An Extreme Case of Interspecific Territoriality: Male Anthidium manicatum (Hymenoptera,Megachilidae) Wound and Kill Intruders

Anthidium manicatum males defend patches of flowering plants as mating territories and copulate with females that enter the territory to obtain nectar or pollen. Territorial males defend their territories not only against conspecific males but also against many other insect species that attempt to utilize the flowers inside the territory. In our study area, a territorial male defended its territory against conspecific males on average only twice per h but it attacked other species of insects on average 70 times per h. Territorial males distinguished between different species of intruders and attacked them unequally. During the hour following experimental removal of the territory holder the number of intruders of some insect species tripled but the number of intruders of those species that were rarely attacked remained approximately the same. A. manicatum males, but not females, bear long spines on the last two segments of the abdomen. Territorial males rammed intruding insects at high speed and shortly before the moment of impact they curved the abdomen forward to hit with the spines. Intruders could be seriously damaged or killed by these attacks. Abdominal spines of A. manicatum males may have evolved as weapons to increase the effectiveness of interspecific territoriality.     

Territorial responses to energy manipulations in the Anna hummingbird

Summary Territorial activity in the Anna hummingbird (Calypte anna) was measured while energy availability on the territory was varied. On days when energy availability was unlimited, residents defended highly exclusive territories primarily by energetically expensive defense behaviors. As energy availability decreased, exclusiveness declined gradually, relative use of energetically inexpensive defense increased, and owners spent less time on the territory.Territorial behavior also varied with short term depressions in energy availability: A lower percentage of intruders was chased and departures of an owner from its territory were more frequent shortly after feeding.When resource dispersion was increased without changing substantially total rewards per territory, chasing by owners increased.     

Matter of Size or Matter of Residency Experience? Territorial Contest in a Green Hairstreak,Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae)

Territorial residents usually win asymmetrical owner‐intruder contests and a variety of hypotheses have been proposed to explain this phenomenon. In the butterfly Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae), male territorial residents defended their territories against intruders during numerous contests and kept them for many successive days. Field observations and experiments were conducted to examine the factors related to this superiority of residents. Forewing length did not differ between residents and intruders, suggesting that body size is not correlated with resource holding potential. Removal–replacement experiments demonstrated that residency did not serve as an arbitrary means for contest settlement, and did not support the recently presented alternative hypothesis that males with higher body temperature are more likely to win. New residents fought longer in defense of the territory as their residence duration in the territory increased. I discuss these results in light of game theory and suggest that the superiority of residents in C. smaragdinus may be based on the asymmetry of resource (territory) value for residents and intruders.     

Chemical signalling and monopolization of nectar resources by territorial Pachymelus limbatus (Hymenoptera Anthophoridae) male bees in Madagascar

Observations on territoriality and the associated odour-marking and foraging behaviours in male Pachymelus limbarus Saussure, a Malagasy endemic anthophorid bee, are reported. Data indicated a monandrous mating system where virgin females are attracted, probably by pheromonal cues, to male-patrolled territories associated with flowering, nectariferous Plectran- rhussensu lato species (Lamiaceae). One male observed in detail applied odour marks every 30–80 minutes on twigs around a food-resource area that contained c. 850 flowers. This marking behaviour always preceded territory residence. The male spent c. 80% of his time patrolling. Meanwhile, any intruding flying insect was inspected or chased, which caused conspecific males, as well as often females, to leave the site. The resident male fed frequently on flowers within his territory (c. 3,600 flower-visits on fine days), except immediately following odour-marking bouts. His flower-visiting was often released by interactions with intruding insects entering the territory. The observations provide the first example of male territoriality based on resource defence polygyny in the bee family Anthophoridae. The importance of resource distribution in the evolution of male territoriality in bees is discussed.     

Intruders in Nests of the Spotless Starling: Prospecting for Public Information or for Immediate Nesting Resources?

The prospective behaviour for nests by conspecific intruders may be a strategy to gather public information for future reproduction or to secure resources for immediate reproduction. Although the relationship between the sex, age and breeding experience of intruders and the sex and age of residents may be crucial for understanding the significance of nest‐prospecting behaviour, a precise determination of these traits has rarely been addressed in field studies. In a free‐ranging population of spotless starlings ( Sturnus unicolor), inexperienced birds were the predominant intruders. The ratio of male‐to‐female intruders was higher among birds without previous breeding experience, although our results did not allow us to determine whether more inexperienced males or females intruded nests. The average annual rate of intrusion of either sex was strongly correlated with the number of fledglings produced in the study colony the previous year, which seems to indicate that the inexperienced intruders were, in general, yearlings. The proportion of the sexes among the intruders in each study year was correlated with the average age of male owners but not with the age of female owners. This result suggests that the presence of intruding males in the nests was affected by an increased opportunity to find vacancies in nesting territories as resident males became older. This finding supports the hypothesis that, in most cases, birds visited nests to obtain personal information about nesting resources. However, the intrusion rates were higher when there were nestlings than during the incubation period, which suggests that collecting public information was also involved in the intruding behaviour. Both sexes regularly attacked intruders, which implies that the intruders inflicted some kind of cost on the owners. Female floaters were more frequently chased, probably because they regularly lay parasitically in conspecific nests.     

Why hummingbirds have such large crops

Summary Male Anna's Hummingbirds (Calypte anna) defend territories that contain a predictable food source, floral nectar. For such a hummingbird, the meal size that maximizes long-term net energy intake is less than the maximal crop volume. Smaller meals must be consumed more frequently, but larger meals increase body mass and therefore flight cost. Individuals without territories or with inadequate territories do not have easy access to nectar and intrude on territories owned by otherC. anna, where they may be chased at nay time. It was predicted that these intruders should minimize the number of potentially risky intrusions necessary for maintenance by ingesting as much nectar as possible whenever they manage to feed without being chased (usually when owners are temporarily absent). Therefore, relative to uninterrupted feeding by owners, uninterrupted intruders should feed longer and take larger meals. Field observations supported these predictions. Intruders apparently filled their crops in all seasons, whereas owners ingested smaller amounts (0.21–0.22 ml) and fed for lengths of time consistent with the prediction of an optimization model (0.21 ml). Thus, owners may energetically optimize meal size whereas intruders fill their crops whenever they are not chased. Under most conditions, hummingbirds only fill their crops one-tenth to one-third full, leading to the question why hummingbirds have such large crops. This study demonstrates that a large crop volume may be of survival value when an individual lacks a territory or has inadequate access to resources and must poach on others' territories.     

Sex Differences in Territorial Behavior Exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta)

Spotted hyenas ( Crocuta crocuta ) are gregarious carnivores that defend group territories against encroachment by neighboring conspecifics. Here we monitored the behavior of members of one clan of free-ranging spotted hyenas during border patrols, 'wars' with neighboring clans, and other interactions with alien intruders, to document differences between the sexes in territorial behavior in this species. We also examined the possibility that the probability or rate of attack on alien hyenas encountered within the clan's territory would vary with the sex of the intruders. Initiation and leadership of most cooperative territorial behaviors were by adult female clan members, although border patrols were occasionally conducted by groups composed exclusively of resident males. The vast majority of alien intruders into the territory of the study clan were males. Resident females were more likely to attack intruding females than intruding males, but hourly rates of aggression directed by females towards aliens did not vary with intruder sex. Resident males were more likely than resident females to attack alien males, and resident males directed significantly higher hourly rates of aggression towards intruding males than females. Although female leadership in most cooperative territorial behaviors distinguishes spotted hyenas from many mammalian carnivores, other sex differences in the territorial behavior of spotted hyenas resemble those documented in other gregarious predators. Sex differences observed in hyena territoriality are consistent with the hypothesis that male and female clan members derive different selective benefits from advertisement and defense of group territories.     

Spawning Behavior and Biparental Egg Care of the Crosshatch Triggerfish, Xanthichthys mento (Balistidae)

Reproductive ecology of the crosshatch triggerfish, Xanthichthys mento (Balistidae) was studied at Hachijojima, Izu Islands, Japan. Males established territories and repeatedly chased females passing nearby. There were 1–3 females in each male's territory before spawning and during egg care. This species spawned in pairs on the sandy bottom. Eggs were scattered and attached to sand particles. Females care for the eggs by blowing water on them and guarding them against intruders, while males helped in guarding. Thus, biparental egg care was observed for 2 days until hatching. Both the males and females disappeared from the territories after the egg care. The reproductive ecology of this species is compared with that of other balistids and the unique features of X. mento are described.     

The Effects of Landmarks on Territorial Behavior in a Convict Cichlid,Amatitlania siquia

Defense of a territory is a primary factor determining the fitness of territorial individuals. Thus, territorial animals use a variety of means to lower the cost of territorial defense. Although landmarks are known to lower the defense costs of territories, little is known about how they could affect fundamental properties of territories, which could ultimately affect the demography, population regulation, individual behavior, and spatial ecology of a territorial species. This field study examined the effects of natural landmarks on territory size, shape, and defensive costs in Amatitlania siquia, a little‐studied member of the convict cichlid group native to Costa Rica and Nicaragua. We investigated the territorial behavior of 103 breeding pairs in a caldera lake in Nicaragua. During 10‐min focal observations, we recorded the locations at which residents chased intruders and then used these locations to outline the territory boundaries. Landmarks affected both size and shape of A. siquia territories. Territory size decreased as the percentage of the boundary at landmarks increased. Territories with <20% of their boundaries at landmarks were approximately circular in shape, while territories with 20–60% of their boundaries at landmarks were on average more elongated. These variations in territory size and shape significantly affected defense costs, with higher numbers of chases per square meter in small and elongated territories. These effects of landmarks on territory size and shape and defense costs are useful to understand the widely seen size and shape variations in naturally occurring territories.     

Behaviour of the female beaugregory damselfish (Stegastes leucostictus)

Much less is known about the behaviour of female beaugregory damselfish than about males of the species. This study was initiated to determine behavioural patterns and interactions of female beaugregories on the back reef of Discovery Bay Marine Laboratory, North shore of Jamaica. Females fed, patrolled, and chased intruders in a significantly larger area and ate significantly more per 15-min observation period than did males. Males spent significantly more time patrolling their territories and chased significantly more total intruders than did females. However, this difference in total chases comes from the fact that males chased the bluehead wrasse, an egg predator, significantly more often than did females while all other species of intruder were chased the same by both sexes. The distance that females travelled from their homesite to court was significantly positively correlated to female body length. This increased distance travelled may relate to mate assessment. Female beaugregory damselfish may be able to be more selective in their choice of mate with increased size and distance travelled.     

Territorial behaviour in an Australian nectar-feeding bird

The red wattlebird, a large Australian honey eater, defends feeding territories from other nectar-feeding birds. One such territory decreased in size and experienced fewer intruders as the density of flowers increased. Daily energy production from the nectar in the territory closely approximated the energy requirements of the territory holder. The bird was probably minimizing energetic costs rather than maximizing energy intake.     

Residency Duration and Shelter Quality Influence Vibratory Signalling Displays in A Territorial Caterpillar

Residents are more likely to win territorial disputes than intruders. One explanation for this prior resident advantage is that residents place a higher value on the resource and are therefore more motivated to win. Although value asymmetry models of animal contests often assume that contestants use information about resource value, information on the proximate cues affecting territorial behaviour is often lacking. We use a simple model system – territorial behaviour in the masked birch caterpillar (Drepana arcuata) to identify factors that affect territorial behaviour. Late instar caterpillars occupy solitary silken leaf shelters, which they defend against wandering conspecifics with a vibratory display. We evaluated how a caterpillar identifies itself as the owner and the factors that influence a resident's motivation to signal. To do so, we conducted three experiments between size‐matched residents and intruders to assess how residency duration and shelter quality independently affected territorial displays during the early stages of a contest. Experiment 1 (Time Exp.) demonstrated that resident signalling rates increase with increased duration on the leaf prior to introducing the intruder. Residents also signal more than intruders after residency periods of 1–3 min and longer, demonstrating that residents gather information about resource value shortly after occupying a leaf. Experiment 2 (Squatter Exp.) aimed to disentangle the effects of time on the leaf and silk accumulation. Squatters (individuals in a shelter made by another) placed for 1–3 min on a leaf containing a full silk shelter signalled more to intruders than did caterpillars placed on a fresh leaf for 1–3 min. Experiment 3 (Shelter Removal Exp.) showed that residents whose shelters had been removed signal less than those occupying an intact shelter, despite an equal length of time investing in them. Our experiment is the first to covary both prior residency duration and territory quality, and we find that the motivation of caterpillars to signal is a function of both of these attributes.     

The social organization of a sexually dimorphic honeyeater: the Crescent Honeyeater Phylidonyris pyrrhoptera,at Wilsons Promontory,Victoria

The social organization of the sexually dimorphic and dichromatic Crescent Honeyeater Phylidonyris pyrrhoptera (Latham 1801) was studied at Wilsons Promontory National Park between January 1994 and January 1997. All breeding attempts at one site were followed during the spring and summer of 1996. Most pairs were multibrooded, with pair-bonds maintained throughout the study. Breeding males held well-defined territories. Females, although generally restricting their activities to within their mate’s territory, visited other males’ territories more frequently than did their mates and did so close to the time of egg laying. Males aggressively chased intruding males, but appeared to permit intrusions by non-mate females. Circumstantial evidence suggests some males may kill neighbour’s nestlings. Female Crescent Honeyeaters are solely responsible for nest construction and incubation. While both sexes attended nestlings, males contributed to parental care significantly less than females. Males also provided significantly less parental care than has been reported for monomorphic species of honeyeater. Single syllable calls are given throughout the year. Peaks in the rate of double and multisyllable calls corresponded with peaks in breeding activity. Only males were observed uttering loud multisyllable calls. In contrast to predictions arising from sexual selection theory and observations of the mating system of sexually dimorphic hummingbirds, the population of Crescent Honeyeaters studied exhibited a socially monogamous mating system. However, the very limited male role in parental care and non-observance of male territory boundaries by females suggest the genetic mating system may not be one of monogamy.     

Territory Defence in Black Redstarts,Phoenicurus ochruros: Effects of Intruder and Owner Age?

European black redstarts, Phoenicurus ochruros, vigorously defend all-purpose territories and exhibit delayed plumage maturation, most subadult males looking exactly female-like in their first breeding season. We tested the hypotheses that such dull subadult male plumages are beneficial in order to reduce aggression of adult males either by deception or by honest signalling of subordinance status, and that, in turn, conspicuous (adult) plumage colorations are able to intimidate contenders because they act as a signal of fighting ability and aggressive motivation. Adult and dull yearling black redstart territory owners were confronted with intruders mimicked by stuffed mounts of either a conspicuous adult or a dull subadult male. Our results do not support the hypotheses tested: dull plumages of young intruders did not reduce aggression from adult territory owners and aggressiveness towards intruders was significantly higher in yearling territory owners as compared with adult owners. Conspicuous intruders did not deter dull territory owners and we found no indications that conspicuous male coloration is a signal of fighting ability or aggressive motivation in this species. Instead, the amount of aggressive response to intruders showed considerable individual variance within age classes regardless of the plumage of the intruder and the status of the owner.     

The social behaviour of the Pin-tailed Whydah Vidua macroura in northern Ghana

This paper describes the pattern of territory usage by male Pin-tailed Whydahs Vidua macroura , and the display postures used by territorial males, intruding males and visiting females are described. Each territory holder used one or two favoured display sites at which he was visited by, and displayed towards females in groups of one to eight. The frequency of display flights and song output reflected the diurnal pattern of visits by females. Each female was courted by up to six males, but the number of copulations observed was too few to indicate the type of mating system used. The operative sex ratio was estimated at 1:6 (females to territory holders), and the results of a removal experiment suggested that a proportion of males was prevented from breeding.     

Effects of familiarity on the behaviour towards intruders in captive common marmosets (Callithrix jacchus)

A study on the behaviour of a group of common marmosets (Callithrix jacchus) towards unrelated intruding conspecifics (juvenile male twins) was performed. The group members' proximity to the intruders during two consecutive study periods was analyzed. During the first period in which the experimental conditions as well as the intruders were unfamiliar, all but one group member were equally likely to stay near the intruders and usually more than three individuals were present simultaneously. During the second period the group members showed different scores of proximity towards the intruders. One individual, an adult male, was in close proximity nearly the whole time. Additionally, the number of individuals simultaneously staying near the intruders was clearly reduced. It is argued that (1) the change in behaviour of the group members was affected by the increasing familiarity and (2) intruder experiments with unfamiliar conspecifics are only of partial value for clarification of the process of immigration.     

Landmark Territoriality in the Neotropical Paper Wasps Polistes canadensis (L.) and P. carnifex (F.) (Hymenoptera: Vespidae)

Male Polistes canadensis and P. carnifex aggregate along crests of prominent ridges in Santa Rosa National Park, Costa Rica. At these sites males of both species defend territories (trees and shrubs) by chasing conspecific rivals. Territories do not contain nests or resources that are collected by females. Chasing by territorial males reduces the amount of time spent by intruders in a territory. I describe and contrast male territorial behavior of both species. Some male P. canadensis are territorial while others in the same area exhibit patrolling behavior, flying from one occupied territory to another. Males of P. carnifex exhibit territoriality only. Patrolling in P. canadensis is an outcome of relatively high male density along the ridge, rendering territories in short supply, as shown by the observation that experimentally vacated territories are seized rapidly by formerly patrolling males. Due to a high intraspecific intrusion rate, territorial male P. canadensis spend less time perching and more time flying and chasing intruders from their territories than do male P. carnifex. Males of these two species also differ in the placement of their territories along the ridgeline; P. canadensis occupy territories in saddles while P. carnifex occupy those at peaktops. I show that this divergent spatial pattern is not maintained by competitive exclusion of either species by the other, and I discuss alternative explanations for their separate spatial distributions. Comparative data suggest that males are territorial because females restrict matings to within territories, and I discuss alternative hypotheses to explain this bias in female behavior.     

Calling as an ownership convention on pied wagtail territories

Winter feeding territories were more profitable to owners than intruders. Even when intruders landed on a territory undetected they failed to achieve a profitable feeding rate because they fed in areas recently depleted by the owner. For intruders therefore, ownership was a sign of unprofitability. Intruder scalled ‘chis-ick’, apparently to find out if a territory was occupied. Owners replied ‘chee-wee’ and this was usually sufficient to cause the intruder to retreat. Only owners called ‘chee-wee’. Owners reacted more aggressively to tape-recorded broadcasts of the ‘chee-wee’ call played on their territory than to broadcasts of ‘chis-ick’.     

Delayed song maturation and territorial aggression in a songbird

Asymmetries in competitive ability can determine the outcome of social interactions in animals and are often expressed through differences in sexual traits. Competitive ability (resource holding potential, RHP), trait expression and ultimately reproductive success may vary with an individual's age or experience. In some species, reproductively mature males delay acquisition of some adult traits and thereby signal their young age. Theory on animal contests predicts that individuals assess the RHP of an opponent relative to their own, such that escalation is more common between evenly matched opponents. Here, we test predictions from this hypothesis that males respond to a territorial intruder based on their RHP relative to the intruder's RHP. We simulated white-crowned sparrows (Zonotrichia leucophrys) intruding into the territory of a recruit or return. Playback of a song repertoire simulating a young male (recruit) elicited a weaker response from established territory holders (return), but a stronger response from recruits. Playback of a single song type simulating an older male elicited the opposite responses. This indicates that males distinguished between simulated young and old intruders based on song, and responded differently depending on their own experience. Our study highlights the possibility that receiver as well as sender traits should be considered when interpreting animal interactions.     

Effect of environmental enrichment upon resource holding power in fish in prior residence situations

Resource holding power (RHP), as expressed by gaining dominance, can be affected by extrinsic and intrinsic factors. Extrinsic factors that increase the RHP include e.g. prior exposure to the contest area. The pay-off asymmetry hypothesis was tested according to which there is an asymmetry in the value the resident's territory has for the resident and intruders, i.e. the resident loses more than intruder opponents when losing the contest over a territory in which it has invested much energy during the exploration of food and shelter resources, during the settlement of conflicts with territorial neighbours, etc. This asymmetry will increase the resident's chance to win the fight over intruders. If the resident-intruder asymmetry is enhanced by presenting resource-rich territories to e.g. fish (structurally diverse or 'rich' aquariums), the hypothesis predicts that the resident's dominance chance (probability to win the fight) will be higher than in structurally 'poor' aquariums. It was found that residents were proportionately more often dominant (fight winners) than intruders in the rich compared to the poor aquariums in all seven tested fish species. This demonstrates that a high territory resource value (aquarium enrichment) significantly facilitates the expression of the resident's dominance advantage (prior residence effect). In contrast to dominance, aggressive behaviour before the dominance settlement did not generally differ between residents and intruders when compared in rich and poor aquariums. This suggests that dominance in itself and aggression prior to dominance settlement are at least partially guided by different motivational systems.