The effect of prey consumption on territorial defense by harriers: differential responses to neighbors versus floaters

Summary Food consumption may reduce fighting intensity of territory owners by decreasing resource value (additional food cannot be consumed) and/or increasing fighting costs (food in the digestive tract may raise injury risks). A territorial harrier's (Circus cyaneus, adult females) decision to reduce its level of aggression should depend upon whether or not the intruder was a competitor for individual prey items (as are smaller male floaters) or for the territory proper (as are female floaters and especially female neighbors). Accordingly, following meals, aggressive intensity of owners was strongly reduced towards male floaters (more were ignored), slightly reduced towards female floaters (more were called at than chased), and remained unchanged towards neighbors (virtually all were chased). Hence, alterations in aggressive behavior of territory owners following food consumption may depend upon the type of intruder and the resource under contest (a food item or a territory).     

There is a limbo under the moon: what social interactions tell us about the floaters’ underworld

The ultimate and proximate causes of natal dispersal have been extensively investigated, but the behaviour of dispersers in relation to social interactions has been largely neglected. Here, we investigated the social organisation of floating individuals during their dispersal by analysing the behaviour of 40 radio-tagged eagle owls Bubo bubo during the wandering and stop phases of dispersal. Unexpectedly, eagle owl floaters mixed with conspecifics independently of their sex, age, phase of dispersal, birthplace, health status and habitat features, showing an ‘underworld’ of interactions characterised by the absence of obvious social organisation or short-term strategies. Non-breeding owls were not transient floaters that occurred at numerous sites for short periods of time but rather had fairly stable home ranges: they attempted to settle as soon as possible within well-defined home ranges. The spatial distribution pattern of floaters and high rates of home range overlap support the prediction that floating individuals are not spatially segregated, challenging the expectation that dominance by size, age and/or health status may determine the exclusive use of some portions of the dispersal area. Finally, (1) the short distances among conspecifics and the extensive home range overlaps allowed us to discard the possibility that neighbouring floaters represent a real cost during dispersal and (2) floater interactions showed a lack of clear mechanisms for avoidance of kin competition among offspring or inbreeding.     

Northern harriers on feeding territories respond more aggressively to neighbors than to floaters

Summary Territory owners often respond less aggressively towards intruding neighbors than towards intruding floaters, an observation termed the dear enemy phenomenon. Comparisons of territory owners' responses to intruding neighbors versus their responses to intruding floaters usually have been made for owners of multi-purpose and/or breeding territories. Here, I describe responses of female northern harriers Circus cyaneus (owners) on winter feeding territories towards three types of intruders (female neighbors, female floaters, and male floaters) and show that the dear enemy phenomenon does not occur. Owners' responses towards neighbors were more intense (mostly flights rather than calls) than responses towards female floaters, which in turn were more intense than responses towards male floaters. The greater intensity of owners' responses towards neighbors compared to owners' responses towards male and female floaters may be related to differences in the threat posed by each of the three intruder types in terms of fighting ability (RHP) and potential losses from intrusion. Hence, whether owners respond more aggressively towards neighbors or floaters, and whether the dear enemy phenomenon is observed, may depend upon the relative magnitude of threat presented by neighbors and floaters to owners in terms of fighting ability and potential losses from intrusion.     

Owner and floater red-winged blackbirds: determinants of status

Summary We used sequential removal experiments to test whether the resource-holding potential (RHP) of territory owner red-winged blackbirds, Agelaius phoeniceus, was superior to that of their first replacements (shallow floaters) and subsequent replacements (deep floaters). Among the removals were secondyear males, which were morphologically inferior to adults and which also tended to be competitively inferior in aviary contests. The highest proportion of secondyear males occurred in the deep floater class. Thus, the RHP of some deep floaters was inferior to that of owners and shallow floaters. However, among adults, owners, shallow floaters, and deep floaters had equivalent morphological and competitive RHP. Furthermore, replacement males that had defended territories for many days were neither morphologically nor competitively superior to males that had defended territories for only a few days. Our results suggest that RHP distinguishes adults from second-year males, but does not separate owners from floaters. The only hypothesis that is potentially supported by our observations is that owners have a greater expected payoff from their territory than intruders, and in this way owners are able to maintain site dominance. The nature of that payoff remains to be determined. Offprint requests to: P.J. Weatherhead     

Spatial scale of local breeding habitat quality and adjustment of breeding decisions

Experimental studies provide evidence that, in spatially and temporally heterogeneous environments, individuals track variation in breeding habitat quality to adjust breeding decisions to local conditions. However, most experiments consider environmental variation at one spatial scale only, while the ability to detect the influence of a factor depends on the scale of analysis. We show that different breeding decisions by adults are based on information about habitat quality at different spatial scales. We manipulated (increased or decreased) local breeding habitat quality through food availability and parasite prevalence at a small (territory) and a large (patch) scale simultaneously in a wild population of Great Tits (Parus major). Females laid earlier in high-quality large-scale patches, but laying date did not depend on small-scale territory quality. Conversely, offspring sex ratio was higher (i.e., biased toward males) in high-quality, small-scale territories but did not depend on large-scale patch quality. Clutch size and territory occupancy probability did not depend on our experimental manipulation of habitat quality, but territories located at the edge of patches were more likely to be occupied than central territories. These results suggest that integrating different decisions taken by breeders according to environmental variation at different spatial scales is required to understand patterns of breeding strategy adjustment.     

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     

Maintenance of variable responses for coping with wetland drying in freshwater turtles

Aquatic animals inhabiting temporary wetlands must respond to habitat drying either by estivating or moving to other wetlands. Using radiotelemetry and capture mark recapture, we examined factors influencing the decisions made by individuals in a population of freshwater turtles (Chelodina longicollis) in response to wetland drying in southeastern Australia. Turtles exhibited both behaviors, either remaining quiescent in terrestrial habitats for variable lengths of time (terrestrial estivation) or moving to other wetlands. Both the proportion of individuals that estivated terrestrially and the time individuals spent in terrestrial habitats increased with decreasing wetland hydroperiod and increasing distance to the nearest permanent wetland, suggesting behavioral decisions are conditional or state dependent (i.e., plastic) and influenced by local and landscape factors. Variation in the strategy or tactic chosen also increased with increasing isolation from other wetlands, suggesting that individuals differentially weigh the costs and benefits of residing terrestrially vs. those of long-distance movement; movement to other wetlands was the near universal strategy chosen when only a short distance must be traveled to permanent wetlands. The quality of temporary wetlands relative to permanent wetlands at our study site varies considerably and unpredictably with annual rainfall and with it the cost-benefit ratio of each strategy or tactic. Residency in or near temporary wetlands is more successful during wet periods due to production benefits, but movement to permanent wetlands is more successful, or least costly, during dry periods due to survival and body condition benefits. This shifting balance may maintain diversity in response of turtles to the spatial and temporal pattern in wetland quality if their response is in part genetically determined.     

Male Iberian rock lizards may reduce the costs of fighting by scent matching of the resource holders

Animals should adopt strategies to minimize the costs of intraspecific aggressive interactions. For example, individuals should be able to identify resource holders in advance and avoid fighting with them because residents are generally more likely than intruders escalate aggression. It has been suggested that scent marks function mainly to allow competitor assessment by conveying the costs of entering a scent-marked area. Individuals may identify territory owners by comparing the scent of substrate marks with the scent of any conspecific they encounter nearby, assessing whether these two scents match or not, a mechanism known as scent matching. Here, we examined the response of male Iberolacerta cyreni lizards to areas scent-marked by other males and the potential role of scent matching in agonistic interactions. We designed a laboratory experiment where we allowed a male to explore the scent-marked substrate of another male, and then we immediately staged agonistic encounters in a nearby clean neutral area with either the male that had produced the scent marks (matching treatment) or with a different non-matching individual male. The higher chemosensory exploratory rates of substrate scent marks in comparison to clean substrates suggested that males detected and spent more time exploring scent marks to obtain information on the donor male. Moreover, this information was later used to decide the fighting strategy. Intruding males delayed time until the first agonistic interaction, reduced the intensity of fights and the number of aggressive interactions, and won less interactions with males which scent matched that of scent marks (because they would be considered as the territory owners) than with other non-matching individuals. Our results show that male I. cyreni lizards use scent matching as a mechanism to assess the ownership status of other males, which could contribute to modulate intrasexual aggression, reducing costs of agonistic interactions.     

Escape from the cell: Spatially explicit modelling with and without grids

This paper is concerned with the representation of individuals embedded in a two- (or three-) dimensional environment, and with the techniques that can be used to simulate the evolution of the spatial patterns both of the populations of those individuals and of their environment. Its scope is therefore that of individual based or agent based modelling, of a general type, including herbivore populations, predator-prey models or any other type that is concerned with the spatial patterning evolving from recruitment, interaction and/or movement of discrete individuals. The aim is to discuss a modelling technique that allows more flexibility in the representation of the positions of individuals than is typically the case for cellular automata (CA), but which also deals efficiently with the problem of searching for neighbours when individual positions can vary nearly continuously. A scaling problem is discussed that arises when the range over which individuals interact is much smaller than the size of the domain. It is argued that validation of CA models involving discrete individuals is made more difficult when the system scale exceeds the size of individuals by a large factor. However, even when the domain size is small, if interaction between individuals is mediated by their size, imposition of a fixed grid upon the dynamics may cause important phenomena to be misrepresented or missed altogether. We suggest that cellular automata, as usually formulated, do not deal adequately with this type of problem, and introduce a particle-in-cell (PIC) method to deal with it in intermediate cases. Alternative data structures are discussed for dealing with more extreme cases, including the possibility of modelling an indefinitely large domain using a changing set of cells (PIC:SI).     

Gap-Crossing Decisions by the Red Squirrel, a Forest-Dependent Small Mammal

Abstract:  Forest-associated species in fragmented landscapes must traverse potentially inhospitable gaps to move between habitat patches. Although conservation biologists advocate connecting patches with corridors or improving the matrix to make it suitable for movement, little is known about the factors influencing gap-crossing decisions for most species. We investigated gap crossing by the red squirrel ( Tamiasciurus hudsonicus ) in logged landscapes in southeastern Alaska, where the species avoids microhabitats associated with gaps created by clearcutting. We released individuals across clearcuts and determined the routes they took home with tracking spools and radio telemetry. Of 36 adult red squirrels translocated across six clearcuts, 14 crossed clearcuts to reach home. Squirrels were more likely to cross clearcuts if the detour efficiency (distance to home crossing gap divided by distance of forested detour) was low, indicating an ability to compare distances along alternate routes and travel costs or risks in different habitats. No other landscape metrics, such as gap size or crossing distance, predicted crossing behavior. Red squirrels of low body mass were more likely to cross clearcuts, where the probability of encountering conspecifics is low. Distance predicted route choice for squirrels detouring around clearcuts. Indirect evidence suggests that perceived predation risk, energetic costs, or both are higher in clearcuts. Detour efficiency reportedly influences the gap-crossing decisions of some forest-associated birds, but this is the first demonstration of its role in gap-crossing decisions by a mammal.     

Extra-pair paternity and the reproductive role of male floaters in the tree swallow (Tachycineta bicolor)

In many avian species, a part of the population is present at the breeding grounds but does not breed. Current theories generally assume that floaters are younger or lower-quality individuals, and empirical data confirm this. However, floating could also arise as an alternative strategy to breeding, if floaters are able to reproduce via extra-pair copulations. Until the present study, there has been no evidence that floaters father offspring. We studied a population of tree swallows (Tachycineta bicolor), a species with one of the highest levels of extra-pair paternity known in birds. Using microsatellite markers, we determined the biological fathers of 65% of the extra-pair young. Of a total of 53 extra-pair young (52% of all offspring), 47% were fathered by local residents, 6% by residents breeding elsewhere (up to 2 km from the focal grid), and 13% by floaters. Residents seemed to be more successful and they were also more likely to return as territory holders in the next breeding season compared to floaters. Extra-pair males were on average in better condition than the within-pair males they cuckolded. Interestingly, resident males that disappeared (possibly to float) during the fertile period were heavier than males that stayed, and floaters were heavier than residents, but not different in any other characteristic. Although alternative interpretations of the data are possible, we propose that floating might be a conditional strategy in tree swallows whereby males in good condition gain more paternity via extra-pair copulations, whereas males in worse condition are more successful by providing parental care.     

Floater mortality within settlement areas can explain the Allee effect in breeding populations

The Allee effect (the positive relationship between population growth rate and population size) is a constraint of some animal populations at low numbers, which increases their likelihood of extinction because of a decrease in reproduction and/or survival. We were able to demonstrate that the Allee effect can be the result of a mortality increase affecting floaters (i.e. dispersing individuals able to enter as breeders in the reproductive population when a breeding territory or a potential mate – owner of a suitable breeding territory – becomes available). Previously, potential mechanisms underlying Allee effects were always related to the breeding portion of a population only. In contrast, our understanding of or solutions to population declines due to the Allee effects can reside elsewhere, away from breeding territories.     

The smell of desperadoes? Beavers distinguish between dominant and subordinate intruders

Olfactory signals can contain information, such as age and social status, and play a vital role in competitor assessment. In many territorial species, subordinates must leave their natal colony to obtain their own territory and mate. These individuals could be aggressive opponents in agonistic encounters, as they will have little to lose (the desperado effect). In this study, we tested the hypothesis that dominance and age are coded in the anal gland secretion (AGS) of the monogamous and highly territorial Eurasian beaver (Castor fiber), and if this information is used by conspecifics to evaluate the potential threat posed by an intruder. Territorial intrusions were simulated by presenting residents with a two-way choice test of AGS from an unknown male territory owner (i.e., dominant) and his son (i.e., subordinate; either 1 or ≥2 years old). Residents spent more time investigating AGS from subordinates than their fathers and responded more aggressively to subordinates than their fathers when subordinates were ≥2 years old. Chemical analyses gas chromatography and multivariate data analysis supported our behavioral findings and revealed differences between chemical profiles of territory owners and subordinates, as well as between the subordinates in different age classes. This study reveals that information about age and social status is coded in AGS of beavers and that this information is used to determine the level of an eventual future response to the signaler.     

Investment in territorial defence depends on rearing environment in brown trout (Salmo trutta)

In many animals, territoriality will arise or cease depending on environmental factors such as intruder rate and resource availability. We investigated the effect of rearing environment on territorial behaviour in ~1.5-month-old brown trout. In the laboratory, wild-caught (reared at a low density) and hatchery-reared (high density) trout were allowed to defend a territory against a size-matched intruder reared in the same or the other environment. Because territorial behaviour should be relaxed at high-rearing densities, we hypothesized that hatchery-reared trout should value their territories less and therefore invest less in defence compared with wild-caught trout. However, in all cases, territory owners were more likely to win the contest and hatchery-reared trout were just as likely as wild-reared to win mixed contests. Furthermore, pairs of hatchery-reared trout initiated contests sooner, fought longer and were more aggressive during the contest compared with pairs of wild trout. When hatchery-reared owners met wild intruders, the contest ended sooner compared with when the roles were reversed. We conclude that territorial behaviour in brown trout is largely innate, but that the hatchery environment has promoted more aggressive individuals. These results suggest that hatchery-reared trout invest more time and energy to obtain the same contest success as wild trout. In conclusion, the lack of experience of territorial defence in a high-density rearing environment seems to reduce the efficiency of territorial behaviour. In turn, this may have negative consequences for the performance of released hatchery fish in the wild.     

Laboratory and field evidence of sex-biased movement in the invasive round goby

Activity levels are modulated by trade-offs between reducing predation risk and the need to move in order to find food or mates. Because these trade-offs affect males and females differently, many species show sex-specific movement, dispersal patterns, and spatial navigation capacities, with the sex that gains the most from territory ownership often dispersing less. Unlike mammals and birds, sex differences in movement among fishes remain poorly studied, and the connections between tests of movement propensity in the laboratory and in the field are rarely made. Here, we examine the differences in movement between male and female round goby (Neogobius melanostomus) in both laboratory and field settings. This fish species is invasive in North America and currently undergoing further range expansions. In the laboratory, round goby males were more active and explored a novel environment more readily than did females. A large-scale mark–recapture study in Lake Ontario over two years revealed that males moved more than females between years, but there were no within-year sex differences. Thus, round goby display male-biased movement patterns, providing a comparison point to dispersal patterns in other taxa. Understanding sex-specific movement of round goby in the field will also help predict dispersal and population dynamics, both in areas where round goby have already become established and where they are continuing to invade.     

Relationships between spacing behavior and growth rates: A field study of lizard feeding territories

Summary Relationships between spacing behavior and growth rates were investigated in a field experiment with juvenile lizards, Anolis aeneus. The behavioral variable most closely related to juvenile growth was distance moved per unit time. This variable had a curvilinear relationship with growth, such that juveniles moving approximately 400 cm/h grew more rapidly than those traveling either larger or shorter distances per unit time. Daily fluctuations in arthropod abundance were also related to growth rates, with restricted growth during periods of low food availability. Temporal fluctuations in prey and distance traveled per unit time had independent effects on growth; together these two variables accounted for 43% of the variance in growth rate for the juveniles in this study.Territory size, overlap and social status appeared to affect growth indirectly, by influencing distance traveled per unit time. Optimal travel distances of around 400 cm/h were most likely when a juvenile had a relatively exclusive territory of about 0.5 m². High ranking juveniles were more apt to achieve this spacing pattern than were low ranking juveniles, but some high ranking juveniles had very large territories, extensive overlap with subordinates, supraoptimal travel distances and relatively low growth rates. Low ranking juveniles tended to fall into two groups: subordinates, with a small home range overlapping that of a more dominant individual and low travel distances, and floaters, with a large home range overlapping several more dominant individuals and high travel distances. Although a few low ranking juveniles achieved travel distances permitting high growth rates, most had either supra or suboptimal travel distances and relatively low growth rates.     

Habitat preference increases territorial defence in brown trout (Salmo trutta)

The striking ability of territory owners to repel intruders has generated a number of theoretical explanations as well as experimental studies in many animal species. However, effects of individual habitat preferences on territorial defence have rarely been studied. From the territory value hypothesis, we predicted that owners of preferred habitats should invest more resources in defence than owners of non-preferred habitats. We tested this prediction with young territorial brown trout in a two-stage experiment. First, trout were allowed to choose individually between gravel and a uniform bright substrate. As expected, they showed a significant (79%) preference for gravel. However, there was considerable variation between individuals in substrate preference, with a few fish preferring the bright substrate. Half of the tested fish were then transferred to a gravel substrate and the rest to a bright substrate, manipulating habitat type in relation to preference. Territory owners were then staged against size-matched intruders whereupon contest aggression was observed and the winner of each contest determined. Overall, owners won most of the contests. Satisfied owners won 86% and owners of less preferred territories, 74% of the contests. Furthermore, more satisfied owners attacked sooner and were more aggressive relative to the intruders. We conclude that brown trout show individual variation in habitat preference, which appears to be linked with their investment in territorial defence. These results suggest that understanding and modelling of animal contests could benefit from considering how territorial defence is influenced by individual habitat preference and specialisation. Received: 25 February 2000 / Revised: 22 May 2000 / Accepted: 25 June 2000     

Individual experience-based foraging can generate community territorial structure for competing ant species

To gain additional territory while defending existing territory, animals must acquire and use information regarding resource characteristics and competitive pressure. For social organisms like ants, individual workers have experiences to acquire information, but territory establishment is a colony level behavior. Colony behavior, in turn, affects community structure. Here, I investigate how an individual ant’s previous experience affects its future foraging behavior and how individual behaviors can scale up to community territorial structure for two coexisting Formica species. To do this, I combine a field survey, a multi-agent computer simulation, and a manipulation experiment. The field survey shows that workers of both species co-occur on many trees early in the season, but ants on trees become segregated by species as the season progresses. The simulation demonstrates how this segregated spatial distribution can result from ants using a foraging strategy in which individuals show a preference for foraging sites based on previous experience. The experiment suggests that these ants are indeed capable of experience-based foraging behavior; ants preferentially return to sites where they have had positive experiences and avoid sites where they have had negative experiences. Results from this study suggest that spatially explicit information can be collected and stored by individuals to facilitate colony territorial structure, and that future investigations of community territory formation should include effects of individual previous experience.     

The effect of conspecifics on habitat selection in territorial species

Summary Despite widespread interest in habitat selection, many of the assumptions about how territorial animals choose habitats have not been tested. This study of juvenile Anolis aeneus lizards focuses on the relationship between the number of previous settlers in a habitat and the subsequent behavior of new arrivals at that habitat. Clearings containing the types of microhabitat preferred by juveniles were established in the field, several juvenile residents were allowed to establish territories in enclosures in the center of each clearing, and then naturally occurring immigrants were allowed access to the empty microhabitat surrounding the enclosures. Arrival rates and the probability of settlement were monitored on a daily basis from the day the first juveniles arrived until several days after the last juvenile had settled (=saturation). In each of seven trials, arrival rates were comparable early and late in the settlement process, and were unrelated to the degree of habitat saturation. Arrival rates did vary on a temporal basis, probably as a result of environmental factors affecting egg laying and hatching schedules, and habitats with high arrival rates saturated more quickly than those with lower arrival rates. All of the individuals arriving at the clearings did not settle, but settlers and non-settlers did not differ with respect to competitive ability, as measured by body size. The probability of settlement increased as settlement proceeded in each of seven trials, up to the day of saturation. These results refute the commonly held assumptions that prospective territory owners avoid entering relatively full habitats, and that they prefer to settle in relatively unsaturated habitats. The discussion considers why assumptions about the behavioral processes of habitat selection are so widely accepted, given the dearth of empirical information on the subject.     

Songs of isolation-reared sparrows function in communication,but are significantly less effective than learned songs

Summary Songs of male song sparrows (Melospiza melodia) and swamp sparrows (M. georgiana) isolated before hatching from conspecific song were played to males and females of both species. Isolate songs of these sparrows resemble natural conspecific song in several aspects of gross structure, but differ from natural song in note structure. Male territory owners of both species responded more to conspecific isolate song than to heterospecific normal song. Captive females of both species, previously treated with estradiol, courted in response to isolate song but not to heterospecific song. We conclude that there is sufficient speciesspecific information in isolate song of both species to allow a degree of normal function. To assess the importance of the structural refinements added through learning, we compared response to isolate and natural conspecific songs. Male territory owners and captive females responded more to natural than to isolate songs in both species. Learning thus significantly increases the potency of song in both intersexual and intrasexual communication.