Shoal composition determines foraging success in the guppy

The composition of an animal group can impact greatly on thesurvival and success of its individual members. Much recentwork has concentrated on behavioral variation within animalpopulations along the bold/shy continuum. Here, we screenedindividual guppies, Poecilia reticulata, for boldness usingan overhead fright stimulus. We created groups consisting of4 bold individuals (bold shoals), 4 shy individuals (shy shoals),or 2 bold and 2 shy individuals (mixed shoals). The performanceof these different shoal types was then tested in a novel foragingscenario. We found that both bold and mixed shoals approacheda novel feeder in less time than shy shoals. Interestingly,we found that more fish from mixed shoals fed than in eitherbold or shy shoals. We suggest that this can be explained bythe fact that nearly all the cases where one fish was followedinto the feeder by another occurred within mixed shoals andthat it was almost always a shy fish following a bold one. Theseresults suggest clear foraging benefits to shy individuals throughassociating with bold ones. Surprisingly, our results also suggestpotential foraging benefits to bold individuals through associatingwith shy individuals. This study highlights a possible mechanismby which interindividual variation in behavioral types is maintainedin a population.     

Boldness,Dominance, and Territoriality in the Color Polymorphic Tree Lizard,Urosaurus ornatus

Social dynamics in territorial species often reflect underlying variation in aggression and other aspects of social dominance among individuals. In ornate tree lizards (Urosaurus ornatus), males differing in dewlap color differ in social dominance: while blue males are the dominant, aggressive morph and always territorial, yellow males tend to exhibit a less‐aggressive satellite behavioral tactic. However, in habitats with fewer available territorial resources, yellow males defend territories and increase in relative abundance. These observations suggest that consideration of social dominance alone may be insufficient to explain U. ornatus' territorial dynamics in the wild. Here, we tested how both dominance and another important behavioral trait, boldness, contribute to the outcome of territorial disputes in tree lizards. We recorded the territorial behavior of blue and yellow male tree lizards (entered in pairs) in an experimental arena. At the end of each trial, we then approached each male and recorded whether it fled (shy) or not (bold) in response to our approach. As expected, dominant blue males exploited the higher quality perch more often than yellow males. However, when approached by a simulated predator, blue males were more likely to flee than yellow males. Thus, while blue males are more dominant, yellow is likely the bolder morph. As a result, this morph may be better equipped to defend territories in riskier environments. We conclude that although dominance asymmetries may predictably drive initial territorial interactions among competing males, variation in other behaviors (like boldness) may perturb the long‐term outcome of these interactions across variable environments.     

Boldness predicts social status in zebrafish (Danio rerio)

This study explored if boldness could be used to predict social status. First, boldness was assessed by monitoring individual zebrafish behaviour in (1) an unfamiliar barren environment with no shelter (open field), (2) the same environment when a roof was introduced as a shelter, and (3) when the roof was removed and an unfamiliar object (Lego® brick) was introduced. Next, after a resting period of minimum one week, social status of the fish was determined in a dyadic contest and dominant/subordinate individuals were determined as the winner/loser of two consecutive contests. Multivariate data analyses showed that males were bolder than females and that the behaviours expressed by the fish during the boldness tests could be used to predict which fish would later become dominant and subordinate in the ensuing dyadic contest. We conclude that bold behaviour is positively correlated to dominance in zebrafish and that boldness is not solely a consequence of social dominance.     

Behavioral Syndromes Break Down in Urban Song Sparrow Populations

Animals in urban habitats face a number of unique stresses, including the necessity of dealing with high levels of human activity. Growing research suggests that: (1) inherent traits, as opposed to learned behavior, influence which species invade urban habitats, and (2) individuals exhibit behavioral syndromes that limit behavioral flexibility. As a result, perhaps only animals with inherently bold personalities successfully settle in areas of high human activity, and such animals may also exhibit correlated variation in other behavioral traits, such as territorial aggression. In this study, we examine boldness and aggression in several urban and rural populations of song sparrows (Melospiza melodia). We found that urban birds were both bolder toward humans and also showed higher levels of aggression. We found a correlation between boldness and aggression in all populations combined, but no correlation within urban populations. Our results agree with other recent studies of song sparrow behavior, suggesting that greater boldness and aggression are general features of urban song sparrow populations, and a lack of a correlation between boldness and aggression in urban habitats is a general phenomenon as well. Urban habitats may select for bold and aggressive birds, and yet the traits can vary independently. These results add to a small number of studies which find that behavioral syndromes break down in potentially high quality habitats.     

Size doesn't matter,sex does: a test for boldness in sister species of Brachyrhaphis fishes

The effect of divergent natural selection on the evolution of behavioral traits has long been a focus of behavioral ecologists. Predation, due to its ubiquity in nature and strength as a selective agent, has been considered an important environmental driver of behavior. Predation is often confounded with other environmental factors that could also play a role in behavioral evolution. For example, environments that contain predators are often more ecologically complex and “risky” (i.e., exposed and dangerous). Previous work shows that individuals from risky environments are often more bold, active, and explorative than those from low‐risk environments. To date, most comparative studies of environmentally driven behavioral divergence are limited to comparisons among populations within species that occur in divergent selective environments but neglect comparisons between species following speciation. This limits our understanding of how behavior evolves post‐speciation. The Central American live‐bearing fish genus Brachyrhaphis provides an ideal system for examining the relationship between selective environments and behavior, within and between species. Here, we test for differences in boldness between sister species B. roseni and B. terrabensis that occur in streams with and without piscivorous predators, respectively. We found that species do differ in boldness, with species that occur with predators being bolder than those that do not. Within each species, we found that sexes differed in boldness, with males being bolder than females. We also tested for a relationship between size (a surrogate for metabolic rate) and boldness, but found no size effects. Therefore, sex, not size, affects boldness. These results are consistent with the hypothesis that complex and risky environments favor individuals with more bold behavioral traits, but they are not consistent with the hypothesis that size (and therefore metabolic rate) drives divergence in boldness. Finally, our results provide evidence that behavioral trait divergence continues even after speciation is complete.     

Differences in aggression,activity and boldness between native and introduced populations of an invasive crayfish

Aggressiveness, along with foraging voracity and boldness, are key behavioral mechanisms underlying the competitive displacement and invasion success of exotic species. However, do aggressiveness, voracity and boldness of the invader depend on the presence of an ecologically similar native competitor in the invaded community? We conducted four behavioral assays to compare aggression, foraging voracity, threat response and boldness to forage under predation risk of multiple populations of exotic signal crayfish Pacifastacus leniusculus across its native and invaded range with and without a native congener, the Shasta crayfish P. fortis. We predicted that signal crayfish from the invaded range and sympatric with a native congener (IRS) should be more aggressive to outcompete a close competitor than populations from the native range (NR) or invaded range and allopatric to a native congener (IRA). Furthermore, we predicted that IRS populations of signal crayfish should be more voracious, but less bold to forage under predation risk since native predators and prey likely possess appropriate behavioral responses to the invader. Contrary to our predictions, results indicated that IRA signal crayfish were more aggressive towards conspecifics and more voracious and active foragers, yet also bolder to forage under predation risk in comparison to NR and IRS populations, which did not differ in behavior. Higher aggression/voracity/boldness was positively correlated with prey consumption rates, and hence potential impacts on prey. We suggest that the positive correlations between aggression/voracity/boldness are the result of an overall aggression syndrome. Results of stream surveys indicated that IRA streams have significantly lower prey biomass than in IRS streams, which may drive invading signal crayfish to be more aggressive/voracious/bold to acquire resources to establish a population.     

Assessing the Effects of Rearing Environment,Natural Selection,and Developmental Stage on the Emergence of a Behavioral Syndrome

Although there is much interest in behavioral syndromes, very little is known about how syndromes are generated in wild populations. Here, we assess the roles of correlated selection and divergent rearing environments in generating a syndrome between foraging aggressiveness and boldness in the spider Agelenopsis pennsylvanica. We first tested for and confirmed the presence of a behavioral syndrome between boldness and foraging aggressiveness in wild penultimate A. pennsylvanica (r = 0.24). Then, to assess the effects of rearing environment on the boldness–aggressiveness syndrome, we compared the behavioral tendencies of field‐ vs. laboratory‐reared spiders over the course of their development. The presence of the boldness–aggressiveness syndrome differed based on spiders' developmental stage and rearing environment: field‐reared juveniles did not exhibit a syndrome between boldness and foraging aggressiveness, but field‐reared penultimates did. In contrast, laboratory‐reared spiders never exhibited a behavioral syndrome, regardless of their developmental stage. Thus, the boldness–aggressiveness syndrome in A. pennsylvanica manifests only when individuals are reared in the field. Selection data from a mark–recapture study failed to indicate any signature of correlated selection, despite our finding that at least one element of the syndrome (foraging aggressiveness) can respond to selection (Heritability h² = 0.27, from mid‐parent breeding study). Thus, contemporary correlated selection does not appear to be a major driver of the boldness–aggressiveness syndrome of A. pennsylvanica. Taken together, our data are consistent with the hypothesis that the boldness–aggressiveness syndrome exhibited by wild A. pennsylvanica develops as a result of environmentally induced phenotypic plasticity, and not correlated selection.     

Personality-dependent inter- and intraspecific foraging competition in the invasive round goby,Neogobius melanostomus

This study examines the impact of boldness on foraging competition of the highly invasive round goby Neogobius melanostomus Pallas 1815. Individual risk tolerance, or boldness, was measured as the time to resume movement after a simulated predation strike. Fish that resumed movement faster were categorized as “bold,” fish that took more time to resume movement were categorized as “shy” and those that fell in between these two categories were determined to have “intermediate” boldness. Competitive impacts of boldness in N. melanostomus were determined in a laboratory foraging experiment in which interspecific (juvenile Atlantic cod Gadus morhua Linnaeus 1758) and intraspecific (intermediate N. melanostomus) individuals were exposed to either bold or shy N. melanostomus competitors. G. morhua consumed fewer prey when competing with bold N. melanostomus than when competing with shy N. melanostomus, whereas intermediately bold N. melanostomus foraging was not affected by competitor boldness. Bold and shy N. melanostomus consumed similar amounts of prey, and the number of interactions between paired fish did not vary depending on the personality of N. melanostomus individuals. Therefore, intraspecific foraging competition was not found to be personality dependent. This study provides evidence that individual differences in boldness can mediate competitive interactions in N. melanostomus; nonetheless, results also show that competition is also governed by other mechanisms that require further study.     

Understanding the adaptive consequences of hatchery-wild interactions in Alaska salmon

About 31% of salmon harvested in Alaska comes from the hatchery production of hundreds of millions of pink and chum salmon and smaller numbers of sockeye, Chinook, and coho salmon. The numbers of hatchery-reared juveniles released in some areas are greater than the numbers of juveniles from wild populations. However, virtually nothing is known about the effects of hatchery fish on wild populations in Alaska. Possible effects of these interactions can be inferred from studies of salmonids in other areas, from studies of other animals, and from theory. Numerous studies show a complex relationship between the genetic architecture of a population and its environment. Adaptive responses to nature and anthropogenic selection can be influenced by variation at a single gene, or more often, by the additive effects of several genes. Studies of salmonids in other areas show that hatchery practices can lead to the loss of genetic diversity, to shifts in adult run timing and earlier maturity, to increases in parasite load, to increases in straying, to altered levels of boldness and dominance, to shifts in juvenile out-migration timing, and to changes in growth. Controlled experiments across generations show, and theory predicts, that the loss of adaptive fitness in hatchery salmon, relative to fitness in wild salmon, can occur on a remarkably short time scale. All of these changes can influence survival and impose selective regimes that influence genetically based adaptive traits. The preservation of adaptive potential in wild populations is an important buffer against diseases and climate variability and, hence, should be considered in planning hatchery production levels and release locations. The protection of wild populations is the foundation for achieving sustained harvests of salmon in Alaska.     

Long-term behavioral repeatability in wild adult and captive juvenile turtles (Terrapene carolina): Implications for personality development

Animal personality can be defined as conspecific individuals consistently differing in behavioral tendencies. Personality is typically identified by behavioral repeatability, which occurs when within-individual variance is low relative to among-individual variance in the population. Intraspecific comparisons of behavioral repeatability in juveniles and adults within and across years are rare, but would be useful for testing hypotheses related to origins of animal personality and whether individuals exhibit stable or diverging behavior with ontogeny. To examine within- and across-year behavioral repeatability for eastern box turtles (Terrapene carolina), we assessed boldness (movement latency after brief confinement) of captive-born juveniles twice within three days when eight months old. We then repeated these tests for the same individuals one year later. Juveniles exhibited repeatable boldness within and across years. Although increasing body temperature was slightly associated with decreased movement latency, test year (1 or 2), or housing experience (being raised in an enriched or unenriched condition) had no effects on boldness. We also assessed across-year repeatability of boldness (head emergence from the shell after brief confinement) for wild adults at 1–3 year intervals. Adults also exhibited repeatable across-year boldness that was of similar magnitude to juveniles. We found no indication that sex class or whether adults had been radio-tracked influenced boldness. Our results suggest eastern box turtles demonstrate consistent individuality in boldness from an early age that is largely unaffected by temporal or environmental variation, and these behavioral differences can be maintained for multiple years in captivity and the wild, contrasting with theoretical expectations for personality development. These findings add to recent accumulating evidence demonstrating juvenile and adult box turtles exhibit multiple repeatable behaviors over the short- and long-term. We suggest this species is quickly gaining traction as a model organism for studying the proximate and ultimate causes of personality development within long-lived animals.     

Interpopulation Variations in Behavioral Syndromes of a Jumping Spider from Insecticide‐Treated and Insecticide‐Free Orchards

Variations in environmental conditions can influence behavioral syndromes (correlated tendencies in behaviors), and understanding the factors that shape trait covariation is particularly relevant when species are challenged by environmental changes. We investigated how behavioral syndromes varied at extremes of a gradient of anthropogenic disturbance, using apple orchards with different histories of insecticidal applications as a model system. Eris militaris (Araneae: Salticidae) jumping spiders were sampled from an insecticide‐free orchard and an insecticide‐treated orchard from Southern Québec. Spiders were tested for activity, aggression, boldness, and voracity under standardized conditions. Behavioral syndrome structure was compared between the two populations using Bayesian multiresponse models and structural equation modeling. Syndrome structure differed significantly between the two populations. The insecticide‐free population showed evidence of a syndrome involving all measured traits, while only aggression, boldness and voracity were correlated in the insecticide‐treated population. The insecticide‐free population showed negative correlations between active and voracious behavioral types vs. aggressive and bold types while the insecticide‐treated population showed a negative correlation between aggression‐boldness and voracity. This research is a first step in investigating the impact of anthropogenic disturbances on behavioral syndromes and demonstrates that behavioral syndromes may vary with respect to insecticidal applications.     

Behavioral syndromes and the evolution of correlated behavior in zebrafish

Studies of "behavioral syndromes" in different populations andspecies of animals can be used to shed light on the underlyingmechanisms of evolution. For example, some personality syndromessuggest the existence of an underlying hormonal link, whereasother relationships between boldness and aggression appear tobe the result of similar selective pressures. Here, we used1 wild-derived and 2 laboratory strains of zebrafish (Daniorerio) to examine relationships among 5 behavioral measures:shoaling, activity level, predator approaches, latency to feedafter a disturbance, and biting to a mirror stimulus. We foundevidence of an activity syndrome, as if underlying metaboliccosts influence variation in multiple forms of behavior. Evidencefor a relationship between boldness and aggression was alsoapparent but depended both on strain and which specific behaviorpatterns were identified as measures of "boldness." Althoughsome comparisons of laboratory and wild-derived strains wereconsistent with a domestication syndrome, others were not. Mostobserved relationships were relatively weak and occasionallyinconsistent, arguing against strong underlying genetic linkagesor pleiotropic effects relating any of the behavioral measures.Instead, it may be more important to study the details of selectivecontext or the long-term impact of linkages between some, butnot all, of a large set of genes influencing complex behavioraltraits. We found profound differences among strains in mostbehavior patterns, but few sex differences. One strain (TM1)was consistently different from the others (SH and Nadia) beingmore social, more likely to approach predators, and taking lesstime to recover from disturbance than were the other 2 strains.     

Identification of the origin of an Atlantic salmon (Salmo salar L.) population in a recently recolonized river in the Baltic Sea

The founder event in a recently recolonized salmon population in the Baltic Sea (Gulf of Finland) was investigated. To identify the origin of the founders, four wild populations and two hatchery stocks were analysed using six microsatellite loci. The results of assignment tests and factorial correspondence analysis suggest that the initial recolonizers of the river Selja originated from the geographically nearest (7 km) wild population (river Kunda) but as the result of stocking activities, interbreeding between recolonizers and hatchery individuals has occurred in subsequent years. Although the hatchery releases are outnumbering the wild salmon recruitment in the Baltic Sea at present, our results suggest that the native populations may still have an important role in colonization processes of the former salmon rivers.     

Behavioural Syndromes in Urban and Rural Populations of Song Sparrows

Animals in urban habitats are often noticeably bold in the presence of humans. Such boldness may arise due to habituation, as urban animals learn, through repeated exposure, that passing humans do not represent a threat. However, there is growing research suggesting that: (1) inherent traits, as opposed to learned behaviour, influence which species invade urban habitats, and (2) individuals exhibit individual personality traits that limit behavioural flexibility, with the possible result that not all individuals would be able to demonstrate an appropriate level of boldness in urban environments. As a result, perhaps only birds with inherently bold personalities could successfully settle in an area of high human disturbance, and further, we might also expect to see the existence of behavioural syndromes, where boldness is correlated with variation in other behavioural traits such as aggression. In this study, we examined boldness and territorial aggression in urban and rural populations of song sparrows. We found that urban birds were bolder towards humans and that urban birds also showed higher levels of territorial aggression. We also found an overall correlation between boldness and territorial aggression, suggesting that urban boldness may be part of a behavioural syndrome. However, we see no correlation between boldness and aggression in the urban population, and thus, more work is needed to determine the mechanisms accounting for high levels of boldness and aggression urban song sparrows.     

Maternal effects in vulnerability to eye‐parasites and correlations between behavior and parasitism in juvenile Arctic charr

Hatchery‐reared fish show high mortalities after release to the wild environment. Explanations for this include potentially predetermined genetics, behavioral, and physiological acclimation to fish farm environments, and increased vulnerability to predation and parasitism in the wild. We studied vulnerability to Diplostomum spp. parasites (load of eye flukes in the lenses), immune defense (relative spleen size) and antipredator behaviors (approaches toward predator odor, freezing, and swimming activity) in hatchery‐reared juvenile Arctic charr (Salvelinus alpinus) using a nested mating design. Fish were exposed to eye‐fluke larvae via the incoming water at the hatchery. Fish size was positively associated with parasite load, but we did not find any relationship between relative spleen size and parasitism. The offspring of different females showed significant variation in their parasite load within sires, implying a dam effect in the vulnerability to parasites. However, the family background did not have any effect on spleen size. In the mean sire level over dams, the fish from the bolder (actively swimming) families in the predator trials suffered higher loads of eye flukes than those from more cautiously behaving families. Thus, the results indicate potentially maternally inherited differences in vulnerability to eye‐fluke parasites, and that the vulnerability to parasites and behavioral activity are positively associated with each other at the sire level. This could lead to artificial and unintentional selection for increased vulnerability to both parasitism and predation if these traits are favored in fish farm environments.     

Correlation between boldness and body mass in natural populations of the poeciliid Brachyrhaphis episcopi

The boldness of individual Brachyrhaphis episcopi , collected from regions of high and low predation, was investigated using two independent assays: (1) the time to emerge from cover and (2) the propensity to leave shoal mates and investigate a novel object. A strong correlation between the two assays was revealed such that fish that emerged from shelter sooner were also more likely to approach a novel object. This is indicative of a boldness personality axis acting across both behavioural contexts. Fish from high-predation areas were bolder than those from low-predation areas and males were bolder than females. A significant correlation between body mass, standard length ( L _S) and boldness score was also found. In general, bold fish had a greater body mass at a given L _S than shy fish. These results suggest that personality traits are strongly influenced by population-specific ecological variables and may have fitness consequences in wild populations.     

First maturity and spawning periodicity of hatchery‐origin pallid sturgeon in the upper Missouri River above Fort Peck Reservoir,Montana

The pallid sturgeon Scaphirhynchus albus conservation propagation program has augmented declining wild populations since the 1990s and the older age classes of hatchery‐origin fish are beginning to reach sexual maturity in the wild. Currently, the majority of the information available on the age and size at first maturity and spawning periodicity for pallid sturgeon in the upper basin is from captive hatchery‐origin pallid sturgeon (i.e. age and size at first maturity and spawning periodicity) or from wild pallid sturgeon artificially spawned in the propagation program (i.e. spawning periodicity). The purpose of this study was to document age and size at first maturity and spawning periodicity of known age hatchery‐origin pallid sturgeon that have reached maturity in the wild. Radio‐tagged pallid sturgeon in the upper Missouri River upstream of Fort Peck Reservoir were serially sampled in the early‐spring over multiple years and assigned to reproductive classifications each year based on sex‐steroid concentrations. The youngest reproductively‐active male hatchery‐origin pallid sturgeon sampled was 14.5 years old and the youngest female was 18. Hatchery‐origin males were observed having annual (N = 3) and biennial (N = 2) reproductive cycles. The observed spawning periodicity was similar to what has been reported elsewhere for the species. The youngest mature fish in this study are older and larger than what has been reported for those retained in captivity, indicating that body size alone is not a reliable predictor of maturity for pallid sturgeon.     

Interaction between hatchery and wild Pacific salmon in the Far East of Russia: A review

We review studies of interactions between hatchery and wild Pacific salmon in the Russian Far East. This includes the role of hatchery practices that result in premature migration to the sea and increased mortality, and data on feeding and territorial competition between juveniles of hatchery and wild origin. In the course of downstream migration many juvenile hatchery salmon are eliminated by wild salmon predation. During the marine period, Japanese hatchery chum salmon (Oncorhynchus keta) distribution overlaps the distribution of Russian wild salmon. Consequently, replacement of wild populations by hatchery fishes, as a result of abundant juvenile hatchery releases combined with extensive poaching in spawning grounds, is apparent in some Russian rivers. Interactions between the populations occur in all habitats. The importance of conservation of wild salmon populations requires a more detailed study of the consequences of interactions between natural and artificially reared fishes.     

Population genetic structure in wild and hatchery populations of white cloud mountain minnow (Tanichthys albonubes): Recommendations for conservation

We used mitochondrial (mt) cytochrome b gene (cyt b) to compare the genetic variability in three hatchery broodstocks of white cloud mountain minnow with the variability in six wild populations sampled in two river drainages. A total of 43 haplotypes in 102 specimens were observed, with no haplotype shared between wild and hatchery populations. The nucleotide diversity of the wild samples (0.048) was significantly higher than that of the hatchery ones (0.007), but the haplotype diversity was almost similar between them. Two major phylogenetic haplotype groups were revealed and estimated to diverge about 6.531 myr (million years) ago. Significant genetic differentiation was revealed between wild and hatchery populations as well as among nine sampled populations, suggesting at chance effect during the founding process for the hatchery population and a subsequent genetic drift. According to the network, the connection between wild and hatchery populations indicates that present hatchery populations originated from single wild population. We suggested that two regions (Pearl River system and Lu River) identified by reciprocal mtDNA monophyly and SAMOVA should be regarded as three different ESUs and two different MUs in South China, respectively.     

Compensatory releases reduce genetic differentiation among Atlantic salmon populations in the Baltic Sea: evidence from the River Ume‐Vindelälven

About 90% of salmon smolts in the Baltic Sea derive from hatchery enhancement programmes designed to compensate for loss of catches and genetic resources due to the damming of salmon rivers. The potential threat of genetic homogenization from extensive hatchery releases, however, has not been thoroughly investigated. We provide evidence that straying from deliberate releases poses a threat to indigenous populations by identifying the origin of 127 fin‐clipped (hatchery) salmon caught in the River Ume‐Vindelälven during 1997 to 2000, using mtDNA and six microsatellite loci. The analysis of eight potential donor stocks revealed that compensatory releases from the R. Ångerman and R. Luleälven hatcheries have resulted in a significant amount of straying to the river Ume‐Vindelälven (at least 10 and 12 migrants per year). As predicted to due to increased migration, the analysis of temporal samples from the wild population of R. Vindelälven showed a decreasing trend in genetic differentiation estimates (measured as Fst) relative to hatchery the hatchery strains of R. Ångermanälven and R. Luleälven. Our results suggest that gene flow from compensatory releases poses a serious threat to the genetic makeup of the existing wild populations in the Baltic.