The effects of previous experience and current condition on status contests in the bluebanded goby (<Emphasis Type="Italic">Lythrypnus dalli</Emphasis>)

Social status can change over a lifetime and affect fitness. Status transitions are often influenced by previous experience such that previous winners tend to dominate in future interactions, while losers tend to remain subordinate. We investigated the role of social status experience on future status contests in the bluebanded goby (Lythrypnus dalli), a highly social, sex-changing fish. Female L. dalli were given long-term experience as alpha (dominant) or beta (subordinate) in a stable social group of three fish (two females, one dominant male). We hypothesized that females with previous dominance experience would be more likely to become dominant in a novel context and change sex, a status-dependent transition. We found that in size-matched pairs, half of the previous alphas and half of the previous betas established and maintained dominance in the novel pair. Previous betas were more likely to become dominant in pairs with small relative differences in physical condition. Betas that initiated more interactions with the male in the stable group were also more likely to become dominant. Experience only had a transient effect on agonistic behavior during status establishment. In pairs in which the previous beta established dominance, individual agonistic efficiency, or the proportion of their approaches that led to a displacement, was lower. There was no long-term effect of experience on the rate of sex change. These data suggest that long-term tenure as a subordinate does not, itself, prevent status ascension and that physical condition and individual behavior are relevant factors that influence L. dalli contests.     

Ontogeny of food-caching site preferences in young Florida Scrub-Jays: evidence of learning or changing social status?

Florida Scrub-Jays (Aphelocoma coerulescens) are cooperative breeders endemic to Florida’s oak scrub. In autumn, Florida Scrub-Jays cache thousands of acorns and exhibit behaviors that appear to balance cache site selection against food degradation or cache robbery. However, both experience and position within a social dominance hierarchy could affect individual cache preferences. We examined the cache site preferences of birds with differing levels of caching experience and at different strata within a complex social dominance hierarchy. Our objective was to determine how experience, social position, and social context when caching influenced microhabitat preferences, and if these change as jays age, gain experience, and their social position changes. Naïve first-year birds preferred to cache in well-hidden, densely vegetated sites with relatively high soil moisture content. Naïve birds also cached farther from provisioning points if observed by a socially dominant bird than when they were alone or in the presence of birds of equal social status. Experienced adults preferred to cache in open, dry sandy sites and social context at the time of caching did not influence their preferences. As naïve birds aged, they gained experience and their social position changed. Experienced second-year birds shifted their preference to more open, drier sites, and did so more often when they remained subordinate within their group rather than becoming dominant breeders. Second-year birds that remained subordinate did not alter their caching behavior if observed by dominant birds. These patterns suggest that after gaining experience, jays learned which sites were more appropriate for caching and shifted their preference, regardless of their changing social status. We suggest that the risk of cache loss to food degradation might be greater than the risk of pilfering for Florida Scrub-Jays, especially for birds in any social strata except the most subordinate, but this requires additional study.     

Social context, competitive interactions and the dynamic nature of antipredator responses of juvenile rainbow trout Oncorhynchus mykiss

The effects of food ration and social context, as well as possible interactions, on the antipredator behaviour of juvenile rainbow trout Onchorhynchus mykiss were investigated in a pair of laboratory studies. In experiment 1, pairs of dominant and subordinate O. mykiss were exposed to conspecific alarm cues when maintained under high or low food rations. Under high food rations, dominant individuals responded to predation risk, whereas subordinates used the opportunity to feed. Under low food ration, however, the opposite pattern was observed, where subordinates responded to predation cues and dominants did not. Experiment 2 consisted of performing the same experiment, however separating the dominant and subordinate O. mykiss 3 h before testing. When tested separately, dominant and subordinate individuals did not differ in their responses to alarm cues, regardless of food ration. These results demonstrate that there is a complex interaction between current energy status and social context on decision making by prey animals.     

Jumping sans legs: does elastic energy storage by the vertebral column power terrestrial jumps in bony fishes?

Despite having no obvious anatomical modifications to facilitate movement over land, numerous small fishes from divergent teleost lineages make brief, voluntary terrestrial forays to escape poor aquatic conditions or to pursue terrestrial prey. Once stranded, these fishes produce a coordinated and effective “tail-flip” jumping behavior, wherein lateral flexion of the axial body into a C-shape, followed by contralateral flexion of the body axis, propels the fish into a ballistic flight-path that covers a distance of multiple body lengths. We ask: how do anatomical structures that evolved in one habitat generate effective movement in a novel habitat? Within this context, we hypothesized that the mechanical properties of the axial skeleton play a critical role in producing effective overland movement, and that tail-flip jumping species demonstrate enhanced elastic energy storage through increased body flexural stiffness or increased body curvature, relative to non-jumping species. To test this hypothesis, we derived a model to predict elastic recoil work from the morphology of the vertebral (neural and hemal) spines. From ground reaction force (GRF) measurements and high-speed video, we calculated elastic recoil work, flexural stiffness, and apparent material stiffness of the body for Micropterus salmoides (a non-jumper) and Kryptolebias marmoratus (adept tail-flip jumper). The model predicted no difference between the two species in work stored by the vertebral spines, and GRF data showed that they produce the same magnitude of mass-specific elastic recoil work. Surprisingly, non-jumper M. salmoides has a stiffer body than tail-flip jumper K. marmoratus. Many tail-flip jumping species possess enlarged, fused hypural bones that support the caudal peduncle, which suggests that the localized structures, rather than the entire axial skeleton, may explain differences in terrestrial performance.     

RESEARCH PAPER: Multiple Sexual Ornamentation Signals Male Quality and Predicts Female Preference in Minnows

Sexual ornamentation often consists of multiple components. Different sexual signals may indicate different aspects of mate quality or reflect quality in different time scales. On the other hand, same signals can have a dual function and are used both in male–male competition and courtship. Many fish species are capable of rapidly altering their colouration (ephemeral colour changes), but this capability is usually ignored in sexual selection studies. Here, we used experimentally manipulated social environments to study the ephemeral colour changes in multicomponent sexual signals of male minnows (Phoxinus phoxinus) during male–male competition and female choice. We found that the dominant males courted the females more actively and had redder and/or darker skin colouration than the subordinate males. Furthermore, darkness difference between subordinate and dominant males increased in the presence of female, which suggests that the male–male competition may increase the honesty of signalling and thus facilitate female choice. In support of this hypothesis, females had a strong behavioural preference towards the more colourful males, which may indicate female choice. As colourful males often had a higher social status than paler individuals, it is possible that females base their preference on male status, not only the colouration per se. In any case, our results suggest that sexual ornamentation of male minnows may signal status, courting activity and superior quality of the males and that these signals may have a dual function in both male–male competition and female choice. Females preferred different ornamental traits (dark and red colour patterns) relatively equally, indicating that mate choice is based on multiple cues.     

Paternity and dominance loss in male breeders: the cost of helpers in a cooperatively breeding mammal

Paternity insurance and dominance tenure length are two important components of male reproductive success, particularly in species where reproduction is highly skewed towards a few individuals. Identifying the factors affecting these two components is crucial to better understand the pattern of variation in reproductive success among males. In social species, the social context (i.e. group size and composition) is likely to influence the ability of males to secure dominance and to monopolize reproduction. Most studies have analyzed the factors affecting paternity insurance and dominance tenure separately. We use a long term data set on Alpine marmots to investigate the effect of the number of subordinate males on both paternity insurance and tenure of dominant males. We show that individuals which are unable to monopolize reproduction in their family groups in the presence of many subordinate males are likely to lose dominance the following year. We also report that dominant males lose body mass in the year they lose both paternity and dominance. Our results suggest that controlling many subordinate males is energetically costly for dominant males, and those unable to support this cost lose the control over both reproduction and dominance. A large number of subordinate males in social groups is therefore costly for dominant males in terms of fitness.     

Sexual behaviour in female pigs

This paper reviews the influence of social mechanisms on oestrus and sexual motivation in pigs. The social relations between the animals and the signals they send out can inhibit as well as encourage their social motivation. Social stimuli from both boars and other sows in oestrus tend to induce and synchronise oestrus and ovulation amongst sows. The courting behaviour of boars is also facilitated by social stimuli from other boars. However, when sows are kept under conditions where the social pressure is high, e.g. due to limited space and/or resources, the social stress experienced by particularly the subordinate individuals may inhibit sexual motivation during oestrus. To a large extent this effect seems to be mediated via specific fear reactions towards dominant individuals of sows that have lost many aggressive encounters. For example, fear reduces the sexual motivation during mating and during sexual interactions amongst sows within a group, and fear may thus inhibit their chances of reproductive success. Similarly, fear of humans caused by innate fearfulness or negative handling procedures reduces sexual motivation in the presence of the human handler even when sows are in standing oestrus.     

The cost of dominance: suppressing subordinate reproduction affects the reproductive success of dominant female banded mongooses

Social species show considerable variation in the extent to which dominant females suppress subordinate reproduction. Much of this variation may be influenced by the cost of active suppression to dominants, who may be selected to balance the need to maximize the resources available for their own offspring against the costs of interfering with subordinate reproduction. To date, the cost of reproductive suppression has received little attention, despite its potential to influence the outcome of conflict over the distribution of reproduction in social species. Here, we investigate possible costs of reproductive suppression in banded mongooses, where dominant females evict subordinates from their groups, thereby inducing subordinate abortion. We show that evicting subordinate females is associated with substantial costs to dominant females: pups born to females who evicted subordinates while pregnant were lighter than those born after undisturbed gestations; pups whose dependent period was disrupted by an eviction attained a lower weight at independence; and the proportion of a litter that survived to independence was reduced if there was an eviction during the dependent period. To our knowledge, this is the first empirical study indicating a possible cost to dominants in attempting to suppress subordinate breeding, and we argue that much of the variation in reproductive skew both within and between social species may be influenced by adaptive variation in the effort invested in suppression by dominants.     

Social context influences aggressive and courtship behavior in a cichlid fish

Social interactions require knowledge of the environment and status of others, which can be acquired indirectly by observing the behavior of others. When being observed, animals can also alter their signals based on who is watching. Here we observed how male cichlid fish (Astatotilapia burtoni) behave when being watched in two different contexts. In the first, we show that aggressive and courtship behaviors displayed by subordinate males depends critically on whether dominant males can see them, and in the second, we manipulated who was watching aggressive interactions and showed that dominant males will change their behavior depending on audience composition. In both cases, when a more dominant individual is out of view and the audience consists of more subordinate individuals, those males signal key social information to females by displaying courtship and dominant behaviors. In contrast, when a dominant male is present, males cease both aggression and courtship. These data suggest that males are keenly aware of their social environment and modulate their aggressive and courtship behaviors strategically for reproductive and social advantage.     

Intermale aggression of subordinate resident long-evans rats

Dominant male rats were separated from their colonies and subordinate residents were tested for aggression after a two week period. Subordinate animals showed significant increases in body weight and aggressive behavior toward intruders. Replacement of the previously dominant male led to rank reversals in three of six colonies. To assess whether two weeks of separation from the dominant male was necessary for subordinate males to exhibit aggression towards conspecifics, another intruder test was given following an additional two weeks of social housing. At the conclusion of this test, the more aggressive resident was removed and a naive intruder was immediately introduced into the colony with the subordinate male resident and again at 1, 5, and 10 day intervals. Aggressive posturing by the subordinate resident increased immediately following the removal of the dominant male, and remained at a high level throughout the subsequent intruder tests. In contrast, biting was initially low but increased steadily throughout the period of separation. These findings demonstrated that social or situational changes within a colony can greatly influence the aggression of subordinate males.     

Reproductive skew can provide a net advantage in both conditional and unconditional social interactions

We revisit a model for the evolution of costly social behaviour in the presence of reproductive skew. The model population is structured into groups, and reproductive skew is captured by assuming individuals adopt one of two social roles (dominant/subordinate). Unlike previous work, we adopt an ultimate perspective by tracking a mutant allele over the entire course of an invasion. Our main analysis applies the theory of branching processes, but a parallel analysis using the inclusive-fitness approach is also provided. Our first two results are modifications of known inequalities describing selective advantages for behaviours expressed conditional upon social status. We find that altruistic subordinate individuals are favoured more readily than previously thought; spiteful dominant individuals, however, are favoured less readily. Secondly, we identify the condition under which unconditional altruism (performed by both dominant and subordinate) will be adaptive. Our third main result shows that increasing the strength of selection can also change the range of parameters over which costly social behaviours are favoured. We find that stronger selection makes it relatively easier for subordinate altruism to emerge, but more difficult for dominant spite and unconditional altruism to occur. We discuss the possible implications of our results for human social evolution.     

Food availability and competition do not modulate the costs of Plasmodium infection in dominant male canaries

Understanding the different factors that may influence parasite virulence is of fundamental interest to ecologists and evolutionary biologists. It has recently been demonstrated that parasite virulence may occur partly through manipulation of host competitive ability. Differences in competitive ability associated with the social status (dominant or subordinate) of a host may determine the extent of this competition-mediated parasite virulence. We proposed that differences between subordinate and dominant birds in the physiological costs of infection may change depending on the level of competition in social groups. We observed flocks of domestic canaries to determine dominant or subordinate birds, and modified competition by providing restricted (high competition) or ad libitum food (low competition). Entire flocks were then infected with either the avian malaria parasite, Plasmodium relictum or a control. Contrary to our predictions we found that the level of competition had no effect on the outcome of infection for dominant or subordinate birds. We found that dominant birds appeared to suffer greater infection mediated morbidity in both dietary treatments, with a higher and more sustained reduction in haematocrit, and higher parasitaemia, than subordinates. Our results show that dominance status in birds can certainly alter parasite virulence, though the links between food availability, competition, nutrition and virulence are likely to be complex and multifaceted.     

Sperm competition within a dominance hierarchy: investment in social status vs. investment in ejaculates

There is increasing recognition that male-male competition can take many forms, but as yet the form is not predictable a priori. Many recent studies have focused attention on how males in disadvantaged mating roles compensate through sperm competition. However, mating systems in which subordinate males are reproductively suppressed, particularly through the stress of social interactions, may limit the ability of males to respond by increasing investment in sperm quality. We examined the interaction between social status and ejaculate tactics in Nauphoeta cinerea, a cockroach that has a mating system with well-characterized dominance hierarchies. Both social experience with other males and social status influenced aspects of ejaculates. The stress of social interactions reduced the size of the ejaculate and number of sperm inseminated. In ejaculates formed prior to social experience, however, males that go on to become dominant inseminated more sperm than males that go on to become subordinate, suggesting innate differences among males. Our results show that though selection for increased success in sperm competition for subordinate males in a hierarchy can occur, both the traits and the way in which the balance between pre- and post-copulatory strategies is negotiated will depend on specific details of the mating system. These details will include how the physiological effects of social interactions may limit selection through male-male competition.     

Plasticity in reproductive phenotypes reveals status-specific correlations between behavioral, morphological, and physiological sexual traits

Reproductive success is determined by a complex interplay between multiple sexual traits that promote mate acquisition and, following copulation, provide control over paternity. The intensity of sexual competition that individuals experience often fluctuates, and here we investigate how this influences the expression of reproductive traits and their relationships. We show in the fowl, Gallus gallus, that males of different social status, which experience different intensities of sexual competition, before and after copulation, have different reproductive phenotypes. Dominant males are more vigilant, feed less, and have larger sexual ornaments than subordinate males. Experimentally manipulating social status revealed that these differences were phenotypically plastic, indicating multiple sexual traits were dependent on the social environment. We integrated these data with previous published findings on changes in sperm numbers and velocity to show that relationships between traits were different for males when they were dominant and when they were subordinate. Furthermore, when males switched status a complex array of negative and positive correlations between the degree traits changed was observed. Our results suggest that variation in the intensity of sexual competition generates reversible plasticity in reproductive phenotypes and that relationships between sexual traits may be variable and influence the evolution of reproductive strategies.     

Modulation of social behavior and dominance status by chronic pain in mice

The potential influence of pain on social behavior in laboratory animals has rarely been evaluated. Using a new assay of social behavior, the tube co‐occupancy test (TCOT), we assess propinquity—the tendency to maintain close physical proximity—in mice exposed to pain using subcutaneous zymosan or spared nerve injury as noxious stimuli. Our previous experience with the TCOT showed that outbred mouse sibling dyads show higher levels of tube co‐occupancy than stranger dyads. We find here that long‐lasting pain from spared nerve injury given to both mice in the dyad abolishes this effect of familiarity, such that strangers also display high levels of propinquity. We performed a separate experiment to assess the effect on dominance behavior of nerve injury to one or both mice of a dyad in which relative dominance status had been previously established via the confrontation tube test. We find that neuropathic pain given only to the dominant mouse reverses the relationship in male but not female mice, such that the previously subordinate mouse becomes dominant. These observations bolster the scant but growing evidence that pain can robustly affect social behavior in animals.     

The effects of dominance on leadership and energetic gain: a dynamic game between pairs of social foragers

Although social behaviour can bring many benefits to an individual, there are also costs that may be incurred whenever the members of a social group interact. The formation of dominance hierarchies could offer a means of reducing some of the costs of social interaction, but individuals within the hierarchy may end up paying differing costs dependent upon their position within the hierarchy. These differing interaction costs may therefore influence the behaviour of the group, as subordinate individuals may experience very different benefits and costs to dominants when the group is conducting a given behaviour. Here, a state-dependent dynamic game is described which considers a pair of social foragers where there is a set dominance relationship within the pair. The model considers the case where the subordinate member of the pair pays an interference cost when it and the dominant individual conduct specific pairs of behaviours together. The model demonstrates that if the subordinate individual pays these energetic costs when it interacts with the dominant individual, this has effects upon the behaviour of both subordinate and the dominant individuals. Including interaction costs increases the amount of foraging behaviour both individuals conduct, with the behaviour of the pair being driven by the subordinate individual. The subordinate will tend to be the lighter individual for longer periods of time when interaction costs are imposed. This supports earlier suggestions that lighter individuals should act as the decision-maker within the pair, giving leadership-like behaviours that are based upon energetic state. Pre-existing properties of individuals such as their dominance will be less important for determining which individual makes the decisions for the pair. This suggests that, even with strict behavioural hierarchies, identifying which individual is the dominant one is not sufficient for identifying which one is the leader.     

Reproductive suppression in female Alpine marmots, Marmota marmota

We studied mechanisms responsible for reproductive skew in female Alpine marmots. In each social group, only dominant females produced weaned young although subordinate females had similar body condition, were fertile as indicated by high levels of oestradiol during the mating season, and occasionally started pregnancies. During the period of gestation, dominant females initiated significantly more agonistic interactions against subordinate females, resulting in significantly increased levels of glucocorticoids and decreased levels of progesterone in subordinates. Results suggested that reproductive suppression in female Alpine marmots is mediated by the negative effects of stress (glucocorticoids) on the activity of the hypothalamic-pituitary-gonadal axis. The strength of competition between subordinate and dominant females was affected by their relatedness. Dominant females attacked unrelated subordinate females more, whereas amicable behaviour was observed mainly between dominant females and their daughters. These differences could be explained by differences in indirect fitness: related subordinate females benefited from warming the offspring of the dominant female during hibernation. Reproductive suppression was apparently costly for dominant females, because their reproductive success decreased as the number of adult subordinate females in a group increased. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Group composition affects male reproductive partitioning in a cooperatively breeding cichlid

Individuals within groups of cooperatively breeding species may partition reproduction, with the dominant pair often taking the largest share. The dominant's ability to reproductively control subordinates may depend on differences in competitive ability, due to, e.g. body size differences, but may also depend on the number of same‐sex competitors inside the group. We tested experimentally whether subordinates reproduce more when these subordinates are large or when a second subordinate of the same sex need to be controlled by the dominants, using the cooperatively breeding cichlid Neolamprologus pulcher. Dominant pairs were assisted by a large and a small unrelated subordinate; sexes of these fish were varied in a full‐factorial design (giving four treatments). Dominant males lost significantly more parentage to the large subordinate male when a small subordinate male was also present, compared to when a small subordinate female was present. However, subordinate paternity was generally low and did not significantly curb total dominant male reproductive output, which was more affected by the sizes and numbers of reproductive females present inside his group. Dominant female maternity, clutch sizes and total output did not depend on the treatments. Subordinate–subordinate reproduction was virtually absent (one out of 874 offspring). Female subordinates were more likely to provide care for their own broods. In contrast, male subordinates did not adjust their level of care to their parentage. Variability in female subordinate alloparental brood care was particularly high, with females showing more care than males in general. We also detected effects of growth rate and food ration on parentage independent of the treatments, most notably: (i) a trade‐off between dominant male growth rate and paternity; (ii) a decrease in dominant male paternity with increasing food ration; (iii) a positive effect of growth rate on paternity in small males. We conclude that dominant males should be sensitive to the number and sizes of subordinate males present in their group, particularly when these subordinates are not helpful or grow fast, and food is plentiful. Dominant females should be less sensitive, because female subordinates do not appear to impose reproductive costs and can be helpful through alloparental brood care.     

Influence of social status on individual foraging and community structure in a bird guild

We investigated the influence of social interactions on individual foraging behavior and community structure of frugivorous birds in southern Costa Rica. Detailed observations of large, heterospecific feeding assemblages at fruiting trees revealed the existence of an interspecific dominance hierarchy, largely consistent with body and bill size. Social status influenced access to food in several ways. First, subordinate species were interrupted more and tended to have shorter foraging bouts than dominant species (n > 1.000 abouts). Second, analysis of over 7,000 videotaped head movements showed that subordinate species spent a smaller fraction of their foraging bouts actually feeding (as opposed to looking about) than did dominants. Third, when many birds were in a tree simultaneously, the foraging bouts of subordinate species were shortened; this effect was less pronounced or absent for species higher in the dominance hierarchy. Fourth, subordinate species foraged less frequently in mixed-species assemblages than did dominant species. Finally, subordinate species fed disproportionately more in the late afternoon at fruiting trees. The influence of social status appeared to manifest itself at the community level. The species composition of foraging assemblages was compared at isolated fruiting trees situated in an agricultural landscape near to (< 0.5 km) and far from (> 6 km) a large tract of primary forest. Whereas the full complement of avian frugivores foraged at the near trees, visitors to the far trees were predominantly of high social status. We discuss reasons why high social status and associated traits might confer an advantage in exploiting human-dominated habitat.     

Canopy structure and nitrogen distribution in dominant and subordinate plants in a dense stand of Amaranthus dubius L. with a size hierarchy of individuals

The objective was to investigate how nitrogen allocation patterns in plants are affected by their vertical position in the vegetation (i.e. being either dominant or subordinate). A garden experiment was carried out with Amaranthus dubius L., grown from seed, in dense stands in which a size hierarchy of nearly equally aged individuals had developed. A small number of dominant plants had most of their leaf area in the highest layers of the canopy while a larger number of subordinate plants grew in the shade of their dominant neighbours. Canopy structure, vertical patterns of leaf nitrogen distribution and leaf photosynthetic characteristics were determined in both dominant and subordinate plants. The light distribution in the stands was also measured. Average N contents per unit leaf area (total canopy nitrogen divided by the total leaf area) were higher in the dominant than in the subordinate plants and this was explained by the higher average MPA (leaf dry mass per unit area) of the dominant plants. However, when expressed on a weight basis, average N contents (LNC_av; total canopy N divided by the total dry weight of leaves) were higher in the subordinate plants. It is possible that these higher LNC_av values reflect an imbalance between carbon and nitrogen assimilation with N uptake exceeding its metabolic requirement. Leaf N content per unit area decreased more strongly with decreasing relative photon flux density in the dominant than in the subordinate plants showing that this distribution pattern can be different for plants which occupy different positions in the light gradient in the canopy. The amount of N which is reallocated from the oldest to the younger, more illuminated leaves higher up in the vegetation may depend on the sink strength of the younger leaves for nitrogen. In the subordinate plants, constrained photosynthetic activity caused by shading might have reduced the sink intensity of these leaves.