins) ( 11). Here I attempt to expand these ideas to examine how sociality may be important for the welfare of a range of vertebrate taxa: exploring the risks and benefits of sociality to animal welfare; investigating the relationship between sociality and the impacts of human-induced rapid environmental change (HIREC) ( 12) on sociality and animal welfare. I then examine this new perspective within the context of the five domains model ( 13) of animal welfare.

Sociality and Living in Groups

E.O Wilson listed 10 "qualities" of sociality ( 10) which are used to understand and classify conspecific groups. These characteristics of animal societies have also been used to provide a framework for evaluating the importance of sociality for individual and group welfare in cetaceans (whales and dolphins) ( 11). Here this framework is used to examine Wilson's 10 qualities of sociality within the context of wildlife welfare, more generally ( Table 1).

Table 1. Wilson's 10 "qualities" of sociality ( 10) and welfare considerations for wild animals [adapted from Brakes ( 11)].

Risks and Benefits of Sociality

Within the vertebrate taxa there are a variety of social structures, types of association between individuals, and transmission pathways for social information. As a result the relationship between sociality and welfare across vertebrate taxa is multifaceted (see Table 2). Undoubtedly, sociality provides benefit to individual welfare, for example in predator defense, alloparental care, or by increasing foraging success through the transmission of information about food sources or other types of public information, or through co-operative foraging. But sociality also has associated risks, such as the increased probability of the spread of disease, or conspicuousness to predators. These risks and benefits can change according to the environment and may also be dynamic across the developmental stages of an organism.

Table 2. Summary of the importance of social living for wild animal welfare [after Brakes ( 11)].

While the risk of spread of disease and parasitic burdens associated with living in social groups has been extensively studied, sociality has, nevertheless, evolved independently in many taxa, demonstrating that these risks are overall offset by the benefits of sociality.

Social Support Hypotheses

The potential benefits of group living are varied ( Table 1), but social support may specifically provide beneficial effects to the recipient, irrespective of whether or not the recipient is being challenged; or social partners may modulate or downregulate the impact of stressors on the recipient's homeostasis [a process known as stress buffering ( 30)]. Stress buffering has been investigated for farm animal welfare and it has been suggested that farmers could exploit animals' natural ability to benefit from their social group to obtain better welfare outcomes ( 16). Arguably, for some highly social species of wildlife it may also be the case that better management can be achieved through strategic use of elements of animal sociality to enhance welfare outcomes.

Social Learning, Animal Culture, and Welfare

Social learning-"learning that is influenced by observation of, or interactions with, another animal (typically a conspecific) or its products" ( 31)-has been observed across a broad range of vertebrate taxa ( 32) and can be important for conservation efforts ( 21, 33, 34). Social learning can act more rapidly than the intergenerational process of Darwinian selection ( 21) and some facets of sociality may also hold important insights for wild animal welfare. For example, social learning can provide mechanisms for resilience to ecological or anthropogenic stressors, such as reduced prey abundance [for example through diversification of foraging strategies ( 22, 35, 36)]. In contrast, it can also result in the transmission of socially learnt conservative foraging specializations, such as those found in killer whales ( Orcinus orca), resulting in the species being less, rather than more, resilient to fluctuations in prey abundance ( 21).

Human Impacts on Sociality and Wildlife Welfare

The impact of human activities on wildlife and habitat is ubiquitous across virtually all ecosystems and HIREC is now a widely acknowledged phenomenon ( 12). From deforestation, to climate change and ocean acidification, wildlife around the globe is challenged by rapid environmental change, but aspects of sociality may provide opportunities for resilience to this rapid change, or increase vulnerabilities ( 45). It is timely to examine how sociality may influence wild animal welfare and responses to HIREC.

HIREC has the potential to influence all behavioral domains associated with social behavior, from breeding to communication, foraging and migration. A method to classify this vast array of threats from an individual welfare perspective is to consider acute and chronic implications for sociality and welfare ( Table 3).

Table 3. Acute and chronic implications of HIREC for sociality and welfare.

Cognition, Sociality, and Wildlife Welfare

Understanding and predicting how individuals behave in response to HIREC may be important for evaluating aspects of sociality relevant to wildlife welfare. For example, understanding how individual innovations arise may be relevant for predicting-and setting up optimal conditions for-the spread of novel behavior through social learning. Wildlife behavioral responses are governed by cognitive processes ranging from perceptual processes to learned behavior. Understanding these cognitive processes and the effect of HIREC associated stress, may also assist in reducing human impacts on wildlife ( 46). It has been argued, using the same rationale, that if cognition underlies a behavior that is relevant to welfare, then understanding these cognitive processes may help in achieving better welfare outcomes for species such as cetaceans ( 11) and this is arguably also the case for other wildlife.

Greggor et al. ( 46) argue that " cognitive theory can thus help predict how best to manipulate and exploit attentional biases, innate responses, and learning tendencies to enhance conservation efforts." For example, they argue that mitigation methods will only be effective if they are reliably perceived by the target species (e.g., preventing birds colliding with human-made structures) and that this perception is rapidly learned. The same argument can been made for a range of wildlife mitigation activities, including: acoustic "pingers" on fishing nets designed to prevent dolphins from becoming entangled and boundaries designed to prevent crop raiding. The success of some mitigation methods may also be contingent on the social transmission of information, which in turn is contingent on other aspects of sociality such as social structure.

Tools for Assessing Sociality and Wildlife Welfare

Asher et al. ( 47) argued for expanding the scope and development of new quantitative methods for the analysis of various aspects of behavioral organization as indicators for animal welfare, including: fractal analysis, temporal methods, social network analysis (SNA), and agent-based modeling and simulation. Social network analysis has now begun to emerge as a tool for assessing animal welfare. Kleinhappel et al. ( 44) argue that SNA is as yet, underused in the field of animal welfare and suggest a number of opportunities for using these methods, particularly in relation to the welfare of captive animals. However, SNA also has important application for assessing the welfare of wild populations.

Extending the Five Domains Model to Incorporate Sociality in Wildlife

There are several models that might be applicable for expanding our consideration of wild animal welfare in relation to sociality. Hinde's framework for classifying aspects of sociality ( 51) might for example provide a basis for attempting to maintain optimal conditions of sociality from a welfare perspective within wild populations. However, trying to ensure optimality in the wild is both challenging to define and difficult in practice.

A more realistic approach may be to use the five domains model of animal welfare ( 13). While generally applied to domestic and captive animals, where some degree of compromise in welfare circumstances is implicit, the five domains model may be a more practical framework for managing welfare issues arising from human impacts on wild animal sociality. It is suggested here that the five domains model may be useful to avoid directly aversive situations resulting from poor management of the sociality of wildlife, for example through the removal of repositories of social knowledge ( 25). Instead, it may be better to try to manage wild environments so that socially complex systems can thrive and avoid situations that will hinder their development. The five domains model itself resulted from recognition of the need to provide positive experiences for animals (beyond the five freedoms framework) ( 52) and this approach may be useful for exploring how we can monitor and potentially assist social systems to flourish in the wild.

The objective of the five domains model is to "draw attention to areas that are relevant to both animal welfare assessment and management" and focus on the "presence or absence of various internal physical/functional states and external circumstances that give rise to welfare-relevant negative and/or positive mental experiences, i.e., affects" ( 13). In formulating the five domains model, Mellor "carefully and cautiously" considered the affects of the first four domains on nutrition, environment, health and behavior, on the fifth domain of mental state.

Considering some of the issues raised in this review, here I have attempted to explore some of the possible implications of sociality for wildlife welfare within the context of the five domains model ( Table 4). This list is not intended to be exhaustive, but instead is a first exploration.

Table 4. Potential interface between sociality and wild animal welfare as it relates to the five domains model ( 13) of animal welfare.

Play and the Five Domains Model

The stated objective of the five domains model is to provide opportunities for positive welfare ( 13) and it follows that one such opportunity might be granted through play in wildlife. Play is variable across species, has a variety of definitions ( 54, 55) and its proximate mechanisms and ultimate functions are not well understood. Play can indicate good welfare conditions, but can also be an indication of stressful situations, for example resulting in reduced parental care ( 56). Play has a variety of roles and it has been argued that play is usually the first behavior to disappear when welfare is compromised: when animals are stressed, anxious, hungry or unwell ( 57, 58). It has been suggested that play may be a reliable indicator of psychological and physical well-being ( 56). However, using play as a definitive indicator for optimal or good welfare can be confounded by several factors including: the heterogeneity of play; the fact that play can be a coping strategy; factors with no obvious relation to animal welfare can influence intrinsic playfulness within individuals; and the fact there are circumstances in which negative welfare can actually result in more play. Thus, play cannot definitely be considered an indicator of optimal or good welfare ( 59). But play can also be dynamic across life stages and arguably may be important for positive mental state in some species (see Table 4). For example, it has been argued that play may be socially contagious and therefore capable of spreading good welfare in groups ( 56).

Play behavior may also be of value in relation to innovation. In addition to having a potentially positive influence on mental states, there is potentially reciprocity between the social aspects of play, innovation and how innovation may then spread in wild populations. Bateson ( 57) argues that numerous functions for play have been proposed but they are not mutually exclusive and there are indications that those individuals who play most are most likely to survive and reproduce.

Potentially, the reciprocity between play and vital rates may manifest through the social transmission of innovative behaviors. Although fitness outcomes are not always synonymous with welfare outcomes, associations and bonds forged through play may influence the spread of information across a network and the trial and error aspects of play may help individuals accumulate latent information, which may facilitate innovation as their environment changes. Thus, it can be argued that play may be an important part of the behavioral repertoire of some wildlife for maintaining resilience.

Conclusion

The interplay between sociality and the welfare of wildlife is multifaceted. The widespread evolution of sociality, social learning, and in some instances even animal culture, among vertebrate taxa is evidence that overall the benefits of social living out-weight the costs. Nevertheless, the individual and group welfare issues associated with the processes of social living are undoubtedly important for the consideration of wild animal welfare.

The principle objective of the five domains model is to move beyond an animal welfare focus of survival, toward individuals thriving. In order to meet the objective of thriving wild vertebrate populations, it is essential to incorporate aspects of sociality into their welfare assessments.

Author Contributions

The author confirms being the sole contributor of this work and has approved it for publication.

Funding

Funding for open access of this article was kindly supplied by the Institutional APC Fund, University of Exeter.

Conflict of Interest Statement

The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Acknowledgments

I would like to thank Sasha Dall for commenting on an earlier version of this text and Andy Butterworth, Mark Simmonds, and Clive Phillips for valuable conversations which have helped shape some of the thoughts presented here.

References

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Sociality and Wild Animal Welfare: Future Directions

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Emergent evidence of aspects of sociality, such as social structure and social learning, across many vertebrate taxa, warrant more detailed consideration of their influence on welfare outcomes for wildlife.