Territorial competition among wild brown trout fry: effects of ownership and body size

The effects of ownership, energy status and relative size on the ability to defend a territory were studied in dyadic contests among wild brown trout fry. A logistic regression showed that the territory owner won 85% of the contests with opponents of equal size, and that a 30% size advantage was required to balance the advantage of ownership. Energy status did not affect the ability to win territorial contests.     

Does prior residency interact with loss? A study of male–male contests in the hermit crab Pagurus minutus

Residency is an important predictor of success in contests with ownership asymmetries. Residency often can interact with a winning experience. However, given that some residents lose a contest even when showing an ownership advantage and that the process leading to loss often determines the loser's subsequent success, prior ownership might also interact with a loss. Here, we staged experimental contests between males of the hermit crab Pagurus minutus with a similar-sized weapon (i.e., cheliped) to examine this possibility. Male–male contests in this species occur between a solitary intruder and an owner guarding a mature female. We evaluated (a) whether resource ownership and female value affect the contest outcome and (b) whether the probability of winning after losing differs depending on the initial role of the loser (i.e., owner or intruder) by using precopulatory guarding pairs of P. minutus collected from the field. In the first fighting trial, we found an ownership advantage and increasing owner success as the body size of his partner increased. Although some owners lost the fight, in contrast to our prediction, the frequency of losing in the second fighting trial did not differ between prior owners and prior intruders. Because losers from the first fighting trial of male–male contests have no female regardless of their initial role, this shared solitary status might be related to the lack of difference in success in the second fighting trial. Moreover, unlike in other animals, resident status might not always assure greater fighting ability in P. minutus males because guarding Pagurus males can avoid male–male contests by climbing up objects in the field. Losers in the first trial, therefore, may have been weaker contestants based on traits other than size, regardless of whether they were owners or intruders.     

Does Host Value Influence Female Aggressiveness,Contest Outcome and Fitness Gain in Parasitoids?

Intraspecific competition for resources is common in animals and may lead to physical contests. Contest outcomes and aggressiveness can be influenced by the resource holding potential of contestants but also by their perception of the resource value (RV). Competitors may assess resource quality directly (real RV) but may also estimate it according to their physiological status and their experience of the habitat quality (subjective RV). In this article, we studied contests between females of the solitary parasitoid Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae) when exploiting simultaneously a host, a Delia radicum L. (Diptera: Anthomyiidae) pupa. We tested the effect of factors modifying host value on the occurrence of agonistic behaviours, contest outcomes and host exploitation. The factors tested were: the quality of the previous habitat experienced by females, female egg load, host parasitism status and the stage reached by the owner female in her behavioural oviposition sequence. Females successfully protected their host against intruders during its exploitation, but not after oviposition, and their aggressiveness did not seem to be influenced by their perception of the RV. The fact that the host is subsequently parasitized by the opponent females appears to mainly depend on the host selectiveness of females.     

Sexual dominance in hermit crab shell fights: asymmetries in owner–intruder status,crab size,and resource value between sexes

We studied sexual dominance and seasonal differences in aggressiveness of individuals in intraspecific competition for shells of the hermit crab Pagurus filholi in terms of size of contestants and duration of the attempt to deprive other crabs of their shell. Experiments were conducted using paired intrasexual and intersexual contests in the pre-breeding and post-breeding seasons. Size ratios between contestants were systematically varied to assess the sexual difference in size and owner advantages. In both intrasexual and intersexual contests intruder crabs tended to win the contests more often as their size increased, that is, size advantage overcame owner advantage. Although we did not recognize a sexual difference in size and owner advantages in contest outcomes, male intruder crabs took a shorter time to deprive female owners of a shell than to deprive male owners. Furthermore, male individuals in the pre-breeding season had significantly longer fight durations. Fighting is costly. Thus males can afford to expend more energy and time fighting, indicating that males are dominant over females in shell fights as both intruders and owners. Electronic Publication     

Asymmetrical Resource Ownership Increases Owners’ and Reduces Non‐Owners’ Motivation to Fight in the Mangrove Rivulus,Kryptolebias marmoratus

Resource ownership often increases an individual's aggressiveness and its probability of defeating a competitor. Individuals contesting resource owners could therefore incur higher costs, making individuals reluctant to compete with owners. We tested the hypothesis that animals use asymmetry in resource ownership as a cue for contest costs and adjust contest decisions accordingly. Using a mangrove rivulus fish (Kryptolebias marmoratus), we staged (1) contests with a randomly assigned asymmetry in resource ownership (one fish was provided with a shelter) and (2) contests in which neither fish had a shelter. Owners that were in their shelters at the contest start showed a greater tendency to fight and won more contests than their intruder opponents; those outside the shelter at the start did not. Compared with fish in contests with no shelters at stake, shelter owners had a higher tendency to fight whether or not they were in their shelters at the start; intruders, however, had a lower tendency to fight only against owners that were inside the shelter at the start. These results demonstrate (1) that ownership status influences both owners’ and intruders’ contest decisions (and in opposite directions), producing a detectable ownership advantage and (2) that intruders required confirmation of their opponents’ ownership status before retreating without challenging them. Ownership status per se is therefore important to the fish's contest decisions.     

Why Do Males of Parnassius imperator Fight for Bare Rocks but Not the Nectar Flower During Mate Selection?

Territoriality is an important male behavioral trait used in mate selection of butterflies, but the factors determining the outcome of territorial contests remain controversial. We studied the foraging and territorial behaviors of males in the Parnassius imperator butterfly. Adult males were selective about nectar sources, but they did not defend nectar flowers. By contrast, they fought over areas of bare rocks because bare rocks were used as lek locations to attract and court receptive females. The ownership of a lek, rather than sex pheromones or wing pigments, played the key role in attracting females. In an owner replacement experiment, the owner of a territory always defeated intruders, indicating that perching order, not relative body size, determined the outcome of territorial contests. Territories are small areas of bare rocks that lack food resources, so intruders do not challenge the ownership of a resident in a location where they can easily find vacant territories. Males that first perch in a territory have an advantage compared to intruders in that they can forage on flowers in the vicinity of the territory. This may help explain why satiated owners always defeated fatigued intruders. Our findings suggest that early perching on bare rock allows male butterflies of P. imperator to occupy a territory, attract receptive females and exclude later‐arriving competitors.     

Male contesis for territories and females in the fly Dryomyza Anilis

Dryomyza anilis males defend the small carcasses on which females oviposit as well as the females themselves during copulation and oviposition. Contests over territories and females, were compared to assess the way different asymmetries affect the outcome of a struggle. In contests over females with a lot of eggs the rate of take-over was lower than in contests over immature females, but the size of the female or her size relative to the owner did not affect the outcome. In territorial disputes and territories was the size difference between the opponents. Fifty-three per cent of the contests over females and 39% of territorial struggles resulted in a take-over. In take-overs, owners were smaller than copulating males or territory owners on average. Take-overs are part of a process in which resource owners continue to change until one is strong enough to defend the resource successfully. Contests over females lasted significantly longer than those over territories, which suggests that for males females are more valuable than territories. This is to be expected as territories are of value because they ultimately increase the probability of obtaining females. A positional advantage or the higher payoff of the owner in disputes over females may be the reason for the lower rate of take-over in female defence than in territorial struggles between opponents of similar size.     

Comparative evidence supports a role for reproductive allocation in the evolution of female ornament diversity

1. Sexually selected ornaments are highly variable, even among closely related species, and the ultimate causes of variation in ornament evolution are unclear, including in rare cases of female ornament expression. One hypothesis is that differences across species in female reproductive allocation may help to explain patterns of female ornament expression among insects with nuptial gifts. 2. Dance flies (Diptera: Empididae: Empidinae) vary considerably among species in the presence and extravagance of female ornaments, which probably evolved through female contests for mates. In most dance flies, adult females appear to acquire all their dietary protein from nuptial gifts provided by males during mating. The importance of nuptial feeding on egg development is not yet known. 3. To test the prediction that the presence of female ornaments reflects differences in the degree to which females rely on nuptial feeding for egg development, egg development was examined in wild females of two species, one ornamented and the other unornamented. An ageing technique based on cuticular bands was validated, which permitted a regression of egg size on adult age. 4. We found that egg development depended on mating status in the ornamented species alone, meaning the eggs of unmated females of the ornamented species did not develop. This contrast across species is consistent with expectations that females of different species vary in their dependence on nuptial gifts for egg development. 5. These findings provide preliminary support for the hypothesis that differences in reproductive allocation mediate the intensity of female contests for nuptial gifts.     

Territory size in the moorhen (Gallinula chloropus): An outcome of RHP asymmetry between neighbours

Territory size can be considered as the outcome of contests between an owner and its neighbours for the resource units in the defended area. Variation in territory size could be dependent upon three possible asymmetries: (1) difference in resource holding potential (RHP); (2) difference in resource value to the competitors; and (3) difference in ‘ownership’ status. These possibilities were tested in a 3-year study of moorhens which defend linear territories along ditches in a grazing marsh habitat. Of the three, relative difference in RHP between an owner and its two neighbours provided the best correlation with territory size. There was no relationship between the amount of resources per unit area and territory size nor between the time of establishing a territory and its later size. Heavier moorhens are more likely to win contests in winter flocks and are thought to have a greater RHP. The ratio of the weight of a male territory owner to the weight of both its male neighbours was highly correlated with territory size. Birds with larger territories may gain more potential nesting sites with good cover from predators.     

Female Contests for Nest Sites and Mates in the Pied Flycatcher Ficedula hypoleuca

Predictions of game theory models about the outcome of animal conflicts have most often been tested using male contests for mates, territories or food. We studied female contests for nest sites and mates in the pied flycatcher, Ficedula hypoleuca, and asked which factors affect the outcome and duration of the contests. Conflicts over residency were experimentally induced, but natural cases in which both females regarded themselves as owners were also included in the analyses. The ensuing escalated contests involved physical fighting, defence of the nestbox of the male and alarm calling. Contrary to expectations from game theory, fighting seemed to occur most often at the start of conflicts, whereas alarm calling occurred towards the end and nestbox visits throughout the conflict. The outcome of the contests was not determined by asymmetries in body size or age, nor by a simple previous-present owner asymmetry. Instead, it depended on the relative residence times of the opponents. The duration of the contests tended to increase with decreasing asymmetry in the residence times of females, whereas body size asymmetries had no influence. Because there is intense competition between females for a mate in the pied flycatcher, we suggest that females do not respect asymmetries in residency and body size, but fight in relation to the value of the mating opportunity. In particular, we point to the possibility that the value of a mating opportunity may increase with residence time because knowledge of other mating options may become outdated.     

Intruder Pressure Affects Territory Size and Foraging Success in Asymmetric Contests in the Water Strider Gerris lacustris

The theory of evolutionarily stable strategies (ESS) in asymmetric contests predicts that the propensity of an individual to expose itself to risk during contests depends on the individual's resource-holding-potential (RHP) and on the value of the disputed resource (V) for the individual compared with that for an opponent. If encounters of a territory owner with individuals of high RHP and high food demands (V) increase in frequency, one should expect a decrease in total aggressiveness of the territory owner, and in consequence a decrease of its territory size. Such a decrease should result in a lower amount of food consumed by the territory owner. Using natural variability in RHP and V in Gerris lacustris, I experimentally tested these predictions. The average prey item has higher value (V) for reproductive female water striders (which probably transform most of their food into eggs), than for nonreproductive females and for males. Because males are smaller, they have lower RHP than females, as RHP depends on size. Thus the reproductive females are the class of individuals of high RHP and high food demands (high V). Most nonreproductive females defend food-based territories. I observed two groups of water striders in a seminatural laboratory setting. As predicted, there was a negative correlation between the rate of encounter with reproductive females and size of the territory, and a positive correlation between territory size and number of Drosophila flies consumed by the owner. Territories were smaller in the group with high rates of encounter between territory owners and reproductive females. Territory owners caught the same number of Drosophila flies as non-territorial individuals in this group. In contrast, in the group with fewer encounters between territory owners and reproductive females, territories were larger, and territory owners gained more food than non territorial water striders.     

Power to Hold Sheltering Burrows by Juveniles of the Signal Crayfish,Pasifastacus leniusculus

Juveniles of the signal crayfish reside during daylight hours in shelters. At twilight they leave for food, at daybreak they either return or find another shelter. We examined conflicts over burrow ownership. At low densities with equal numbers of shelters and crayfish 1/5 of the burrows were occupied. Increasing both the number of crayfish and shelters improved the occupancy close to 50%. Doubling the number of crayfish in relation to the number of shelters increased the occupancy up to 75%. In an experiment with 30 randomly selected crayfish and 15 sheltering holes available the burrow holders were about 1–2 mm longer in carapace length than those found freely moving in the aquarium. Shelter owners were rarely newly moulted individuals. Size asymmetry (2 mm difference in carapace length) between owner and intruder affected the outcome of the contest over burrow ownership. When intruders were larger than owners, takeovers occurred in about 80% of the cases tested. If the owner was large and the intruder small the takeover frequency was about 20%. When the owner and the intruder were of matching size takeovers were still observed in about 45% of the cases. The outcome of the ownership contest has a true meaning. In a nursery-pond experiment, where low-protein food was randomly scattered all around, no size differences were found in carapace lengths of crayfish juveniles residing in different shelters. However, when high-protein food was introduced in a single spot, owners of burrows were significantly larger and more numerous in the nearby shelters than in the shelters furthest away from the food source. Therefore, when food is unevenly distributed the burrow ownership contests may potentially lead to size asymmetries between individuals. This may lead to large individuals nearby the food growing faster than small individuals further away from the food source, a process likely to further enhance size differences.     

Effect of Population Density and Female Body Size on Number and Size of Offspring in a Species with Size‐Dependent Contests over Resources

When size‐dependent contests over resources influence reproductive success, the trade‐off between number and size of offspring depends on the frequency of contests. Under these circumstances, clutch size should decrease and offspring size should increase as contests become more frequent. We tested these predictions with the burying beetle Nicrophorus pustulatus through manipulation of rearing densities. Burying beetles reproduce on small vertebrate carcasses, a rare but high quality food source for the larvae. Large beetles are more likely to win contests over carcasses and gain exclusive access to a carcass. The winner of a contest kills eggs and larvae already present on a carcass. As a result of the rarity of carcasses, burying beetles are unlikely to breed more than once. As predicted, brood size of N. pustulatus decreased with increasing rearing density. Despite a negative correlation between brood size and larval mass, larval mass did not increase with increasing rearing density. This may be due to the special biology of N. pustulatus which can use snake eggs for reproduction. Potentially larger supply of resources and generally small population densities of N. pustulatus may weaken selection on body size and thus the correlation between brood size and larval mass. As size‐dependent constraints can limit reproductive phenotypes, we examined whether female size influenced reproductive phenotype. Small females produced larger broods with smaller, but more variable, offspring than large females. Mechanical constraints of egg size seem an unlikely explanation for the differences because burying beetles can compensate for small egg size through parental care. Energetic constraints may impact small females because body mass and brood size of small females decreased with increasing density. Yet, at all density levels small females produced larger, not smaller, broods than large females. The larger and more variable broods of small females seem to be in agreement with a bet‐hedging strategy.     

The life history continuum hypothesis links traits of male ants with life outside the nest

Biological control of bruchid beetles, Callosobruchus maculatus (Fabricius) (Coleoptera: Bruchidae), infesting cowpea seeds, Vigna unguiculata (L.) Walpers (Fabaceae), can be performed via augmentative releases of Dinarmus basalis Rondani (Hymenoptera: Pteromalidae) parasitoid wasps. Females of the latter species are therefore likely to experience intense intraspecific competition: they should encounter numerous previously parasitized hosts but also conspecific competitors, with which they may fight to secure hosts on which to lay their eggs. Such contests might therefore disrupt biological control programs. Here, we studied aggressive behavior that D. basalis females show toward conspecific competitors and subsequent host exploitation strategies. We further investigated factors that classically affect contest intensity and outcomes in animals, such as the effect of ownership status, by manipulating the residency period before the intruder's arrival. In addition, we tested the effect of the size of female reproductive tissue (measured in terms of egg load) and the quality of the habitat previously experienced by females (either rich or poor in hosts). These two factors are expected to influence the value that females place on the host and therefore the costs they are willing to pay to win it. Finally, we discussed the consequences of agonistic behaviors on females' host exploitation strategies. Our results suggest that contest competition may actually enhance host control by favoring parasitoid dispersion, rather than disrupting it.     

Agonistic Behavior in the Squid Loligo plei (Loliginidae,Teuthoidea): Fighting Tactics and the Effects of Size and Resource Value

Observations of small schools of squids in captivity suggested that dominance relationships among males were based upon major differences in the frequency or duration of their agonistic behavior, but staged contests showed few differences. During staged contests, squids exhibited up to 21 separate behaviors. Some contests included a complex array of visual signals and side-by-side posturing (Lateral Display) followed by physical contact during Fin beating. There was behavioral variability and step-wise escalation during the contests: squids performed either 1. long sequences of visual signaling followed by Chasing and Fleeing; or 2. short sequences of visual signaling followed by physical Fin beating and ending with Chasing and Fleeing. Size influenced outcome in all contests; larger males were more likely to win the contest. Size had no effect on contest duration, but contest duration was shorter when resource value was high, especially when a male established temporary ownership of a female. We speculate that when the perceived resource value is high, male squids are more likely to engage in a shorter yet riskier fighting tactic.     

Early Comers Become Owners: Effect of Residency Experience on Territorial Contest Dynamics in a Lycaenid Butterfly

A positive relationship between ownership and contest success is widespread among territorial butterflies. We tested the hypothesis that owners win simply because they have a longer experience occupying a given territory than do intruders in territorial butterflies. Using the lycaenid butterfly Chrysozephyrus smaragdinus , we staged contests between two males that had experienced different residency periods at the same contested territory, similar to owners and intruders in natural contests. We found that males that had occupied the contested territory for a longer period (a day) always defeated the shorter occupancy males (15 min). Therefore, residency period plays a key role in settling territorial contests. As residency confers advantages in a time-dependent manner, the mechanism of owner dominance could be an increase in motivation to fight as local familiarity increases. Moreover, this result predicts a population-level territorial dynamics. If residency period per se confers advantages in each contest, males arriving at a territory early in their adult season should acquire territorial dominance because they can remain there for a certain period with only a few rivals; late-appearing males do not have this advantage. As predicted, we found that males appearing at territory sites early in the season were more likely to hold their territories. This finding provides new evidence for the evolution of protandry in butterflies.     

Don't count your eggs before they're parasitized: contest resolution and the trade-offsduring patch defense in a parasitoid wasp

Although aggressive conflicts over hosts occur among femalesin many species of insect parasitoids, few studies have examinedthe mechanisms by which these contests are resolved. In Trissolcusbasalis, a parasitoid of pentatomid bug egg masses, femalesco-exploiting an egg mass (patch) encounter one another repeatedlyand fight for possession of the patch. We investigated the resolutionof pairwise contests by experimentally varying the release timeof females and the size of the patch. Logistic regression showedthat the female arriving first on the patch was more likely towin both the first agonistic encounter and to retain overallpossession of the patch. This advantage to the first femalesuggests a resource-correlated asymmetry in favor of the firstfemale, due to her having invested more offspring in the patch.Although escalations were more common when the asymmetry inarrival times was small, the majority of encounters within all contestswere nevertheless resolved without escalated fighting, withthe resident attacking and the intruder backing down. Thus contestresolution basically followed a "bourgeois" rule. Residentstolerated intruders more frequently when the patch size waslarger and the second female was released later, illustratingthe trade-off faced by residents between defending the patchand continuing to exploit it.     

Non‐sex‐biased Dominance in a Sexually Monomorphic Electric Fish: Fight Structure and Submissive Electric Signalling

Males and females commonly compete for limited resources. When interaction costs are similar for both sexes and there are no sexual differences in resource value estimation, a non‐sex‐biased dominance is expected. Moreover, only non‐sex‐biased assessment of contenders fighting ability (Resource Holding Potential, RHP) should influence contest decisions. To test these predictions, we evaluated non‐breeding agonistic intra‐ and intersexual dyadic interactions in the weakly electric fish, Gymnotus omarorum. During the non‐breeding season, resource value is not expected to depend on individuals’ reproductive status and should thus be equal for males and females. In addition, as G. omarorum presents no sexual differences in body size, interaction costs can be considered symmetric between sexes. We confirmed that body size differences, but not individuals’ gender, is the best predictor of dominance. We correlated RHP asymmetries with contest duration and evidenced that body size but not sex influences assessment in intrasexual and intersexual encounters. All dyads tested engaged in agonistic interactions (N = 33) in which a clear dominant emerged. The analysis of conflict phases evidenced the submissive role of electric displays. Electric organ discharge (EOD) interruptions appear early in the contest as an electric hiding attempt, whereas chirps are post‐resolution signals of subordinate status. Interestingly, the decision of interrupting the EOD was also influenced by RHP asymmetries, whereas chirping activity was influenced by the intensity of the attacks received. Our results confirm that body size is the best RHP proxy in non‐breeding intra‐ and intersexual contests of this monomorphic species and demonstrated a sequential pattern of submissive signalling by means of two different electric displays.     

Maternal effects and their consequences for offspring fitness in the Yellow Dung Fly

1. Maternal adult diet and body size influence the fecundity of a female and possibly the quality and the performance of her offspring via egg size or egg quality. In laboratory experiments, negative effects in the offspring generation have often been obscured by optimal rearing conditions.
2. To estimate these effects in the Yellow Dung Fly, Scathophaga stercoraria , how maternal body size and adult nutritional status affected her fecundity, longevity and egg size were first investigated.
3. Second, it was investigated how female age and adult nutritional experience, mediated through the effects of egg size or egg quality, influenced the performance of offspring at different larval densities.
4. Maternal size was less important than maternal adult feeding in increasing reproductive output. Without food restriction, large females had larger clutch sizes and higher oviposition rates, whereas under food restriction this advantage was reversed in favour of small females.
5. Offspring from mothers reared under nutritional stress experienced reduced fitness in terms of egg mortality and survival to adult emergence. If the offspring from low-quality eggs survived, the transmitted maternal food deficiency only affected adult male body size under stressful larval environments.
6. Smaller egg sizes due to maternal age only slightly affected the performance of the offspring under all larval conditions.     

Contest dynamics for food and reproductive resources are defined by health condition in a dung beetle

Male dung beetles compete to obtain food and females, and early resource recognition and appropriation increase the probability of mating. The outcome of such encounters is expected to be defined by self‐ and the opponent’s health condition. In this study, we analyzed the effect of body condition and immune defense on the contest dynamics between males that rolled a food ball with a partner female (owner males) and intruder males of dung beetle Canthon cyanellus. Body condition was measured as body size, body dry mass, lipid mass, and muscle mass; immune defense was estimated via phenoloxidase activity. Owner males with higher lipid mass contacted the food ball significantly earlier than owner males with lower lipid mass. Individuals with lower phenoloxidase activity started to roll food balls earlier than individuals with higher phenoloxidase activity. Owner males that had higher body dry mass, compared to female partners, began to roll the food ball significantly earlier than male–female pairs with lower differences in dry mass. Heavier males won significantly more contests than lighter males. Our results suggest that the health condition is a key factor related to the dynamics and outcome of male–male contests for resources and females in C. cyanellus. Consequently, differences in individual condition are main determinants of contest outcomes in dung beetles.