The correlation between subordinate fish eye colour and received attacks: a negative social feedback mechanism for the reduction of aggression during the formation of dominance hierarchies

Eye darkening has been linked to social status in fish. The subordinate's eyes darken, while the eyes of the dominant fish become pale. Although this phenomenon has been described in salmonid fishes and in the African cichlid Nile tilapia Oreochromis niloticus, it is unclear whether eye darkening correlates with a reduction in aggressive behaviour. Thus, we evaluated the link between social status and eye darkening. We evaluated whether the eye colours of subordinate fish correlate with the frequency of received attacks in a neotropical fish, the pearl cichlid Geophagus brasiliensis. We paired pearl cichlids and quantified both the aggressive behaviour and the eye darkening of each fish. As has been described for Nile tilapia and Atlantic salmon, a clear-cut hierarchical relationship formed, where dominance and subordination were associated with pale and dark eye colours, respectively. Initially, eye colour darkening was positively correlated with the frequency of received attacks; however, a negative association occurred following eye darkening, in which the intensity of aggressive interactions decreased. Thus, fish that initially received a high number of attacks signalled subordination more rapidly and intensely (rapid and dramatic eye darkening), thereby inducing a negative social feedback mechanism that led to reduced aggression.     

Preliminary study on eye colour in Japanese macaques <Emphasis Type="Italic">(Macaca fuscata)</Emphasis> in their natural habitat

Eye colour in Japanese macaques shows apparent differences between individuals, continuously ranging from orange (bright), through shades of yellow and hazel-blue to dark blue (dark). We arbitrarily classified them into either 'yellow' eyes or 'blue' eyes based on the yellow area occupying in the iris' peripupillary ring. Most Japanese macaques have yellow eyes after infant phase, whilst 19, 17, 12, and 15% of monkeys (>6 months, sexes combined) have blue-eye in studied two groups of Shodoshima and two groups of Takasakiyama, respectively. Frequency of eye colour did not differ between males and females, but significantly differed in each age class. Blue eyes significantly more frequently occurred in newborns, infants and aged monkeys than in juveniles and prime adults. Data from mother-infant pairs indicated eye colour could be inherited from their parents. A case of asymmetric eye colour in Japanese macaques was found from a sample of 1962 individuals. Eye colour variation of Japanese macaques was discussed in relation to those of humans and rhesus macaques. A possible evolutionary model of eye colour in Japanese macaques was discussed.     

Endocrine differences among colour morphs in a lizard with alternative behavioural strategies

Alternative behavioural strategies of colour morphs are expected to associate with endocrine differences and to correspond to differences in physical performance (e.g. movement speed, bite force in lizards); yet the nature of correlated physiological and performance traits in colour polymorphic species varies widely. Colour morphs of male tawny dragon lizards Ctenophorus decresii have previously been found to differ in aggressive and anti-predator behaviours. We tested whether known behavioural differences correspond to differences in circulating baseline and post-capture stress levels of androgen and corticosterone, as well as bite force (an indicator of aggressive performance) and field body temperature. Immediately after capture, the aggressive orange morph had higher circulating androgen than the grey morph or the yellow morph. Furthermore, the orange morph maintained high androgen following acute stress (30 min of capture); whereas androgen increased in the grey and yellow morphs. This may reflect the previously defined behavioural differences among morphs as the aggressive response of the yellow morph is conditional on the colour of the competitor and the grey morph shows consistently low aggression. In contrast, all morphs showed an increase in corticosterone concentration after capture stress and morphs did not differ in levels of corticosterone stress magnitude (CSM). Morphs did not differ in size- and temperature-corrected bite force but did in body temperature at capture. Differences in circulating androgen and body temperature are consistent with morph-specific behavioural strategies in C. decresii but our results indicate a complex relationship between hormones, behaviour, temperature and bite force within and between colour morphs.     

Status-dependent foraging behaviour in coral reef wrasses

Field observations using underwater video were used to reveal differences in the duration and frequency that fish engaged in daily behaviours such as chasing, searching, feeding, and travelling, according to their social patterns [passive or aggressive terminal phases (TPs), pair-spawning, or promiscuous groups] and intraspecific status (terminal or initial colour phases). Marked differences were apparent according to status, but this was not consistent among the three species; TP Cheilinus fasciatus tended to swim in longer bouts with less frequent searching or feeding than initial phase (IP) conspecifics; TP Cirrhilabrus punctatus exhibited less frequent feeding and travelling than IP conspecifics, while the most aggressive species Stethojulis bandanensis displayed no significant intraspecific differences. We highlight the importance of social context and individual status when examining fish foraging activities and the utility of underwater video for recording the duration and frequency that fish engage in essential daily activities.     

Does darkening signal submission in territorial contests between juvenile Atlantic salmon, Salmo salar ?

Communication by means of visual signals occurs during the competitive, aggressive and sexual interactions of many animals. Some animals such as fish are able to change their body coloration rapidly, and there is evidence that this is used as a means of signalling. However, the precise meaning of such signals is rarely understood. We examined whether (1) darkness in juvenile Atlantic salmon is associated with submission, and (2) changing to a darker colour could act as a signal to the opponent and hence induce a change in its behaviour. We found that both the sclera of the eye and the overall body coloration tended to darken in fish that were losing territorial encounters, while victors retained their original coloration. The darkening was rapid and usually occurred during a period of sustained attacks by the opponent; however, the aggression level decreased as soon as the losing fish had become darker. We suggest that the darkening of the losing fish was associated with submission and may result in a change in the behaviour of its opponent, so minimizing the potential risk of injury during unnecessarily prolonged fights. Copyright 1999 The Association for the Study of Animal Behaviour.     

Levels of social tolerance between snow volesChionomys nivalis during over-wintering periods

We examined winter social behaviour of the snow voleChionomys nivalis Martins, 1842, a rock-dwelling microtine found principally in high-mountain biotopes. We conducted dyadic encounters among wild-caughtC. nivalis to determine patterns of intraspecific tolerance and the degree of aggressiveness between and within sexes during wintering periods. Our data show a high frequency of agonistic behaviours in all social encounters. Male pairs spent less time being aggressive than female pairs but more time in investigative behaviours. In contrast, the intensity of aggression displayed during interactions was significantly higher in male-male encounters than in female-female encounters, with an intermediate level of aggression in male-female encounters. The low degree of social tolerance found seems to be in agreement with the available data on the social organization of this species and further suggests that individuals may not spatially associate during over-wintering periods. Although both males and females were socially intolerant, the different patterns observed between the sexes may underlie different strategies.     

Host behaviour alteration by its parasite: from brain gene expression to functional test

Many parasites with complex life cycles modify their intermediate hosts'' behaviour, presumably to increase transmission to their final host. The threespine stickleback (Gasterosteus aculeatus) is an intermediate host in the cestode Schistocephalus solidus life cycle, which ends in an avian host, and shows increased risky behaviours when infected. We studied brain gene expression profiles of sticklebacks infected with S. solidus to determine the proximal causes of these behavioural alterations. We show that infected fish have altered expression levels in genes involved in the inositol pathway. We thus tested the functional implication of this pathway and successfully rescued normal behaviours in infected sticklebacks using lithium exposure. We also show that exposed but uninfected fish have a distinct gene expression profile from both infected fish and control individuals, allowing us to separate gene activity related to parasite exposure from consequences of a successful infection. Finally, we find that selective serotonin reuptake inhibitor-treated sticklebacks and infected fish do not have similarly altered gene expression, despite their comparable behaviours, suggesting that the serotonin pathway is probably not the main driver of phenotypic changes in infected sticklebacks. Taken together, our results allow us to predict that if S. solidus directly manipulates its host, it could target the inositol pathway.     

Aggressive behaviour of underyearling rainbow trout in simulated winter concealment habitat

The behaviour of wild underyearling rainbow trout Oncorhynchus mykiss in concealment habitat in a laboratory stream aquarium at 2 and 6°C was recorded daily with an infrared video camera for 90 min over dawn. Aggressive behaviours (threat nips, nips and chases) were frequent during this time as fish entered concealment habitat. Aggressiveness varied widely among fish groups, with a range of 1 to 45 aggressive acts being initiated during a 90-min filming period. Larger fish initiated most of the encounters and removed a higher proportion of fish from concealment than did smaller aggressors. Thirteen per cent of the aggressive acts resulted in the recipient being completely removed from concealment habitat. The highest combined frequency of aggressive acts was 0.64 per visible fish per 10 min period and occurred at relatively low light levels (300 lx). The data suggest that when fish density is high, such as when habitat is limited, aggressive behaviour in winter may cause some underyearling salmonids to be excluded from concealment.     

Swimming abilities of juvenile estuarine fishes: implications for passage at water control structures

The capability of early life history stage fishes to access nursery habitat within managed salt marshes is dependent on their ability to negotiate water control structures (WCSs). Knowledge of swimming ability and hydrodynamic preferences is essential to assess the impact of WCSs on fish movement in managed marshes. These data, however, are lacking for many common estuarine fishes, and the utility of the data for the few species examined thus far is limited. We examined critical swimming speeds and derived linear relationships between fish size and swimming speed for juveniles of six common estuarine fish species of the southeast U.S. and northern Gulf of Mexico coasts. White mullet Mugil curema displayed the greatest swimming ability among these six species and was able to swim against currents ≥ 30 cm s^?1 higher than the other species examined at the same size. The remaining species displayed lower critical swimming speeds and were classified into groups of moderate (pinfish Lagodon rhomboides, striped mullet Mugil cephalus) or slow (silver perch Bairdiella chrysoura, spotfin mojarra Eucinostomus argenteus, spot Leiostomus xanthurus) swimmers. Our results suggest that high-flow conditions at WCSs would likely preclude the passage of all but the largest juvenile fishes, and passage for most juveniles would occur under low-flow conditions; these flows at WCSs are dictated largely by site-specific tidal and weather conditions.     

Inter-male aggression with regard to polygynous mating system in Pampean grassland mouse, Akodon azarae (Cricetidae: Sigmodontinae)

Based on the hypothesis that, in Akodon azarae, polygyny operates through female defence, we studied inter-male aggression in order to test the following predictions: during the breeding period (1) resident males are more aggressive than intruder males in the presence of females (FP), and (2) aggressive behaviour is independent of male condition (resident or intruder) in the absence of females (FA). To test our predictions, we used the resident male behavioural response towards an intruder male in relation to FP or FA. We conducted 30 encounters in FP and 27 in FA in 0.79-m² round enclosures placed in the Espinal Reservation. Our results support the prediction that, in FP, the intensity of aggressive behaviour exhibited by males varied in relation to resident or intruder condition. Resident males showed high levels of aggression towards intruders, and intruders exhibited the greatest values of submissive behaviours with residents. In FA, the intensity of aggressive behaviour did not vary in relation to resident or intruder condition. Both resident and intruder males exhibited low aggressive behaviour and inter-male encounters resulted mainly in non interactive behaviours. Our results support the hypothesis that, in A. azarae, the polygynous mating system operates through female defence.     

Ontogenesis of agonistic vocalizations in the cichlid fish Metriaclima zebra

While acoustic communication has been described in adults of various fish species, our knowledge about the ontogeny of fish sound production is limited. In adults, sound signals are known to be involved during aggressive interactions. However, aggressive behaviour may appear early in the life of fishes due to the possible competition for food and space. If acoustic signals are used to send information to competitors, sounds are likely to play a role during interactions between juvenile fish as well. The apparition and evolution of sound production were monitored in a group of juveniles of the cichlid fish Metriaclima zebra from hatching to 4months of age. In addition, the link between vocalizations and agonistic behaviour was studied during dyadic interactions at three different ages. Sounds production appeared to be present early in the development of this fish and increased along with the number of aggressive behaviours. Recorded sounds consisted, in juveniles, in isolated pulses showing a decrease in frequency and duration as the fish grew. In adults, sounds became bursts of pulses but the transition from isolated to repetitive pulses was not observed. These results are compared to the existing literature on sound production ontogeny in fishes.     

Field study of Haplochromis burtoni: Quantitative behavioural observations

Haplochromis burtoni observed in their natural habitat behave differently from those observed in aquaria in several respects. The majority of the territorial males observed in Lake Tanganyika living in a colony did not have the characteristic black eye bar as a part of their bright coloration, although otherwise the colour markings were identical. Those animals with eye bars appeared to be at a disadvantage in competition with barless fish for territories. The non-territorial males, females and juveniles are uniformly cryptically coloured and remained together in two or three large schools on the edge of the colony. These schools drifted slowly through the colony area, attempting to feed in the territories. Territorial males solicited females and chased non-territorial males. Sequences of behavioural events performed during courtship varied, depending on the local population density. Sound recordings showed no purposeful sounds associated with any behavioural acts.     

Lateralisation of visual function in yellow-eyed mullet (Aldrichetta forsteri) and its role in schooling behaviour

Lateralisation of cognitive function is a common phenomenon found throughout the animal kingdom. In fish, lateralisation is involved in visually mediated behaviours such as schooling, predator avoidance and foraging. This study sought to quantify visual laterality in yellow-eyed mullet (Aldrichetta forsteri), to determine visual asymmetries and the significance of lateralisation, and the factors which influence functional behaviours in schooling fish. Tank-based experiments investigated individual and population level lateralisation, schooling behaviour, and optic lobe asymmetry. Variation in direction and strength of lateralisation in individuals was found in juveniles, but at a population level significant bias was found in adults. Right eye biased fish had a larger right optic lobe than control fish. Strongly lateralised fish showed a tendency to occupy positions of safety within the school more frequently than control fish. In combination with observed differences in schooling behaviour, the possibility of ontogenetic plasticity in behavioural lateralisation and optic lobe morphology is suggested.     

Behavioural plasticity in a native species may be related to foraging resilience in the presence of an aggressive invader

Competition between invasive and native species can result in the exploitation of resources by the invader, reducing foraging rates of natives. However, it is increasingly recognized that multiple factors can enhance the resilience of native species competing for limiting resources with invaders. Although extensively studied in terrestrial species, little research has focused on behavioural plasticity in aquatic ecosystems and how this influences native species resilience. Here, we examined the role of behavioural plasticity in interactions between a native Australian fish, Pseudomugil signifer, and a widespread invasive fish, Gambusia holbrooki. To determine whether P. signifer displays behavioural plasticity that may mitigate competition with G. holbrooki, we first quantified social behaviours (aggression, submission and affiliation) and shoal cohesion for each species in single- and mixed-species groups. Second, we compared the feeding rates of both species in these groups to ascertain if any modulation of social behaviours and cohesion related to foraging success. We found that aggressive and submissive behaviours of G. holbrooki and P. signifer showed plasticity in the presence of heterospecifics, but social affiliation, shoaling and, most importantly, foraging, remained inflexible. This variation in the degree of plasticity highlights the complexity of the behavioural response of a native species and suggests that both behavioural modulation and consistency may be related to sustaining foraging efficiency in the presence of an invader.     

The scaling of locomotor performance in predator-prey encounters: from fish to killer whales.

During predator-prey encounters, a high locomotor performance in unsteady manoeuvres (i.e. acceleration, turning) is desirable for both predators and prey. While speed increases with size in fish and other aquatic vertebrates in continuous swimming, the speed achieved within a given time, a relevant parameter in predator-prey encounters, is size independent. In addition, most parameters indicating high performance in unsteady swimming decrease with size. Both theoretical considerations and data on acceleration suggest a decrease with body size. Small turning radii and high turning rates are indices of maneuverability in space and in time, respectively. Maneuverability decreases with body length, as minimum turning radii and maximum turning rates increase and decrease with body length, respectively. In addition, the scaling of linear performance in fish locomotion may be modulated by turning behaviour, which is an essential component of the escape response. In angelfish, for example, the speed of large fish is inversely related to their turning angle, i.e. fish escaping at large turning angles show lower speed than fish escaping at small turning angles. The scaling of unsteady locomotor performance makes it difficult for large aquatic vertebrates to capture elusive prey by using whole-body attacks, since the overall maneuverability and acceleration of small prey is likely to be superior to that of large predators. Feeding strategies in vertebrate predators can be related to the predator-prey length ratios. At prey-predator ratios higher than approximately 10(-2), vertebrate predators are particulate feeders, while at smaller ratios, they tend to be filter feeders. At intermediate ratios, large aquatic predators may use a variety of feeding methods that aid, or do not involve, whole body attacks. Among these are bubble curtains used by humpback whales to trap fish schools, and tail-slapping of fish by delphinids. Tail slapping by killer whales is discussed as an example of these strategies. The speed and acceleration achieved by the flukes of killer whales during tail slaps are higher and comparable, respectively, to those that can be expected in their prey, making tail-slapping an effective predator behaviour.     

The effect of territorial damselfish (family Pomacentridae) on the space use and behaviour of the coral reef fish, Halichoeres bivittatus (Bloch, 1791) (family Labridae)

Most demersal species of damselfish (family Pomacentridae) are territorial herbivores that aggressively chase other fishes away from their nests. This study investigates whether the aggressive territorial damselfish, Stegastes leucostictus, modifies behaviours and home range area use in a less aggressive, non-territorial species, the slippery dick wrasse, Halichoeres bivittatus (family Labridae). Damselfishes and wrasses are ubiquitous and abundant members of coral reef fish assemblages around the world; hence, this study has broader implications beyond the two Caribbean species chosen for this study. A manipulative field experiment consisted of transplanting one or three S. leucostictus into artificial shelters positioned within adult or adjacent to juvenile H. bivittatus home ranges. The introduction of damselfish had little effect on the size of home range areas of juvenile or adult wrasses, but had a significant effect on the location of their home ranges. The damselfish also affected adult microhabitat use but did not affect use by juveniles. In addition, there was an increase in damselfish-wrasse interactions that resulted in a significant reduction in the amount of time adult wrasses could spend foraging or in a foraging assemblage. It was concluded that aggressive interactions with territorial damselfish affect individual H. bivittatus space use and may consequently affect the spatial distribution of H. bivittatus populations.     

Territorial behaviour reflects sexual status in groups of false clown anemonefish (Amphiprion ocellaris) under laboratory conditions

Anemonefishes (genus Amphiprion) are well known as protandrous sex-changing fish. They live symbiotically with sea anemones and form a social unit that consists of a breeding pair and several sexually immature individuals with ambisexual gonads. In general, the hierarchy of the social rank is strictly maintained in a group, and this hierarchy is considered to be one of the critical factors that induce sex change or sex differentiation in anemonefishes. Field observations have shown that behavioural characteristics of anemonefishes vary depending on the environment. In this study we examined the differences in territorial aggression directed at conspecific intruders according to sexual status in the false clown anemonefish, Amphiprion ocellaris, under laboratory conditions. Resident–intruder tests were conducted on groups consisting of three anemonefish: a dominant female, a second-ranked male and a subordinate ambisexual individual. Females and males showed more aggressive behaviour than did ambisexual individuals, and behaviours were specifically directed at intruders of the same sexual status, not at those of the opposite sex. Notably, the female residents displayed fierce aggressive behaviours particularly toward female intruders, whereas male residents engaged more often in side-by-side swimming with male intruders than with female or ambisexual intruders. Ambisexual intruders were threatened by all residents, rarely attacked. We observed only few aggressive acts towards tank mates. These results suggest that sexually mature resident anemonefish in a confined space, such as under laboratory conditions, perceive intruders of the same sexual status as competitors for reproductive status and thus, dominant social rank.     

Social conditioning and dominance in male betta splendens

To determine whether social conditioning modifies relative dominance in the Siamese fighting fish ($\text{Betta splendens}$), individual fish were matched for gill cover erection frequency (GCEF) to a mirror. One member of each pair experienced a series of four winning encounters (dominants) while the other of the pair experienced four losing encounters (subordinates). After the fighting experience, matched pairs were placed together and aggressive responses of both fish were recorded. Fish experiencing wins displayed significantly more and were more aggressive than fish experiencing losses which showed significantly higher levels of avoidance. However, no change in mirror GCEF occurred as a result of differential experience.     

Secondary poisoning of stoats (Mustela erminea), feral ferrets (Mustela furo), and feral house cats (Felis catus) by the anticoagulant poison,brodifacoum

Abstract

The extent of darkening of melanin‐based plumages in birds has previously been linked with increasing aggressive encounters between individuals. The North Island robin (Petroica longipes) is a territorial New Zealand endemic passerine that displays delayed plumage maturation (darkening of the plumage with age). Aggressive boundary interactions in the robin are relatively common during the breeding season, when territories are protected and juveniles are dispersing. This study tests the hypothesis of aggression‐mediated plumage darkening in a population of North Island robins by examining if males and older (darker) birds are either (1) involved in a higher number of aggressive interactions, or (2) are more often the aggressor than females and younger birds. When sex and age are accounted for, darker individuals will be either (3) involved in a higher number of interactions or (4) more often the aggressor in encounters with other individuals. Data were collected by scoring the plumage darkness of 32 individuals in the field, and observing (1) interaction behaviours, and (2) age and sex of the birds involved in each interaction. The results show no support for any aggression‐mediated plumage darkness in the robin; males and older birds were not involved in more aggressive interactions, and were not more often the aggressor; and neither the frequency of interactions or the number of aggressive interactions were correlated with a darker plumage. Other more complex mechanisms may explain delayed plumage darkness in the North Island robin.     

Energy utilization during swimming and cost of locomotion in larval and juvenile fish

Oxygen consumption and ammonia excretion rates increased in an accelerated manner in larvae and juveniles of whitefish (Coregonus sp.) as a function of swimming speed. The three-dimensional patterns of fish metabolic rates (expressed as energy consumed or nitrogen excreted) versus body weight and swimming speed show that the total standard metabolic rate (i. e. at extrapolated zero swimming speed) increased during early life of whitefish, followed by the expected decrease. This phenomenon might be due to the profound changes in oxidative and glycolytic enzyme activities during fish “metamorphosis”. Standard metabolic rate of ammonia excretion, as a principal product of protein catabolism in fish, decreased by one order of magnitude in early coregonid ontogenesis. This means that protein utilization as an energy source decreases as far as standard metabolism is concerned, but increases with swimming speed. This trend is opposite that in adult fish, where protein utilization in the overall energy supply is diminished at increasing swimming speed. The cost of locomotion offish larvae and juveniles demonstrates that the energy expenditure increases logarithmically with decreasing fish size but at an accelerated rate as compared to adult fish. This contradicts earlier estimates of lower cost of swimming in fish larvae than cost of paddle-propulsion swimming in small invertebrates or cost of flying in insects.