Tristan L. Guttridge

Social preferences of juvenile lemon sharks, Negaprion brevirostris

Group living in sharks is a widespread phenomenon but relatively little is known about the composition and organization of these groups. In binary choice field experiments juvenile lemon sharks were attracted to conspecifics presumably to form groups. Experiments investigating size assortment preferences indicated that lemon sharks aged 2–3 years (but not 0–1 years) preferred to spend more time with a group of size-matched individuals than unmatched ones. Furthermore, in species association tests lemon sharks spent significantly more time associating with conspecifics than with a sympatric heterospecific, the nurse shark, Ginglymostoma cirratum. These findings enhance our knowledge of groupjoining decisions in sharks indicating that active mechanisms can play a role in the formation and composition of shark groups. 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Novel Acoustic Technology for Studying Free-Ranging Shark Social Behaviour by Recording Individuals' Interactions

Group behaviours are widespread among fish but comparatively little is known about the interactions between free-ranging individuals and how these might change across different spatio-temporal scales. This is largely due to the difficulty of observing wild fish groups directly underwater over long enough time periods to quantify group structure and individual associations. Here we describe the use of a novel technology, an animal-borne acoustic proximity receiver that records close-spatial associations between free-ranging fish by detection of acoustic signals emitted from transmitters on other individuals. Validation trials, held within enclosures in the natural environment, on juvenile lemon sharks Negaprion brevirostris fitted with external receivers and transmitters, showed receivers logged interactions between individuals regularly when sharks were within 4 m (∼4 body lengths) of each other, but rarely when at 10 m distance. A field trial lasting 17 days with 5 juvenile lemon sharks implanted with proximity receivers showed one receiver successfully recorded association data, demonstrating this shark associated with 9 other juvenile lemon sharks on 128 occasions. This study describes the use of acoustic underwater proximity receivers to quantify interactions among wild sharks, setting the scene for new advances in understanding the social behaviours of marine animals.

Social learning in juvenile lemon sharks, Negaprion brevirostris

Social learning is taxonomically widespread and can provide distinct behavioural advantages, such as in finding food or avoiding predators more efficiently. Although extensively studied in bony fishes, no such empirical evidence exists for cartilaginous fishes. Our aim in this study was to experimentally investigate the social learning capabilities of juvenile lemon sharks, Negaprion brevirostris. We designed a novel food task, where sharks were required to enter a start zone and subsequently make physical contact with a target in order to receive a food reward. Naive sharks were then able to interact with and observe (a) pre-trained sharks, that is, ‘demonstrators’, or (b) sharks with no previous experience, that is, ‘sham demonstrators’. On completion, observer sharks were then isolated and tested individually in a similar task. During the exposure phase observers paired with ‘demonstrator’ sharks performed a greater number of task-related behaviours and made significantly more transitions from the start zone to the target, than observers paired with ‘sham demonstrators’. When tested in isolation, observers previously paired with ‘demonstrator’ sharks completed a greater number of trials and made contact with the target significantly more often than observers previously paired with ‘sham demonstrators’. Such experience also tended to result in faster overall task performance. These results indicate that juvenile lemon sharks, like numerous other animals, are capable of using socially derived information to learn about novel features in their environment. The results likely have important implications for behavioural processes, ecotourism and fisheries.

Social network analysis and valid Markov chain Monte Carlo tests of null models

Analyses of animal social networks derived from group-based associations often rely on randomisation methods developed in ecology (Manly, Ecology 76:1109–1115, 1995) and made available to the animal behaviour community through implementation of a pair-wise swapping algorithm by Bejder et al. (Anim Behav 56:719–725, 1998). We report a correctable flaw in this method and point the reader to a wider literature on the subject of null models in the ecology literature. We illustrate the importance of correcting the method using a toy network and use it to make a preliminary analysis of a network of associations among eagle rays.

Philopatry and Regional Connectivity of the Great Hammerhead Shark, Sphyrna mokarran in the U.S. and Bahamas

A thorough understanding of movement patterns of a species is critical for designing effective conservation and management initiatives. However, generating such information for large marine vertebrates is challenging, as they typically move over long distances, live in concealing environments, are logistically difficult to capture and, as upper-trophic predators, are naturally low in abundance. As a large bodied, broadly distributed tropical shark typically restricted to coastal and shelf habitats, the great hammerhead shark Sphyrna mokarran epitomizes such challenges. Highly valued for its fins, it suffers high bycatch mortality coupled with conservative fecundity, and as a result, is vulnerable to over-exploitation and population depletion. Although there is very little species specific data available, the absence of recent catch records give cause to suspect substantial declines across its range. Here, we used biotelemetry techniques (acoustic and satellite), conventional tagging, laser-photogrammetry, and photo-identification to investigate; the level of site fidelity, and or residency for great hammerheads to coastal areas in the Bahamas and U.S. and the extent of movements and connectivity of great hammerheads between the U.S. and Bahamas. Results revealed large scale return migrations (3030 km), seasonal residency to local areas (some for 5 months), site fidelity (annual return to Bimini and Jupiter for many individuals) and numerous international movements. These findings enhance the understanding of movement ecology of the great hammerhead shark and have the potential to contribute to improved conservation and management.

Rate of movement of juvenile lemon sharks in a novel open field, are we measuring activity or reaction to novelty?

Personality differences are widespread throughout the animal kingdom and can have important ecological and evolutionary consequences. Despite a rapidly increasing body of literature, large (marine) vertebrates remain underrepresented in personality research. Given their unique life history traits (e.g. slow growth rate, slow reproduction rate, long life span) and their pivotal role in ecosystem processes, this is an important gap in our current knowledge. Here we investigated consistency and plasticity in movement behaviour of wild juvenile lemon sharks, Negaprion brevirostris , by repeatedly subjecting sharks to open field tests. First, we investigated the presence of interindividual differences in movement behaviour in a novel open field. Second, we investigated the effect of trial repetition on movement behaviour to understand whether movement in a novel open field reflects a reaction to novelty, or general activity. Third, we estimated individual differences in habituation/sensitization rates over trial repetition and studied how the habituation rate was predicted by the initial movement rate. We found consistent individual differences in movement behaviour during the open field tests. Sharks showed habituation in movement behaviour (i.e. decrease) over repeated trials indicating that the movement behaviour during the first trials is a reaction to novelty, and not general activity. Individuals, however, differed in their rate of habituation (i.e. plasticity) and this rate was negatively related to an individuals movement behaviour in the first open field trial. In addition to showing individual differences in consistency and plasticity in juvenile lemon sharks, our study emphasizes the importance of examining the validity of personality tests when adapting them to new species.

Long-term migration patterns and bisexual philopatry in a benthic shark species

Knowledge of the broad-scale movement patterns of sharks is essential to developing effective management strategies. Currently there is a large bias in studies focusing on species that are either large apex predators or found in tropical to subtropical regions. There is limited knowledge of the movements and migrations of benthic and temperate shark species. The present study used passive acoustic telemetry to investigate the movement patterns of a benthic shark species, the Port Jackson shark (Heterodontus portusjacksoni). Individuals were tagged with acoustic transmitters between 2012 and 2014 and their movements were monitored within Jervis Bay and along the east Australian coastline for up to 4 years. Male and female Port Jackson sharks demonstrated high levels of philopatry to both Jervis Bay and their tagging location across multiple years. Although males and females did not differ in their arrival times, females departed from Jervis Bay later than males. Approximately half the tagged individuals migrated in a southward direction, with individuals being detected at Narooma, Bass Strait and Cape Barron Island. This study provides conclusive evidence of bisexual philopatry in a benthic temperate shark species, confirming previous hypotheses, and presents the most detailed migration route for Port Jackson sharks to date.

Risky business for a juvenile marine predator? Testing the influence of foraging strategies on size and growth rate under natural conditions

Mechanisms driving selection of body size and growth rate in wild marine vertebrates are poorly understood, thus limiting knowledge of their fitness costs at ecological, physiological and genetic scales. Here, we indirectly tested whether selection for size-related traits of juvenile sharks that inhabit a nursery hosting two dichotomous habitats, protected mangroves (low predation risk) and exposed seagrass beds (high predation risk), is influenced by their foraging behaviour. Juvenile sharks displayed a continuum of foraging strategies between mangrove and seagrass areas, with some individuals preferentially feeding in one habitat over another. Foraging habitat was correlated with growth rate, whereby slower growing, smaller individuals fed predominantly in sheltered mangroves, whereas larger, faster growing animals fed over exposed seagrass. Concomitantly, tracked juveniles undertook variable movement behaviours across both the low and high predation risk habitat. These data provide supporting evidence for the hypothesis that directional selection favouring smaller size and slower growth rate, both heritable traits in this shark population, may be driven by variability in foraging behaviour and predation risk. Such evolutionary pathways may be critical to adaptation within predator-driven marine ecosystems.

Does detection range matter for inferring social networks in a benthic shark using acoustic telemetry

Accurately estimating contacts between animals can be critical in ecological studies such as examining social structure, predator–prey interactions or transmission of information and disease. While biotelemetry has been used successfully for such studies in terrestrial systems, it is still under development in the aquatic environment. Acoustic telemetry represents an attractive tool to investigate spatio-temporal behaviour of marine fish and has recently been suggested for monitoring underwater animal interactions. To evaluate the effectiveness of acoustic telemetry in recording interindividual contacts, we compared co-occurrence matrices deduced from three types of acoustic receivers varying in detection range in a benthic shark species. Our results demonstrate that (i) associations produced by acoustic receivers with a large detection range (i.e. Vemco VR2W) were significantly different from those produced by receivers with smaller ranges (i.e. Sonotronics miniSUR receivers and proximity loggers) and (ii) the position of individuals within their network, or centrality, also differed. These findings suggest that acoustic receivers with a large detection range may not be the best option to represent true social networks in the case of a benthic marine animal. While acoustic receivers are increasingly used by marine ecologists, we recommend users first evaluate the influence of detection range to depict accurate individual interactions before using these receivers for social or predator–prey studies. We also advocate for combining multiple receiver types depending on the ecological question being asked and the development of multi-sensor tags or testing of new automated proximity loggers, such as the Encounternet system, to improve the precision and accuracy of social and predator–prey interaction studies.

Three decades of longlining in Bimini, Bahamas, reveals long‐term trends in lemon shark Negaprion brevirostris (Carcharhinidae) catch per unit effort

In Bimini, Bahamas, the consistent employment of longlines, beginning in 1982, provided a rare opportunity to explore population trends for large resident sharks. This study assessed three shallow water longline survey periods at this location; 1982-1989, 1992-2002 and 2003-2014, with the aim of determining trends in annual catch per unit effort (CPUE) for an IUCN listed near-threatened species, the lemon shark Negaprion brevirostris. A general additive model (GAM) was used to analyse the non-linear annual CPUE values over the entire 32-year research period. The GAM displayed high variability of annual CPUE, with a peak value of 0·026 N. brevirostris per hook day (hooks day(-1) ) in 2000. The temporal pattern of CPUE indicated an abundance trend with a complete cycle, from trough to trough, occurring over a period of approximately 18 years. The 1982-1989 survey period saw the highest proportion of mature individuals (19·8%) and the smallest average pre-caudal length (LPC ; 124·8 cm). The 1992-2002 survey period had the highest average annual CPUE (0·018 hooks day(-1) ), while the 2003-2014 research period saw largest average LPC size (134·8 cm) and the lowest average CPUE values (0·009 hooks day(-1) ) of the entire research period. The long-term trend identified in this study provides a baseline for future assessment.