Genetics of social behaviour in charrs (Salvelinus species)

Juvenile brook charr (Salvelinus fontinalis) are territorial and have high levels of agonistic behaviour. Juvenile lake charr (S. namaycush) rarely show any indication of territoriality and have very low levels of agonistic behaviour. The inheritance of behaviour in their reciprocal hybrids was investigated by multivariate stepwise discriminant analysis. Lateral display, charge, chase, and forage composed the first canonical axis (87.2% of total dispersion). Pooled hybrid means were closer to that of brook charr, suggesting either directional dominance or modification of the behaviour of hybrids in response to water flow. The mean values of reciprocal hybrids were skewed towards the maternal parent, suggesting a maternal effect in the phenotypic expression of behaviour in these fish.     

Agonistic behaviour of age 0, age 1 and non-breeding adult Arctic grayling Thymallus arcticus (Pallas)

Age 0 Arctic grayling begin to show agonistic behaviour 3 weeks after emergence and are territorial by their fourth week. Age 1 fish are territorial throughout the summer feeding period. In adults, territoriality is restricted to larger males during the early spring breeding season while after the breeding season both smaller adult males and adult females also hold territories.
In sub-adults, larger fish and resident fish tend to win most fights. In adults, males usually win against females. In fights between adults of the same sex, larger and resident fish usually win.
The lateral display becomes more important in the agonistic repertoire of Arctic grayling as they mature and the characteristic large dorsal fin develops. Arctic grayling appear to lack an appeasement display.     

Genetic analysis of male reproductive success in relation to density in the zebrafish, Danio rerio

Background

We used behavioural and genetic data to investigate the effects of density on male reproductive success in the zebrafish, Danio rerio. Based on previous measurements of aggression and courtship behaviour by territorial males, we predicted that they would sire more offspring than non-territorial males.

Results

Microsatellite analysis of paternity showed that at low densities territorial males had higher reproductive success than non-territorial males. However, at high density territorial males were no more successful than non-territorials and the sex difference in the opportunity for sexual selection, based on the parameter I _mates, was low.

Conclusion

Male zebrafish exhibit two distinct mating tactics; territoriality and active pursuit of females. Male reproductive success is density dependent and the opportunity for sexual selection appears to be weak in this species.     

Territoriality and male-biased sexual size dimorphism in Argia reclusa (Odonata: Zygoptera)

In Odonata, many species present sexual size dimorphism (SSD), which can be associated with male territoriality in Zygoptera. We hypothesized that in the territorial damselfly Argia reclusa, male–male competition can favor large males, and consequently, drive selection pressures to generate male-biased SSD. The study was performed at a small stream in southeastern Brazil. Males were marked, and we measured body size and assessed the quality of territories. We tested if larger territorial males (a) defended the best territories (those with more male intrusions and visiting females), (b) won more fights, and (c) mated more. Couples were collected and measured to show the occurrence of sexual size dimorphism. Results indicated that males are larger than females, and that territorial males were larger than non-territorial males. Larger territorial males won more fights and defended the best territories. There was no difference between the mating success of large territorial and small non-territorial males. Although our findings suggest that male territoriality may play a significant role on the evolution of sexual size dimorphism in A. reclusa, we suggest that other factors should also be considered to explain the evolution of SSD in damselflies, since non-territorial males are also capable of acquiring mates.     

Does part reproductive history predict competitive interactions and male mating success in pupfish?

Two experiments were performed to determine the effect of past territoriality and mating success on subsequent dominance and ability to attract females. First, in 14 staged, pair-wise encounters, the development of nuptial coloration and agonistic behaviour was examined in two types of males: previously territorial (PT) and previously non-territorial (PNT) males. Previously territorial males developed a more intense nuptial coloration and won all contests. Second, to test the males' ability to attract females, other factors were controlled that could have affected female mate choice, such as male size, quality of breeding substrate and male-male interactions. In 40 trials, PT males were more active, developed a more intense nuptial coloration, courted females more vigorously, spawned sooner and had a higher mating success than PNT males. Thus, in pupfish, Cyprinodon pecosensis, previous reproductive history is a good indicator of both dominance and mating success. Only males in good physical condition achieve high mating success, and both intra- and inter-sexual selection are important in maintaining the close correlation between expression of nuptial coloration, a second sexual trait, and other male attributes, such as physical condition and vigour, that allow a male to secure and defend a territory against rivals and to attract and spawn with females.     

Territorial behaviour in corn mice, <Emphasis Type="Italic">Calomys musculinus</Emphasis> (Muridae: Sigmodontinae), with regard to mating system

We studied territorial behaviour of Calomys musculinus, a promiscuous-polygynous species, in order to test the predictions that during the breeding period, home-range-owner females are more aggressive than home-range intruders and that males do not display territorial behaviour. At the core of the breeding season, we conducted 136 intrasexual and 66 intersexual encounters. The study was carried out in four 0.25-ha enclosures, each containing one independent population. We studied territorial behaviour utilising focal animal sampling to record behaviour. Encounters were conducted in a circular opaque arena (COA) near the home-range’s activity mathematical centre (AMC). Amicable behaviour was not observed in any of the 78 trials among females. The rates of aggressive and submissive behaviour per female–female encounters varied in relation to whether they were home-range owners or intruders, the former being aggressive, the latter submissive. So we could link these drives to the territorial behaviour of C. musculinus. This result fits well with the already known territoriality of C. musculinus during the breeding season. In contrast, the 58 intrasexual male encounters did not show direct aggressive interactions between them. The rates of the different behaviours between males did not vary in relation to whether they were home-range owners or intruders. In our study, which took place during the breeding period, C. musculinus males were nonterritorial, and they never exhibited aggressiveness towards females.     

Genetic versus Rearing-Environment Effects on Phenotype: Hatchery and Natural Rearing Effects on Hatchery- and Wild-Born Coho Salmon

With the current trends in climate and fisheries, well-designed mitigative strategies for conserving fish stocks may become increasingly necessary. The poor post-release survival of hatchery-reared Pacific salmon indicates that salmon enhancement programs require assessment. The objective of this study was to determine the relative roles that genotype and rearing environment play in the phenotypic expression of young salmon, including their survival, growth, physiology, swimming endurance, predator avoidance and migratory behaviour. Wild- and hatchery-born coho salmon adults (Oncorhynchus kisutch) returning to the Chehalis River in British Columbia, Canada, were crossed to create pure hatchery, pure wild, and hybrid offspring. A proportion of the progeny from each cross was reared in a traditional hatchery environment, whereas the remaining fry were reared naturally in a contained side channel. The resulting phenotypic differences between replicates, between rearing environments, and between cross types were compared. While there were few phenotypic differences noted between genetic groups reared in the same habitat, rearing environment played a significant role in smolt size, survival, swimming endurance, predator avoidance and migratory behaviour. The lack of any observed genetic differences between wild- and hatchery-born salmon may be due to the long-term mixing of these genotypes from hatchery introgression into wild populations, or conversely, due to strong selection in nature—capable of maintaining highly fit genotypes whether or not fish have experienced part of their life history under cultured conditions.     

A test of the model of the economic defendability of a resource and territoriality using youngEtroplus maculatus andPelmatochromis subocellatus kribensis

YoungEtroplus maculatus andPelmatochromis subocellatus kribensis were reared on diets of low, medium, or high quantities of food and the ontogeny of behavior documented until fish matured. Data were categorized into: (1) from the beginning of the experiment to the midpoint (100 days forE. maculatus and 118 days forP. s. kribensis); and (2) from the midpoint to maturity. For both species the tendency to be territorial, as deduced by the numbers of territories defended, or number of charges or rams performed, was explained by a unimodal relationship between territorial tendency and food available per gram of fish. Young fish receiving low quantities of food were highly territorial, but as they grew and food available per gram of fish decreased, they gave up territorial behavior. Young fish receiving high quantities of food were not territorial initially, but as they grew became increasingly territorial. Fish receiving intermediate quantities of food always defended territories. Lateral displays were performed most often by fish receiving intermediate quantities of food apparently because they were ambivalent being close to either the lower or upper thresholds for territoriality.     

From breeding pairs to fox towns: the social organisation of arctic fox populations with stable and fluctuating availability of food

Food availability can impact group formation in Carnivora. Specifically, it has been suggested that temporal variation in food availability may allow a breeding pair to tolerate additional adults in their territory at times when food abundance is high. We investigate group occurrence and intraspecific tolerance during breeding in a socially flexible canid, the arctic fox (Vulpes lagopus). We compare Iceland and Sweden where resource conditions differ considerably. A breeding pair was the most common social unit in both populations, but as predicted, groups were more frequent where food abundance varied substantially between years (Sweden: 6 %) than where food availability was stable (Iceland: ≤2 %). Within Sweden, supplemental feeding increased group occurrence from 6 to 21 %, but there was no effect of natural variation in lemming (Lemmus lemmus) availability since group formation was rare also at lemming highs. Thus, additional factors appeared to influence the trade-off between intraspecific territoriality and tolerance. We report two cases where related females showed enduring social relationships with good-neighbour strategies. Related females also engaged in alloparental behaviour in a ‘fox town’ with 31 foxes (4 adults, 3 litters). In contrast, when unrelated foxes bred close to each other, they moved or split their litters during summer, presumably because of territorial conflict. We suggest that fluctuating food availability is linked to group formation in this Arctic carnivore, but also when food availability increases, additional factors such as relatedness, alloparental benefits, competition and predator defence appear necessary to explain group formation.     

Territorial aggression can be sensitive to the status of heterospecific intruders

Territorial animals are known to be able to differentiate between intruding individuals posing a low or high threat and adjust their aggressive response accordingly. However, plastic territorial aggression based on recognising individuals with different attributes is typically assumed to be relevant only in the context of conspecific interactions. In this study, we investigated territorial aggression of neotropical cichlid fish in their natural habitat to assess whether responses to different types of individuals of another species can also be plastic. We show that arrow cichlids (Amphilophus zaliosus) adjusted their territorial aggression regarding the status of heterospecific intruders: breeding individuals of Amphilophus astorquii received a lower level of aggression than non-breeders. The same pattern was also found for the two different types of A. astorquii individuals intruding into conspecific territories. These results suggest that heterospecific individuals should not be ignored when considering selection pressures shaping plasticity of aggressive behaviour in territorial animals.     

Seasonal changes in the territorial behaviour of the satyrine butterfly Lethe diana are mediated by temperature

The territorial behaviour of butterflies often changes with temperature. The satyrine butterfly Lethe diana has three generations a year, and males display territorial behaviour in the May–June and September–October generations, but not in the July–August generation. This study investigated the relationship between this seasonal change in mate-locating behaviour and thermoregulation. When L. diana was able to hold a territory, thoracic temperature ranged from 23.8 to 33.6°C. This temperature was mainly influenced by environmental temperature based on air temperature, solar radiation, and wind, and metabolic heat was estimated to increase thoracic temperature by about 5°C in the May–June generation. When environmental temperature at a territorial site was within this range of the thoracic temperature minus the metabolic heat (approximately 5°C), L. diana males held territories. Since territorial sites were selected irrespective of the temperature, L. diana could not hold a territory when the temperature of the territorial site exceeded the threshold. In July–August, the temperature of the territorial site was almost always above the suitable range. These results suggest that seasonal change in territoriality of L. diana is due to behavioural thermoregulation. Electronic Publication     

Male territoriality and ‘sex confusion’ in recently adapted lizards at White Sands

The evolution of intersexual interactions, like mate choice, during ecological speciation has received widespread attention. However, changes in intrasexual interactions, like male territoriality, during ecological divergence are largely unexamined. We conducted field experiments with adaptively diverged populations of the eastern fence lizard (Sceloporus undulatus) to determine whether territorial males behaved differently towards ecologically similar vs. dissimilar intruders. We performed trials with light‐coloured males from White Sands, New Mexico and dark‐coloured males from the surrounding desert. We found that intruders from White Sands elicited more aggression than intruders from dark‐soil habitat. We also documented a case of ‘sex confusion’ where white‐sand males courted dark‐soil intruders. We found population differences in signalling patch size that can explain both aggression bias and sex misidentification. We argue that direct selection (for population recognition or optimal signal transmission) and indirect selection (by‐products of ecological adaptation) should influence both intersexual and intrasexual interactions during ecological speciation.     

Introduced Goldfish Affect Amphibians through Inhibition of Sexual Behaviour in Risky Habitats: an Experimental Approach

The introduction of alien species is one of the major causes of current and global biodiversity loss. The introduction of fish can be a particular threat to native amphibian populations, which are declining worldwide. One way for amphibians to persist in such altered environments is to adopt anti-predator strategies especially at the behavioural level. However, although it has been shown that avoidance behaviour may decrease the probability of being detected by a potential predator, little is known on the consequences on sexual behaviour. In this study, we tested the hypothesis that adult Alpine newts (Ichthyosaura alpestris) use shelters more often and exhibit less sexual activity in the presence of goldfish (Carassius auratus) and that they reduce sexual activity more in risky micro-habitats than in safe environments. To this end, we assessed behavioural patterns of adult newts in a replicated laboratory design. Goldfish were present in direct contact with newts in half of the tanks. Consistently throughout the study period, significantly more newts used shelter in the presence of fish than in their absence. Newts also significantly decreased their sexual activity level overall, but specially outside the shelter when they were in direct contact with fish. These results show that fish presence can affect newts in complex ways, such as through inhibition of their reproduction. Our work highlights that integrating behaviour in conservation studies is essential to understanding the patterns of coexistence and exclusion between introduced fish and amphibians.     

Estrogen receptor alpha is required in GABAergic,but not glutamatergic,neurons to masculinize behavior

Masculinization of the altricial rodent brain is driven by estrogen signaling during a perinatal critical period. Genetic deletion of estrogen receptor alpha (Esr1/ERα) results in altered hypothalamic-pituitary-gonadal (HPG) axis signaling and a dramatic reduction of male sexual and territorial behaviors. However, the role of ERα in masculinizing distinct classes of neurons remains unexplored. We deleted ERα in excitatory or inhibitory neurons using either a Vglut2 or Vgat driver and assessed male behaviors. We find that Vglut2-Cre;Esr1^lox/lox mutant males lack ERα in the ventrolateral region of the ventromedial hypothalamus (VMHvl) and posterior ventral portion of the medial amygdala (MePV). These mutants recapitulate the increased serum testosterone levels seen with constitutive ERα deletion, but have none of the behavioral deficits. In contrast, Vgat-Cre;Esr1^lox/lox males with substantial ERα deletion in inhibitory neurons, including those of the principal nucleus of the bed nucleus of the stria terminalis (BNSTpr), posterior dorsal MeA (MePD), and medial preoptic area (MPOA) have normal testosterone levels, but display alterations in mating and territorial behaviors. These mutants also show dysmasculinized expression of androgen receptor (AR) and estrogen receptor beta (Esr2). Our results demonstrate that ERα masculinizes GABAergic neurons that gate the display of male-typical behaviors.     

Territorial aggression does not feed back on testosterone in a multiple-brooded songbird species with breeding and non-breeding season territoriality,the European stonechat

Testosterone mediates reproductive behaviours in male vertebrates. For example, breeding season territoriality depends on testosterone in many species of birds and in some, territorial interactions feed back on testosterone concentrations. However, the degree to which territorial behaviour and testosterone are associated differs even between species with seemingly similar life histories, especially between species that also defend territories outside the breeding season. Here, we investigate the link between territorial behaviour and testosterone in European stonechats. Previous studies found that territorial aggression in stonechats depends on testosterone in a breeding, but not in a non-breeding context. We investigated whether stonechats show a rise in testosterone during simulated territorial intrusions (STI) during the breeding season. Post-capture testosterone concentrations of males caught after an STI were not higher than those of males caught in a control situation regardless of breeding stage. However, most of the males would have been able to mount a testosterone response because the same individuals that did not increase testosterone during the STI showed a substantial increase in testosterone after injections of gonadotropin-releasing hormone (GnRH). GnRH-induced and post-capture testosterone concentrations were positively correlated and both decreased with successive breeding stages. Further, territory owners with a short latency to attack the decoy expressed higher post-capture testosterone concentrations than males with a longer latency to attack the decoy. Thus, there is no evidence for behavioural feedback on testosterone concentrations during male-male interactions in stonechats. In combination with previous studies our data suggest that testosterone functions as an on/off switch of high intensity territorial aggression during the breeding season in stonechats. The among-species variation in the androgen control of territorial behaviour may be only partly a result of environmental differences. Instead, potential differences in how territoriality evolved in different species may have influenced whether and how a reproductive hormone such as testosterone was co-opted into the mechanistic control of territorial behaviour.     

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.     

Defence, intrusion and the evolutionary stability of territoriality

Territorial behaviour can only be adaptive if its costs are outweighed by its benefits. Territorial individuals incur costs by defending their territories against intruders. Usually these intruders are assumed to be non-territorial floaters attempting to take over the whole territory or neighbours trying to extend the borders of their own territory. We instead investigate how costs and benefits of territorial behaviour are affected by neighbours which invade to steal resources on a territory.We show analytically that in the absence of defence intrusion into neighbouring territories always pays and that even if territories are defended intrusion levels can still be high. Using a more detailed simulation model we find that territory defence usually disappears from the population even if owners have a strong advantage over intruders in terms of fighting costs or foraging efficiency. Defence and thus territoriality can only be evolutionarily stable if fighting costs for the intruder relative to the productivity of the territory are very high or if crossing the borders between territories carries additional costs.Our results show that stealing of resources by neighbours can have a considerable effect on the evolutionary stability of territory defence and thus territoriality itself. A more mechanistic model of territorial behaviour is needed to incorporate these kinds of mechanisms into a general theory on the evolution of territoriality.     

Manipulations of the AVT system shift social status and related courtship and aggressive behavior in the bluehead wrasse

Arginine vasotocin (AVT) and its mammalian homologoue arginine vasopressin (AVP) influence male sexual and aggressive behaviors in many species. We tested the effects of AVT and an AVP-V(1a) receptor antagonist on the display of alternative male tactics in a tropical coral reef fish, the bluehead wrasse Thalassoma bifasciatum. We gave AVT injections to territorial and nonterritorial males of the large and colorful phenotype (terminal phase) and an AVP-V(1a) receptor antagonist, Manning compound, to territorial males in the field. AVT increased courtship independent of status, while its effects on territoriality and aggression were dependent upon male status. In territorial males, AVT increased courtship and tended to decrease the number of chases toward initial phase individuals. In nonterritorial males, AVT increased courtship, chases toward initial phase individuals, and territorial behavior while decreasing feeding. These are all behaviors rarely seen in nonterritorial males, so AVT made these males act like territorial TP males. The AVP-V(1a) receptor antagonist had opposite effects. It decreased courtship and territorial defense, making these males act more like nonterritorial males. Manipulations of the AVT system shifted males within a single phenotype from the nonterritorial social status to the territorial social status and vice versa. Since the entire suite of behaviors related to territoriality was affected by AVT system manipulations, our results suggest that the AVT system may play a key role in motivation of behaviors related to mating.     

Evolution of territoriality in Hylinae treefrogs: Ecological and morphological correlates and lineage diversification

Given the diverse nature of traits involved in territorial defence, they may respond to different selective pressures and then exhibit distinct patterns of evolution. These selective pressures also may cause territorial behaviour to be associated with environmental and morphological variables. Such associations, however, have mostly been studied at the intraspecific level, being phylogenetic analyses of territoriality in a broad taxonomic framework rare in the literature. We used the anuran subfamily Hylinae to test (1) whether two territorial-behaviour traits with different levels of aggression—territorial call and physical combat—are evolutionarily more labile than a morphological trait used in physical combat—the spine-shaped prepollex; (2) whether reproduction in lentic waters and phytotelmata, as well as resource scarcity, might favour the occurrence of territoriality; (3) if physical combat is more important than territorial call for the evolution of body size and sexual size dimorphism and (4) the relationships between territorial-behaviour traits and lineage diversification. We mainly used the literature to build two datasets with different levels of certainty. Territorial-behaviour traits exhibited intermediate levels of phylogenetic signal in Hylinae, whereas the phylogenetic signal for the presence of the spine-shaped prepollex was strong. We found support for the hypothesis that reproduction in lentic water favours the occurrence of territorial behaviour, because the expression of territorial-behaviour traits was more associated with reproduction in lentic than in lotic waters. Territorial-behaviour traits were not correlated with annual precipitation nor with habitat complexity. Body size and sexual size dimorphism were not correlated with the presence of territorial call nor with physical combat. We identified negative correlations between diversification rates and physical combat. Relationships of territorial call and physical combat with diversification rates suggest that these territorial behaviours influence evolutionary processes in different ways.