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.     

Floater interference reflects territory quality in the Spanish Imperial Eagle Aquila adalberti: a test of a density‐dependent mechanism

We report on an 11‐year study of floater interference in a population of Spanish Imperial Eagles Aquila adalberti. We analysed changes over the years in the productivity of 15 territories to test predictions of two hypotheses of density‐dependent productivity in relation to the presence of floaters (birds without territories). According to the ‘interference' hypothesis, the frequency of intrusion by floaters increases with density, resulting in a decrease in productivity. Thus, in a high‐density population a negative relationship between floater intrusions and productivity of the territory is expected. In contrast, under the ‘habitat heterogeneity' hypothesis, as density increases a higher proportion of individuals is forced to occupy lower quality habitats. Support of this hypothesis requires that floaters detect differences in quality among territories and preferentially visit the better quality territories. Consequently, a positive relationship between floater intrusions and productivity is expected. Results showed that floaters tended to visit their natal area at the beginning of the breeding season. Among floater Eagles, males made significantly more intrusions per day than did females, but females stayed in the natal population for longer each year than males. Floater intrusions and productivity were highly positively correlated, supporting the ‘habitat heterogeneity' hypothesis; individuals were apparently able to assess the quality of a territory and, at the frequencies observed, their interference with the breeding pair had no obvious negative effect on productivity.     

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 cost of being alone: the fate of floaters in a population of cooperatively breeding pied babblers Turdoides bicolor

The occurrence of group-living behaviour has often been explained by the benefits individuals receive through cooperation; including increased reproductive output, vigilance against predators, and load-lightening behaviour. However, to fully understand the benefits of group-living, it is important to quantify the costs of living alone. Here, we look at the fate of floaters (individuals who have no fixed territory and remain alone for extended periods) in a population of cooperatively breeding pied babblers Turdoides bicolor . We found that individuals spent less time foraging and more time vigilant for predators when found as a floater compared to when they were in a group. Consequently, they suffered a continuous loss of body mass, with long-term floaters suffering the highest losses. This had a long-term effect: floaters that eventually did regain a position in a group usually entered as helpers, in contrast to dispersers, who usually entered a new group as breeders. This high cost of living alone highlights the benefits of group-living and may help to understand patterns of delayed dispersal in some social species.     

When individuals senesce: the 'Florida effect' on stable populations of territorial, long-lived birds

Senescence can be defined as the entire set of age-related changes that affect both vitality and function, one of which is within-individual age-related decline in reproduction. This factor is crucial for population persistence, because the senescence of individuals of a population can increase the likelihood of local extinction. Using simulations based on long-term information on a small metapopulation of a long-lived bird species, we highlight two mechanisms able to engender senescence in both breeders and floaters (i.e. non-breeding individuals) of the same population. We define 'floater shortage senescence' as breeder senescence due to low juvenile replacement rates because of high floater mortality during dispersal. Less obviously, senescence can also occur with very low floater mortality rates, and when breeding populations are remarkably free from factors that could cause catastrophic population decimation. In this scenario, low mortality in reproductive areas results in territory owners in breeding populations being characterised by progressively older breeding individuals, and old individuals waiting for a breeding opportunity: a phenomenon we refer to as the 'Florida effect'. Consistent with current views that adaptive death plays a crucial role in population dynamics, it seems reasonable to suppose that, under stable environmental conditions, the evolution of some mating mechanisms could limit the senescence of breeding individuals in a population, allowing the pool of breeding individuals to be refreshed by selection of younger breeders.     

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.     

Basal song rate variation in male red-winged blackbirds: sound and fury signifying nothing?

Male red-winged blackbirds (Agelaius phoeniceus) vary substantiallyin how much they sing. We tested whether song rate reflectedthe male’s quality, the quality of his territory, or hisfrequency of interaction with potential competitors. Song ratedeclined with the seasonal decline in nest initiations, suggestingthat song is used to attract females. However, males with highersong rates did not obtain larger harems. This suggests thatfemales do not use male song rate to assess either male qualityor territory quality. If song rate signals male quality to malerivals, and if territory owners are of higher quality than floaters,then owners' song rates should be higher than those of floaters.However, when we removed territory owners temporarily, the songrate of their floater replacements was similar to that of theoriginal owners. Finally, a male’s song rate was not affectedby how frequently he chased intruders, or by the proximity ornumber of neighbors he had. In short, the only information thatbasal song rate of redwinged blackbirds appears to provide toconspecifics is that the territory is occupied. In simulatedterritory takeover attempts, however, song rate was significantlyhigher than the basal rate. Thus, more may be revealed by thesinger in these situations. [Behav Ecol 1991;2:123-132]     

The resident's dilemma: a female choice model for the evolution of alternative mating strategies in lekking male ruffs (Philomachus pugnax)

Previous models for the evolution of alternative male matingbehavior have virtually ignored the role of female choice. Wepresent a model in which female choice favors the evolutionand maintenance of alternative mating strategies in male ruffs,Philomachus pugnax. Resident male ruffe establish and defendcourts on leks against other residents, while non-territorialsatellite males move between leks and among courts on a lek.Residents appear to actively recruit satellites to their courts,even though satellites may mate with females once there. Residentbehavior toward satellites and data on female behavior suggestthat residents benefit from a satellite's presence due to somefemale preference for mating on co-occupied courts. However,if all residents accept satellites, none gains any relativeadvantage, yet all pay the costs of having satellites on theircourt. We present a game theoretical model that shows that therelative nature of female choice places residents in an evolutionarydilemma with respect to satellite acceptance. Although all residentswould benefit if satellites could be cooperatively excludedfrom leks, the only evolutionarily stable strategy for individualresidents is to defect and accept satellites. The model alsodemonstrates that this "resident's dilemma" likely exists onlyin a local sense, since the failure of residents to cooperativelyexclude satellites from leks need not result in globally lowerpayoffs, due to frequency-dependent selection on the proportionof satellites in the population. Our analysis suggests thatthe resident-satellite relationship in ruffs, despite its obviouscompetitive elements, is fundamentally a cooperative associationfavored by female choice. Female choice has also been proposedas the primary mechanism selecting for male association to formleks in ruffe. In this context, resident-satellite associationsmay be thought of as transitory "leks within a lek     

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).     

Behavioral syndromes influence mating systems: floater pairs of a lizard have heavier offspring

Variation in correlated behaviors or behavioral syndromes couldhave interesting effects on mating systems, especially if thevariation in syndrome exists in both sexes. Both males and femalesof the lizard Eulamprus heatwolei display two behavioral typesof a behavioral syndrome, defined by correlations between territorial,exploratory, and predator avoidance behaviors. We tested howthis variation in behavioral syndrome influences reproductivesuccess, pairing patterns, and offspring weight. We used spatialbehavior and residency in the field to identify territorialand floater individuals. Females were relocated to the laboratoryto give birth, and all offspring, dams, and potential sireswere genotyped to determine offspring paternity. During fieldsurveys, 164 lizards were caught of which 27.5% were territorialand the rest were floaters. Paternity was assigned to 66% ofthe 104 offspring produced by 33 dams. Territorial sires fathereda greater proportion of the offspring of territorial dams thanfloater sires. Larger territorial males were more likely tosire the entire clutch or share paternity with fewer additionalsires than smaller territorial males. Floater sire size, however,did not influence the number of fathers per litter. Floaterfemales produced heavier offspring than their territorial counterparts,and offspring fathered by floaters were heavier than maternalhalf-sibs fathered by territorial males. We speculate that differencesin offspring weight may be the result of differences in yolkprovisioning by females and parent genetic compatibility.     

The Evolution of Reliable and Unreliable Badges of Fighting Ability

SYNOPSIS. When a population may be characterized by interferencecompetition for resources, variation in fighting ability amongindividuals, and repeated confrontations between individuals,together with difficulty of individual recognition, badges ofstatus should invade as recognition marks that render good fightersmemorable. Reliability of such badges can be maintained by negativefrequency-dependent selection when individuals of differentappearance (and status) either play mutually beneficial rolesor employ alternate competitive tactics. In territorial socialsystems intraspecific mimicry of recognition badges should evolvebecause, in contrast to group-living situations, the cost toa cheat of being discovered is low when individuals are dispersed.The general result of such mimicry is that good and poor fightersbecome similar in appearance. From the theoretical treatmentof status recognition badges I derive a number of predictionsthat pertain both to interand intraspecific differences in conspicuouscoloration and to the evolution of local song dialects in birds.     

The Relationship Between the Presence of Satellite Males and Nest-Holders' Mating Success in the Azorean Rock-Pool Blenny Parablennius sanguinolentus parvicornis

In the Azorean rock-pool blenny, sexually active males may adopt alternative reproductive tactics. In the present paper the relationship between the presence of satellite males and the reproductive success of nest-holders was investigated by comparing nests with and without an associated satellite male. Males with an associated satellite male suffered more conspecific intrusions but they did not display a higher frequency of attacks towards conspecifics. Nest-holder males were more aggressive towards other conspecifics than towards satellites and the tolerance of nest-holders towards satellites was inversely correlated with the time spent by the satellites in the breeding territory, which suggests control by the nest-holder male of the satellite investment in shared territorial defence. Nest-holders with an associated satellite male had higher condition factors and received more female visits and more spawnings. These results bear two possible interpretations. (1) Nest-holders benefit from the presence of a satellite male by increased attractiveness of their nests to females; satellite males are mutualists helping to defend the nest-owner's territory and to attract females, which is why they are tolerated. (2) Satellite males associate preferentially with more successful nest-holder males which have higher condition factors, and by doing so have more opportunities to achieve parasitic fertilizations. Only experiments will allow these two hypotheses to be distinguished.     

Strained energy budget of winter floaters in the Willow Tit as indicated by ptilochronology

Most Willow Tits Parus montanus live within flock territories during the non-breeding season, but many juveniles roam around or switch between several flocks. Such floaters behave submissively and spend time watching other flock members in addition to scanning for predators. During September–December 1989–2000, I studied individually colour-ringed Willow Tits living in a subalpine forest area in central Norway. When among flock members, the floaters foraged less in favourable parts of trees and were more vigilant than the lowest-ranked juvenile flock members. The nutritional condition of the birds was assessed by examining the growth bar widths of their tail feathers. Birds in better condition have wider growth bars than those in poorer condition. The daily growth bars of induced feathers of male and female floaters, laid down under winter conditions, were significantly narrower than those of the lowest ranked flock members. These results suggest that juvenile floater Willow Tits maintain a poorer nutritional status than low-ranking juvenile flock members during winter.     

The feeding ecology of Bonelli’s eagle (<Emphasis Type="Italic">Aquila fasciata</Emphasis>) floaters in southern Spain: implications for conservation

Although many studies have investigated the feeding habits of Palaearctic raptors, few have analysed non-breeding populations during dispersal. Bonelli’s eagle (Aquila fasciata), a threatened species in Western Europe, has a relatively long and critical dispersal period. We studied feeding habits, prey selection, and the influence of prey density on floater abundance in this species during its dispersal period in southern Spain. Differences were found between the diet of floaters and that of the closest breeding populations. Diet diversity was rather low for floaters, with European wild rabbits (Oryctolagus cuniculus) being the main prey and the only prey positively selected. Moreover, the number of floater Bonelli’s eagles observed in the dispersal areas was positively associated with rabbit abundance. Other prey included red-legged partridges (Alectoris rufa) and pigeons (Columba sp.). We propose measures to match suitable prey availability for floater eagles in settlement areas and the use of such areas as human hunting fields.     

Sanctions and mutualism stability: when should less beneficial mutualists be tolerated?

Why do mutualists perform costly behaviours that benefit individuals of a different species? One of the factors that may stabilize mutualistic interactions is when individuals preferentially reward more mutualistic (beneficial) behaviour and/or punish less mutualistic (more parasitic) behaviour. We develop a model that shows how such sanctions provide a fitness benefit to the individuals that carry them out. Although this approach could be applied to a number of symbioses, we focus on how it could be applied to the legume‐rhizobia interaction. Specifically, we demonstrate how plants can be selected to supply preferentially more resources to (or be less likely to senesce) nodules that are fixing more N₂ (termed plant sanctions). We have previously argued that appreciable levels of N₂ fixation by rhizobia are only likely to be selected for in response to plant sanctions. Therefore, by showing that plant sanctions can also be favoured by natural selection, we are able to provide an explanation for the stability of the plant‐legume mutualism.     

A marginal model of tolerated theft

Using marginal analysis to represent Blurton Jones's concept of tolerated theft, I show how equilibrium resource transfers among individuals might be affected by foraging behavior, resource qualities, and number of participants. The model applies to hominids and other species that exchange or share food or other resources. Among the results: Tolerated theft enhances the value to be derived from resources, packets intermediate in size are most likely to be subjected to tolerated theft, packet division is more likely to be unequal than equal, division is a function of group size, and tolerated theft is most likely in small groups. The model also suggests that among reciprocators the widest possible exchange or sharing is in the self-interest of the individual procuring the resource. In general, evolutionary cost-benefit accounting should track marginal changes in the value (fitness or utility) of resources. Marginal valuation is conceptually primary and may produce results that differ from direct measures of quantity.     

Controlling infestations of Parkinsonia aculeata in a riparian zone at the landscape scale

Abstract To prioritize weed management activities it is necessary to predict the extent of future infestations of individual weed species. Management effort should focus on populations which are likely to spread and produce new satellite populations. Deciding which aspects of an invading population to manage, new emerging distant satellites or the existing core from which most seed originates has been the subject of much debate. We use extensive survey data to define the core and satellite populations and dispersal function of Parkinsonia aculeata in the semi‐arid rangelands of the Northern Territory of Australia. We described populations in terms of three size classes, classified on the basis of plant height (small (<0.4 m), medium (>0.4 and <2.5 m) and large (>2.5 m)). Data were recorded as a spatial point pattern and a dispersal function was derived from the distribution of the distance of each medium plant to the nearest P. aculeata neighbour that was greater than 2.5 m. All large and medium P. aculeata were classified as members of core or satellite populations. From these classifications, we tested critical assumptions of Moody and Mack's model, namely (i) the satellites form multiple disjunct foci that do not overlap; (ii) the system is homogenous and there are no abiotic or biotic restrictions to the weeds expansion enabling it to expand radially in any direction; (iii) the densities of all foci are uniform; (iv) there is no extinction through stochastic processes; and (v) there are no threshold effects. When the medium size class was considered, the population density and area occupied by the weed increased. Two satellite populations were comprised solely of medium plants, suggesting the number of satellites increased marginally from eight to 10 when data from both size classes were considered. Most individuals were part of the core population. The core population increased in density and we argue this core population is most likely to give rise to new satellites and should be managed in preference to emerging satellite populations. We suggest the invasion is behaving like a source‐sink system, where the core population is the source of most new medium plants.     

Reproductive parasitism: male and female responses to conspecific and heterospecific intrusions at spawning in a mouth‐brooding cichlid Ophthalmotilapia ventralis

A rare form of alternative reproductive behaviour without simultaneous parasitic spawning was observed in Ophthalmotilapia ventralis, a lekking mouth‐brooding cichlid from Lake Tanganyika. Floater males attempted to sneak opportunistically into the territory to actively court the female, while the owner (bourgeois male) defended the territory against other potential intruders. Floater males had more body fat than territory owners and generally higher condition factors. In field experiments, the response of bourgeois males and courted females was tested towards floaters and egg predators (a catfish Synodontis multipunctatus) present in the territories. Territory owners responded aggressively particularly to floaters, and female responsiveness to bourgeois male courtship tended to decline when floaters were present. The potential influence of reproductive parasitism on sexual selection in mouth‐brooding cichlids is discussed.     

Implications of goat eradication on the survivorship of the Galapagos hawk

Non-native mammals cause ecological disasters in island ecosystems and their eradication is usually considered beneficial to native biodiversity. Goats (Capra hircus) were introduced to Santiago Island, Galapagos, Ecuador, in the early 1800s, and their numbers increased to about 100,000 by 1970. A goat eradication campaign initiated in 2002 was successful, eliminating the last individuals in 2006. To evaluate the effects of goat eradication, between 1998 and 2010 we studied the Galapagos hawk (Buteo galapagoensis) population on Santiago Island before, during, and after eradication. We used a 12-year data set in a capture–mark–recapture analysis to estimate the apparent survivorship of territorial adults in 33 breeding territories, and a 5-year data set to estimate the population sizes of the floater (non-territorial) fraction of the population. Juvenile floaters showed a drastic decline starting in 2006 and continuing in 2007, 2008, and 2010, which we attribute to the completion of goat eradication in 2006, and subsequent habitat changes. We found a significant decline in adult survivorship after the goat eradication program. Additionally, group size positively affected adult survivorship in this cooperatively polyandrous raptor, presumably reflecting the benefit of shared defense and offspring provisioning during harsher conditions. The changes in the hawk population after goat eradication are an example of unforeseen consequences of a restoration program, and we hypothesize that these changes are adjustments towards a new equilibrium under the current ecosystem characteristics and capacity. © 2012 The Wildlife Society.     

Effects of androgens on social behavior and morphology of alternative reproductive males of the Azorean rock-pool blenny

In the Azorean rock-pool blenny Parablennius sanguinolentus parvicornis two sequential reproductive tactics occur. Larger and older males establish breeding territories, while some of the smaller males become attached to nest-holder territories, acting as satellites on these territories, which they help to defend while trying parasitic fertilizations when females go in the nests to spawn. In the present paper we tested the effects of the androgens 17alpha-methyltestosterone (MT) and 11-ketotestosterone (KT) in the expression of male secondary sex characters and bourgeois behavior in satellite males. One week after satellites were implanted with Silastic tubes containing MT, KT, or castor oil (control), androgen-treated satellites had developed male secondary sex traits such as longer and wider male-type genital papilla and anal glands that secrete a sex pheromone, both traits being less expressed or absent, respectively, in satellite males. Androgen treatment had no effect on the gonadosomatic index or on the development of the testicular gland. KT treatment had a positive effect on relative liver weight. In terms of behavior, androgen-implanted individuals were less aggressive both in a mirror test and toward females when these were introduced into their tanks. MT-treated individuals spend more time inside the provided nests. Only androgen-implanted satellites managed to have the females entering their nests. When given a chance in a group tank either to try to attract females to their own nest or to act as satellites of an already established nest-holder's nest, MT-implanted males spent significantly more time in their own nest than near the nest-holder nest. These data suggest that androgens, particularly testosterone, may be involved in mating tactic switching in this species.