Understanding how individual behavior shapes the structure and ecology of populations is key to species conservation and management. Like many elasmobranchs, manta rays are highly mobile and wide-ranging species threatened by anthropogenic impacts. In shallow water environments, these pelagic rays often form groups and perform several apparently socially mediated behaviors. Group structures may result from active choices of individual rays to interact or passive processes. Social behavior is known to affect spatial ecology in other elasmobranchs, but this is the first study providing quantitative evidence for structured social relationships in manta rays. To construct social networks, we collected data from more than 500 groups of reef manta rays (Mobula alfredi) over 5 years in the Raja Ampat Regency of West Papua. We used generalized affiliation indices to isolate social preferences from non-social associations, the first study on elasmobranchs to use this method. Longer lasting social preferences were detected mostly between female rays. We detected assortment of social relations by phenotype and variation in social strategies, with the overall social network divided into two main communities. Overall network structure was characteristic of a dynamic fission-fusion society, with differentiated relationships linked to strong fidelity to cleaning station sites. Our results suggest that fine-scale conservation measures will be useful in protecting social groups of M. alfredi in their natural habitats and that a more complete understanding of the social nature of manta rays will help predict population responses to anthropogenic pressures, such as increasing disturbance from dive tourism. In social animals, relationships between individuals have important implications for species conservation. Like many other sharks and rays, manta rays are threatened species, and little is known about their natural behavior or how their populations are structured. This study provides evidence of social structure in a wild, free-ranging population of reef manta rays. We show for the first time that individual manta rays have preferred relationships with others that are maintained over time, and structured societies. This study extends our knowledge of elasmobranch ecology and population structuring. Results suggest that understanding social relationships in manta rays will be important in protecting populations from human impacts and developing sustainable, localized conservation and management initiatives.

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Social preferences and network structure in a population of reef manta rays

In many animal populations, individuals exhibit repeatable behavioral traits across a range of contexts, and similarly, individuals differ in ecological traits such as habitat use, home range sizes, growth rates, and mating success. However, links between an individual's positions on behavioral vs. ecological axes of variation remain relatively unstudied in the wild. In the course of fieldwork on a remote floodplain in tropical Australia, we quantified boldness and ecological traits in 86 free-ranging (radio-tracked) monitor lizards (Varanus panoptes). These large (up to 7 kg) lizards exhibited a spectrum of boldness, as reflected in correlated scores of responses to approach, handling, and novel prey. Bolder lizards had larger core home ranges and higher mating success and spent more time in areas of high predator abundance, and their seasonal regimes of predation-induced mortality differed from those of shyer lizards. Thus, behavioral differences among lizards underpin much of the variation in ecological traits and individual fitness within this population. Analyses of ecology and microevolution in natural populations cannot afford to ignore the complex covariation between behavior, ecology, and evolution in the wild.

https://esajournals.onlinelibrary.wiley.com/doi/pdf/10.1002/ecs2.2125

The ecological and life history correlates of boldness in free‐ranging lizards

Despite substantial research interest in understanding individual-level consistency in behavioral attributes, significant knowledge gaps remain across traits and taxa. For example, relatively few studies have looked at social personality in large marine species such as elasmobranchs and whether or not individual differences in behavior are maintained in unstable social groups (i.e., fission-fusion dynamics). However, it is important to investigate this topic in other model species than the usually small species with short generation times typically investigated in these areas of behavioral ecology. Indeed, studies on ecologically diverse taxa could provide mechanistic insights into the emergence and maintenance of animal personality and dynamics of social groups in animals. In addition, understanding social behavior at the group- and individual-level could improve conservation management of these large animals with long generation times (e.g., removal of particular behavioral types by fisheries practices). Here, we investigated consistent individual differences in sociability in wild juvenile lemon sharks (Negaprion brevirostris) over both short- (4 to 18 days) and long-term (4 months) sampling periods. Individual sharks were observed in social groups and scored according to the number of social interactions performed during observations. Despite variable individual group compositions between repeated trials, sharks showed consistent individual differences in their social behavior over both time scales. These results suggest reduced plasticity and highlight individuality as an important explanatory variable for the social dynamics of juvenile lemon sharks. In addition, long-term stability observed in this wild population demonstrates the importance of personality in the daily behavioral repertoire of juvenile lemon sharks. Our results are discussed in the context of other shark studies and taxonomic groups and potential avenues for future research are proposed. This study investigated the social personality axis in a wild population of juvenile lemon sharks. First, we demonstrated consistent individual differences in their tendency to socialize. Second, we showed that individuals maintained their differences over a four-month period in the wild. Finally, we found that individual social behaviors were maintained despite being tested in variable group compositions. These results highlight the importance of individuality in the social dynamic of a poorly investigated animal and suggest personality as an important aspect of juvenile lemon sharks’ everyday life over a relatively long-term period.

/pdf/are-some-sharks-more-social-than-others-short-and-long-term-549uxhn09h.pdf

Are some sharks more social than others? Short- and long-term consistencies in the social behavior of juvenile lemon sharks

The notion that men are more variable than women has become embedded into scientific thinking. For mental traits like personality, greater male variability has been partly attributed to biology, underpinned by claims that there is generally greater variation among males than females in non-human animals due to stronger sexual selection on males. However, evidence for greater male variability is limited to morphological traits, and there is little information regarding sex differences in personality-like behaviours for non-human animals. Here, we meta-analysed sex differences in means and variances for over 2100 effects (204 studies) from 220 species (covering five broad taxonomic groups) across five personality traits: boldness, aggression, activity, sociality and exploration. We also tested if sexual size dimorphism, a proxy for sex-specific sexual selection, explains variation in the magnitude of sex differences in personality. We found no significant differences in personality between the sexes. In addition, sexual size dimorphism did not explain variation in the magnitude of the observed sex differences in the mean or variance in personality for any taxonomic group. In sum, we find no evidence for widespread sex differences in variability in non-human animal personality. 

A meta-analysis of sex differences in animal personality: no evidence for the greater male variability hypothesis.

Individual differences in numerical skills are influenced by brain lateralization in guppies (Poecilia reticulata)

The field of animal personality has received considerable attention in past decades, yet few studies have examined personality in the wild. This study investigated docility, a measure of boldness, in two Port Jackson shark ( Heterodontus portusjacksoni ) populations using field tests, and if laterality differences explained docility levels. We developed a struggle test as an assay for docility, which is particularly amenable to field studies. The struggle test was effective, and repeatable inter-individual docility differences were observed. Sex, but not population, influenced docility scores, with male sharks being less docile than females. This difference is likely due to the contrasting role each sex plays during mating. We also found individualized lateralization. However, no individual-level relationship between lateralization and docility was detected. Despite reported links between laterality and some personality traits, the relationship between laterality and boldness remains inconclusive in sharks. Further studies will prove essential to clarify the mechanisms behind personality traits in vertebrates.

Into the wild: developing field tests to examine the link between elasmobranch personality and laterality

Following a lack of detected change in white shark Carcharodon carcharias L. 1758 diet and nutritional condition attributed to the interaction with the cage-diving industry, Lusseau and Derous (Tourism Management, 2019, 75, 547-549) cautioned the use of muscle lipids and fatty acids in this context, advocating for other biomarkers. This study provides additional evidence from peer-reviewed literature to contend the usefulness of elasmobranch muscle fatty acid profiles to detail diet and habitat use. It also presents findings from a controlled experiment on captive Port Jackson sharks Heterodontus portusjacksoni (Meyer 1793) whereby long-term (daily for 33 days) 3 min exhaustive chase exercise changed muscle lipid class profiles, supporting its use to infer nutritional condition after activities such as interactions with wildlife tourism operators. Conversely, the unaltered muscle fatty acid and lipid content suggests their use in trophic ecology is not confounded by activities such as interacting with tourism operators, remaining useful biomarkers to investigate diet and habitat use.

Sherrie L. Chambers

Papers

Into the wild: developing field tests to examine the link between elasmobranch personality and laterality

The use of muscle lipids and fatty acids to assess shark diet and condition.