The Effect of Social Hierarchy on Captive Coyote (Canis latrans) Foraging Behavior

Foraging efficiency of individuals in pack forming species may be influenced by social dynamics within a pack. The effects of social hierarchy in particular may influence individual foraging behavior in canids, such as coyotes (Canis latrans). To examine the impact of social hierarchy on foraging behavior, we tested 16 captive coyotes in eight naturally established dominant–subordinate pairs, using the guesser–knower paradigm. We measured the efficiency of subordinate coyotes to relocate a food resource when alone and then allowed pairs to forage together, such that subordinates had prior knowledge of food location but dominants did not. To determine whether (1) subordinates used a direct or discursive strategy to obtain food in the presence of a dominant and (2) dominants used an exploitative or independent strategy to obtain food in the presence of a subordinate with previous knowledge, we measured their search efficiency (e.g., correct choice of area, feeder, and latency to correct feeder). Results showed subordinates learned to relocate food and increase efficiency when alone. In a social context, however, subordinate efficiency decreased. That is, subordinates approached the correct area, but searched more feeders before finding the correct one. Dominants initially used an independent search strategy but then quickly displaced the subordinate and monopolized the resource, reducing subordinate efficiency further. Despite continual displacement and reduction in efficiency, subordinates did not alter their foraging strategy over time. Our results suggest prior information can improve individual foraging advantage, but that social status strongly impacts individual foraging efficiency in social species such as coyotes.     

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.     

Social Status and Feeding in Atlantic Salmon Salmo salar Parr: The Effect of Visual Exposure to a Dominant

Groups of four Atlantic salmon parr were established and the dominant individual identified. Feeding responses were then observed in the remaining (subordinate) fish housed in one of three conditions: in the same compartment as the dominant fish (full exposure), separated from it by a transparent partition (visual exposure) or behind an opaque partition (isolated). Subordinate parr in the full exposure condition responded to and ate fewer food pellets than did the dominant fish. However, subordinate parr in the visual exposure condition and in the isolation condition fed actively, indicating that neither visual intimidation nor long-term stress-induced reduction of appetite were responsible for the reduced food intake of subordinate parr in direct contact with dominants. Dominant parr and subordinates in the visual exposure condition, but not subordinates in the exposed condition, moved further from their feeding station when attacking food items that were contested by a competitor compared with the distance moved for uncontested items. The presence of a competitor therefore forced both dominant and visually exposed subordinates to pay higher costs while foraging.     

Growth depensation and aggression in laboratory reared coho salmon: the effect of food distribution and ration size

Groups of recently emerged coho salmon fry Oncorhynchus kisutch were reared for 3 months on food that appeared either asynchronously at a single location (localized) or synchronously and spatially dispersed (dispersed). Groups were further subdivided into those receiving low (1%) or high (3% body weight per day) rations, with five replicate groups for each treatment combination. At low ration there was greater growth depensation, i.e. growth variation, in groups receiving localized as compared to dispersed food. At high ration there was no difference. There was no effect of food distribution upon mean fish weight, but groups receiving high rations had greater mean fish weights than groups receiving low rations. There was no overall difference in the frequency of chasing between any of the treatment combinations. However, in localized food groups, dominants defended positions close to where food entered the tank, giving them greater access than subordinates. In dispersed food groups, while dominants also defended particular areas, this did not result in greater access to food. These results demonstrate that although feeding methodology may not directly influence the frequency of aggressive interactions, feeding methods which facilitate food monopolization by dominants can accelerate the growth of these individuals at the expense of subordinates. In aquacultural applications where greater size is desirable, or otherwise selected for, this may result in the unintentional selection for increased aggressiveness.     

Predator-induced changes in morphology of a prey fish: the effects of food level and temporal frequency of predation risk

In a series of experiments, we investigated the effects of food availability and risk frequency on the dynamics of predator-induced changes in growth and morphology of prey fish using goldfish (Carassius auratus) as our test species. In experiment 1, we fed goldfish high or low food rations and exposed them to either alarm cues from conspecifics, cues from swordtails or a water control. After 60 days, goldfish in the alarm cue treatment significantly increased their body depth and body weight but had smaller body length than goldfish exposed to swordtails cues or water, likely reducing their vulnerability to gape-limited predators. Importantly, food level had an impact on the amplitude of the morphological changes. In experiment 2, goldfish were exposed to two different frequencies of predation cues or a water control for 50 days. The cues were either continued or discontinued from day 51 to 100, and all cues were resumed from day 101 to 150. We found that goldfish exposed to predation cues increased their depth and weight at a faster rate than did the goldfish exposed to water, and of particular significance was the fact that frequency of risk had an effect on the amplitude of the change. When the cues were interrupted, the increase in growth rate parameters was reduced to the level of the goldfish exposed to water. However, when the cues were resumed, the rate increased to match the growth rate of the goldfish that were continuously exposed to the cues. Finally, we staged encounters between goldfish of differing morphologies and yellow perch (Perca flavescens) and found that deep-bodied goldfish had better survival than the shallow-bodied ones. These experiments illustrate the dynamic nature of inducible morphological defences.     

The effect of ration level and social rank on the development of fin damage in juvenile rainbow trout

The development and severity of fin damage was examined in groups of juvenile rainbow trout Oncorhynchus mykiss of different strength of feeding hierarchies. The development of dorsal and caudal fin damage over time was compared between four groups fed different ration levels (0·25, 0·5, 1·0 and 1·5% body weight day⁻¹) and between individuals of different feeding rank within each group. Dominance hierarchies were assessed from repeated daily measurements of food consumption of individuals using radiography. The feeding and growth data indicated that the strength of the social hierarchy weakened with increasing ration. Caudal fin damage developed with time in all groups whereas dorsal fin damage developed only under limited rations. The severity of both dorsal and caudal fin damage was significantly dependent on the ration size fed to the group, with lower ration groups sustaining more fin damage. The severity of dorsal fin erosion was greater than for the caudal fin. Within the two lower ration groups, subordinate fish suffered the most dorsal fin damage. The results suggested that the severity of dorsal fin damage within groups of juvenile rainbow trout can be used as an indicator of hierarchy Strength.     

Do capuchin monkeys, Cebus apella, know what conspecifics do and do not see?

Capuchin monkeys were tested in five experiments in which two individuals competed over food. When given a choice between retrieving a piece of food that was visible or hidden from the dominant, subordinate animals preferred to retrieve hidden food. This preference is consistent with the hypotheses that either (1) the subordinate knew what the dominant could and could not see or (2) the subordinate was monitoring the behaviour of the dominant and avoiding the piece of food that it approached. To test between these alternatives, we released subordinates with a slight head start forcing them to make their choice (between a piece of food hidden or visible to the dominant) before the dominant entered the area. Unlike chimpanzees, Pan troglodytes, subordinates that were given a head start did not preferentially approach hidden pieces of food first. Therefore, our experiments provide little support for the hypothesis that capuchin monkeys are sensitive to what another individual does or does not see. We compare our results with those obtained with chimpanzees in the same paradigm and discuss the evolution of primate social cognition. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Dominance and neophobia in coyote (Canis latrans) breeding pairs

Previous research suggests that within social animals, subordinate individuals are less neophobic than dominant individuals. We investigated the effect of social status on neophobic responses using 10 captive coyote breeding pairs. Social status was determined from observations of feeding behavior and agonistic interactions during a series of reference trials. Once dominance was established, we surrounded an experimental area with novel stimuli (ropes adorned with interspersed flags) to create a novel context around a familiar food source. Contrary to hypotheses, dominant coyotes were first to feed, showed more interest toward novel stimuli, and eventually crossed the barrier, which subordinates never did. Our results indicate that dominant coyotes are less neophobic of novel settings that contain familiar food than subordinates are. Since a reduction in neophobia can be interpreted as an increase in risk taking, our results support previous observations that dominant (alpha) coyotes take more risks than subordinates. Our results also suggest reasons for differential observations of coyote behavior in the field: artificial selection against bold behavior in populations undergoing predator control.     

Do chimpanzees know what conspecifics know?

We conducted three experiments on social problem solving by chimpanzees, Pan troglodytes. In each experiment a subordinate and a dominant individual competed for food, which was placed in various ways on the subordinate's side of two opaque barriers. In some conditions dominants had not seen the food hidden, or food they had seen hidden was moved elsewhere when they were not watching (whereas in control conditions they saw the food being hidden or moved). At the same time, subordinates always saw the entire baiting procedure and could monitor the visual access of their dominant competitor as well. If subordinates were sensitive to what dominants did or did not see during baiting, they should have preferentially approached and retrieved the food that dominants had not seen hidden or moved. This is what they did in experiment 1 when dominants were either uninformed or misinformed about the food's location. In experiment 2 subordinates recognized, and adjusted their behaviour accordingly, when the dominant individual who witnessed the hiding was replaced with another dominant individual who had not witnessed it, thus demonstrating their ability to keep track of precisely who has witnessed what. In experiment 3 subordinates did not choose consistently between two pieces of hidden food, one of which dominants had seen hidden and one of which they had not seen hidden. However, their failure in this experiment was likely to be due to the changed nature of the competition under these circumstances and not to a failure of social-cognitive skills. These findings suggest that at least in some situations (i.e. competition with conspecifics) chimpanzees know what conspecifics have and have not seen (do and do not know), and that they use this information to devise effective social-cognitive strategies. Copyright 2001 The Association for the Study of Animal Behaviour.     

Olfactory cues in discrimination among individuals in dominance hierarchies in the cockroach, Nauphoeta cinerea

ABSTRACT. The purpose of this study was to explore the olfactory recognition mechanisms used by individual subordinate male cockroaches, Nauphoeta cinerea (Olivier), in a dominance hierarchy. Results of two independent sets of experiments, one using an olfactometer and the other using an arena, suggest that olfactory cues are important to males. Subordinate males did not avoid dominant male odour in an olfactometer but could discriminate between the odours of familiar and unfamiliar individual dominant males, preferring the odour of the dominant male with which they had previous social contact. Since test subordinates did not avoid dominant male odours in the olfactometer, it is possible that the distance an individual remains from a dominant is important. To test this, individual dominant and subordinate males were tethered in the centre of an arena. Subordinate males were introduced into the arena and allowed to approach the tethered male. They approached tethered dominants and subordinates differently, and tended to keep away from the dominants. Tethered males unfamiliar to the test animals were avoided as well.     

Chimpanzees know what conspecifics do and do not see

We report a series of experiments on social problem solving in chimpanzees, Pan troglodytes. In each experiment a subordinate and a dominant individual were put into competition over two pieces of food. In all experiments dominants obtained virtually all of the foods to which they had good visual and physical access. However, subordinates were successful quite often in three situations in which they had better visual access to the food than the dominant, for example, when the food was positioned so that only the subordinate (and not the dominant) could see it. In some cases, the subordinate might have been monitoring the behaviour of the dominant directly and simply avoided the food that the dominant was moving towards (which just happened to be the one it could see). In other cases, however, we ruled out this possibility by giving subordinates a small headstart and forcing them to make their choice (to go to the food that both competitors could see, or the food that only they could see) before the dominant was released into the area. Together with other recent studies, the present investigation suggests that chimpanzees know what conspecifics can and cannot see, and, furthermore, that they use this knowledge to devise effective social-cognitive strategies in naturally occurring food competition situations. Copyright 2000 The Association for the Study of Animal Behaviour.     

The building-up of social relationships: behavioural types, social networks and cooperative breeding in a cichlid

Consistent individual differences in behavioural types may not only cause variation in life-history decisions, but may also affect the choice of social partners and sociality in general. Here, we tested whether and how behavioural type influences the establishment of social ties using the cooperatively breeding cichlid, Neolamprologus pulcher. In a habitat saturation experiment with individuals pre-tested for behavioural type, we first analysed whether behavioural type affected the likelihood of settlement (i.e. social status), group sizes, and the types of dominant and subordinate individuals accepted as group members. Corrected for effects of body size and sex, the behavioural type did not affect settlement. However, bold dominant males only accepted smaller females, and grouped with bold subordinates, while shy dominant males accepted larger females than themselves, and grouped with shy subordinates. Second, we analysed the relationships between behavioural type and the aggressiveness or affiliation social network. Behavioural type significantly affected the number and quality of connections within the two networks. We show that behavioural types affect group composition, social networks and status achieved, in interaction with body size. Thus, the interactions within groups may depend not only on age, size and sex, but also on the behavioural type of the individuals involved.     

The effect of physical condition and shoalmate familiarity on proliferation of alarm substance cells in the epidermis of fathead minnows

Ostariophysan fishes have specialized epidermal alarm substance cells (ASCs). In this study, metabolic cost to the production and maintenance of these cells was demonstrated. Fathead minnows Pimephales promelas were maintained on high or low food rations and, to test for the effect of social context on the number of ASCs, they were held with either zero or two shoalmates. Shoalmates were familiar (from the focal fish's shoal), or unfamiliar (from a shoal separated by 1 km from the focal fish's shoal). After 16 days, epidermal thickness, number of mucus cells and number of ASCs were all significantly greater for minnows on the high ration than for those on the low ration. Within the high ration group, physical condition was positively correlated with epidermal thickness and ASC number. The shoalmate treatment had no measurable effect on the epidermis. It was concluded that food availability determines investment into ASCs, inferring a trade-off between the cost of ASCs and the fitness benefits they accrue.     

The effect of feeding hierarchy on individual variability in daily feeding of rainbow trout, Oncorhynchus mykiss (Walbaum)

Three groups of rainbow trout, Oncorhynchus mykiss , (initial weight 40 g) were fed one of three rations (low, medium or high) for 73 days. Consumption by individual fish within the three ration groups was measured on four occasions (days 27, 55, 64 and 72) using radiography. Food intake by individual fish varied between days and this variability was expressed using the coefficient of variation (CV). Dominant fish within each ration group were defined as individuals with the greater share of the group meal and these fish had low CVs for food intake indicating relatively little variation in daily consumption. By contrast, the fish which had consumed a low mean proportion of the group meal displayed high CVs for food intake, indicating considerable variability in the sizes of individual meals consumed by these fish. As group ration increased, the range of meal sizes and individual CVs in daily feeding decreased, suggesting that the strength of the feeding hierarchy and the variability in individual consumption decreased as food availability increased. It is suggested that radiography can be used to assess social relationships within groups of fish allowing the assessment of feeding hierarchies in larger groups of fish than would be possible by observational techniques.     

Perception of alarm cues influences the outcome of shelter competition in crayfish20

The effects of predators on prey populations may significantly alter many aspects of prey biology, including spatial distribution, foraging activities, and social interactions. In aquatic habitats, chemosensation is an important mode of communication and has been shown for many taxa, including crayfish, to be used in detection of predator and/or conspecific alarm cues. Here, we report on an experiment to test the hypothesis that detection of alarm cues results in greater individual investment in contests over shelters. We tested this hypothesis through dyadic contests between sex- and size-matched, non-reproductive individuals of Faxonius virilis. We found that crayfish responded to exposure to alarm cues by spending more time inside a shelter. We also report that in contests between pairs in which one crayfish had been exposed to alarm cues and the other had not, exposed individuals were significantly more likely to win ownership of a single shelter. However, we did not detect any differences in the contest parameters we recorded between exposed and unexposed crayfish. These impacts on both individual and social behavior indicate that the presence of predators is likely to have large effects on the distribution and structure of crayfish populations.     

Fattening strategies of wintering great tits support the optimal body mass hypothesis under conditions of extremely low ambient temperature

1.  In a seasonal environment, subcutaneous energy reserves of resident animals often increase in winter and decline again in summer reflecting gradual seasonal changes in their fattening strategies. We studied changes in body reserves of wintering great tits in relation to their dominance status under two contrasting temperature regimes to see whether individuals are capable of optimizing their body mass even under extreme environmental conditions.
2.  We predicted that dominant individuals will carry a lesser amount of body reserves than subordinate great tits under mild conditions and that the body reserves of the same dominant individuals will increase and exceed the amount of reserves of subordinates under conditions of extremely low ambient temperatures, when ambient temperature dropped down to −37 °C.
3.  The results confirmed the predictions showing that dominant great tits responded to the rising risk of starvation under low temperatures by increasing their body reserves and this was done at the expense of their safety.
4.  Removal experiments revealed that lower body reserves of subordinate flock members are due to the increased intraspecific competition for food under low ambient temperatures.
5.  Our results also showed that fattening strategies of great tits may change much quicker than previously considered, reflecting an adaptive role of winter fattening which is sensitive to changes in ambient temperatures.     

Seizing the Opportunity: Subordinate Male Fowl Respond Rapidly to Variation in Social Context

Dominance affects mating and reproductive success in many group-living species. Potential mechanisms include subordinates being inherently less attractive and social constraints imposed by dominant individuals. To test the former possibility, we measured morphology in 45 male fowl, Gallus gallus , prior to group formation. Males were then assigned to social groups (three males and three females in each). None of the measured traits predicted subsequent social status, suggesting that subordinates were not inherently unattractive. We then manipulated social constraints in each group to test if subordinates were socially constrained. We removed either the alpha (experimental) or the gamma male (control) for 40 min and observed the effect on the beta male's reproductive behavior. Controls accounted for putative group size and disturbance effects, and ensured that the only difference between treatments was the relative dominance of the remaining male. In each trial, we measured the beta male's courtship effort and his mating success, as well as his proximity to females and to the remaining male. Results show that social context did not affect mating success, but had a significant impact on courtship behavior. Beta males courted significantly more often when they had exclusive access to a female, as opposed to when another male was nearby. Furthermore, their courtship effort was higher if the nearby male was a fellow subordinate, as opposed to the dominant male. We conclude that both the proximity and social status of nearby males affects, either directly or indirectly, the courtship efforts of subordinate male fowl.     

Damage released prey alarm substances or predator odours? Risk assessment by an aquatic oligochaete

Although the abilities of prey to detect and respond to chemical substances associated with a predator have been widely reported, the factors promoting the evolution of responses to prey alarm cues vs. predator odours are still vague. In this article, we combined field research with laboratory experiments to explore which chemical substance associated with predator activity (predator odour, conspecific or heterospecific alarm substances) induces defence responses in the aquatic oligochaete Stylaria lacustris, which is vulnerable to common littoral predators. The field results indicated that predators injure the oligochaetes and a great proportion, up to 45% of individuals in the population, were found to be damaged. The results of the laboratory experiments revealed that chemical odours from damselfly larvae feeding on S. lacustris did not induce the defence response in the oligochaetes. On the contrary, oligochaetes detected and responded to alarm substances from damaged conspecifics alone and substances from damaged cladoceran Daphnia magna. We discussed conditions favouring the responses to damage released prey alarm cues instead of predator odours in Stylaria lacustris. Our data suggest that the selection of responses to alarm cues from damaged prey vs. predator odours may be dependent on three factors: (1) non-species-specific predation, (2) divergence of food niche of the different stages of the predator and (3) complex food web with multiple predators. Handling editor: S. Declerk     

Changes in sperm quality and numbers in response to experimental manipulation of male social status and female attractiveness

In promiscuous species, male reproductive success is determined by the interaction between the ability to access and choose females of the highest reproductive quality and, after copulation, the ability to outcompete the ejaculates of rival males. Disentangling the factors regulating the interplay between traits conferring a reproductive advantage before and after copulation is therefore crucial to understanding how sexual strategies evolve. Here we show in the fowl Gallus gallus, where social status determines copulation success, that dominant males produce more sperm than subordinates but that the quality of dominant males' sperm decreases over successive copulations, whereas that of subordinates remains constant. Experimentally manipulating male social status confirmed that ejaculate quality (the number and quality of sperm produced) was a response to the social environment rather than the result of intrinsic differences between dominant and subordinate males. We further show that dominant males responded to variation in female sexual ornamentation, which signals reproductive quality, by adjusting the number and quality of sperm they transferred, whereas subordinate males did not: they transferred ejaculates of similar quality to females with different ornament sizes. These results indicate that trade-offs between traits influencing reproductive success before and after copulation, combined with variation in social dynamics and female quality, may favor the evolution of phenotypically plastic alternative reproductive strategies.     

The number of subordinates moderates intrasexual competition among males in cooperatively breeding meerkats

For dominant individuals in cooperatively breeding species, the presence of subordinates is associated with both benefits (i.e. increased reproductive output and other group-living benefits) and costs (i.e. intrasexual competition on reproduction). The biological market theory predicts that dominant individuals are tolerant to same-sex group members when there are only a few subordinates, so as to maximize their own reproductive success. We investigated factors affecting aggression by dominant males and submission by subordinate males for a cooperatively breeding mammal, meerkats, Suricata suricatta. In this species, reproductive conflict occurs between the dominant male and the non-offspring males. As predicted, the number of subordinates in a group was positively associated with the aggression frequency by the dominant male and with the submission frequency by the subordinate males. Relative to the aggression frequency against male offspring, the frequency of aggression against non-offspring males was comparable in small groups, but was higher in large groups. These results indicate that reproductive conflict is present between the dominant male and the non-offspring males but is moderated in groups with small numbers of subordinates. This study provides an empirical data agreeing with the biological market theory in the context of intrasexual competition in cooperatively breeding species.