Stable isotopes and foraging behaviors support the role of antipredator benefits in driving the association between two marine fishes.

Research from terrestrial communities shows that diminished predation risk is a principal driver of heterospecific grouping behavior, with foraging ecology predicting the roles that species play in groups, as more vulnerable foragers preferentially join more vigilant ones from whom they can benefit. Meanwhile, field studies examining the adaptive significance of heterospecific shoaling among marine fish have focused disproportionately on feeding advantages such as scrounging or prey-flushing. Juvenile bonefish (Albula vulpes) occur almost exclusively among mojarras (Eucinostomus spp.) and even elect to join them over conspecifics, suggesting they benefit from doing so. We evaluated the roles of risk-related and food-related factors in motivating this pattern of affiliation, estimating: (1) the relative levels of risk associated with each species' search and prey capture activities, via behavioral vulnerability traits discerned from in situ video of heterospecific shoals, and (2) resource use redundancy, using stable isotopes (δ13C, δ15N, and δ34S) to quantify niche overlap. Across four distinct metrics, bonefish behaviors implied a markedly greater level of risk than those of mojarras, typified by higher activity levels and a reduced capacity for overt vigilance; consistent with expectations if their association conformed to patterns of joining observed in terrestrial habitats. Resource use overlap inferred from stable isotopes was low, indicating that the two species partitioned resources and making it unlikely that bonefish derived substantive food-related benefits. Collectively, these findings suggest that the attraction of juvenile bonefish to mojarras is motivated primarily by antipredator advantages, which may include the exploitation of risk-related social cues.

No Evidence for Long-Term Carryover Effects in a Wild Salmonid Fish.

AbstractEarly-life experiences can shape life histories and population dynamics of wild animals. To examine whether stressful stimuli experienced in early life resulted in carryover effects in later life stages, we conducted several experimental manipulations and then monitored wild fish with passive integrated transponder tags during juvenile out-migration and adult return migration. In total, 3,217 juvenile brown trout (Salmo trutta) were subjected to one of six manipulations: chase to exhaustion, thermal challenge, food deprivation, low-concentration cortisol injection, high-concentration cortisol injection, and sham injection, plus a control group. Cortisol and food deprivation treatments were previously shown to have short-term effects on juveniles, such as lower survival to out-migration and changes in migration timing. However, it remained unknown whether any of the six manipulations had effects that carried over into the adult phase. We therefore investigated whether these extrinsic manipulations, as well as intrinsic factors (size and condition), affected probability of return as adults and time spent at sea. Of the 1,273 fish that out-migrated, 146 returned as adults. We failed to detect any effect of treatments on return rates, while high-concentration cortisol weakly affected time spent at sea in one tagging event. We also found that juvenile condition was positively correlated to likelihood of adult return in only one tagging event. Overall, our findings did not identify either intrinsic factors or extrinsic stressful early-life experiences that have strong effects on fish that survive to adulthood. This suggests that some species may be more resilient than others to stressful stimuli encountered early in life.

Effects of predator exposure on baseline and stress-induced glucocorticoid hormone concentrations in pumpkinseed Lepomis gibbosus.

We compared baseline and maximal cortisol concentrations between predator exposure and prey blood samples in pumpkinseed Lepomis gibbosus, captured using a standardised fishing event underneath osprey Pandion haliaetus nests and away from osprey nests. We did not detect differences in cortisol or glucose between sites. These findings suggest that predictable sources of predation risk may not confer stress-related costs in teleosts.

If and when: intrinsic differences and environmental stressors influence migration in brown trout (Salmo trutta).

Partial migration is a common phenomenon, yet the causes of individual differences in migratory propensity are not well understood. We examined factors that potentially influence timing of migration and migratory propensity in a wild population of juvenile brown trout (Salmo trutta) by combining experimental manipulations with passive integrated transponder telemetry. Individuals were subjected to one of six manipulations: three designed to mimic natural stressors (temperature increase, food deprivation, and chase by a simulated predator), an injection of exogenous cortisol designed to mimic an extreme physiological challenge, a sham injection, and a control group. By measuring length and mass of 923 individuals prior to manipulation and by monitoring tagged individuals as they left the stream months later, we assessed whether pre-existing differences influenced migratory tendency and timing of migration, and whether our manipulations affected growth, condition, and timing of migration. We found that pre-existing differences predicted migration, with smaller individuals and individuals in poor condition having a higher propensity to migrate. Exogenous cortisol manipulation had the largest negative effect on growth and condition, and resulted in an earlier migration date. Additionally, low-growth individuals within the temperature and food deprivation treatments migrated earlier. By demonstrating that both pre-existing differences in organism state and additional stressors can affect whether and when individuals migrate, we highlight the importance of understanding individual differences in partial migration. These effects may carry over to influence migration success and affect the evolutionary dynamics of sub-populations experiencing different levels of stress, which is particularly relevant in a changing world.

The Evolution of Lateralization in Group Hunting Sailfish.

Lateralization is widespread throughout the animal kingdom [1-7] and can increase task efficiency via shortening reaction times and saving on neural tissue [8-16]. However, lateralization might be costly because it increases predictability [17-21]. In predator-prey interactions, for example, predators might increase capture success because of specialization in a lateralized attack, but at the cost of increased predictability to their prey, constraining the evolution of lateralization. One unexplored mechanism for evading such costs is group hunting: this would allow individual-level specialization, while still allowing for group-level unpredictability. We investigated this mechanism in group hunting sailfish, Istiophorus platypterus, attacking schooling sardines, Sardinella aurita. During these attacks, sailfish alternate in attacking the prey using their elongated bills to slash or tap the prey [22-24]. This rapid bill movement is either leftward or rightward. Using behavioral observations of identifiable individual sailfish hunting in groups, we provide evidence for individual-level attack lateralization in sailfish. More strongly lateralized individuals had a higher capture success. Further evidence of lateralization comes from morphological analyses of sailfish bills that show strong evidence of one-sided micro-teeth abrasions. Finally, we show that attacks by single sailfish are indeed highly predictable, but predictability rapidly declines with increasing group size because of a lack of population-level lateralization. Our results present a novel benefit of group hunting: by alternating attacks, individual-level attack lateralization can evolve, without the negative consequences of individual-level predictability. More generally, our results suggest that group hunting in predators might provide more suitable conditions for the evolution of strategy diversity compared to solitary life.

Individual-level behavioral responses of immature green turtles to snorkeler disturbance.

Despite many positive benefits of ecotourism, increased human encounters with wildlife may have detrimental effects on wild animals. As charismatic megafauna, nesting and foraging sea turtles are increasingly the focus of ecotourism activities. The purpose of our study was to quantify the behavioral responses of immature green turtles (Chelonia mydas) to disturbance by snorkelers, and to investigate whether turtles have individual-level responses to snorkeler disturbance. Using a standardized disturbance stimulus in the field, we recorded turtle behaviors pre- and post-disturbance by snorkelers. Ninety percent of turtles disturbed by snorkeler (n = 192) initiated their flights at distances of ≤3 m. Using principal component analysis, we identified two distinct turtle personality types, 'bold' and 'timid', based upon 145 encounters of 19 individually identified turtles and five disturbance response variables. There was significant intra-individual repeatability in behavioral responses to disturbance, but bolder turtles had more behavioral plasticity and less consistent responses than more timid individuals. Bolder individuals with reduced evasion responses might be at a higher risk of shark predation, while more timid turtles might have greater energetic consequences due to non-lethal predator effects and repeated snorkeler disturbance. Over the longer term, a turtle population with a mix of bold and timid individuals may promote more resilient populations. We recommend that snorkelers maintain >3 m distance from immature green turtles when snorkeling, and that ecotourism activities be temporally and spatially stratified. Further, turtle watching guidelines need to be communicated to both tour operators and independent snorkelers to reduce the disturbance of turtles.

Do physical habitat complexity and predator cues influence the baseline and stress-induced glucocorticoid levels of a mangrove-associated fish?

As human populations continue to expand, increases in coastal development have led to the alteration of much of the world's mangrove habitat, creating problems for the multitude of species that inhabit these unique ecosystems. Habitat alteration often leads to changes in habitat complexity and predation risk, which may serve as additional stressors for those species that rely on mangroves for protection from predators. However, few studies have been conducted to date to assess the effects of these specific stressors on glucocorticoid (GC) stress hormone levels in wild fish populations. Using the checkered puffer as a model, our study sought to examine the effects of physical habitat complexity and predator environment on baseline and acute stress-induced GC levels. This was accomplished by examining changes in glucose and cortisol concentrations of fish placed in artificial environments for short periods (several hours) where substrate type and the presence of mangrove roots and predator cues were manipulated. Our results suggest that baseline and stress-induced GC levels are not significantly influenced by changes in physical habitat complexity or the predator environment using the experimental protocol that we applied. Although more research is required, the current study suggests that checkered puffers may be capable of withstanding changes in habitat complexity and increases in predation risk without experiencing adverse GC-mediated physiological effects, possibly as a result of the puffers' unique morphological and chemical defenses that help them to avoid predation in the wild.

Proto-cooperation: group hunting sailfish improve hunting success by alternating attacks on grouping prey.

We present evidence of a novel form of group hunting. Individual sailfish (Istiophorus platypterus) alternate attacks with other group members on their schooling prey (Sardinella aurita). While only 24% of attacks result in prey capture, multiple prey are injured in 95% of attacks, resulting in an increase of injured fish in the school with the number of attacks. How quickly prey are captured is positively correlated with the level of injury of the school, suggesting that hunters can benefit from other conspecifics' attacks on the prey. To explore this, we built a mathematical model capturing the dynamics of the hunt. We show that group hunting provides major efficiency gains (prey caught per unit time) for individuals in groups of up to 70 members. We also demonstrate that a free riding strategy, where some individuals wait until the prey are sufficiently injured before attacking, is only beneficial if the cost of attacking is high, and only then when waiting times are short. Our findings provide evidence that cooperative benefits can be realized through the facilitative effects of individuals' hunting actions without spatial coordination of attacks. Such 'proto-cooperation' may be the pre-cursor to more complex group-hunting strategies.

Maximum swimming speeds of sailfish and three other large marine predatory fish species based on muscle contraction time and stride length: a myth revisited.

Billfishes are considered to be among the fastest swimmers in the oceans. Previous studies have estimated maximum speed of sailfish and black marlin at around 35 m s-1 but theoretical work on cavitation predicts that such extreme speed is unlikely. Here we investigated maximum speed of sailfish, and three other large marine pelagic predatory fish species, by measuring the twitch contraction time of anaerobic swimming muscle. The highest estimated maximum swimming speeds were found in sailfish (8.3±1.4 m s-1), followed by barracuda (6.2±1.0 m s-1), little tunny (5.6±0.2 m s-1) and dorado (4.0±0.9 m s-1); although size-corrected performance was highest in little tunny and lowest in sailfish. Contrary to previously reported estimates, our results suggest that sailfish are incapable of exceeding swimming speeds of 10-15 m s-1, which corresponds to the speed at which cavitation is predicted to occur, with destructive consequences for fin tissues.

Puffed and bothered: Personality, performance, and the effects of stress on checkered pufferfish.

Although consistent individual-level differences in behaviour are widespread and potentially important in evolutionary and ecological processes, relatively few studies focus on the physiological mechanisms that might underlie and regulate these individual-level differences in wild populations. We conducted experiments to determine whether checkered pufferfish (Sphoeroides testudineus), which were collected from a dynamic (in terms of depth and water temperature) tidal mangrove creek environment in The Bahamas, have consistent individual-level differences in locomotor activity and the response to a simulated predator threat, as well as swimming performance and puffing in response to stressors. The relationships between personality and performance traits were evaluated to determine whether they represented stress-coping styles or syndromes. Subsequently, a displacement study was conducted to determine how personality and performance in the laboratory compared to movements in the field. In addition, we tested whether a physiological dose of the stress hormone cortisol would alter individual consistency in behavioural and performance traits. We found that pufferfish exhibited consistent individual differences in personality traits over time (e.g., activity and the duration of a response to a threat) and that performance was consistent between the lab and the natural enclosure. Locomotor activity and the duration of startled behaviour were not associated with swimming and puffing performance. Locomotor activity, puffing performance, and swimming performance were not related to whether fish returned to the tidal creek of capture after displacement. Similarly, a cortisol treatment did not modify behaviour or performance in the laboratory. The results reveal that consistent individual-level differences in behaviour and performance were present in a population from a fluctuating and physiologically challenging environment but that such traits are not necessarily correlated. We also determined that certain individual performance traits were repeatable between the lab and a natural enclosure. However, we found no evidence of a relationship between exogenous cortisol levels and behavioural traits or performance in these fish, which suggests that other internal and external mechanisms may underlie the behaviours and performance tested.

How long is too long in contemporary peer review? Perspectives from authors publishing in conservation biology journals.

Delays in peer reviewed publication may have consequences for both assessment of scientific prowess in academics as well as communication of important information to the knowledge receptor community. We present an analysis on the perspectives of authors publishing in conservation biology journals regarding their opinions on the importance of speed in peer-review as well as how to improve review times. Authors were invited to take part in an online questionnaire, of which the data was subjected to both qualitative (open coding, categorizing) and quantitative analyses (generalized linear models). We received 637 responses to a total of 6,547 e-mail invitations sent. Peer-review speed was generally perceived as slow, with authors experiencing a typical turnaround time of 14 weeks while their perceived optimal review time is six weeks. Male and younger respondents seem to have higher expectations of review speed than females and older respondents. Majority of participants attributed lengthy review times to the 'stress' on the peer-review system (i.e., reviewer and editor fatigue), while editor persistence and journal prestige were believed to speed up the review process. Negative consequences of lengthy review times appear to be greater for early career researchers and can also have impact on author morale (e.g. motivation or frustration). Competition among colleagues were also of concern to respondents. Incentivizing peer review was among the top suggested alterations to the system along with training graduate students in peer review, increased editorial persistence, and changes to the norms of peer-review such as opening the peer-review process to the public. It is clear that authors surveyed in this study view the peer-review system as under stress and we encourage scientists and publishers to push the envelope for new peer review models.

Does Angling Technique Selectively Target Fishes Based on Their Behavioural Type?

Recently, there has been growing recognition that fish harvesting practices can have important impacts on the phenotypic distributions and diversity of natural populations through a phenomenon known as fisheries-induced evolution. Here we experimentally show that two common recreational angling techniques (active crank baits versus passive soft plastics) differentially target wild largemouth bass (Micropterus salmoides) and rock bass (Ambloplites rupestris) based on variation in their behavioural tendencies. Fish were first angled in the wild using both techniques and then brought back to the laboratory and tested for individual-level differences in common estimates of personality (refuge emergence, flight-initiation-distance, latency-to-recapture and with a net, and general activity) in an in-lake experimental arena. We found that different angling techniques appear to selectively target these species based on their boldness (as characterized by refuge emergence, a standard measure of boldness in fishes) but not other assays of personality. We also observed that body size was independently a significant predictor of personality in both species, though this varied between traits and species. Our results suggest a context-dependency for vulnerability to capture relative to behaviour in these fish species. Ascertaining the selective pressures angling practices exert on natural populations is an important area of fisheries research with significant implications for ecology, evolution, and resource management.

Not So Fast: Swimming Behavior of Sailfish during Predator-Prey Interactions using High-Speed Video and Accelerometry.

Billfishes are considered among the fastest swimmers in the oceans. Despite early estimates of extremely high speeds, more recent work showed that these predators (e.g., blue marlin) spend most of their time swimming slowly, rarely exceeding 2 m s(-1). Predator-prey interactions provide a context within which one may expect maximal speeds both by predators and prey. Beyond speed, however, an important component determining the outcome of predator-prey encounters is unsteady swimming (i.e., turning and accelerating). Although large predators are faster than their small prey, the latter show higher performance in unsteady swimming. To contrast the evading behaviors of their highly maneuverable prey, sailfish and other large aquatic predators possess morphological adaptations, such as elongated bills, which can be moved more rapidly than the whole body itself, facilitating capture of the prey. Therefore, it is an open question whether such supposedly very fast swimmers do use high-speed bursts when feeding on evasive prey, in addition to using their bill for slashing prey. Here, we measured the swimming behavior of sailfish by using high-frequency accelerometry and high-speed video observations during predator-prey interactions. These measurements allowed analyses of tail beat frequencies to estimate swimming speeds. Our results suggest that sailfish burst at speeds of about 7 m s(-1) and do not exceed swimming speeds of 10 m s(-1) during predator-prey interactions. These speeds are much lower than previous estimates. In addition, the oscillations of the bill during swimming with, and without, extension of the dorsal fin (i.e., the sail) were measured. We suggest that extension of the dorsal fin may allow sailfish to improve the control of the bill and minimize its yaw, hence preventing disturbance of the prey. Therefore, sailfish, like other large predators, may rely mainly on accuracy of movement and the use of the extensions of their bodies, rather than resorting to top speeds when hunting evasive prey.

Effects of emergence time and early social rearing environment on behaviour of Atlantic salmon: consequences for juvenile fitness and smolt migration.

Consistent individual differences in behaviour have been well documented in a variety of animal taxa, but surprisingly little is known about the fitness and life-history consequences of such individual variation. In wild salmonids, the timing of fry emergence from gravel spawning nests has been suggested to be coupled with individual behavioural traits. Here, we further investigate the link between timing of spawning nest emergence and behaviour of Atlantic salmon (Salmo salar), test effects of social rearing environment on behavioural traits in fish with different emergence times, and assess whether behavioural traits measured in the laboratory predict growth, survival, and migration status in the wild. Atlantic salmon fry were sorted with respect to emergence time from artificial spawning nest into three groups: early, intermediate, and late. These emergence groups were hatchery-reared separately or in co-culture for four months to test effects of social rearing environment on behavioural traits. Twenty fish from each of the six treatment groups were then subjected to three individual-based behavioural tests: basal locomotor activity, boldness, and escape response. Following behavioural characterization, the fish were released into a near-natural experimental stream. Results showed differences in escape behaviour between emergence groups in a net restraining test, but the social rearing environment did not affect individual behavioural expression. Emergence time and social environment had no significant effects on survival, growth, and migration status in the stream, although migration propensity was 1.4 to 1.9 times higher for early emerging individuals that were reared separately. In addition, despite individuals showing considerable variation in behaviour across treatment groups, this was not translated into differences in growth, survival, and migration status. Hence, our study adds to the view that fitness (i.e., growth and survival) and life-history predictions from laboratory measures of behaviour should be made with caution and ideally tested in nature.

The evolutionary and ecological consequences of animal social networks: emerging issues.

The first generation of research on animal social networks was primarily aimed at introducing the concept of social networks to the fields of animal behaviour and behavioural ecology. More recently, a diverse body of evidence has shown that social fine structure matters on a broader scale than initially expected, affecting many key ecological and evolutionary processes. Here, we review this development. We discuss the effects of social network structure on evolutionary dynamics (genetic drift, fixation probabilities, and frequency-dependent selection) and social evolution (cooperation and between-individual behavioural differences). We discuss how social network structure can affect important coevolutionary processes (host-pathogen interactions and mutualisms) and population stability. We also discuss the potentially important, but poorly studied, role of social network structure on dispersal and invasion. Throughout, we highlight important areas for future research.

The influence of water temperature and accelerometer-determined fight intensity on physiological stress and reflex impairment of angled largemouth bass.

Release of fish captured by recreational anglers is a common practice due to angler conservation ethics or compliance with fisheries regulations. As such, there is a need to understand the factors that influence mortality and sub-lethal impairments to ensure that catch-and-release angling is a sustainable practice. Longer angling times generally contribute to increased stress and mortality in fish such that reducing these times putatively reduces stress and improves survival. However, the relative importance of fight intensity (rather than simply duration) on fish condition is poorly understood. The objective of this research was to examine the effects of fight intensity on physiological stress and reflex impairment of largemouth bass (Micropterus salmoides). The largemouth bass were angled using conventional recreational fishing gear in May (water temperature ∼12°C) and June (∼22°C) of 2014 in Lake Opinicon, Ontario, Canada. Fight intensity was quantified using tri-axial accelerometer loggers mounted on the tips of fishing rods. Upon capture, reflex impairment measures were assessed, and fish were held for 1 h prior to blood sampling for measurement of physiological stress (blood glucose and lactate concentrations and pH). Physiological stress values showed a negative trend with fight duration and total fight intensity, but a positive trend with average fight intensity. Water temperature emerged as the most important predictor of the stress response in largemouth bass, while fight duration and intensity were not strong predictors. Reflex impairment was minimal, but higher reflex impairment scores were associated with elevated blood glucose. Overall, the findings of this study suggest that angling for largemouth bass at colder temperatures (<15°C) causes greater physiological stress than at warmer temperatures (>20°C). Based on our findings, we conclude that fight intensity is likely not to be a major driver of physiological stress in this species using typical largemouth bass angling gear, owing to the relatively short fight times (i.e. <2 min).

Metamorphosis and animal personality: a neglected opportunity.

Developmental perspectives represent an underutilized area of animal personality research, in spite of their obvious importance in biology. Animals that undergo metamorphosis are particularly neglected and represent a unique in situ experimental opportunity to study how personality differences are associated with physiological, morphological, or ecological traits over development.