Bibliometric investigation of the integration of animal personality in conservation contexts

Consistent individual differences in behavior, commonly termed animal personality, are a widespread phenomenon across taxa that have important consequences for fitness, natural selection, and trophic interactions. Animal personality research may prove useful in several conservation contexts, but which contexts remains to be determined. We conducted a structured literature review of 654 studies identified by combining search terms for animal personality and various conservation subfields. We scored the relevance of personality and conservation issues for each study to identify which studies meaningfully integrated the 2 fields as opposed to surface-level connections or vague allusions. We found a taxonomic bias toward mammals (29% of all studies). Very few amphibian or reptile studies applied personality research to conservation issues (6% each). Climate change (21%), invasive species (15%), and captive breeding and reintroduction (13%) were the most abundant conservation subfields that occurred in our search, though a substantial proportion of these papers weakly integrated conservation and animal personality (climate change 54%, invasive species 51%, captive breeding and reintroduction 40%). Based on our results, we recommend that researchers strive for consistent and broadly applicable terminology when describing consistent behavioral differences to minimize confusion and improve the searchability of research. We identify several gaps in the literature that appear to be promising and fruitful avenues for future research, such as disease transmission as a function of sociability or exploration as a driver of space use in protected areas. Practitioners can begin informing future conservation efforts with knowledge gained from animal personality research.     

Variation in plasticity of personality traits implies that the ranking of personality measures changes between environmental contexts: calculating the cross-environmental correlation

Studies increasingly explore whether there is variation between individuals in how they adjust their behavior to different environmental contexts using random regression analysis (RR). RR estimates the between-individual variance in elevation (expected behavior in the mean environment) and variance in plasticity (individual-specific adjustment of behavior to the environment) and the covariance between these properties. These (co)variances allow deriving both environment-specific repeatability estimates and the individual-level correlation in behavior between environmental conditions, both of which are of key interest for understanding consistency in behavior over space and time, a core issue in animal personality research. However, very few studies use the RR estimates to produce this information. In this paper, I outline how to extract this information (including example code in R) and apply the approach to 16 estimates based on nine published RR studies. Despite the fact that the behaviors have a decent repeatability under different environmental conditions, I find that in half the cases, the cross-environmental correlation is low (<0.4). This implies that individuals differ in their behavioral trait values but that the ranking of these individual values tends to change between environmental conditions. This level of inconsistency in animal personality traits appears not to have been fully appreciated, and future studies on plasticity using RR should explicitly include it.     

Underaddressed animal-welfare issues in conservation

Much progress has been made toward assessing and improving animal welfare in conservation. However, several glaring knowledge gaps remain where animal-welfare concerns exist but animal-welfare studies have not been performed in politically sensitive contexts. Based on contemporary issues in Australia, we identified 4 topics that require more research: animal-welfare oversight for operations designated as management (as opposed to research); animal-welfare impacts of biological agents used to control invasive animals; welfare of animals hunted recreationally; and animal-welfare impacts associated with indigenous wildlife use. Animal-welfare science may be applied to these sensitive topics through simple quantitative studies (e.g., quantifying the frequency of adverse animal-welfare events). Several such studies have effectively addressed animal-welfare concerns in similarly contentious contexts, including feral camel (Camelus dromedarius) culling in Australia, recreational hunting in Scandinavia, and indigenous whale hunting in the United States. For discussions of animal welfare in conservation to be evidence-based, courageous research is required in the 4 key areas we identified.     

Habitat structure helps guide the emergence of colony-level personality in social spiders

Studying the environmental factors that guide the emergence of collective behaviors is instrumental to understanding the ecology and evolution of animal societies. Although recent work has provided insights into the demographic factors that influence inter-colony variation in collective behavior (i.e., colony-level personality or collective personality), relatively few studies have investigated how the physical environment (e.g., habitat structure) affects colony-level personality. Here, we study the emergence of collective personality in prey capture behavior in the social spider, Stegodyphus dumicola. We measured collective prey capture behavior four times over 36 days in a classic repeated measures design. We used four different artificial habitat (web support) structures in three different treatments: habitat structure was either (1) fixed and undisturbed, (2) disturbed with a complete removal of webbing between each measurement, or (3) disturbed with changes of habitat structure between each measurement. Our results revealed that repeatability in colony-level personality was retained as long as habitat structure was not altered. However, the repeatability of colony-level personality declined precipitously when groups were forced to build their webs on novel habitat structures. Furthermore, habitat structure affected collective capture behavior, that is, latency to attack and the number of attackers differed among colonies on different habitat structures. Collectively, our data demonstrate that habitat structure is instrumental in shaping both the mean and repeatability of the collective behavior of colonies and may influence overall foraging success.     

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.     

Animal agency in wildlife conservation and management

Wildlife conservation and management (WCM) practices have been historically drawn from a wide variety of academic fields, yet practitioners have been slow to engage with emerging conversations about animals as complex beings, whose individuality and sociality influence their relationships with humans. We propose an explicit acknowledgement of wild, nonhuman animals as active participants in WCM. We examined 190 studies of WCM interventions and outcomes to highlight 3 common assumptions that underpin many present approaches to WCM: animal behaviors are rigid and homogeneous; wildlife exhibit idealized wild behavior and prefer pristine habitats; and human–wildlife relationships are of marginal or secondary importance relative to nonhuman interactions. We found that these management interventions insufficiently considered animal learning, decision-making, individuality, sociality, and relationships with humans and led to unanticipated detrimental outcomes. To address these shortcomings, we synthesized theoretical advances in animal behavioral sciences, animal geographies, and animal legal theory that may help conservation professionals reconceptualize animals and their relationships with humans. Based on advances in these fields, we constructed the concept of animal agency, which we define as the ability of animals to actively influence conservation and management outcomes through their adaptive, context-specific, and complex behaviors that are predicated on their sentience, individuality, lived experiences, cognition, sociality, and cultures in ways that shape and reshape shared human–wildlife cultures, spaces, and histories. Conservation practices, such as compassionate conservation, convivial conservation, and ecological justice, incorporate facets of animal agency. Animal agency can be incorporated in conservation problem-solving by assessing the ways in which agency contributes to species’ survival and by encouraging more adaptive and collaborative decision-making among human and nonhuman stakeholders.     

Research Training for Releasable Animals

Abstract:  Restrictions on training potentially releasable animals such as those undergoing rehabilitation care or wild-caught captives have limited our understanding of sensory processes, cognition, and physiology important for conservation of species. It is common practice among several U.S. federal agencies to limit training of animals available for release. The behavioral argument justifying this practice is that training habituates subjects to people and conditions them to associate people with rewards such as food; habituation to and positive associations with people will lead animals into dangerous situations after their release. If under special circumstances research training is permitted, all trained behaviors must be extinguished before release because behaviors will transfer to the natural setting. Research on animal learning and memory indicates that these may not be accurate scenarios. A review of the literature on habituation, classical and instrumental conditioning, and compound conditioning suggests that learning within a research setting does not add to learning that already occurs in procedures associated with basic feeding and care. In fact, animals probably learn less about people in a training setting. Furthermore, context-specific effects on memory limit behavior transfer from captive to natural settings. Extinction is strongly susceptible to context effects, which suggests that extinction does not effectively transfer to the postrelease setting. Counterintuitively, extinction of responses to experimental stimuli under some circumstances may enhance undesirable learning about humans. Under those circumstances in which isolation from human contact is difficult or undesirable, behavioral research can present an ideal format for minimizing learning about humans and provide biological information important for conservation.     

Learning outcomes measured in zoo and aquarium conservation education

Communicating the topic of conservation to the public and encouraging proenvironmental behaviors can mitigate loss of biodiversity. Thus, the evaluation of educational efforts is important to ascertain the educational effects and provide high-quality conservation education. The learning outcomes of conservation education are diverse (e.g., attitudes, knowledge, and behavior). Considering the specific characteristics of these different outcomes and the factors that influence them is crucial to delivering successful conservation education. We reviewed 29 peer-reviewed articles published in English from January 2011 to April 2020 on empirical studies of learning outcomes of on-site conservation education in zoos and aquaria, institutions that seek to educate the public about conservation. We examined the range of learning outcomes, their definitions, and factors that influenced them. Cognitive outcomes were most frequently investigated (37%) in comparison with other outcomes (e.g., affective outcomes, 31%). The articles did not use explicit definitions for learning outcomes, and implicit or explorative definitions provided were inconsistent. Outcomes were influenced by various factors (e.g., prior experiences, staff interaction, animal behavior). Our results suggest the agenda of conservation education research should be broadened by examining all learning outcomes relevant to behavior change. Educational and behavior change theories should be used as a background for conservation education research to ensure clear and consistent definitions, derive appropriate instruments to measure learning outcomes, and relate learning outcomes to influencing factors. We recommend conservation education researchers and practitioners to treat conservation education holistically and acknowledge its learning outcomes’ full complexity.     

Campesino hunting and conservation in Latin America

Hunting presents a paradox for biodiversity conservation. It is both a problem and a solution to species declines and poverty. Yet, conservation scientists hold different assumptions about the significance and sustainability of hunting based on the cultures and identities of hunters. In Latin America, conservationists largely sort hunters as either indigenous or campesino. Indigenous hunters are often characterized as culturally driven stewards of wildlife sustainability. Campesino hunters, by contrast, are described as peasants—cultureless, uneducated, and uncaring toward wildlife sustainability. Although such ethnically fueled hunting discourse promotes hunting research, campesino hunters remain underrepresented in most comparative hunting reviews. Moreover, there are no targeted syntheses on the current state of knowledge about campesino hunting, nothing to guide conservation research and practice with and for the largest group of hunters in Latin America. We reviewed 334 articles published from 1937 to 2018 in English (55%) and Spanish (45%)—mostly published in 145 peer-reviewed journals—on the meanings, motivations, and sustainability of campesino hunting in Latin America. Although studies spanned 17 countries, 7 ecosystems, and >75 indigenous and nonindigenous demographics in 30 research contexts, they predominantly focused on nonindigenous campesinos for species-specific conservation and protected area management in tropical broadleaf forests of Mexico, Peru, and Colombia. Authors used 12 methods to collect campesino hunting data, primarily interviews, surveys, and questionnaires, and drew from 10 local and traditional knowledge themes about wildlife trends and uses. Eighteen drivers, 14 constraints, and 10 conflicts—mainly subsistence, income, ethics, regulations, and crop or livestock protection—shaped whether campesino hunters pursued 799 species, 70% of which were least concern species. Yet, only 25 studies (8%) empirically assessed sustainability. Our results show the need for increased interdisciplinary and geographic engagement with campesino hunting across Latin America.     

Personality trait differences between mainland and island populations in the common frog (Rana temporaria)

Understanding and predicting species range expansions is an important challenge in modern ecology because of rapidly changing environments. Recent studies have revealed that consistent within-species variation in behavior (i.e., animal personality) can be imperative for dispersal success, a key process in range expansion. Here we investigate how habitat isolation can mediate differentiation of personality traits between recently founded island populations and the main population. We performed laboratory studies of boldness and exploration across life stages (tadpoles and froglets) using four isolated island populations and four mainland populations of the common frog (Rana temporaria). Both tadpoles and froglets from isolated populations were bolder and more exploratory than conspecifics from the mainland. Although the pattern can be influenced by possible differences in predation pressure, we suggest that this behavioral differentiation might be the result of a disperser-dependent founder effect brought on by an isolation-driven environmental filtering of animal personalities. These findings can have important implications for both species persistence in the face of climate change (i.e., range expansions) and ecological invasions as well as for explaining rapid speciation in isolated patches.     

Limited contributions of released animals from zoos to North American conservation translocations

With the loss of biodiversity accelerating, conservation translocations such as reintroductions are becoming an increasingly common conservation tool. Conservation translocations must source individuals for release from either wild or captive-bred populations. We asked what proportion of North American conservation translocations rely on captive breeding and to what extent zoos and aquaria (hereafter zoos) fulfill captive breeding needs. We searched for mention of captive breeding and zoo involvement in all 1863 articles included in the North American Conservation Translocations database, which comprises journal articles and grey literature published before 2014 on conservation translocations in Canada, the United States, Mexico, the Caribbean, and Central America before 2014 as identified by a comprehensive literature review. Conservation translocations involved captive breeding for 162 (58%) of the 279 animal species translocated. Fifty-four zoos contributed animals for release. The 40 species of animals bred for release by zoos represented only 14% of all animal species for which conservation translocations were published and only 25% of all animal species that were bred for releases occurring in North America. Zoo contributions varied by taxon, ranging from zoo-bred animals released in 42% of amphibian conservation translocations to zero contributions for marine invertebrates. Proportional involvement of zoos in captive-breeding programs for release has increased from 1974 to 2014 (r = 0.325, p = 0.0313) as has the proportion of translocation-focused scientific papers coauthored by zoo professionals (from 0% in 1974 to 42% in 2013). Although zoos also contribute to conservation translocations through education, funding, and professional expertise, increasing the contribution of animals for release in responsible conservation translocation programs presents a future conservation need and opportunity. We especially encourage increased dialogue and planning between the zoo community, academic institutions, and governments to optimize the direct contribution zoos can make to wildlife conservation through conservation translocations.     

Fighting without weaponry: a review of male-male contest competition in butterflies

The males of many butterfly species compete via pair-wise intrasexual contests, yet lack any obvious morphological traits conventionally associated with animal aggression. The evolution of contest behavior in this group is therefore interesting because the means by which individuals are able to inflict costs upon each other during combat are unclear. Here we review the relevant empirical information within a game-theoretical context in order to analyze contest-related costs and mechanisms of dispute settlement. Territorial butterfly contests carry obvious role asymmetries, and individuals in the 'resident' role are more likely to win across all studied species. We use this phenomenon as a framework for gaining further insight into the evolution of contest settlement in this group. Four principal hypotheses are evaluated: (1) that residency is used as an arbitrary convention, (2) that residents stand to gain a greater payoff if successful, (3) that residency bestows higher resource-holding potential (RHP) on males in that role, and (4) that residents are males of intrinsically higher RHP. We found most support for the latter explanation (the 'superior-competitor' hypothesis), partly because the alternatives require a restrictive set of ecological conditions, but also due to positive support for the idea of intrinsic RHP. The strongest evidence is provided by species in which repeated interactions (between the same individuals) always have the same outcome, regardless of changes in residency status. However, we also found a consistent effect on contest duration due to apparent 'confusion over residency,' which suggests a conventional element to contest behavior. Although butterfly contests must be costly in some way, the literature provides limited insight into how costs accrue, and hence the determinants of contest RHP remain obscure. We suggest that careful experimentation, using 'natural' manipulations where possible, should prove most beneficial in identifying costs and exploring schedules of contest settlement in this group. Moreover, future investigations should be sensitive to the fact that butterfly territoriality and associated male-male interactions are seated within a broader ecological and life-historical context.     

Making literature reviews more reliable through application of lessons from systematic reviews

Review articles can provide valuable summaries of the ever‐increasing volume of primary research in conservation biology. Where findings may influence important resource‐allocation decisions in policy or practice, there is a need for a high degree of reliability when reviewing evidence. However, traditional literature reviews are susceptible to a number of biases during the identification, selection, and synthesis of included studies (e.g., publication bias, selection bias, and vote counting). Systematic reviews, pioneered in medicine and translated into conservation in 2006, address these issues through a strict methodology that aims to maximize transparency, objectivity, and repeatability. Systematic reviews will always be the gold standard for reliable synthesis of evidence. However, traditional literature reviews remain popular and will continue to be valuable where systematic reviews are not feasible. Where traditional reviews are used, lessons can be taken from systematic reviews and applied to traditional reviews in order to increase their reliability. Certain key aspects of systematic review methods that can be used in a context‐specific manner in traditional reviews include focusing on mitigating bias; increasing transparency, consistency, and objectivity, and critically appraising the evidence and avoiding vote counting. In situations where conducting a full systematic review is not feasible, the proposed approach to reviewing evidence in a more systematic way can substantially improve the reliability of review findings, providing a time‐ and resource‐efficient means of maximizing the value of traditional reviews. These methods are aimed particularly at those conducting literature reviews where systematic review is not feasible, for example, for graduate students, single reviewers, or small organizations.     

Social personality traits in chimpanzees: temporal stability and structure of behaviourally assessed personality traits in three captive populations

Animals of many species show consistency in behaviour across time and contexts that differs from other individuals' behaviour in the same population. Such ‘personality’ affects fitness and has therefore become an increasingly relevant research topic in biology. However, consistent variation in social behaviour is understudied. In socially living species, behaviour occurs in a social environment and social interactions have a significant influence on individual fitness. This study addressed personality in social behaviour of 75 captive chimpanzees in three zoos by coding observed behaviour. Fifteen behavioural variables were significantly repeatable (range 0.21–0.93) in at least two of the three zoos. The behaviours showed considerable long-term stability across 3 years, which did not differ from the short-term repeatability. The repeatable behaviours were then analysed with factor analyses. They formed five independent factors, three of which consisted of social traits and were labelled ‘sociability’, ‘positive affect’ and ‘equitability’. The two non-social behaviour factors were labelled ‘anxiety’ and ‘activity’. The factor scores were analysed for sex and population differences. Males had higher factor scores in all traits except ‘sociability’. The factor scores differed also between the zoos, implying considerable external effects in trait expression. The results show that chimpanzees show personality in a broad range of social and non-social behaviours. The study highlights the importance of assessing personality in the social behaviour, especially in cohesive social species, as only then can we understand the consequences of personality in socially living species.     

A Framework for Prioritizing Domestic Animal Breeds for Conservation Purposes at the National Level: a Norwegian Case Study

Abstract: Although encompassing only a handful of species, domestic animals have had profound effects on food production for humankind and on human societies. On a global basis, a large proportion of breeds are currently in danger of becoming extinct in the near future. At this critical time, resources are limited and only a selected number of breeds may be supported by concrete conservation programs. I present a framework to facilitate decision making on a national level as to which breeds to support for conservation. These decisions should be made by a national committee of experts experienced with the different breeds and species. First, they must define the species of interest. Second, they should collate as much data as possible and score each breed for key criteria: degree of endangerment, presence of traits of current economic value, presence of traits of current scientific value, agroecological value in a special landscape, cultural-historical value, and genetic uniqueness. The degree of endangerment is the most important criterion because great uncertainty about the future means that all breeds may have traits of future economic or scientific value, so the main aim should therefore be to minimize loss of breeds. To illustrate use of the framework, I considered breed prioritization in Norway. I compared and scored 45 breeds from 17 domestic animal species for these key criteria and thus identified Norwegian breeds of high priority for conservation.     

Behavioral repeatability and personality in pit-building antlion larvae under differing environmental contexts

Over the last decades, there has been growing interest among behavioral ecologists in exploring animal personalities. However, while the foraging behavior of active foragers has been extensively studied, only little is known about that of sit-and-wait predators within the personality framework. We investigated the existence of repeatability and personality in pit-building antlion larvae in the context of foraging (pit construction) and habitat selection (relocation distance and direction) over time and under three environmental contexts: thermal conditions, sand depth, and soil type. Over time, repeatability was much stronger for relocation distance than for movement directionality. Additionally, we observed positive correlations across the two levels of sand depth and soil type but not between thermal conditions. Change in substrate type may induce faster decision-making in these sand-dwelling insects or could be perceived by such insects as a more drastic alteration in their habitat. We suggest that different individuals indeed possess distinct personalities. We also suggest that repeatability should be measured at two levels: the amount of energy expenditure (distances and pit construction) and the pattern of energy expenditure (directionality). Finally, our study illustrates how differing environmental conditions can result in differing levels of plasticity, while largely preserving individual personalities.     

Conservation and social outcomes of private protected areas

Government administered protected areas (PAs) have dominated conservation strategies, discourse, and research, yet private actors are increasingly managing land for conservation. Little is known about the social and environmental outcomes of these privately protected areas (PPAs). We searched the global literature in English on PPAs and their environmental and social outcomes and identified 412 articles suitable for inclusion. Research on PPAs was geographically skewed; more studies occurred in the United States. Environmental outcomes of PPAs were mostly positive (89%), but social outcomes of PPAs were reported less (12% of all studies), and these outcomes were more mixed (65% positive). Private protected areas increased the number or extent of ecosystems, ecoregions, or species covered by PAs (representativeness) and PA network connectivity and effectively reduced deforestation and restored degraded lands. Few PPA owners reported negative social outcomes, experienced improved social capital, increased property value, or a reduction in taxes. Local communities benefited from increased employment, training, and community-wide development (e.g., building of schools), but they reported reduced social capital and no significant difference to household income. The causal mechanisms through which PPAs influence social and environmental outcomes remain unclear, as does how political, economic, and social contexts shape these mechanisms. Future research should widen the geographical scope and diversify the types of PPAs studied and focus on determining the casual mechanisms through which PPA outcomes occur in different contexts. We propose an assessment framework that could be adopted to facilitate this process.     

Network position: a key component in the characterization of social personality types

In recent years, animal social interactions have received much attention in terms of personality research (e.g. aggressive or cooperative interactions). However, other components of social behaviour such as those describing the intensity, frequency, directedness and individual repeatability of interactions in animal groups have largely been neglected. Network analysis offers a valuable opportunity to characterize individual consistency of traits in labile social groups and therein provide novel insights to personality research in ways previously not possible using traditional techniques. Should individual network positions be consistently different between individuals under changing conditions, they might reflect expressions of an individual's personality. Here, we discuss a conceptual framework for using network analyses to infer the presence of individual differences and present a statistical test based on randomization techniques for testing the consistency of network positions in individuals. The statistical tools presented are useful because if particular individuals consistently occupy key positions in social networks, then this is also likely to have consequences for their fitness as well as for that of others in the population. These consequences may be particularly significant since individual network position has been shown to be important for the transmission of diseases, socially learnt information and genetic material between individuals and populations.