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.     

Estimation and limitation of numbers of floaters in a Eurasian Sparrowhawk population

Over a 20‐year period, the numbers of Eurasian Sparrowhawk Accipiter nisus nests found in a 200 km² area in south Scotland remained relatively stable (mean 33.3 pairs, CV = 10.6%). Nest numbers fluctuated from year to year in a manner expected of a population subject to density‐dependent regulation. The numbers of non‐breeders (floaters) could not be counted directly, but the number of female floaters was estimated, using known mortality rates, from the numbers of females recruited to the breeding population each year at different ages. Female floater numbers were estimated by two methods: Method A assumed that birds bred for the first time in their first, second or third year, in the same ratio as they were found breeding for the first time in the study area; and Method B assumed that all third‐year birds found breeding for the first time in the study area had bred previously, unknown to us, outside the area. Under Method A, floaters consisted of some 1‐year and some 2‐year birds; while under Method B, floaters consisted only of some 1‐year birds. Under both methods, the estimated number of female floaters fluctuated greatly from year to year, but under Method A they averaged 0.90 per female breeder, and under Method B they averaged 0.28 per female breeder. The Method B estimate was most consistent with other data and with the finding that some birds found breeding in the study area were likely to have bred previously outside the area (because of territory changes). Moreover, the mean and variance in female floater numbers estimated by Method B were similar in magnitude to the values obtained in a simulation model. In this model, breeding density was given a fixed ceiling, while breeding success was allowed to vary from year to year within the limits observed in the study area. It was concluded that (a) recruitment of floaters to the breeding sector was density dependent with respect to breeder numbers, i.e. broadly speaking, floaters filled gaps in the territorial system left by the deaths and movements of established breeders; and (b) floater numbers themselves were probably not regulated in a density dependent manner, but depended on whatever was the balance between annual additions (from reproduction and immigration) and subtractions (from mortality, emigration and entry to the breeding sector).     

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.     

Territorial defense, territory size, and population regulation

The carrying capacity of an environment is determined partly by how individuals compete over the available resources. To territorial animals, space is an important resource, leading to conflict over its use. We build a model where the carrying capacity for an organism in a given environment results from the evolution of territorial defense effort and the consequent space use. The same evolutionary process can yield two completely different modes of population regulation. Density dependence arises through expanding and shrinking territories if fecundity is low, breeding success increases gradually with territory size, and/or defense is cheap. By contrast, when fecundity is high, breeding success sharply saturates with territory size, and/or defense is costly, we predict fixed territory sizes and regulation by floaters. These "surplus" individuals form a buffer against population fluctuations. Yet floaters can also harm breeder performance, and by comparing population growth of a territorial population to a nonterritorial (and individually suboptimal) alternative, we can quantify the harmful effect of evolutionary conflict on population performance. Territoriality has often been found to increase population stability, but this may come at a cost of reduced equilibrium densities.     

Interactions between Males of the Lizard Liolaemus tenuis: Roles of Familiarity and Memory

For territorial organisms, recognition of familiar individuals can reduce the frequency and intensity of aggressive encounters (‘dear enemy’ phenomenon), stabilize social systems, and reduce the cost of territory maintenance. Here, we investigated the behavioural events displayed during contests between familiar and unfamiliar individuals in the lizard Liolaemus tenuis (Liolaemidae), a species in which males are territorial. The behaviours recorded were attack, warning, evasion, and submission, and the latencies to the first aggressive (attack or warning) behaviour. Additionally, we assessed the ability of individuals to remember a familiar conspecific after a period without social interaction. Individual males reduced and delayed aggressive behaviour directed towards socially familiar individuals compared with unfamiliar ones. These results suggest that males distinguished between familiar and unfamiliar conspecific males and are in agreement with the ‘dear enemy’ phenomenon. Other behaviours were similar in the contests between familiar and unfamiliar individuals. Recognition of familiar conspecifics was lost after 20 d without social interactions. This may be relevant for interactions with floater males or with neighbours that lose their territory and subsequently attempt to fight for their ex‐neighbour's territory.     

Experimental tests of endocrine function in breeding and nonbreeding raptors

Many long-lived avian species defer reproduction for several years, often displaying a "floating" behavior characterized by the lack of mates and exclusive territories. Understanding the proximate mechanisms regulating floating behavior is a relevant topic of research for physiologists, behavioral ecologists, and population biologists because a prolonged period of nonbreeding can negatively affect lifetime fitness and change population dynamics. Here we tested two hypotheses linking endocrine function to floating status: (a) floaters undergo a period of sexual immaturity characterized by lower gonadal function (hypothesis of sexual immaturity), and (b) floating status is socially imposed by dominant conspecifics and revealed by the adrenocortical response to stress (hypothesis of social subordination). The two hypotheses were tested in a population of free-living black kites Milvus migrans in Do?ana National Park (southwest Spain), where breeders coexist with young floaters that defer reproduction for 3-7 yr. Hypophysial-gonadal function, estimated as androgen production in response to experimental challenge with gonadotropin-releasing hormone (c-GnRH-I), was similar in magnitude and timing between floating and breeding males. The same treatment was, however, unable to elicit any response in terms of increasing estradiol or total androgen levels in females regardless of their breeding status. Following experimental capture and restraint, the adrenocortical response to stress (estimated as circulating corticosterone levels) was higher in floating than in breeding males, while females showed the opposite pattern (i.e., lower response to stress in young floaters compared with breeders). Contrary to the hypothesis of sexual immaturity, our results suggest that floating males are physiologically capable of reproducing. The reported differences in adrenocortical function support the idea that floaters are socially subordinate to breeders, and corticosterone responses reflect the sex-specific roles during competition in socially monogamous species.     

Demography of Slate‐throated Redstarts (Myioborus miniatus): a non‐migratory Neotropical warbler

Most wood‐warblers (Parulidae) are non‐migratory residents of the Neotropics and subtropics, and the demographic characteristics of these species are poorly known. I examined the annual survival, reproductive output, dispersal, age of first breeding, and other demographic characteristics of a permanently territorial non‐migratory tropical warbler, the Slate‐throated Redstart (Myioborus miniatus), based on a 5‐yr study of a color‐banded population in Monteverde, Costa Rica. Territorial males showed strong site fidelity, but 26% of females engaged in short‐distance between‐year breeding dispersal. Estimated annual survival of territory holders, corrected for undetected female breeding dispersal, was 0.56 for males and 0.43 for females, values lower than expected and comparable to survival estimates for North American migrant warblers. The lower annual survival of females had two demographic consequences; unpaired territorial males were present in 3 of 5 yr, and some 1‐yr‐old males appeared to be floaters. Unpaired females or female floaters, however, were not observed. Mean natal dispersal distance was significantly greater for females (935 m) than males (485 m). Estimated first‐year survival was 0.29, but this is almost certainly an underestimate because of undetected long‐distance, female‐biased natal dispersal. Annual fecundity (fledglings per female) was 1.8, less than that of temperate warblers and attributable to small mean clutch sizes and a low incidence of double brooding. Estimated population growth rate (λ) was <1 for both males and females, suggesting that the study population was a demographic sink, most likely due to lower‐than‐expected adult survival.     

Seasonal Changes of Vocal Rates and Their Relation to Territorial Status in Male Galápagos Sea Lions (Zalophus wollebaeki)

Male vocal displays play an important role in sexual selection through both male–male competition and female choice. While this is supported by numerous studies in birds, much less attention has been paid to the role of such displays in mammals. We investigated the function of vocal displays in a polygynous mammal, the Galápagos sea lion (Zalophus wollebaeki). In our study population a large proportion of mature males are unable to establish a territory, providing the opportunity to compare the vocal behaviour between territorial and non‐territorial males. We examined how seasonal patterns of vocalizations differed between territorial and non‐territorial males and how the number of females present in a territory influenced behaviour of territorial males. We found that territorial males vocalized at higher rates than non‐territorial males, and territorial males vocalized more at the onset than towards the end of territory tenure. During the onset of territory‐tenure vocalizations of territorial males were directed more often towards other males than females. Furthermore, we found that vocalizations of territorial males were not only given in male–male interactions, but were also actively directed towards females. Territorial males vocalized at higher rates when more females were present in their territory. Our results suggest that vocalizations are important in male–male interactions, are relevant in territorial disputes and are used as a proxy for females to assess a male’s quality.     

Acoustic Monitoring Reveals Congruent Patterns of Territorial Singing Behaviour in Male and Female Tropical Wrens

Many animals defend territories against conspecific individuals using acoustic signals. In birds, male vocalizations are known to play a critical role in territory defence. Territorial acoustic signals in females have been poorly studied, perhaps because female song is uncommon in north‐temperate ecosystems. In this study, we compare male vs. female territorial singing behaviour in Neotropical rufous‐and‐white wrens Thryothorus rufalbus, a species where both sexes produce solo songs and often coordinate their songs in vocal duets. We recorded free‐living birds in Costa Rica using an eight‐microphone Acoustic Location System capable of passively triangulating the position of animals based on their vocalizations. We recorded 17 pairs of birds for 2–4 consecutive mornings and calculated the territory of each individual as a 95% fixed kernel estimate around their song posts. We compared territories calculated around male vs. female song posts, including separate analyses of solo vs. duet song posts. These spatial analyses of singing behaviour reveal that males and females use similarly sized territories with more than 60% overlap between breeding partners. Territories calculated based on solo vs. duet song posts were of similar size and similar degrees of overlap. Solos and duets were performed at similar distances from the nest for both sexes. Overall, male and female rufous‐and‐white wrens exhibit very similar spatial territorial singing behaviour, demonstrating congruent patterns of male and female territoriality.     

Young But Not Old Adult African Striped Mice Reduce Their Activity in the Dry Season When Food Availability is Low

An individual′s survival and fitness depend on its ability to effectively allocate its time between competing behaviors. Sex, social tactic, season and food availability are important factors influencing activity budgets. However, few field studies have tested their influences. The African striped mouse (Rhabdomys pumilio) lives in highly seasonal habitats in southern Africa, and individuals can adopt different social tactics. We investigated seasonal changes in activity budgets of different tactics and predicted that individuals will reduce their activity in the non‐breeding season to save energy when food availability is low and that young non‐breeding adults (‘philopatrics’) invest mainly in activities related to gaining body mass to increase survival probability. We predicted old adults (‘breeders’), which bred during the previous breeding season, to invest mainly in maintenance of their social status. We conducted 90 focal observations during the non‐breeding season and 73 during the breeding season. Activity budgets of striped mice were season and tactic specific, with philopatrics, but not breeders, reducing activity when food availability was low, possibly to decrease energy expenditure. Philopatrics of both sexes foraged and basked more in the breeding season than during the non‐breeding season. Male philopatrics gained body mass and female philopatrics maintained their body mass in both seasons. Sex‐specific differences occurred during the breeding season, when female breeders foraged more than male breeders, while male breeders chased other individuals more than female breeders. These findings indicate that individuals adopting different social tactics display distinct behaviors to fulfill tactic‐specific energetic needs .     

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.     

Territorial intrusion risk and antipredator behaviour: a mathematical model

In territorial animals that hide to avoid predators, a predatory attack creates a conflict because a hiding animal cannot defend its territory from conspecific intruders. When intruders are persistent, a past conspecific intrusion informs a territorial resident that future intrusions by the same animal are likely. Using a mathematical model, I examine the effects that past territorial intrusions can have on antipredator behaviour. Past territorial intrusions rarely affect a resident animal's time to hide (the optimal behaviour is to hide as soon as the predator initiates its attack). In contrast, past intrusions should shorten the length of time during which territory holders remain in hiding, with the magnitude of this effect depending on the time of the predator's attack, the re-intruder's pattern of return, and the intrusion rates of other conspecifics. The results of the model show that we need more information on patterns of re-intruders' behaviour, and emphasize that a similar functional explanation could underlie other behavioural changes following territorial and/or aggressive encounters (such as winner/loser effects or changes in display frequency and territorial vigilance). Differences between my findings and those from previous studies suggest that the trade-off between antipredator behaviour and territorial defence can involve different costs from the trade-off between antipredator behaviour and foraging.     

The Effects of Social Context on the Food-Caching Behavior of Florida Scrub-Jays (Aphelocoma coerulescens)

Florida scrub‐jays are cooperative breeders that live in family groups consisting of a breeding pair, often with several non‐breeding helpers. Florida scrub‐jays cache food by scatter‐hoarding items for later consumption. Within family groups, members have the opportunity to observe and pilfer the caches of other members. We observed jays harvesting experimentally provisioned peanuts alone and in the presence of other family members, to determine whether jays modify their food‐handling behavior relative to social context. Non‐breeding helpers were less likely to cache in the presence of the dominant male breeder than when alone and all jays tended to cache out of sight when observed by another jay. These changes in caching behavior are consistent with cache protection strategies employed by other species. However, the adaptive value of such cache protection within a sedentary cooperatively breeding family group on a year‐round territory is unclear.     

An Experimental Study of Chick Provisioning in the Cooperatively Breeding Acorn Woodpecker

Acorn woodpeckers (Melanerpes formicivorus) are cooperative breeders in which groups consist of a variable number of cobreeding males, joint‐nesting females, and non‐breeding helpers of both sexes that are offspring from prior nests. We temporarily manipulated brood size of nests to determine the feeding response of birds in relation to their status (breeder or non‐breeding helper) and sex. All categories of birds responded similarly to brood size increases, adjusting their feeding rate upwards so as to maintain approximately the same per‐nestling feeding rate. Breeders, however, exhibited more flexibility with respect to brood size reductions, decreasing their feeding rate while helpers did not. This suggests that the ‘feeding rules’ of helpers are less flexible than those of breeders, a result not previously detected in other cooperative breeders that have been studied to date. Particularly surprising was the finding that helpers maintain their feeding rates when brood demand is decreased rather than when it was increased, suggesting that the flexibility they exhibit is not a result of birds using the opportunity afforded by reduced brood demand to engage in other less cooperative activities.     

Effects of age,territoriality and breeding on survival of Bonelli’s Eagle Aquila fasciata

Survival typically contributes most to population trends in long‐lived birds and its accurate estimation is therefore vital for population management and conservation. We evaluated the effects of age, territoriality and reproduction on survival in Bonelli’s Eagle Aquila fasciata through multistate capture‐mark‐recapture analyses on a long‐term dataset. Monitoring was carried out in southeast France (1990–2008) and involved the surveying of territorial Eagles, the marking of fledged chicks, and the recording of resightings and recoveries of marked non‐territorial and territorial birds. Survival improved with age, but territoriality was not retained in the best model; yearly survival was estimated at 0.479 for fledglings (to 1 year of age), 0.570 for 1‐ and 2‐year‐olds, and 0.870 for 3‐year‐old and older individuals. The second best model supported a further increase in survival from 3‐year‐olds (0.821) to older individuals (0.880). In the third best supported model, territoriality enhanced survival, but only in 2‐year‐olds (0.632 vs. 0.562 for non‐territorial). We found no correlation between the previous breeding stage and future survival, consistent with the long lifespan of the study species. Nevertheless, 4‐year‐old and older successful breeders were more likely to breed the following year than failed adult breeders (0.869 vs. 0.582), suggesting that the cost of reproduction is small in comparison with the variation in quality among individuals or their territories.     

The Effects of Breeder Loss on Wolves

Abstract Managers of recovering wolf (Canis lupus) populations require knowledge regarding the potential impacts caused by the loss of territorial, breeding wolves when devising plans that aim to balance population goals with human concerns. Although ecologists have studied wolves extensively, we lack an understanding of this phenomenon as published records are sparse. Therefore, we pooled data (n = 134 cases) on 148 territorial breeding wolves (75 M and 73 F) from our research and published accounts to assess the impacts of breeder loss on wolf pup survival, reproduction, and territorial social groups. In 58 of 71 cases (84%), ≥1 pup survived, and the number or sex of remaining breeders (including multiple breeders) did not influence pup survival. Pups survived more frequently in groups of ≥6 wolves (90%) compared with smaller groups (68%). Auxiliary nonbreeders benefited pup survival, with pups surviving in 92% of cases where auxiliaries were present and 64% where they were absent. Logistic regression analysis indicated that the number of adult-sized wolves remaining after breeder loss, along with pup age, had the greatest influence on pup survival. Territorial wolves reproduced the following season in 47% of cases, and a greater proportion reproduced where one breeder had to be replaced (56%) versus cases where both breeders had to be replaced (9%). Group size was greater for wolves that reproduced the following season compared with those that did not reproduce. Large recolonizing (>75 wolves) and saturated wolf populations had similar times to breeder replacement and next reproduction, which was about half that for small recolonizing (≤75 wolves) populations. We found inverse relationships between recolonizing population size and time to breeder replacement (r= —0.37) and time to next reproduction (r= —0.36). Time to breeder replacement correlated strongly with time to next reproduction (r=0.97). Wolf social groups dissolved and abandoned their territories subsequent to breeder loss in 38% of cases. Where groups dissolved, wolves reestablished territories in 53% of cases, and neighboring wolves usurped territories in an additional 21% of cases. Fewer groups dissolved where breeders remained (26%) versus cases where breeders were absent (85%). Group size after breeder loss was smaller where groups dissolved versus cases where groups did not dissolve. To minimize negative impacts, we recommend that managers of recolonizing wolf populations limit lethal control to solitary individuals or territorial pairs where possible, because selective removal of pack members can be difficult. When reproductive packs are to be managed, we recommend that managers only remove wolves from reproductive packs when pups are ≥6 months old and packs contain ≥6 members (including ≥3 ad-sized wolves). Ideally, such packs should be close to neighboring packs and occur within larger (≥75 wolves) recolonizing populations.     

The Role of Wing Coloration in Sex Recognition and Competitor Recognition in Rubyspot Damselflies (Hetaerina spp.)

The decision rules that animals use for distinguishing between conspecifics of different age and sex classes are relevant for understanding how closely related species interact in sympatry. In rubyspot damselflies (Hetaerina spp.), the red wing coloration of mature males is hypothesized to be a key trait for sex recognition and competitor recognition within species and the proximate trigger for interspecific male–male aggression. We tested this hypothesis by manipulating the wing coloration of tethered conspecific intruders and measuring the responses of territory holders of three species in the field. As predicted, covering the red spots of mature males with black ink nearly eliminated territorial responses, and in some cases, territorial holders clasped the blackened males as if they were females. Adding red spots to female wings triggered territorial responses and nearly eliminated sexual responses. Immature males with artificial red spots were attacked at the same rate as mature male intruders, and much more frequently than were immature male controls. The results varied somewhat by species. In H. titia, the only species of Hetaerina with substantial black wing pigmentation, the effects of blackening the red spots of intruders varied both geographically and seasonally. But even when blackening the red spots of male intruders did not reduce the aggressive response of H. titia territory holders, adding artificial red spots to female wings elicited aggressive responses and nearly eliminated sexual responses. The results of this study further strengthen the evidence that interspecific aggression in Hetaerina results from overlap in territorial signals and that the derived black wing pigmentation of H. titia reduces interspecific aggression.     

Egg investment in response to helper presence in cooperatively breeding Tibetan ground tits

Life‐history theory predicts a trade‐off between current and future reproduction to maximize lifetime fitness. In cooperatively breeding species, where offspring care is shared between breeders and helpers, helper presence may influence the female breeders’ egg investment, and consequently, survival and future reproductive success. For example, female breeders may reduce egg investment in response to helper presence if this reduction is compensated by helpers during provisioning. Alternatively, female breeders may increase egg investment in response to helper presence if helpers allow the breeders to raise more or higher quality offspring successfully. In the facultatively cooperative‐breeding Tibetan ground tit Pseudopodoces humilis, previous studies found that helpers improve total nestling provisioning rates and fledgling recruitment, but have no apparent effects on the number and body mass of fledglings produced, while breeders with helpers show reduced provisioning rates and higher survival. Here, we investigated whether some of these effects may be explained by female breeders reducing their investment in eggs in response to helper presence. In addition, we investigated whether egg investment is associated with the female breeder's future fitness. Our results showed that helper presence had no effect on the female breeders’ egg investment, and that egg investment was not associated with breeder survival and reproductive success. Our findings suggest that the responses of breeders to helping should be investigated throughout the breeding cycle, because the conclusions regarding the breeders’ adjustment of reproductive investment in response to being helped may depend on which stage of the breeding cycle is considered.