Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

Influence of territoriality on adult density in two rockfishes of the genus Sebastes

Nonterritorial Sebastes carnatus and S. chrysomelas existed, along with territorial individuals, at 3 tagging sites off southern California, USA, which were monitored for nearly 1 yr. To test the hypothesis that territoriality affected adult density in these species, territorial fish were removed and the subsequent utilization of vacated territories by other fish was monitored. Intrusion of other fish into vacated territories increased significantly in 90% of the removals. Other fish colonized both the feeding and sheltering parts of the vacated territories, indicating that the previous owners had successfully defended both parts of their territories. Many of the colonizers had already possessed territories; they expanded their territories or moved into presumably better havitat. Several previously-nonterritorial fish also moved into vacated areas, and at least some of them appeared to establish territories. These fish, then, had previously been capable of establishing territories, but were prevented from doing so by resident territory holders. Thus territoriality, rather than such other factors as predation or low recruitment, limited the number of territorial fish at each site. However, territorial fish did not inhibit the settlement of larval recruits, and the relative mortality rates of older territorial vs nonterritorial fish were not determined. Thus the question of whether territoriality was a major factor controlling total density remains unresolved.     

Site fidelity and territorial movements of males in a rapidly declining population of yellow-headed blackbirds

We tested several hypotheses to explain low between-year territory fidelity in a breeding population of yellow-headed blackbirds (Xanthocephalus xanthocephalus). During a 5-year study the population of territorial males declined by two-thirds and some of the marshes that supported territories significantly deteriorated. Individual males held territories and bred for an average of 1.9 years. Of males that bred for at least 2 years, 30% skipped owning a territory in the study area during at least 1 year of their breeding lifetimes. Our information suggests that they may have bred outside of the area in those years. Of males with territories in two or more breeding seasons, 60% changed breeding marshes at least once. Males changed territories during 42.9% of between-year opportunities to do so. We found no support for the hypotheses that male yellow-headed blackbirds: (1) are more likely to move when territory density is low; (2) are likely to abandon territories that are deteriorating; or (3) change territories to improve their reproductive success. We suggest three non-mutually exclusive explanations for the yellow-headed blackbird's weak site fidelity: (1) it is a response to habitat deterioration and to other factors that may be causing the population's decline; (2) the males, being migratory, make fresh settlement decisions each year after they arrive on the breeding grounds in the general vicinity of their previous year's breeding; (3) yellow-headed blackbirds may have evolved in, and be adapted to, highly unstable habitats, moving frequently in response to changes in local breeding site conditions. Correspondence to: L.D. Beletsky     

Territorial behavior of the black and yellow rockfish and gopher rockfish (Scorpaenidae,Sebastes)

Tagged black-and-yellow rockfish Sebastes chrysomelas and gopher rockfish S. carnatus were observed in situ for 1 yr at 3 study sites off California, USA in order to determine the nature of their home ranges and territories. Three basic patterns of space use were identified: (1) a shelter hole located within a larger area of activity (in which the fish presumably fed, since its size increased with size of fish and decreased with prey density); (2) discontinuous home ranges, in which the shelter holes and feeding areas were separte; and (3) dispersed home ranges with little or no exclusive area. Fish in the first category were regarded as territorial, since their home ranges were defended and relatively exclusive. Because individuals did not patrol borders, however, territories often overlapped at their peripheries. In addition, some territory holders appeared to tolerate the sheltering of other fish within their territories, as long as these intruders left the territory during periods of activity. Some fish with discontinuous home ranges (commuters) may have been territorial, but others may have been forced to occupy suboptimal feeding areas. Fish in the third category (floaters) were regarded as nonterritorial; their activities were apparently limited to areas where they could escape the aggression of territory holders. The varying patterns appeared to develop through defense and accommodation during the turnover of individuals, with each fish securing shelter and feeding areas as these became available.     

Territory quality and territorial behavior in two sympatric species of hummingbirds

Summary Changes in territorial behavior of blackchinned hummingbirds (Archilochus alexandri) in response to experimental changes in territory quality were investigated using artificial feeders and simultaneous, pair-wise observations of owners. Some of the responses of A. alexandri were similar to those documented by a previous study of the Anna's hummingbird (Calypte anna): A. alexandri defending rich territories spent more time on their territories, encountered a greater percentage of intruders, and chased more intruders than did A. alexandri defending poor territories. In contrast to C. anna, A. alexandri supplemented chases with energetically inexpensive threat vocalizations more extensively when territory quality was increased. This difference may be related to A. alexandri's more tenuous control of rich territories. When both species were observed on very poor territories, A. alexandri chased a greater percentage of intruders, consumed a greater proportion of available food, and obtained a greater net energy gain from their territories. When observed simultaneously on territories with ad lib food, both species defended highly exclusive territories but A. alexandri suffered higher intrusion pressure and spent more time and energy in defense. These interspecific differences in territorial behavior may facilitate sympatric coexistence of the two species through aggressive partitioning of flower patches according to productivity: the greater payoff received by C. anna on rich territories and A. alexandri on poor territories should favor a corresponding monopolization of rich areas by C. anna and poor areas by A. alexandri.     

Territory quality affects female preference in a Lake Victoria cichlid fish

Mate preferences on male colour have been implicated in generating and maintaining species diversity among haplochromine cichlid fish. Their lek-like mating system suggests that not only male colour but also territory quality is instrumental in mate choice. We assessed the relative importance of territory quality and male colour in mate choice by testing whether territory quality can override the female preference for males of her own colour in the Lake Victoria cichlid genus Pundamilia. First, we showed in experimental groups that the dominant male preferentially monopolised a large tube relative to a small tube. The situation mimics quality difference in rocky crevices that serves as a focal point for male courtship display. Second, in mate-choice tests, Pundamilia nyererei females were allowed to choose between closely related P. nyererei and P. pundamilia males; these species differ strikingly in male nuptial coloration, but little else. We gave either both males the same small tube or one of them a large tube. The preference of P. nyererei females for P. nyererei males in the control situation (where both males had a small tube) was significantly diminished in favour of P. pundamilia males when the latter had the large tube. The results provide experimental evidence that differences in territory quality can override the female preference for males of her own colour. This finding is critical for a recent hypothesis proposing that male competition for mating territories can facilitate the process of sympatric speciation by sexual selection.     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Relative importance of male and territory quality in pairing success of male rock ptarmigan (Lagopus mutus)

We studied pairing success in male rock ptarmigan (Lagopus mutus) in northern Alaska to learn whether males obtaining more females possessed phenotypic traits that influenced female choice directly, whether these traits permitted males to obtain territories favored by females, or whether both processes occurred. The number of females per male varied from zero to three. Several male and territory traits were significantly correlated with number of females per male. We used multiple regression to obtain a single measure of male quality and a single measure of territory quality. These measures of male and territory quality correlated with each other and with male pairing success. We used path analysis to separate direct effects of male quality on pairing success from indirect effects due to high-quality males obtaining high-quality territories. Both direct and indirect pathways had significant effects on pairing success, and direct and indirect effects of male traits on pairing success were about equal. This study illustrates an analytical approach for estimating the relative importance of direct and indirect causal relationships in natural systems. Received: 13 January 1998 / Accepted after revision: 7 November 1998     

Long-term mate and territory fidelity in neotropical buff-breasted wrens (Thryothorus leucotis)

Among sedentary species, mate and site fidelity may be imposed by year-round territoriality and high annual survivorship, both of which theoretically lead to few opportunities to move between mates and breeding territories. We examined mate and territory fidelity in sedentary neotropical buff-breasted wrens (Thryothorus leucotis) in relation to the opportunities to divorce (vacancies on neighbouring territories), breeding experience and site of origin (within vs outside local breeding population). Overall, buff-breasted wrens showed high rates of mate and territory fidelity between years, with individuals significantly more faithful to their territories than to their partners. Mate and territory fidelity was not forced upon buff-breasted wrens, as most individuals had opportunities to switch mates and territories. However, this was not uniform among all individuals. Individuals that had bred together displayed 100% mate and territory fidelity despite the fact that males and females in almost all experienced pairs had opportunities to switch mates and territories. Local recruits (philopatric offspring that acquired territories in the local breeding population) also displayed mate and territory fidelity, but fidelity occurred because recruits rarely had the opportunity to divorce prior to initiating their first breeding attempt. By contrast, almost half of all territorial individuals that originated outside the local breeding population (immigrants) had opportunity to divorce, with approximately two thirds of them switching mates and territories. Divorces occurred within the first 5 months of pairing, suggesting that immigrants may have sampled mates and territories before forming permanent partnerships.     

Predation risk and the territory value of cover: an experimental study

In an experiment on territorial brown trout (Salmo trutta), we addressed the novel hypothesis that protective cover increases the value of a territory in relation to the perceived level of predation risk. We predicted that territory holders should invest more resources defending territories with cover than territories without cover, and that defence should increase as predation risk increases. First, trout were allowed to establish ownership in territories with or without overhead cover. Second, predation risk was manipulated by simulating aerial predator attacks in half of the territories of each type, whereupon the preference for cover was estimated. Third, owners of the four types of territories were staged in dyadic contests against size-matched intruders. Territory owners showed a preference for cover, which increased further after simulated predator attacks. In subsequent contests, conflicts over territories with cover were settled faster than conflicts over territories without cover, which may suggest that the value of cover increases the motivational asymmetry between owner and intruder. Consistent with our hypothesis, owners of territories with cover were much more aggressive if they had been subjected to predator attacks the day before the conflict. These results suggest that territory owners are able to estimate the value of protective cover in response to variation in the level of predation risk in the habitat.Communicated by J. Krause     

Deceptive behavior in pied flycatchers

Summary One explanation for polygyny in pied flycatchers (Ficedula hypoleuca) is the deception hypothesis, which proposes that females mate with already-mated males only because they are unaware that such males already have a mate on another territory. Recently this hypothesis has been criticized on the grounds that already-mated and unmated males differ sufficiently in their behavior that human observers can discriminate between the two classes fairly easily. Here we test whether male pied flycatchers change their behavior when visited by a female so as to make this discrimination more difficult. In our experiments we presented a caged female near the nestbox of an advertising male, in order to mimic the situation in which a female investigates the male and his nest site. During control periods, when only an empty cage was presented, we found that already-mated males were present on their secondary territories significantly less and sang significantly less than unmated males on their primary territories, confirming the earlier results. When we presented the stimulus female, all behaviors we measured changed significantly relative to control periods for both mated and unmated males: both classes of males increased the time spent on the territory, decreased singing rates, and increased various courtship behaviors. During the experimental periods there were no significant differences between mated and unmated males on any of the behavioral measures. Discriminant analysis was more successful in classifying males as to mating status using data from the control periods (without female) than using data from the experimental periods (with female). We conclude that when a female is present male pied flycatchers change their behavior in ways that make it more difficult to discriminate mated from unmated males.Offprint requests to: W.A. Searcy     

Nepotistic access to food resources in cooperatively breeding carrion crows

Offspring delayed dispersal is the principal mechanism leading to formation of kin-based societies. It has been suggested that parents promote offspring philopatry by providing them with preferential access to the food resources of the territory and that parental tolerance may be affected by territory quality. However, few studies have addressed this hypothesis in kin-living vertebrate species. Here, we show that in cooperative breeding groups of carrion crows (Corvus corone corone) containing retained offspring and immigrants, dominant breeding males behaved nepotistically on an experimental source of food by (1) attacking immigrants with more frequency and intensity than offspring and (2) associating preferentially with their offspring on the feeding spot and sharing food with them. This parental facilitation allowed the offspring to spend more time feeding than higher-rank immigrants. We also found that a year-round experimental food supplementation neither increased breeding males’ tolerance nor relented the overall aggressiveness in the groups. This indicates that higher natal philopatry observed on fed territories compared to unfed ones is not a consequence of a more benign social environment. Rather, it suggests that offspring value territory resource wealth and adjust the timing of dispersal accordingly.     

Alternative mating strategies in the water strider Gerris elongatus (Heteroptera,Gerridae)

Summary Males of the water strider Gerris elongatus established territories which included copulation and oviposition sites (small pieces of fallen bamboo). Males were aggressive and competition for territory and females was observed frequently. Male midlegs were more developed than female midlegs and were used as weapons. Reproductive behaviour changed as the breeding season advanced. Early in the season immature females were attracted by male surface wave courtship signals, then copulated white floating on the water surface without ovipositing (type 1). In midseason, males established territories, produced calling signals and attracted females which copulated and oviposited there with male postcopulatory guarding (type 2). In late season, many females oviposited without postcopulatory guarding on pondweed mats near fallen bamboo. Non-territorial males waiter for the arrival of these females and copulated without courtship, but mating success was low (type 3). These alternative mating strategies appeared to depend on differences in male size. Larger males were superior to smaller males in many ways (establishing territory, fighting, mating etc.). The largest males defended territories and had higher mating success than small non-territorial males. Medium sized males used all three strategies according to the number of empty territories and seasonal femald distribution.     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Use of space by Gobius cruentatus and Thorogobius ephippiatus (Teleostei: Gobiidae) in south-west Ireland

The mechanisms of spatial organisation exhibited by two co-existing species of Gobiidae, Gobius cruentatus Gmelin, 1789 and Thorogobius ephippiatus (Lowe, 1839) were examined to determine the nature of, and the factors contributing to, these mechanisms. This study was carried out in Lought Hyne, south-west Ireland, during 1990. Both species were found to occupy territories exclusive to conspecifics, whilst G. cruentatus was also found to maintain a territory exclusive to T. ephippiatus. This and other interspecific differences in the spatial patterns exhibited by these species are suggested to be related to size-based dominance abilities and their differential requirements of space-related resources.     

Ontogeny of space use and diet of two temperate damselfish species, <Emphasis Type="Italic">Parma microlepis</Emphasis> and <Emphasis Type="Italic">Parma unifasciata</Emphasis>

Studies of reef fish herbivory have mainly focused on the impacts and behaviour of adults of tropical species. In this study, the ontogenetic shifts in home range, aggression, feeding rate, diet and gut morphology in juveniles and adults of two temperate territorial damselfishes, Parma microlepis and Parma unifasciata, were determined. Both P. microlepis and P. unifasciata juveniles under 80 mm TL exhibited no aggressive chases towards conspecifics or other species, while above 80 mm TL aggressive chase frequency increased in conjunction with an increase in home range, defended as a territory. Ontogenetic diet shifts, characterised by an increase in herbivory (P. unifasciata: juveniles: 64% plant material, adults: 95% plant material; P. microlepis: juveniles: 43% plant material, adults: 67% plant material) were observed for both species. The ratio of digestive tract length to body length, which often accompanies a switch to herbivory, increased significantly with ontogeny for both species. Compared to tropical confamilial grazers, these temperate damselfish species feeding rates were lower, and they had larger territories which were not as strongly defended (fewer aggressive chases).     

Territory characteristics and coexistence with heterospecifics in the Dartford warbler <Emphasis Type="Italic">Sylvia undata</Emphasis> across a habitat gradient

The study of successional gradients may help to understand the relative influence of habitat structure and competition on territory characteristics. Here, we evaluate the effects of vegetation cover, conspecific and heterospecific densities, and distance to the nearest neighbor on territory size, shape, and overlap in insectivorous birds. We studied these effects along a gradient of postfire habitat regeneration in which foliage cover and densities of focal species varied several-fold. We delineate 197 territories (minimum convex polygons) of the shrub-dwelling Dartford warbler (Sylvia undata) and 255 of the syntopic Sardinian (Sylvia melanocephala), subalpine (Sylvia cantillans) and melodious (Hippolais polyglotta) warblers at three plots in NE Catalonia (Spain and France) in 1987–2005. After accounting for the effect of the number of locations used to delineate polygons, generalized linear mixed models (GLMM) showed a reduction in territory area of the Dartford warbler as conspecific density increased and distance to nearest neighbor decreased, in accordance with the contender pressure hypothesis for territory size regulation. Heterospecific density was not included in the final model of territory size and the effect of habitat structure was marginal. Territory roundness was positively correlated with its size and with conspecific density, probably in relation to energetic constraints, and negatively with heterospecific density. Territorial exclusion was almost complete among Dartford warblers, whereas interspecific territory overlap was extensive and tended to increase with heterospecific density and with structural diversity along the gradient. Our results support the hypothesis that Mediterranean warbler coexistence derives from ecological segregation and not from interspecific territoriality.     

Determinants of male mating success in the red bishop (Euplectes orix)

We studied sexual selection in the red bishop, Euplectes orix, a colonial, polygynous weaverbird widely distributed over sub-Saharan Africa. Male reproductive success measured in terms of the number of nests accepted by females and the number of eggs and nestlings in all the nests on a male's territory varied considerably. The standardized variance (variance/mean²) in male reproductive success ranged from 0.505 to 1.737 in different years, indicating a high potential for sexual selection in this species. An analysis of genetic parentage for 432 nestlings by non-radioactive, multilocus DNA fingerprinting confirmed that male reproductive success (number of young sired on the territory) in this species can be reliably estimated by the measures introduced above. In all 4 study years there was a strong positive correlation between male mating success and the total number of nests that males built in their territories. The number of nests built can be partitioned into the number of weeks a male held a territory and his nest-building performance. Both factors exert a significant positive effect on male mating success and in combination explained between 53.3 and 86.3% of the variation in male reproductive success. Male morphological characters were found to be of no importance. Males that established a territory in the following season built more nests and held their territories for longer than males that did not establish a territory in the following season, suggesting that these measures might be indicators of male condition and quality. Male nest-building performance (number of nests built per week) seems to be unrelated to male condition or quality. Received: 8 January 1999 / Received in revised form: 7 June 1999 / Accepted: 13 June 1999     

Territory defence in tropical birds: are females as aggressive as males?

Much of our knowledge concerning the functions of territorial behaviour and how territories are defended by individuals comes from research on birds. The vast majority of this work has focused on temperate zone breeding territoriality in which territories are defended most obviously by males. Our understanding of the female role in territory defence is limited because they are less conspicuous and much harder to observe. We studied sex roles in territory maintenance and defence in a duetting, resident neotropical passerine, the white-bellied antbird (Myrmeciza longipes). This species maintains territories and pair bonds year round and both sexes sing and actively participate in territory defence. We performed a series of playback experiments throughout the dry (non-breeding) and wet (breeding) seasons. We exposed territorial pairs to three types of stimuli including: (1) single sex, male only songs, (2) single sex, female only songs, and (3) both sex songs/duets. Contrary to findings for most other tropical species, individuals defended their territories with equal levels of aggression regardless of stimuli. Furthermore, sex roles were very different, with males responding more aggressively than females to all stimuli throughout both seasons. Both males and females consistently responded more aggressively to territorial intrusions during the dry season than during the wet season, likely because food abundance is low in the dry season and territory value is high. Our analysis of duetting behaviour suggests that duets do not serve a significant role in mate guarding, or territory defence.     

Territory choice during the breeding tenure of male sedge warblers

A territorial male can shift the location of its territory from year to year in order to increase its quality. The male can base its decision on environmental cues or else on its breeding experiences (when territory shift is caused by breeding failure in previous seasons). We tested these possible mechanisms of territory choice in the sedge warbler (Acrocephalus schoenobaenus), a territorial migrating passerine that occupies wetlands. This species bases its territory choices on an environmental cue: tall wetland vegetation cover. We found that the magnitude of territory quality improvement between seasons (measured as the area of tall wetland vegetation) increased throughout the early stages of a male's breeding career as a result of territory shifts dependent on the earliness of arrival. The distance the territory was shifted between seasons depended negatively on the previous year's territory quality and, less clearly, on the previous year's mating success. On the other hand, previous mating or nesting success had no influence on territory quality improvement between seasons as measured in terms of vegetation. The results imply that tall wetland vegetation is a long-term, effective environmental cue and that a preference for territories in which this type of landcover prevails has evolved into a rigid behavioral mechanism, supplemented by short-term individual experiences of breeding failure.