Deferred agonistic behavior in a long-lived scincid lizard Eumeces laticeps

Summary A laboratory experiment with the broad-headed skink (Eumeces laticeps) involving staged agonistic encounters demonstrates that larger males have an advantage over smaller ones in agonistic bouts. Field data on head wounds produced by intraspecific fighting during the breeding season show a much higher frequency of new wounds among males over 100 mm in snout-vent-length than in smaller males. The significant difference in new-wound frequency strongly suggests avoidance of fights by the small males, which is corroborated by laboratory and field observations. Access by males to reproductively active females depends on the ability to defeat other males in aggressive contests virtually always involving head biting if the males are of nearly equal size. Because the probability of winning agonistic encounters increases with size, young males avoid fights with older males. Aggressive contests with larger males and reproductive attempts other than courtship in the absence of larger males are deferred.Aggressive behavior in E. laticeps may be employed in direct defense of females, but might also be expressed in defense of specific sites and/or territories. In the laboratory, males in their home cages were significantly more likely to win encounters with males of similar size than were males fighting in the home cages of opponents. This suggests that encounter site could be important in determining encounter outcome and that field study of possible site defense or territoriality is needed.     

Patterns of range use,range defense,and intergroup spacing in moustached tamarin monkeys (Saguinus mystax)

In this paper we examine patterns of group spacing and habitat utilization in neighboring groups of marked free-ranging moustached tamarin monkeys (Saguinus mystax) inhabiting Padre Isla, a small island in the Amazon Basin of northeastern Peru, and describe the set of behavioral mechanisms used by tamarins to maintain the spatial isolation and social integrity of individual groups. Specifically, we address a series of questions regarding the importance of resource defense, mate defense, and territorial defense in intergroup interactions. From June through November 1990, we recorded 67 intergroup interactions involving members of our two main study groups. These interactions occurred at a rate of .14/observation hour and were of two general types. Vocal battles averaged 18 min in duration and were characterized by a series of high frequency, short syllable, long calls that were exchanged between groups separated by distances of greater than 25 m. Aggressive encounters averaged 26 min in duration and involved visual contact, alarm calls, scent marking, and sequences of chases and retreats. Intergroup confrontations did not cluster around the perimeter of a group's home range, and there was no evidence that moustached tamarins patrolled range borders. Our data indicate that 35% of aggressive encounters occurred in the vicinity of major feeding trees. Priority access to these sites is likely to have an important influence on tamarin foraging success. Mate defense and the exploration of new breeding opportunities also appear to be important functions of intergroup conflicts. Not only did the frequency of aggressive encounters increase during breeding periods, but three-fourths of all observed copulations occurred during or within 30 min of an encounter. Given the high degree of reproductive competition reported among tamarin females and the time and energy group members devote to intergroup aggression, maintaining access to a stable home range and the resources contained within that range appear to be critical functions of moustached tamarin social interactions.     

The effects of luteinizing hormone, oestrogen and ovariectomy on the agonistic behaviour of female Quelea quelea

Previous work with male Quelea showed that agonistic behaviour in relation to individual distance is controlled by luteinizing hormone (LH), rather than by testosterone, and that males are more aggressive than females. Experiments with female groups are reported which show that: (a) LH injections increase aggressive encounter frequency; (b) ovariectomy in the breeding season (but not outside it) also increases encounter frequency; and (c) oestrogen injections decrease encounter frequency. The effects of LH were shown to be specific to agonistic responses rather than mediated through changes in activity. Correlations between changes in natural hormone levels and encounter frequency support the injection findings. These results are consistent with the hypothesis that LH controls aggressive encounters over individual distance in the female as in the male and that oestrogenic inhibition of this LH-mediated aggressiveness is a cause of female subordination and the lower encounter frequency found in female groups. The annual cycle of encounter frequency is described and the significance of different systems of hormonal control is discussed.     

The Equivocal Relationship Between Territoriality and Scent Marking in Wild Saddleback Tamarins (<Emphasis Type="Italic">Saguinus fuscicollis</Emphasis>)

Researchers have often assumed that scent marking serves a territorial function in callitrichines, although some controversy exists. To fulfill such a function, scent marks should 1) prevent intrusions, 2) ensure access to feeding resources, 3) enable avoidance of intergroup encounters, or 4) play an important role in the aggressive encounters between groups. We studied 13 saddleback tamarins (Saguinus fuscicollis) belonging to 3 free-ranging groups, which formed mixed-species troops with moustached tamarins (S. mystax) in the Amazonian rain forest of Peru. None of the predictions were confirmed. The tamarins used a border-marking strategy, marking more on the periphery of their territory. However, feeding trees in overlap and encounter areas received more scent marking but were still visited by neighboring groups. Intergroup encounters occurred more often than expected, and scent-marking frequency was not higher during them than when no other group was present. It appears that instead of defending a territory in the classic sense, the tamarins are optimizing signal transmission by depositing their scents where the probability of detection by neighbors is higher. Saddleback tamarins may use shared areas of their home ranges to exchange information with neighboring groups, perhaps regarding reproductive opportunities.     

Food conditions, competitive regime, and female social relationships in Japanese macaques: within-population variation on Yakushima

Feeding conditions, competitive regime, and female social relationships of Japanese macaques (Macaca fuscata) on Yakushima were compared between the two habitats at two different altitudes (coniferous forest, 1,000–1,200 m and coastal forest, 0–200 m). Fruit availability was higher in the coastal forest. There was no consistent difference in the frequency of agonistic interactions within a group during feeding between the two habitats. The coastal forest evoked stronger inter-group contest competition compared to the coniferous forest as evidenced by a higher inter-group encounter rate and a higher proportion of aggressive encounters to non-aggressive ones. Birth rate was higher in larger groups compared to smaller ones in the coastal forest, but did not differ in the coniferous forest. In spite of these differences in competitive regime, no variation in female social relationships was observed, such as direction and concentration on particular individuals in grooming, linearity in dominance rank, counter-attack, and support of juvenile kin during agonistic interactions. The present results indicate that the female social relationships of Japanese macaques are robust and do not change according to changes in the current environment.     

Increased Frequency of Intergroup Encounters in Wild Bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>) Around the Yearly Peak in Fruit Abundance at Wamba

Intergroup interactions in primates vary from nonagonistic to severely aggressive. Food resources and fertile females cause intergroup aggression when groups defend resources and mates from other groups. Peaceful intermingling during intergroup encounters is rare but has been reported in several primates, including bonobos (Pan paniscus). Although intergroup encounters in bonobos occur at both nonprovisioned and provisioned sites, provisioning may be one factor responsible for frequent intergroup encounters. We studied intergroup encounters between one bonobo group (PE) and its neighboring groups, one semihabituated and two habituated groups, under nonprovisioned conditions in 2010–2015 at Wamba, Democratic Republic of Congo. We examined whether fruit abundance and females with sexual swellings influenced encounter frequency. PE group encountered the other groups on 7.1 days (mean, range: 0–19) per month. Up to four different groups met simultaneously at one location. Encounter frequency was highest around the yearly peak in fruit abundance, suggesting that reduced food competition is linked to increased encounter frequency. During periods of relatively low fruit abundance, the probability of an encounter was higher when a higher number of adult females with maximum sexual swellings were present, suggesting that the attractiveness of females with sexual swellings promoted group encounters during periods of relatively low fruit abundance. These findings suggest that both competition for food and the availability of females with sexual swellings influence encounter frequency in bonobos.     

Interaction effects of corticosterone and experience on aggressive behavior in the green anole lizard

Aggressive encounters are accompanied by a release of stress hormone, and this corticosterone (CORT) secretion could influence aggressive behavior in subsequent encounters. We investigated the modulating effects of CORT on aggressive behavior in the context of a 5-day social experience in male green anole lizards. In Experiment 1, we measured plasma CORT levels in animals that were exposed for different times to aggressive males. In Experiment 2, using metyrapone, a CORT synthesis blocker, we tested whether CORT secretion in response to the aggressive stimulus plays a role in experience-dependent facilitation of aggressive behavior. We hypothesized that aggressive encounters would increase plasma CORT levels, and that blocking CORT synthesis with metyrapone treatment during the aggressive encounter would cause an animal to become more aggressive. We also tested whether blocking CORT would interfere with the influence of 5-day social experience on animals' behavior in a subsequent aggressive encounter. Animals that were exposed to another male showed higher plasma CORT levels immediately after the 10 min encounter than animals exposed to the non-social video, and this high level was maintained through day 5. Within the aggressive video groups, in Experiment 2, there was a distinctly different pattern in displays depending on drug condition: vehicle-injected animals showed gradual increases followed by decreases in aggressive behavioral responses to the video as the five days proceeded (habituation), while animals injected with metyrapone started out with high aggressive behavior and did not decrease behavioral responses at later trials (no habituation). Finally, when tested with a novel conspecific on day 6, animals previously injected with metyrapone showed no higher aggression than did animals previously injected with vehicle and exposed to the aggressive video. These results suggest that blocking CORT synthesis during the exposure to the aggressive video induced animals to remain aggressive toward the repetitive stimulus without habituating, while not becoming more aggressive than controls toward a novel challenger.     

Fruitful and Futile Encounters along the Association Reaction between Proteins

The association reaction between pairs of proteins proceeds through an encounter complex that develops into the final complex. Here, we combined Brownian dynamics simulations with experimental studies to analyze the structures of the encounter complexes along the association reaction between TEM1-β-lactamase and its inhibitor, β-lactamase-inhibitor protein. The encounter complex can be considered as an ensemble of short-lived low free-energy states that are stabilized primarily by electrostatic forces and desolvation. For the wild-type, the simulation showed two main encounter regions located outside the physical binding site. One of these regions was located near the experimentally determined transition state. To validate whether these encounters are fruitful or futile, we examined three groups of mutations that altered the encounter. The first group consisted of mutations that increased the experimental rate of association through electrostatic optimization. This resulted in an increase in the size of the encounter region located near the experimentally determined transition state, as well as a decrease in the energy of this region and an increase in the number of successful trajectories (i.e., encounters that develop into complex). A second group of mutations was specifically designed to either increase or decrease the size and energy of the second encounter complex, but either way it did not affect k_on. A third group of mutations consisted of residues that increased k_on without significantly affecting the encounter complexes. These results indicate that the size and energy of the encounter regions are only two of several parameters that lead to fruitful association, and that electrostatic optimization is a major driving force in fast association.     

Why do male chimpanzees defend a group range?

Male chimpanzees, Pan troglodytes, cooperate to defend a community range within which resident females range in smaller core areas. There has been debate over exactly what males are defending, whether mates, territory or both. One hypothesis holds that males are defending mates, and that an increase in community range size will lead directly to the acquisition of more females. However, males frequently attack females as well as males at the edge of the community range. We examined 18 years of observational data on the Gombe chimpanzees to determine the behaviour of males during extragroup encounters, and the consequences of changes in community range size on the number of adult females and indirect measures of food availability. Males were always aggressive to males from other communities, and often attacked adult females, especially those that were not sexually receptive, were older, and/or had more than one offspring. The number of females did not increase with range size, but several measures suggested an increase in food availability with range size. These measures include more time spent in large foraging parties, higher encounter rates with resident females, more encounters with sexually receptive females and higher female reproductive rates. These findings suggest that males defend a feeding territory for their resident females and protect them from sexual harassment. Although a large range may eventually attract more females, this is not an immediate consequence of range expansion. Male number was not correlated with community range size.     

Aggressive intergroup encounters in two populations of Japanese macaques (Macaca fuscata)

It is predicted that variation in intergroup relationships in group living primates reflects the cost and benefit of resource defense. We tested the applicability of the model by examining population difference, group difference, and seasonal difference in behaviors during intergroup encounters in two populations of Japanese macaques (Macaca fuscata), one of six groups from Yakushima Island, and the other of three groups from Kinkazan Island. We found that the nature of intergroup encounter varied with group identity, reproductive seasonality, and population. Yakushima groups showed aggressive behaviors more frequently than did Kinkazan groups and the difference was consistent with the food competition model, both because of the involvement of females, and because home ranges were smaller on Yakushima than on Kinkazan, and thus more defensive. Both sexes of animals participated in aggressive interactions, but males were more aggressive than females. Furthermore, Yakushima population showed more agonistic intergroup behaviors during the mating season than the non-mating season. Also during the encounters, intergroup mating was observed, but only in Yakushima. It is concluded that intergroup relationships reflect the mate guarding behavior by group males. However, the agonistic relationship during non-mating season, especially that of among females, is also consistent with the food competition model. It is also noted that males' behavior toward other groups can also be interpreted as a form of investigative behavior before possible transfer into a new group.     

Inter-male aggression in relation to female availability and residence status in corn mice <Emphasis Type="Italic">Calomys musculinus</Emphasis>

We conducted an experimental study to test the hypothesis that, at low female availability, males show intrasexual aggressive behaviour and home range owners are more aggressive than home range intruders. Using field dyadic arena test, we carried out 35 male–male trials in four 0.25 ha enclosures, two male-biased (experimental enclosures) and two unbiased (control enclosures). Dyadic encounters were conducted between unrelated and sexually mature males of similar weight and age which established home ranges in the same enclosure at the same trapping session. Each inter-male encounter was performed in the home range of one of the opponents between the home range owner and a home range intruder. When sex ratios were male-biased, inter-male amicable behaviour was absent and we found significant rates of inter-male aggressiveness, being home range owners much more aggressive than intruders. In the unbiased enclosures, inter-male encounters resulted mainly in noninteractive or amicable interactions. We found that inter-male aggression varied in relation to female availability being more evident in home range owners.     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

Increased host aggression as an induced defense against slave-making ants

Slave-making ants reduce the fitness of surrounding host colonies through regular raids, causing the loss of brood and frequently queen and worker death. Consequently, hosts developed defenses against slave raids such as specific recognition and aggression toward social parasites, and indeed, we show that host ants react more aggressively toward slavemakers than toward nonparasitic competitors. Permanent behavioral defenses can be costly, and if social parasite impact varies in time and space, inducible defenses, which are only expressed after slavemaker detection, can be adaptive. We demonstrate for the first time an induced defense against slave-making ants: Cues from the slavemaker Protomognathus americanus caused an unspecific but long-lasting behavioral response in Temnothorax host ants. A 5-min within-nest encounter with a dead slavemaker raised the aggression level in T. longispinosus host colonies. Contrarily, encounters with nonparasitic competitors did not elicit aggressive responses toward non-nestmates. Increased aggression can be adaptive if a slavemaker encounter reliably indicates a forthcoming attack and if aggression increases postraid survival. Host aggression was elevated over 3 days, showing the ability of host ants to remember parasite encounters. The response disappeared after 2 weeks, possibly because by then the benefits of increased aggression counterbalance potential costs associated with it.     

Random walks and the gas model: spacing behaviour of Grey-Cheeked Mangabeys

1. Simple mechanical models (random walks and the 'gas model') were used to investigate the movement patterns and intergroup encounter rates of a high- and low-density subpopulation of Grey-Cheeked Mangabeys ( Cercocebus albigena johnstoni, Lydeker) at two sites in the Kibale Forest, Western Uganda.
2. Random walk simulations showed that the presence of conspecific groups could act as 'social barriers' which constrained group movements, and promoted high levels of site attachment to a specific home range area.
3. Encounter rate models showed that in the low-density subpopulation (Kanyawara), intergroup encounters occurred no more frequently than expected if groups were moving randomly and independently of each other. This was in contrast to previous analyses which suggested that Grey-Cheeked Mangabeys employed a social spacing mechanism (mediated by male loud calls) in order to reduce the frequency of encounter to below chance levels.
4. Encounters in the high-density subpopulation (Ngogo) were found to occur less frequently than expected at short range (within 100 m). This was suggested to be due to the operation of a territorial mate defence strategy by males at this site.     

Winning fights elevates testosterone levels in California mice and enhances future ability to win fights

The 'winner effect' has been studied in a variety of species, but only rarely in mammals. We compared effects of winning three, two, one, or zero resident-intruder encounters on the likelihood of winning a subsequent aggressive encounter in the California mouse (Peromyscus californicus). During the training phase, we ensured that resident males won all encounters by staging contests with mildly sedated, smaller intruders. During the test phase, the resident male encountered an unfamiliar, more evenly matched intruder that had experience winning an encounter and was larger than the resident. Testosterone (T) plasma levels significantly increased after the final test when they had experienced two prior winning encounters, and the probability of winning a future encounter increased significantly after three prior wins independent of intrinsic fighting ability. We hypothesize a 'winner-challenge' effect in which increased T levels serve to reinforce the winner effect in male California mice.     

Variation in Intergroup Encounters in Two Populations of Japanese Macaques

The nature of intergroup encounters differed between two populations of wild Japanese macaques (Macaca fuscata): the Yakushima and Kinkazan populations. In the Yakushima population, intergroup encounters were more likely to result in the displacement of one group, intergroup agonistic interaction was common, and intergroup dominance was usually distinct. When displacement occurred at Yakushima, larger groups tended to dominate smaller ones. Conversely, in the Kinkazan population, intergroup encounters rarely resulted in displacement, intergroup agonistic interaction was rare, and intergroup dominance was usually unclear. Thus, monkeys in Yakushima appear to defend resources actively during encounters, while those in Kinkazan usually did not defend resources. The frequency of encounters was significantly higher in Yakushima than in Kinkazan. The two populations had very different group densities and traveling speeds, both of which directly influence the chance of encounters. Taking these differences into account, we compared the observed frequency with those predicted by the ideal gas model. The observed frequencies in both populations were about one-third of the number expected with the model, which suggests that the differences in encounter frequency were caused by differences in group density and traveling speed. We discuss this intraspecific variation in light of economic defendability in connection to habitat differences and the evolutionary significance of resource defense behavior.     

Biomass encounter rates limit the size scaling of feeding interactions

The rate that consumers encounter resources in space necessarily limits the strength of feeding interactions that shape ecosystems. To explore the link between encounters and feeding, we first compiled the largest available dataset of interactions in the marine benthos by extracting data from published studies and generating new data. These data indicate that the size‐scaling of feeding interactions varies among consumer groups using different strategies (passive or active) to encounter different resource types (mobile or static), with filter feeders exhibiting the weakest feeding interactions. Next, we used these data to develop an agent‐based model of resource biomass encounter rates, underpinned by consumer encounter strategy and resource biomass density. Our model demonstrates that passive strategies for encountering small, dispersed resources limits biomass encounter rates, necessarily limiting the strength of feeding interactions. Our model is based on generalisable assumptions, providing a framework to assess encounter‐based drivers of consumption and coexistence across systems.     

Group encounters and territoriality in wild Alaotran gentle lemurs (Hapalemur griseus alaotrensis)

During a 3 month field study, 18 group encounters between four groups of Alaotran gentle lemurs (Hapalemur griseus alaotrensis) were observed in the Lake Alaotra marshland in Madagascar. Behaviors observed during group encounters are described, and quantitative data on intergroup interactions and ranging patterns are discussed in the context of territoriality. Intergroup interactions varied in their intensity, ranging from visual monitoring, scent marking, and display locomotion to penetrating the neighboring range. A quarter (27%) of the encounters were aggressive, involving chases and confrontation displays, and no affiliative interactions were observed. Both adult males and females were involved in intergroup encounters, with males playing more active roles. All encounters took place in the small overlapping areas of neighboring home ranges. The outcome of an aggressive encounter was determined by its location, with the resident group always driving out the intruding one. The Alaotran gentle lemur actively advertised and consistently defended a large area of its home range against intrusion of neighboring groups and can therefore be considered territorial. Am. J. Primatol. 46:251–258, 1998. © 1998 Wiley-Liss, Inc.     

Morphological and Behavioural Traits Affecting the Intensity and Outcome of Male Contests in Gallotia galloti galloti (Family Lacertidae)

This study describes the sequence of behaviour during aggressive encounters between male ‘tizón’ lizards (G. galloti galloti) and assesses the effect of morphological and behavioural traits on the outcome and intensity of staged aggressive encounters between males. Aggressive encounters ranged from only throat extension to escalated fights with biting and rolling over. Winners were heavier, had longer heads, and performed tongue-flicking, throat extension and biting at a higher rate than losers. The rate of aggressive behaviour increased with decreasing difference in snout-to-vent length, head length and head width of the contestants. The results are in agreement with some predictions of the sequential assessment game model in that probability of victory increased with the difference in fighting ability and that the rate of aggressive behaviour was higher in contests between animals of similar size.     

Effect of soil hardness on aggression in the solitary wasp Mellinus arvensis

1. Two alternative nesting strategies are exhibited by soil‐nesting Mellinus arvensis females, digging a new nest (diggers) and searching for an old unoccupied burrow (searchers). Wasps appear unable to distinguish between occupied and unoccupied nests, and aggressive interactions between searchers and nest owners at nest entrances are frequent. 2. In aggressive encounters, there is an advantage in size and residency status. 3. The costs associated with the two nesting strategies vary across geographically separated populations: nest digging incurs costs in terms of time, and these vary according to the hardness of the soil substrate; nest searching is variably costly in terms of risk of injury in aggressive encounters with nest‐owning females. 4. Individual female wasps can switch between nesting strategies, and thus soil hardness, by affecting the cost of nest construction, affects the relative frequencies of the two nesting strategies within a population, favouring an increase in the searching strategy. This, in turn, affects the frequency and intensity of aggression between females at a nesting aggregation. 5. Female body size is correlated with soil hardness. As aggressive encounters are more frequent in sites with hard soil substrates, there is increased selective advantage in having large body size at these sites. 6. Body size is determined primarily by the availability of food resources during larval development, which is, to a degree, a function of the size of the adult female. There is a trade‐off between provisioning a few cells with many provisions in each, leading to the development of few but large adults, as opposed to many cells with few provisions, leading to many small offspring. The relative advantage of these two provisioning strategies is, at least in part, a function of the hardness of the soil substrate.