Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Optimal floating and queuing strategies: consequences for density dependence and habitat loss

Field studies of many vertebrates show that some individuals (floaters) do not defend territories even when there is space for them to do so. We show that the evolutionarily stable strategy (ESS) for the threshold territory quality at which floating takes place is that which maximizes the size of the floating population (but not the total population, breeding population, or reproductive output). The ESS is solved separately for two assumptions: whether individuals wait to occupy a single territory or multiple territories and whether queuing rules are strict or if all waiting individuals are equally likely to obtain the next territory. The four combinations of these assumptions all give the same evolutionarily stable population size of both floaters and breeders. At the ESS, only territories with expected lifetime reproductive success (LRS) exceeding 1 should be occupied, which introduces a limit to ideal habitat selection. The behavioral decision to float alters the shape of the density-dependent response, reduces the equilibrium population size, and affects the response of the population to habitat loss. Specifically, the floater: breeder ratio is directly related to average breeding habitat quality, and the floater population size will decrease more than the breeding population size if better than average quality habitat is lost.     

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.     

Social status influences microhabitat selection: breeder and floater Eagle Owls Bubo bubo use different post sites

Social status can be reflected in many aspects of an individual’s behaviour and ecology, including habitat use and conspecific interactions. In territorial species where at least two social groups – breeding birds and non‐territorial floaters – are recognized, the diverse tasks associated with territorial ownership can lead territory holders to behave differently from the non‐territorial part of the population. Territory holders defend their breeding area and reproduce, whereas floating individuals are dispersing and lead a more transient life, during which they do not show any territorial behaviour even when settling in a more or less fixed area (known as the stop phase). As social interactions are based on visual and vocal cues, the use of specific sites for sending and/or receiving signals can be a crucial choice in an animal’s life. By analysing the post‐site selection of Eagle Owl Bubo bubo breeders and floaters during their nocturnal activity, we found that: (1) territory holders selected more visible and dominant posts than non‐territorial floaters; (2) the choice of posts made by floating individuals did not differ between the wandering and stop phases of dispersal; and (3) floating females intruded more frequently than floating males within a breeder’s home‐range. These findings highlight the fact that two social strategies are possible within the same species, depending on an individual’s social status and its related tasks. Breeders could take advantage of visible locations to declare their status as territory holders, whereas floaters could benefit from a more secretive life to wander unnoticed among occupied territories. This secretive life would help floaters to reduce the risks associated with conspecific aggression. Finally, the greater occurrence of floating females within breeders’ home‐ranges can be explained by the fact that female incursions in a breeder’s home‐range are less risky than male intrusions.     

Density-dependent age at first reproduction in the eastern kingbird

Theory predicts that maximal fitness is obtained by individuals who begin to breed immediately upon reaching sexual maturity. However, delayed breeding occurs regularly in some taxa, and in birds and mammals is most often associated with long lifespan and/or limited access to suitable habitats. Delayed breeding is not expected among relatively short-lived species such as migratory passerine birds, but this assumption remains untested in many species. Here we quantify age at first reproduction in an eastern kingbird Tyrannus tyrannus population breeding in an ecological island, and through both observational and experimental approaches, investigate the potential causes for the high frequency of delayed breeding that occurs in this population. Nearly half of the fledged nestlings that returned to the breeding grounds did not breed in their first potential breeding season. Some non-breeders occupied territories, for at least some period, but most remained as non-territorial 'floaters'. Parentage analysis failed to show any reproductive success for female floaters, and only limited success for male floaters, indicating that floating was not a successful reproductive tactic. On the other hand, a strong negative relationship existed between population size and the proportion of young birds that bred in their first year, and non-territorial birds of both sexes quickly filled territory vacancies created by experimental adult removals. Limited breeding habitat and territorial behavior of older birds thus appear to be the main causes of delayed breeding in kingbirds. The frequency of delayed breeding in most species is unknown but of potential significance because failure to incorporate accurate estimates of age at first reproduction in population models may lead to flawed population projections.     

Dispersal, migration, and offspring retention in saturated habitats

We examine the evolutionary stability of year-round residency in territorial populations, where breeding sites are a limiting resource. The model links individual life histories to the population-wide competition for territories and includes spatial variation in habitat quality as well as a potential parent-offspring conflict over territory ownership. The general form of the model makes it applicable to the evolution of dispersal, migration, partial migration, and delayed dispersal (offspring retention). We show that migration can be evolutionarily stable only if year-round residency in a given area would produce a sink population, where mortality exceeds reproduction. If this applies to a fraction of the breeding habitat only, partial migration is expected to evolve. In the context of delayed dispersal, habitat saturation has been argued to form an ecological constraint on independent breeding, which favors offspring retention and cooperative breeding. We show that habitat saturation must be considered as a dynamic outcome of birth, death, and dispersal rates in the population, rather than an externally determined constraint. Although delayed dispersal often associates with intense competition for territories, life-history traits have direct effects on stable dispersal strategies, which can often override the effect of habitat saturation. As an example, high survival of floaters selects against delayed dispersal, even though it increases the number of competitors for each breeding vacancy (the "habitat saturation factor"). High survival of territory owners, by contrast, generally favors natal philopatry. We also conclude that spatial variation in habitat quality only rarely selects for delayed dispersal. Within a population, however, offspring retention is more likely in high-quality territories.     

Density dependence in Golden Eagle Aquila chrysaetos fecundity better explained by individual adjustment than territory heterogeneity

Density-dependence effects acting on fecundity can be explained by two competing hypotheses. The individual adjustment hypothesis (IAH) states that, as population density increases, interference among individuals negatively affects their breeding performance. The second hypothesis, the habitat heterogeneity hypothesis (HHH), proposes that, as more individuals occupy the space available, lower quality habitats are increasingly used, causing average population fecundity to decline. In territorial species, it is often predicted that interference mechanisms (IAH) should be of less importance than spatial heterogeneity (HHH). Here, we test this prediction in Golden Eagles, using 35 years of reproduction monitoring data from a population that has been recolonizing the grounds of a French National Park (Ecrins) in the Alps. During the study period, the Eagle population increased from c. 11 to 41 territorial pairs, providing a good opportunity to explicitly assess fecundity across a gradient of densities. Under the IAH, we expect the fecundity of all territories to diminish as density rises. Under strict HHH, older territories should maintain higher fecundity across time, and a positive relationship between fecundity and the seniority of a territory should be observed. A density-dependent pattern was clearly detected at the population level. At the territory level, the decrease of fecundity was strongly related to population size but not to territory seniority. Fecundity decreased similarly in all territories, suggesting that the IAH better explains the observed pattern. Two alternative mechanisms, related to the IAH, could be at play in this population: (1) negative interference by neighbours and non-territorial Eagles and (2) the contraction of individual territories over time. Our results provide new insights into density dependence in territorial raptors, suggesting that, in addition to habitat heterogeneity, interference mechanisms might actually also play an important role.     

Communal defense of territories and the evolution of sociality

The evolution of group living has attracted considerable attention from behavioral ecologists working on a wide range of study species. However, theoretical research in this field has been largely focused on cooperative breeders. We extend this line of work to species that lack alloparental care (hereafter termed "noncooperative species") but that may benefit from grouping by jointly defending a common territory. We adopt a demographically explicit approach in which the rates of births and deaths as well as the dispersal decisions of individuals in the population determine the turnover rates of territories and the competition for breeding vacancies thus arising. Our results reveal that some of the factors thought to affect the evolution of cooperative breeding also affect the evolution of group living in noncooperative species. Specifically, high fecundity and low mortality of resident individuals both increase the degree of habitat saturation and make joining an established group more profitable for nonresidents (floaters). Moreover, if floaters can forcefully take over territories, the degree of habitat saturation also affects the chance that residents become targets of takeovers. In this situation, communal defense of territories becomes an important benefit that further promotes the evolution of group living.     

Effects of age and experience on the responses of territorial and floater male Red‐winged Blackbirds to models of receptive females

The ability of sexually mature non‐territorial floaters to sire offspring affects the success of floating as a breeding strategy. Red‐winged Blackbirds (Agelaius phoeniceus) have second‐year (SY) and after‐second‐year (ASY) floater males, and genetic studies suggest that floaters may gain paternity. Despite these studies, we still know little about the fitness costs and benefits of floating in this species. By presenting taxidermic models of females in soliciting, precopulatory postures in territories of experienced (previously attracted at least one mate in the study area) and inexperienced (did not previously defend a territory in the study area) males, I was able to examine the copulation behavior and success of floater male Red‐winged Blackbirds as well as the effect of experience for territorial males. Floaters trespassed during 66.1% of presentations and 85.4% of trespassers were SY males. Experienced territorial males (92.5%) and neighbors (87.5%) were most successful in attempts to copulate with models, inexperienced territorial males (62.5%) and ASY floaters (50.0%) had intermediate success, and SY floaters (6.9%) were least successful. Experienced territorial males were more likely to approach models than inexperienced males, and floaters were more likely to approach models in territories of experienced than inexperienced males. These results provide further evidence that floaters trespass frequently, suggest that floaters sire offspring, and demonstrate that prior breeding experience affects the behavior and reproductive success of territorial male Red‐winged Blackbirds. Floating appears to be a conditional strategy for ASY male Red‐winged Blackbirds, but, because it is still not known if SY floaters sire offspring, these males may be trespassing to gain information or experience.     

Defence, intrusion and the evolutionary stability of territoriality

Territorial behaviour can only be adaptive if its costs are outweighed by its benefits. Territorial individuals incur costs by defending their territories against intruders. Usually these intruders are assumed to be non-territorial floaters attempting to take over the whole territory or neighbours trying to extend the borders of their own territory. We instead investigate how costs and benefits of territorial behaviour are affected by neighbours which invade to steal resources on a territory.We show analytically that in the absence of defence intrusion into neighbouring territories always pays and that even if territories are defended intrusion levels can still be high. Using a more detailed simulation model we find that territory defence usually disappears from the population even if owners have a strong advantage over intruders in terms of fighting costs or foraging efficiency. Defence and thus territoriality can only be evolutionarily stable if fighting costs for the intruder relative to the productivity of the territory are very high or if crossing the borders between territories carries additional costs.Our results show that stealing of resources by neighbours can have a considerable effect on the evolutionary stability of territory defence and thus territoriality itself. A more mechanistic model of territorial behaviour is needed to incorporate these kinds of mechanisms into a general theory on the evolution of territoriality.     

Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae)

The tropical damselfly Paraphlebia zoe has two male morphs: a black-winged (BW) male which is associated with territorial defense of oviposition sites; and a hyaline-winged (HW) male similar in appearance to females, and, compared to the black morph, less frequently found defending territories. In a wild population of this species, we first assessed the relationship between phenotypic traits [male morph, size and territorial status (being territorial or non-territorial)], their role on mating success, and the degree to which a particular territory may contribute to male mating success. Second, to relate a physiological basis of being territorial we compared both morphs in terms of muscular fat reserves and thoracic muscle, two key traits related to territory defense ability. Males of both morphs defended territories although the BW males were more commonly found doing this. BW males were larger than HW males and size predicted being territorial but only within HW males (territorial males were larger) but not in BW males. Male mating success was related to territorial status (territorial males achieved a higher mating success), but not to morph or size. Furthermore, territory identity also explained mating success with some territories producing more matings than others. The BW morph stored more fat reserves which may explain why this morph was more likely to secure and defend a place than the HW morph. However, the HW morph showed higher relative muscle mass which we have interpreted as a flexible strategy to enable males to defend a territory. These results are distant to what has been found in another male dimorphic damselfly, Mnais pruinosa, where the advantage of the non-territorial morph relies on its longevity to compensate in mating benefits compared to the territorial morph.     

Responses of territorial and floater male Red‐winged Blackbirds to models of receptive females

Red‐winged Blackbirds (RWBL; Agelaius phoeniceus) have a polygynous mating system and, because territorial males commonly have harems of two to five females, some second‐year (SY) and after‐second‐year (ASY) males do not establish nesting territories, but become floaters. Previous studies have revealed high rates of extra‐pair copulations in this species and that sexually mature male floaters and territory owners do not differ in size, testosterone levels, or reproductive capability, suggesting that floaters may occasionally gain paternity. During May and June 2008, we observed the behavioral responses of floater males to taxidermic mounts (models) of female RWBL placed in a precopulatory position. Floaters intruded into territories during 46% of model presentations, with 20% of intrusions by ASY floaters and 80% by SY floaters. During intrusions, ASY floaters attempted to copulate with models 93% of the time compared to 80% for SY floaters. Copulations were successful during 30% of attempts by ASY males and 25% of attempts by SY floaters. The frequency of intrusions by ASY and SY floaters, attempted copulations by SY floaters, and successful copulations by ASY floaters increased as territorial males spent more time off their territories. Responses of floater males toward models in our study suggest that floater male RWBL attempt to exploit available breeding opportunities. The lack of evidence for extrapair young (EPY) fathered by floater male RWBL in previous studies, combined with our results indicating that the presence of territorial males limits floater intrusions, copulation attempts, and successful copulations, suggests that the reproductive success of floater males is limited in part by the aggressive behavior of territorial males.     

Facing the crowd: intruder pressure,within‐group competition,and the resolution of conflicts over group‐membership

Recent theory in social evolution has been mainly concerned with competition and cooperation within social groups of animals and their impact on the stability of those groups. Much less attention has been paid to conflicts arising as a result of solitary floaters (outsiders) attempting to join groups of established residents (insiders). We model such conflicts over group‐membership using a demographically explicit approach in which the rates of births and deaths in a population determine the availability of group‐vacancies and the number of floaters competing over these vacancies. We find that the outcome of within‐group competition, reflected in the partitioning of reproduction among group members, exerts surprisingly little influence on the resolution of insider‐outsider conflict. The outcome of such conflict is also largely unaffected by differences in resource holding potential between insiders and outsiders. By contrast, whether or not groups form is mainly determined by demographic factors (variation in vital rates such as fecundity and mortality) and the resulting population dynamics. In particular, at high floater densities territory defense becomes too costly, and groups form because insiders give in to the intruder pressure imposed on them by outsiders. We emphasize the importance of insider‐outsider conflicts in social evolution theory and highlight avenues for future research.     

A general model of site-dependent population regulation: population-level regulation without individual-level interactions

In this paper, we use a modeling approach to explore the population regulatory consequences of individual choices for where to breed in heterogeneous environments. In contrast to standard models, we focus on individuals that interact only indirectly through their choices of breeding sites (i.e., individuals preempt the occupation of a breeding site by others when they choose to breed there). We consider the consequences of individuals choosing breeding sites either randomly or sequentially from best to worst. Our analysis shows that average per-capita fecundity of the population is independent of the number of occupied breeding sites if individuals choose sites at random and that variation in average per-capita fecundity increases as population size declines. In contrast, if individuals choose breeding sites sequentially from highest to lowest quality, then as population size increases average per-capita fecundity declines and variation in average per-capita fecundity increases. Consequently, aggregate population-level demographic rates can change in ways that generate population regulation, even when change in population size does not change the demographic performance of any individual on any particular breeding site. However, such regulation occurs only when individuals make adaptive choices of where to breed. Because variation in average per-capita fecundity decreases when population size declines, populations regulated in a site-dependent manner should be much less susceptible to the vicissitudes of small population size than those which choose breeding sites at random.     

Form,function and consequences of density dependence in a long‐distance migratory bird

The density dependence of demographic parameters and its implications for population regulation have long been recognized. Recent work has revealed potential effects of density on mating systems and sexual selection, but few studies concurrently assess the consequences of density on both demography and sexual selection. Such an approach is important because population processes and individual behaviors can interact to influence population growth and evolutionary trajectories. In this study, we tested the density dependence of breeding success, extra‐pair paternity, and the opportunity for sexual selection in a population of American redstarts Setophaga ruticilla using two different measures of density. To evaluate temporal patterns, we analyzed annual territory density, based on the total number of territories at our study site each year. To evaluate spatial patterns, we analyzed local territory density within years, based on the number of territories surrounding a focal territory. Greater annual density was associated with fewer offspring fledged per female, a reduced mean population rate of fledging success, and a lower relative contribution of extra‐pair paternity to male fitness. Greater local density was associated with fewer offspring fledged, reduced fledgling success, higher rates of nest loss, and higher rates of paternity loss on focal territories. Interestingly, greater local density was also associated with greater nestling mass on focal territories, which could imply that more densely‐packed territories contain superior resources. Overall, our results suggest that the effects of crowding via greater territory density reduce fecundity through increased nest predation, rather than reduced food availability, and increase rates of extra‐pair paternity. Thus, the selective pressures faced by individuals and their reproductive behaviors are likely to differ based on the annual and local density they experience, which may then feed back into population demography.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.     

Survival of the Orange-tufted Sunbird,Nectarinia osea

Cape Sugarbirds are southern African endemics, found in fynbos vegetation of the Western and Eastern Cape Provinces of South Africa, and they feed almost exclusively on the nectar of Protea inflorescences. Breeding male Sugarbirds are highly territorial and defend resources for themselves, their mates and their offspring. Sugarbirds, however, often leave their breeding territories in search of food during the dry season and return to the same breeding sites each year, thus experiencing an annual cycle of movement from one food source to another. Male territory size as well as breeding success, were determined over two breeding seasons for a population in the Helderberg Nature Reserve in the Western Cape Province. Sugarbird fledgling success was significantly greater for males defending large territories. Males moved or increased the size of their territories between seasons, possibly to improve breeding success, but no changes in territory size were observed during the breeding season.     

Effect of competitor abundance on feeding territoriality in a grazing fish,the ayuPlecoglossus altivelis

The grazing fish, ayu,Plecoglossus altivelis Temminck & Schlegel, establishes feeding territorialiry during the young stage. The population density fluctuates from year to year by more than a hundredfold, but the determinant of territory size is less well known. The feeding territoriality of ayu was examined under simulated habitat conditions where fish density was manipulated and food resources were renewable. Fish competed for algae attached to the substrata and were divided into residents with territories, and floaters without territories. By experimental alteration of fish density the number of residents increased with density and rerritory size decreased with density. Floaters intruded into territories in a school to feed on algae, which induced overt aggression of the resident and reduced the productivity of algae growing there. Both the intruding frequency of floaters over territorial areas and their feeding pressure on algae increased at higher floater density. Floaters functioned to shift cost-benefit relationships for various territory sizes. They acted as food competitors to restrict territory size below a maximum through competitive interference. Although the growth rate of residents was inversely related to fish density, residents grew faster than floaters in each group. Under a given set of competitor abundances, economic defensibility determined territory size.     

Multiple density-dependence mechanisms regulate a migratory bird population during the breeding season

The mechanisms regulating bird populations are poorly understood and controversial. We provide evidence that a migratory songbird, the black-throated blue warbler (Dendroica caerulescens), is regulated by multiple density-dependence mechanisms in its breeding quarters. Evidence of regulation includes: stability in population density during 1969-2002, strong density dependence in time-series analyses of this period, an inverse relationship between warbler density and annual fecundity, and a positive relationship between annual fecundity and recruitment of yearlings in the subsequent breeding season. Tests of the mechanisms causing regulation were carried out within the Hubbard Brook Experimental Forest, New Hampshire, during 1997-1999. When individuals from abutting territories were experimentally removed in a homogeneous patch of high-quality habitat, the fecundity of focal pairs nearly doubled, revealing a locally operating crowding mechanism. A site-dependence mechanism was indicated by an inverse relationship between population size and mean territory quality, as well as by greater annual fecundity on the sites that were most frequently occupied and of highest quality. These site-dependence relationships were revealed by intensive monitoring of territory quality and demography at the landscape spatial scale. Crowding and site-dependence mechanisms, therefore, acted simultaneously but at different spatial scales to regulate local abundance of this migratory bird population.