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1 June 2016 The Unknown Life of Floaters: The Hidden Face of Sexual Selection
Juan Moreno
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Sexual selection, as a form of social selection based on reproductive resources, is a crucial driver of evolutionary change. Many studies on sexual selection identify potential targets only within the reproductive fraction of populations. Floaters constitute the non-territorial fraction of the population, according to the usual definitions. Floaters have been identified through exhaustive capture and marking programmes, removal and nest-box addition experiments, extra-pair paternity studies, acoustic marking and genetic studies. The literature shows that floaters may represent a considerable fraction of populations, especially among males. There is no clear evidence that size, condition or testosterone level is necessary for explaining floater status generally. However, the literature suggests that ornament size and expression are involved in territorial exclusion and may be either its cause or one of its consequences. There is some evidence that floaters survive and reproduce less well than territorials, and that changes from floater to territorial status are accompanied by changes in survival and reproductive rates. However, certain male floaters may obtain some reproductive success through extra-pair copulations. The possibility that floating constitutes a successful alternative strategy in some species cannot be excluded, although the current preliminary consensus is that floaters are ‘making the best of a bad job’. Floater status may be imposed by limitations in the availability of mates or breeding space resulting in skewed population sex ratios, polygamous mating systems, high population densities and increased demand for specific breeding requirements such as space in colonies or adequate nesting cavities. Predictions concerning the effects of these factors have not been conducted to date. Few studies have been able to clarify the duration of floater status in any population. For short-lived species, floater status in a single breeding season may in fact imply zero lifetime reproductive success. In males, the existence of a considerable fraction of floaters attempting to breed may select for intense territorial behaviour and competitive mate guarding tactics in territory holders and in aggressive extrapair copulation and territory acquisition tactics in floaters. Interference competition from floaters may lead to density-dependent declines in reproductive success. In females, the attempts by floaters to attain breeding opportunities may have contributed to the observed propensities for female prospecting and for female-female aggression and the signalling of female dominance towards other females. Moreover, there may exist selection in females for signalling quality to mates in order to avoid being evicted by rivals. Excluding floaters from the analysis of sexually selected traits may severely affect sexual selection estimates because of biased sampling for large or more intensely expressed ornamentation. The importance of sexual selection may be negated or underestimated when in fact its action on floaters could be maintaining current levels of expression in the territorial fraction. Existing phenotypes should express, in their morphology, physiology and behaviour, the relentless drive through evolutionary time to avoid becoming a floater.


Sexual selection as a form of social selection based on reproductive resources is a crucial driver of evolutionary change (Darwin, 1871; West-Eberhard, 1983). There is some current debate concerning the fitness components linked to sexual selection (Cain and Rosvall, 2014). Some authors define sexual selection as only related to pure mating advantages (Lyon and Montgomerie, 2012), regarding other forces linked to intraspecific competitive processes as social selection (following West-Eberhard, 1983). However, for simplicity we here define sexual selection as based also on advantages related to fecundity. Given the overlap between traits involved in acquiring and defending mates and other resources necessary for reproduction, the difference between the two forms of selection may be rather semantic (Lyon and Montgomerie, 2012). Although the origin of sexual selection in its widest meaning lies in competition for access to gametes in anisogamous organisms, it has been transformed during evolution into a complex force scrutinising each and every organismic trait for its potential effect on mating and breeding success (Andersson, 1994). Many studies have been devoted to identifying the traits that are being sexually selected and to clarifying the strength of selection based on acquisition of reproductive resources. These resources include not only gametes and adequate pair mates but also the territories or nesting sites that are essential for reproduction. Traits that promote success in the acquisition of reproductive resources can be morphological, physiological or behavioural, or express a combination of aspects of phenotypes. They can serve to exclude competitors from available resources through aggression or signals of social dominance, but also to attract potential mates in a social or purely sexual context. They can be expressed in both sexes, depending on the relative strength of competition for reproductive resources operating in males and females (Tobias et al., 2012). Although males have traditionally been considered as the sex that experiences the strongest sexual selection, due to the intrinsic constraint arising from different rates of gamete production, females may also have to compete for resources other than the gametes necessary for successful reproduction, such as mates, territories or nesting sites. Thus, it is not obvious that the traits expressed by females in contests for reproductive resources are merely carry-over effects from selection on males (Tobias et al., 2012).

Many studies on sexual selection identify potential targets in the reproductive fraction of populations. Typically, they compare the success obtained in mating or offspring production by adult phenotypes in a breeding population and estimate selection differentials (e.g., Garant et al., 2004; Møller and Szep, 2005; Hegyi et al., 2006). This is the correlative or descriptive approach that has supported the importance of sexual selection for the evolution of morphological, physiological and behavioural phenotypes in many populations. Alternatively, other workers have conducted experiments in which they have manipulated the expression of morphological or behavioural attributes to try to register their importance for mating and reproductive success (Andersson, 1982; Møller, 1988). Usually, individuals have been caught in their breeding territories or at their active nest sites in order to manipulate their phenotypes. In both the descriptive and experimental studies, research has dwelt on the phenotypes of breeders or at least of territorial individuals. However, there is a fraction of adult individuals in most populations that do not reproduce during a proportion of their lifetimes (Newton, 1998). In birds such as crows, swans and waders, nonbreeders occur separately in flocks and can be readily observed and counted (Carrick, 1963; Harris, 1970; Holmes, 1970; Patterson, 1980). In others, such as many songbirds and raptors, some individuals live secretive lives in and around the territories of breeders or move continuously from one place to another (Kendeigh, 1941; Delius, 1965; Smith, 1978; Newton, 1979; Rohner, 1997; McNabb et al., 2007; Tanferna et al., 2013). This fraction of unmated, non-territorial individuals that are difficult to observe and count is defined by the term ‘floaters’ in the literature (Winker, 1998). The importance of floaters for population processes and conservation biology has been recently reviewed in the literature (Lenda et al., 2012; Penteriani et al., 2011). There is agreement that the usual neglect of this population fraction in the ecological literature may seriously undermine our understanding of many aspects of natural populations. Sexual selection is no exception in this context, as floaters may constitute the strongest expression of its strength in not merely reducing but annulling the reproductive success of certain individuals. Just comparing the phenotype-dependent success of the breeding or territorial fraction of populations in comparative or experimental studies is insufficient for understanding the force of a process in promoting competitiveness in reproductive contexts. This systematic undervaluation of sexual selection has been covered by recent reviews (e.g. Lenda et al., 2012) but has not yet received full attention from researchers. Here we concentrate on avian populations, for which knowledge on floaters remains scant. We only analyse breeding season floating, excluding all studies on winter floating (reviewed by Brown and Long, 2007). Given the existence of recent reviews on the importance of floaters for population dynamics (Newton, 1998) and conservation biology (Penteriani et al., 2011), this review only examines the implications for studies of sexual selection. No phylogenetically controlled comparative analyses have been conducted due to the paucity of information available and the strong taxonomic bias in the literature towards certain taxonomic groups, such as passerines.

Floaters and non-breeders

Floaters constitute the non-territorial fraction of the population, according to the usual definitions. Territorial status is used here in its widest meaning, including ownership of any resource necessary for breeding such as adequate space in a breeding colony, display space in a lek or an adequate nesting cavity. However, this fraction is composed partly by juveniles who have not attained the condition or experience necessary for initiating breeding activities (Cooper et al., 2009; Bayne and Hobson, 2001; Rivera et al., 2011; Mumme, 2015). These juveniles may routinely end up breeding as they mature (Newton and Rothery, 2001; Sergio et al., 2011; Rivera et al., 2011; Loewenthal et al., 2015) and hence are only temporarily excluded from the breeding fraction of the population. Moreover, being juvenile floaters may be the general pattern for most individuals and not a characteristic of a particular fraction (Delgado et al., 2009; Ryder et al., 2011). Thus, excluding such floaters from analyses may not seriously affect an evaluation of the strength of sexual selection. However, floaters also include adult individuals that do not breed for part or all of their lifetimes (Shutler and Weatherhead, 1991; Reitsma et al., 2008; Villavicencio et al., 2013). This fraction is defined by the non-acquisition of territories, which leads to their floating behaviour. For logistic reasons most studies deal with territorial birds that may be located and captured for phenotypic measurements. It is thus obvious that they are missing part of the picture by excluding floaters. The question is whether all floaters are non-reproductive, as emphasised by certain definitions. Assigning uncategorised birds to the resident territorial fraction on the basis of adult plumage (e.g., Morton et al., 2000) is questionable as it assumes that all floaters must be juveniles. We hereafter refer only to non-juvenile floaters. Future studies should attempt to identify the fraction of adult floaters as others (e.g., Butchart, 2000) have done.

To exclude cases in which floaters obtain reproductive success by means other than territory acquisition and pair-bond formation (see section “Is floating a strategy or the ‘best of a bad job’?”), we define non-reproductives (NR) as adult birds that are temporarily or permanently excluded from the breeding pool. To be regarded as such, floaters must be present in breeding areas. Hence individuals that are alive but not present at breeding colonies are NR but should not be considered as floaters. Not all individuals in seabird populations experiencing ‘sabbaticals’ or breeding intermittently (e.g., Aebischer and Wanless, 1992) can be considered floaters as some may not be present at the breeding colonies. So not all NR are floaters nor are all floaters NR (see section “Is floating a strategy or the ‘best of a bad job’?”). Some studies have found that NR non-floaters are less common than NR-floaters (9% and 91% of NR, Harris and Wanless, 1995). Future long-term studies should attempt to separate the NR non-floater and floater categories. The discussion below deals with floaters irrespective of whether or not they are NR.

How are floaters detected?

Exhaustive individual marking and identification of all resident and transient birds is difficult but may enable a clear insight into the lives of floaters (Penteriani and Delgado, 2012). Non-invasive genetic techniques may offer a viable approach for detecting and estimating the abundance of floaters through sampling of faeces or the feathers of moulting individuals (Rudnick et al., 2008). The abundance of floaters can be inferred from the speed or frequency of mate replacement or territory reoccupation after the loss of an individual or a pair (Driscoll et al., 1999; Fedy and Stutchbury, 2004; Schweizer and Whitmore, 2013; Mumme, 2015). There are also experimental ways to detect floaters. The rapid reoccupation of territories vacated through owner mortality may indicate the presence of numerous floaters (Vili et al., 2013). The most common way with which floaters have been identified relies on experimental removal of territorial individuals and the detection of subsequent territory occupation by individuals not associated with any known territory or nest site. Newcomers should breed in the same year to confirm that they are mature and capable of breeding. Moreover, it is necessary that removal and replacement should involve both sexes as otherwise the experiment may reveal only an unequal sex ratio. A large fraction of avian studies incorporating floaters into the population picture have been based on experimental removals. Newton (1998) reviewed extensively the literature on removal experiments. From his table 4.1, one can deduce that one sex was replaced in 62% of 34 removal studies on 23 songbird species in spring, and in 23% both sexes were replaced. Replacements appeared in six of 12 studies of seven grouse species. For all species combined (74 studies on 53 species), there were replacements in 43%, indicating a non-breeder surplus.

The literature on replacement experiments for the last two decades (table 1) shows that 0–100% of removed males are replaced within periods ranging from a few hours to several weeks, with an average rate of replacement of 60% ± (SD) 34% (13 studies on 11 species, table 1). Few studies have removed female territory owners (table 1) but they suggest that female floaters are less common (43% ± 33% replacement, n = 6 studies on six species). These data indicate that there are more male than female floaters (Marra and Holmes, 1997), although the difference between sexes in replacement rates is not significant (Mann-Whitney U-test, U = 31, P = 0.16). They also show that male floaters can occupy a high proportion of vacancies, with replacements being more rapid and frequent in high quality, continuous habitats.

Table 1

Removal studies in which floaters have been detected (only involving reoccupation by floaters, not by neighbours or territory switchers).

[Estudios de retirada de territoriales en que se han detectado flotantes (solo reocupación por flotantes, no vecinos o individuos que cambian de territorio.]


However, these studies offer only a partial view of the floating fraction as not all existing floaters may be able to occupy experimentally vacated territories. This has been shown for several species of forest grouse (Fischer and Keith, 1974; Lewis and Zwickel, 1980; Szuba and Bendell, 1988). The occupation of vacated territories may depend on territory quality, with floaters only colonising high quality areas (Manuwal, 1974; Bowman and Bird, 1986; Porter and Coulson, 1987; Newton and Marquiss, 1991; table 1). Some vacated territories may be occupied by neighbours and switching territory owners and not by floaters (Butchart et al., 1999; Pryke and Andersson, 2003; Fedy and Stutchbury, 2004; Villavicencio et al., 2013). Moreover, removal studies may only give a minimum estimate of floaters where 100% replacement occurs. The floater fraction can only be accurately estimated in these cases when the same number of territory owners as of existing floaters is removed. In removal studies on nest-box breeding populations, prompt occupation by presumed floaters has been detected after removal of nest-box owners (Heusmann and Belville, 1978; Alatalo et al., 1983). The absence of replacements in some experiments where pairs or females are removed may be due to a lack of female floaters (Marra and Holmes, 1997).

In species that are limited by nest-site availability, such as many cavity-nesters, the provision of artificial sites such as nestboxes often leads to an immediate increase in breeding density (Newton, 1994, 1998; Wiebe, 2011). Wiebe (2011) reviewed 31 studies of 20 species where the density of cavity nests in mature forest habitat was manipulated. Changes in breeding density on treatment plots were reported in ten experiments (32%), but statistically significant effects analysed by species were reported only in six cases (19%). With the exception of Bortolotti (1994), who added nest-boxes late in the breeding season to test for the presence of floaters in the population, none of the studies attempted to determine whether NR occupied boxes. No researchers in studies in which changes in breeding density were found tracked the movements of individually marked birds before and after boxes were installed, and none estimated breeding densities in buffer zones surrounding the treatment plots to control for movements of individuals across plot boundaries. The conclusion by Wiebe (2011) is that reviewed studies provide no strong evidence that there is in fact a surplus of NR imposed by cavity limitation. Wiebe (2011) offers recommendations for the design of future studies wanting to approach this matter. However in some studies of non-forest birds, the occupation by floaters was confirmed through rings (Stutchbury and Robertson, 1985; Village, 1990). Saitou (2001) put up additional nestboxes for grey starlings Sturnus cineraceus and these were quickly occupied by floaters in the early part of the breeding season. The intensity of intraspecific brood parasitism (IBP) was significantly reduced. The removal of boxes had the opposite effect. Floater females were thus involved in IBP before manipulation. As in removal experiments, the possibility that newcomers arrive from breeding territories elsewhere should be excluded before a rise in breeding density can be attributed to the presence of floaters. Moreover, as in removal experiments, the numbers of floaters may be underestimated.

Another form of floater detection has involved studies on extra-pair paternity in populations of genotyped territorial birds. Commonly in these studies, the sires of many extra-pair offspring are not identified genetically among territorial genotyped males (table 2). In general, almost two-thirds of extra-pair offspring (64 ± 20%, n = 7 studies on 6 species) in recent studies of extra-pair paternity are not assigned to a sire among resident territorial males (table 2). This fraction may be overestimated if some territorial males escape control by researchers. The options are that these sires are male floaters or that they are territorials outside the study area (Peer et al., 2000; Kempenaers et al., 2001). Given that the first possibility appears more plausible based on the presumed costs of looking for extra-pair copulations (EPCs) far from the territory (Dunn et al., 1994; but see Leisler et al., 2000; Kempenaers et al., 2001), the evidence of floater existence derived from extra-pair paternity studies appears robust. The same can be said for genetically identified cases of IBP where the parasitic embryos or nestlings cannot be genetically connected with any identified breeding female in the study area. These cases appear much less frequently, which may signify that in most populations female floaters constitute a relatively smaller fraction than male floaters. In studies in which the floater fraction has been genetically identified, one in four extra-pair offspring are sired by identified floaters (23 ± 15%, n = 5 studies on 5 species, table 2). The lower rate of floater involvement recorded in these studies compared with participation in extrapair activity by unidentified males may be due to incomplete floater identification or to incomplete monitoring of territorial residents in the studies mentioned above. Alternatively, unidentified sires may be resident in unmonitored adjacent habitats. In any case, these studies imply scant reproductive success for the fraction of floaters involved in EPC, thereby strengthening the ‘best of a bad job’ hypothesis (see section “Is floating a strategy or the ‘best of a bad job’?”). The main fraction of floaters may not reproduce at all (Cooper et al., 2009).

Table 2

Proportion of extra-pair young (EPY) sired by floater males (*, considering broods with genetically identified floaters) or by unidentified males not recorded as territorial (no floaters identified as such), and the proportion of broods containing some EPY. The numbers of nestlings and broods studied are also presented.

[Proporción de pollos extrapareja (EPY) producidos por machos flotantes (*, considerando nidadas con flotantes identificados genéticamente) o por machos no identificados y no registrados como territoriales (sin flotantes identificados como tales), y la proporción de nidadas con algún EPY. Se presentan también las cifras de pollos y nidadas estudiados.]


Some studies have identified floaters through territory take-overs after nestbuilding and egg-laying (Butchart et al., 1999; Moreno, 2015) or during repeated visits to leks (Westcott and Smith, 1994). The birds taking over territories at this stage may induce floater or NR status in the evicted individuals, which have no time to establish a territory of their own or to form a pair bond (Piper et al., 2000; Fedy and Stutchbury, 2004). However, the possibility remains that evicted birds are successful in establishing themselves away from the study area where the take-over took place and thus remain undetected as late territorials.

Acoustic marking (Voegeli et al., 2008; Kirschel et al., 2011) and recordings of spontaneous calls together with broadcast of male territorial songs in owls (Martínez and Zuberogoitia, 2002) have also been useful in detecting floaters. Estimates of numbers of floaters based on visual estimation may seriously underestimate their frequency when compared with non-invasive genetic sampling (Katzner et al., 2011).

How large is the floater and NR fraction?

If floaters and NR constitute very small fractions of avian populations, their role in clarifying the strength of sexual selection may be irrelevant. Therefore, it is essential to know the size of the pool of floaters and NR in any populations for the purposes of this review. The traditional focus of the literature on avian floaters is on the importance of density-dependent population regulation. Territoriality is described as a process regulating population numbers by excluding certain individuals from breeding when density approaches a critical level (Newton, 1998). The argument involved occasionally a groupselection aspect, now discredited, portraying reproductive exclusion as an adaptation for promoting higher success for the population as a whole (Wynne-Edwards, 1962). However, group selection is unnecessary in this context as individuals may achieve the exclusion of others through their own competitive behaviour that is favoured by individual selection (Newton, 1998). The emphasis has been put on food resources as the main driver of the size of any population. Individuals are excluded from breeding by territorial birds wanting to ensure sufficient food resources for breeding successfully. This argument does not apply to colonial birds whose territories only include a small area around their nest site without any food resources. Nevertheless, there also floaters in colonial seabird populations (Young, 1972; Manuwal, 1974; Pierotti, 1980). Thus exclusion seems to involve something else than just food supplying territories. Reproductive exclusion is directly related to the capacity to acquire mates or space for breeding in competitive contexts. Stronger limitations on these resources are likely to result in a larger floating or NR population.

It has been assumed that floaters constitute a larger fraction of the population in large long-lived birds, given the existence of a large pool of young, immature birds and the occurrence of delayed breeding in these species (Newton, 1998). However, if we exclude the immature fraction from the nonbreeding pool, the contention remains questionable. It could be based on the difficulty of observing and counting floaters in small, short-lived species. Brown (1969) estimated a theoretical maximum ratio of non-breeders to breeders for a range of bird species with different reproductive and mortality rates. In the most extreme case, non-breeders could outnumber breeders by two or more times. Thus, in theory, competition for breeding resources (including participation in leks) could be excluding a large fraction of individuals from reproduction.

The literature shows that floaters may represent in fact a considerable fraction of the population for both sexes. Newton (1998) reviewed published papers up to 1996. Based on his table 3.2, we may conclude that in 20 studies on 17 species, ranging in size from wrens to swans, on average 39% ± 22% (range 3%–72%) of individuals were nonterritorial non-breeders. In eight studies in which fractions were estimated separately for both sexes, the value was higher for males in six cases and higher for females in two cases. Values for females ranged from 0 in the song sparrow Melospiza melodia to 62 in the northern goshawk Accipiter gentilis (mean 28% ± 26 %, n = 10 studies). This indicates that, although less common than male floaters, female floaters are present in some avian populations and may constitute an important fraction of females. Although values are generally higher for large species in Newton's review, they may include immature birds which are not dealt with here. It is therefore possible that mature non-breeders are as common in large-bodied as in smallbodied species.

Studies from the last two decades in which the floater fraction has been identified, and some papers not cited by Newton (1998), show that 41% ± 26% of individuals behave as floaters (eight studies on seven species) (table 3). In studies in which the floater fraction has been estimated separately for the two sexes, 40% ± 23% of males (14 studies on 14 species) and 23% ± 18% of females (nine studies on nine species) behave as floaters (table 3). Again, we find that female floaters are less abundant in avian populations than male floaters, although the difference is not quite significant (Mann-Whitney U-test, U = 33, P = 0.058). However, floaters may still represent a considerable fraction of individuals even for females. These figures do not change appreciably if we exclude studies in which it is specified that the floater fraction includes juveniles (table 3). Unfortunately, most studies do not report the proportion of floaters made up of juveniles. The fact that between a quarter to almost half of the population in these studies is made up of non-territorial floaters supports the importance of considering this fraction in studies on sexual selection.

Are floaters phenotypically different from residents?

Excluding situations in which it is predominantly young birds that are prevented from breeding, it would be interesting to know the traits characterising floaters when compared to territorials and the factors that may facilitate the transition between floater and territorial status. In some long-lived birds, site tenacity appears to be more important than age in determining an individual's success in establishing a territory (Sergio et al., 2009; Loewenthal et al., 2015). Some studies have shown that floaters were drawn from among the most dominant individuals in the non-territorial fraction of the population (Knapton and Krebs, 1974; Smith and Arcese, 1989). In several studies in which removed territory owners were held captive and later released, such birds mostly managed to displace their replacements to regain their territories and mates, either in the same year or the next (Watson and Jenkins, 1968; Harris, 1970; Smith, 1978; Szuba and Bendell, 1988; Village, 1990). This indicates that replacing floaters are subdominant to the original territory owners. In the great tit Parus major, however, the probability that replacement pairs would be able to retain their territories increased with the time elapsed before the original owners were encountered again, supporting a role for ‘owner’ effects (Krebs, 1982). Size or condition is often linked to non-territorial status (Alisauskas, 1987; Richner, 1989). A defining characteristic of floaters in migratory species may be delayed arrival at the breeding grounds (Sergio et al., 2009). However, delayed arrival is probably the consequence of certain physiological or behavioural attributes, so future studies should clarify the underlying basis for late initiation of migration or migration speed, although delayed arrival must sometimes be due to extrinsic factors, notably weather. Breeders and floaters may also show a different habitat use (Campioni et al., 2010, 2012) and diet (Caro et al., 2011). These differences may be a consequence, not the source, of floater status.

Table 3

Presence (Y = yes, N = no) and fractions of floaters (number of individuals in parenthesis) in avian populations (ads includes specifically only non-juveniles, juv includes young birds that reproduce later, m&f presents joint data for both sexes, when several years are presented averages or data for the year with most data are given).

[Presencia (Y = si, N = no) y fracciones de flotantes (número de individuos en paréntesis) en poblaciones de aves (ads incluye específicamente solo no juveniles, juv incluye jóvenes que se reproducen luego, m&f presenta datos para el conjunto de los dos sexos, cuando se presentan varios años se ofrecen medias o datos del año con más datos).]




Table 4

Phenotypic differences (= similar, > < different, > greater, < smaller) between floaters (F) and territorials (T) and method used for the identification of floaters (R = removal, B = Banding and territory mapping).

[Diferencias fenotípicas (= similar, >< diferente, > mayor, < menor) entre flotantes (F) y territoriales (T) e identificación de flotantes (R = retirada, B = marcaje y mapeo de territorios).]




Excluding age, several other traits have been related to floater status. I have revised the literature for the last two decades for studies in which an attempt has been made to compare traits of territory occupants and floaters (table 4). Studies that clearly confound floaters with dispersing juveniles have not been included here. These studies present a mixed picture in which floaters do not differ from residents at all or only with respect to specific traits (table 3). The most common traits analysed are mass and body condition (ten studies on eight species), structural body size (six studies on five species) and ornament size (seven studies on seven species). Floaters are lighter in three cases and smaller in two cases, with two studies showing floaters in better condition than residents. Thus, there is no clear evidence that either size or condition is necessary for explaining floater status generally. However, in six of seven studies (86%) male floaters were less ornamented than residents and in three of four cases (75%) female floaters were less ornamented. Thus, the literature suggests that ornament size and expression are involved in territorial exclusion and may be either its cause or one of its consequences. Experiments in which ornament expression is manipulated and its consequences for acquisition of floater status evaluated are sorely needed. Testosterone levels have been measured in four studies on three species (table 4) and were lower in floaters than in resident territorials in only one case. Floaters proved to be less aggressive in only one of two studies on dominance and aggression. Thus, testosterone does not seem to be involved in determining floater status, although data are scant. One conclusion from this review is that excluding floaters from an analysis of sexually selected traits may severely bias distributions towards larger ornament sizes or more intense expressions.

Is floating a viable strategy or the ‘best of a bad job’?

There is evidence that floaters survive less well than territorials, and that changes in territorial status are accompanied by changes in survival rates (e.g., Carrick, 1963; Smith, 1976; Watson, 1985; Harris and Wanless, 1995; Rohner, 1995; Cam et al., 1998; Dwyer et al., 2012). Together with the reduction in breeding opportunities (Smith and Arcese, 1989; Stutchbury and Robertson, 1985, 1987) this supports the contention that floaters are low-quality birds following a conditional strategy that, in general, implies reduced fitness (Newton, 1998). In several species, replacers achieve significantly lower reproductive success than the original territorial occupants, a probable effect of their younger age (Manuval, 1974; Village, 1990; Newton and Marquiss, 1991; Komdeur and Edelaar, 2001). The evidence on breeding performance excluding age effects is scant (Linz et al., 2011). Intermittent breeding in seabirds may be an indication of poor ability to raise progeny (Bradley et al., 2000). However, there is some evidence that certain male floaters may obtain some reproductive success through extra-pair copulations (EPC) (Ewen et al., 1999; Johnson et al., 2000; Leisler et al., 2000; Peer et al., 2000; Conrad et al., 2001; Kempenaers et al., 2001; table 2). In some cases, floaters enjoy a better body condition than territory owners (Andersson, 1994; Kempenaers et al., 2001) which could allow them to pursue EPC successfully. Floating may also constitute a temporary conditional strategy in some cases (Fedy and Stutchbury, 2004). The possibility that floating may constitute a successful alternative strategy in some species, as shown for ‘sneaker’ strategies in some fish (Taborsky, 1994), remains a tantalising possibility to be explored. However, it should be shown that floaters on average and not just in a fraction of cases attain similar lifetime fitness as territory owners on average. For that, we require data on survival and lifetime reproduction of most of the floaters in a population.

Although there is some disagreement concerning the relative success of these floater strategies compared with territorial ones, the preliminary consensus at present, based on as yet scant studies, is that floaters are ‘making the best of a bad job’ (Rohner, 1995, 1997; Newton, 1998; Cam et al., 1998; Johnson et al., 2000; Cooper et al., 2009). Thus, although adult male floaters appear capable of engaging in extra-pair copulations in the red-winged blackbird Agelaius phoeniceus (Moulton et al., 2013), no direct genetic evidence exists to indicate that they produce extra-pair young (Weatherhead and Boag, 1995; Gray, 1996; Yasukawa et al., 2009). In fact, male floaters appear to be waiting in most cases for ‘real’ reproductive options through territory acquisition (Smith, 1978; Ens et al., 1995; Bruinzeel and van de Pool, 2004), something unexpected if floating is a stable evolutionary strategy. Involuntary movement between nest sites or territories may be followed by floating during several years (Kokko et al., 2004), which suggests that floating is also involuntary. Moreover, floaters have to compete with territorial individuals, who may be responsible for most extra-pair affairs (Zilberman et al., 1999; table 2). The general pattern is for territorial males to sire the majority of offspring (Jonson et al., 2000).

Female floaters could compensate for their lack of pair bonding and territoriality through intense egg dumping (Sandell and Diemer, 1999; Saitou, 2001). Parasitic females have been shown to be of high quality and to survive better in the cliff swallow Petrochelidon pyrrhonota (Brown and Brown, 2004). However, full compensation of their floater status would probably require intensities of IBP only found in some species (Zhang et al., 2011). Moreover, there is evidence that intraspecific egg dumping in ducks is in fact mediated through kin selection (Andersson, 2001). The evolutionary stability of alternative reproductive strategies in birds is therefore not ensured. However, the possibility of floating as a successful strategy remains suggestive.

Limitations on reproductive resource acquisition

Shortages of mates or breeding space may involve skewed population sex ratios, polygamous mating systems, high population densities and specific breeding requirements such as space in colonies or adequate nesting cavities. Population density is thus only one factor promoting the existence of floaters and NR. Some studies have linked reproductive exclusion of males in monogamous systems to a scarcity of females in the population induced by ecological factors. Females may be more vulnerable to several mortality factors, such as starvation or predation on the nest (Breitwisch, 1989). However, the population sex ratio is frequently deduced from the territorial fraction of the population without considering the floater fraction. Nevertheless, there may be a considerable fraction of female floaters excluded from reproduction due to lack of mates or resources, whose inclusion could change the sex ratio estimates derived by researchers. Until this fraction is included in sex ratio computations, the sex ratio basis for male floating remains in doubt.

The opportunity for sexual selection has frequently been linked to operational sex ratios. The operational sex ratio (OSR) is usually defined as the ratio of fertilisable females to sexually active males and its derivation in practice excludes individuals whose status is either unknown or uncertain regarding their capacity to reproduce. The ratio of male to female floaters may be different from the OSR and may have consequences for sexual selection. A strong bias towards male floaters may promote competitive behaviours in males while the opposite bias would favour female signalling and competition. The mating system may critically impinge on the sex ratio of the floater population. In strongly polygynous systems, many males are necessarily excluded from reproduction (Shutler and Weatherhead, 1991, 1994; Moulton et al., 2013), while the opposite may happen in socially polyandrous systems (Butchart, 2000; Emlen and Wrege, 2004). Given an unbiased population sex ratio, we should expect more male than female floaters in the former case and the opposite in the latter. The degree of polygamy may thus constitute a critical factor in explaining the frequency of floaters in avian populations. Floating is thus a product of sexual selection that may in turn promote further sexual selection to evade this fate in a vicious circle, whose evolutionary outcome we can observe today.

The main cause of floating has traditionally been considered to be habitat crowding in conjunction with territoriality (Newton, 1998). Accordingly, habitat limitation should promote floating (Komdeur, 1996; Snetsinger et al., 2005). Rigid territoriality and high life expectancy may induce floater behaviour depending on prey density (Barraquand et al., 2014). Even in lekking species, opportunities for acquiring territories may be limited (Ryder et al., 2011). Many colonial birds are restricted to breeding at specific locations with adequate conditions with respect to predator avoidance, ease of access or proximity to food sources. Adequate space or nest sites could be as limiting in colonial species as in those with food-based territories. We should expect considerable floater fractions in species with the most restricted habitat requirements for breeding (reviewed in Newton, 2008). Other birds require cavities for nesting that they cannot excavate themselves (secondary cavity nesters). Access to these cavities when in short supply may be limited and may exclude a fraction of potential breeders from reproduction (Newton, 1994, 1998; Wiebe, 2011). We should expect a higher floater fraction in cavity nesters than in open nesters in otherwise similar conditions i.e. within the same study area. To my knowledge, no study has approached this issue with marked floaters. There is evidence that floaters prefer to wait for vacancies in good breeding areas rather than occupying low quality areas (Manuwal, 1974; Lewis and Zwickel, 1980; Porter and Coulson, 1987; Newton and Marquiss, 1991; Rutz and Biljsma, 2006). They may also confront the choice of accepting a poor territory now or waiting another year or years before a better territory becomes available (Ens et al., 1995; Holt and Martin, 1997). Floaters may in effect be queuing for good territories (Smith, 1978; Bruinzeel et al., 2006). Thus, competition for good sites may be determining the size of the floater fraction. Without good knowledge of the determinants of habitat quality for breeding, the presence of a considerable fraction of floaters may be difficult to understand.

Exclusion: temporary or permanent?

Few studies have been able to clarify the duration of floater status in any population. Floaters may either acquire a territory or disappear without breeding (Piper et al., 2000; Piper, 2001). Evicted territory owners may become floaters that may later return to resident status (Westcott and Smith, 1994; Butchart, 2000). Temporary floater status may not imply permanent exclusion from the breeding pool in long-lived birds (Saitou, 2001; Fedy and Stutchbury, 2004; Schmutz et al., 2014). Thus, so-called ‘sabbaticals’ or extended non-breeding periods are common in seabirds and may only involve temporary floater status (Aebischer and Wanless, 1992; Bradley et al., 2000). However, being a floater in one year may significantly reduce the chances of breeding the next year (Cam et al., 1998). Site tenacity may contribute to success in establishing a territory rather than age per se (Loewenthal et al., 2015). For short-lived species, NR status in a single breeding season may in fact imply zero lifetime reproductive success. Thus there should be stronger selection for avoiding floater status in short-lived birds. This should be expressed through the extent and intensity of aggressive territory take-over behaviour in both males and females of these species. Lethal levels of aggression have been observed in passerines in contexts of territory take-overs by females (Moreno, 2015). Strikingly little is known on the levels of lethal intraspecific aggression and the frequency of deaths in either sex in territorial or mate-acquisition contexts. Deaths during competition for reproductive resources may be the expression of selection for avoidance of floater status.

The literature on NR in social species is covered under the term “reproductive skew”. When not linked to age-dependent processes, NR in avian social groups experience the same fitness reduction as in other species (Stacey and Koenig, 1990; Ridley et al., 2008). Avian sociality may be favoured by the costs of living alone as a floater in social species (Ridley et al., 2008), but may be constrained by adaptations for avoidance of NR status. The absence of any floaters has been detected in at least one species of cooperative breeder (Eguchi et al., 2007), which may suggest that floaters become subordinate helpers in these species in order to avoid permanent exclusion from breeding resources. The increased aggressiveness promoted by NR may inflate the costs of coexisting in social groups and favour solitary living or family groups where kin selection may soften competition for breeder status to some degree. Although reproductive skew has received a great deal of attention recently, it has not been clearly linked to studies on floaters in avian populations. It is doubtful if any social grouping can sustain permanent exclusion of breeding status in some of its members without breaking apart. We should not therefore expect drastic and permanent measures of reproductive exclusion affecting adult birds in groupliving birds.

Competition or choice?

Since Darwin (1871), the two main mechanisms behind sexual selection are considered to be competition for reproductive resources and choice by the operationally limiting sex, usually females. In males, the existence of a considerable fraction of NR attempting to breed may select for intense territorial behaviour and competitive mate guarding tactics in territory holders and in aggressive territory acquisition tactics in floaters (Arcese, 1987; Westcott and Smith, 1994; Zilberman et al., 2001; Carmen, 2004; Gruell et al., 2007; Moulton et al., 2013; Turrin and Watts, 2014). A higher density of floaters may require a higher level of signalling in territory owners throughout the season (Stutchbury, 1992; Sunde and Bolstad, 2004; Penteriani and Delgado, 2008). Floaters may prospect for territorial vacancies intensively, explaining the frequent appearance of intruder males at active nests (Tobler and Smith, 2004; Dwyer et al., 2013; Veiga et al., 2013; Turrin and Watts, 2014; Wischhoff et al., 2015). They may look out for weak or senescent territory owners (Westcott and Smith, 1994; Bornschein et al., 2015). Only high quality owners may be able to keep away intruders (Moreno et al., 2013). Increasing interference competition from floaters may explain why some territories experience a density-dependent reduction in reproductive success (Bretagnolle et al., 2008; Grunkorn et al., 2014). Floater pressure may impose the formation of polyandrous trios in some cases (Carrete et al., 2006). Floaters may use the presence of nestlings or fledglings to target territories for future attempt at territorial take-over (Piper et al., 2006). The high level of male-male aggression (Sunde and Bolstad, 2004) and of attacks on chicks by floaters (Kazama et al., 2012) observed in some populations may be difficult to understand without considering the need to obtain a territory and such behaviour may compromise the reproductive success of the breeding fraction (e.g., Carrete et al., 2006; Kazama et al., 2012).

Floater males may be also selected to try to force copulations or harass females in relentless attempts to reproduce (Moulton et al., 2013). Females may obtain direct benefits from mating with dominant males that can keep harassing floaters away (Moreno et al., 2013). The existence of intense male aggression towards other males, and towards breeding females, may be the consequence of the existence of a considerable floater fraction. Only aggressive floaters in the past may have contributed genes to future generations. Females may have been selected to avoid male harassment and forced copulations, which may explain the rarity of extrapair fertilisations in many studies - most workers explain this rarity by female choice mechanisms. More work is required to clarify the importance of floaters in the evolution of extra-pair paternity.

In females also, the desperate attempts by floaters to attain breeding opportunities may have contributed to the observed propensities for prospecting by females (Veiga et al., 2013) and for female-female aggression and the signalling of female dominance towards other females (Stutchbury and Robertson, 1987; Moreno et al., 2014). Female floaters may frequently be chased away by territory owners to preclude IBP tactics (Veiga et al., 2012) and this may lead to increased androgen levels in females in high density situations with the presence of more floaters (Pilz and Smith, 2004). The need to avoid such aggression may have selected for delayed plumage maturation in females of some species (Coady and Dawson, 2013). Although female aggression towards other females has received much less attention than male aggression, evidence is accumulating that females may compete aggressively for breeding opportunities and that they may signal their aggressive dispositions (Tobias et al., 2012; Cain and Rosvall, 2014). Including avoidance of NR status in analyses may help to explain the current distribution of social signals during the breeding season in female birds.

Floater males may try to attract fertile females for EPCs by exhibiting signals of quality (Gruell et al., 2007). Given the low social status of floaters observed in several studies (see below), their success in this endeavour appears doubtful (Moulton et al., 2013). Some studies report a high proportion of unidentified EPC perpetrators which suggests that EPCs may be driven to a large extent by floater activity (see above). The relative importance of extra-pair female mate choice versus male drive to obtain EPCs at any cost is currently under debate. Under the first scenario, females could be attempting to ameliorate their initial pair choice concerning social bonds by seeking EPCs. Including floaters in this scenario of female choice is the unavoidable consequence of the presence of floaters among extra-pair sires. However, it is at present risky to assume that all floater-dependent EPCs rely on female mate choice. The aggressive scenario of male harassment and forced copulation attempts remains a plausible alternative (Westneat and Stewart, 2003).

In the case of female floaters, they may be less attractive to territorial males and thereby receive less male support in competitive interactions. This possibility has not been well covered by the literature, although there is evidence that females are more aggressive towards intruders when mates are absent and when mates have low testosterone levels (Morales et al., 2014; Moreno et al., 2015). These results suggest that females rely partly on their mates to conserve their mating status. Being attractive may strengthen mate support in territorial contests with intruding female floaters. Thus, there may exist selection in females for signalling quality to mates in order to avoid being evicted and converted into female floaters. Female floaters may be more prone to intrude in occupied territories than male floaters due to the lower risk of suffering attacks by male owners (Campioni et al., 2010).

The underestimation of sexual selection

Relating the mating and reproductive success of territorial birds to the expression of potential signals or weapons is the typical approach of studies on sexual selection. Expression of sexually selected traits can be experimentally manipulated or just observed in natural conditions. In studies excluding floaters, trait expression by the NR fraction of the population is unknown. Therefore, the full population range of variation of the traits studied is insufficiently understood (see above). Existing trait expression in the territorial fraction may be difficult to interpret without considering the range of values removed by selection acting on floaters (fig. 1). Current selection acting on certain traits may be underestimated when the fraction of individuals with low reproductive success imposed by floater status is excluded from the analyses (fig. 1). The importance of sexual selection may be negated when, in fact, its action on floaters may be maintaining current levels of trait expression in the territorial fraction. Phenotypic variation in the breeding population may be unrelated to mating or breeding success but its existence may be explained by selection against unidentified phenotypes in the coexisting floater population. We should be therefore remiss to disclaim sexual selection processes operating in populations when the floater fraction has not been identified. Studies estimating sexual selection differentials with and without including floaters should be conducted to judge the degree of underestimation of sexual selection in the literature.

Fig. 1.

Upper graph: Including floaters (broken line) may displace the distribution of a sexually selected trait towards lower expressions than observed in territorials (continuous line). Lower graph: This implies that the fitness difference between maximum and mean values (dotted lines) is greater when floaters are included in the study and selection is accordingly stronger (arrows).

[Gráfico superior: Incluir a los flotantes (línea a trazos) puede desplazar la distribución de rasgos sexualmente seleccionados hacia expresiones más bajas que lo observado en los individuos territoriales (línea continua). Gráfico inferior: Ello implica que las diferencias en eficacia biológica entre los valores máximo y medio (líneas punteadas) es mayor cuando se incluye a los flotantes en el estudio y la selección correspondientemente más fuerte (flechas).]


The existence of female floaters raises the question of determinants of the strength of sexual selection forces. If these females could breed, the OSR would be less malebiassed and thereby could soften inter-male competition for breeding opportunities. Thus, egg dumping apart, the exclusion of certain females from the reproductive fraction should be studied as a driver of sexual selection whose importance depends on the fraction of NR females. The extent of female floating should depend on female competition for reproductive resources which prevents males from attracting other females to their territories and determines the lower limits on territory size. These processes, considered by some authors to be covered by the term ‘social selection’ but here included in a wide conception of sexual selection, may be essential in explaining the expression of territorial behaviour, aggression and signalling in birds, as well as in other animals. Female-female competition for breeding resources may be limiting the female breeding pool and inducing strong competitiveness and attractiveness in males to ensure their breeding status. The existence of floaters confirms the vital role of female competitiveness in driving sexual selection.

An assumption in population biology and conservation biology is that floaters constitute a buffer that may dampen increases in breeder mortality, thereby reducing the extinction risk of populations (Newton, 1998; Sarah et al., 2004; Blanco et al., 2009; Penteriani et al., 2011; Hockey et al., 2011). This assumption is based on the supposed phenotypic similarity of breeder and floater fractions, possibly only separated by age (Penteriani et al., 2009). However, the substitution of competitively superior, highquality breeders by floaters may in fact represent a decline in the mean phenotypic quality of breeders, based on available evidence on non-juvenile floaters (Cam et al., 1998). If floaters constitute a geneticallybased alternative strategy (see above), their occupation of vacated territories may even imply a reduced breeding performance as territorial breeders given their poor adaptation for this role. In general, colonisation by floaters of empty territories should not be considered an unmitigated plus for the conservation of endangered species. The buffer may to a high degree be made up by the poorer phenotypes in the population.

Table 5

Identification of questions and tasks for future research on the role of floaters in sexual selection.

[Identificación de preguntas y tareas para investigación futura sobre el papel de los flotantes en la selección sexual.]


Conclusions and future prospects

Several important issues require further research in order to clarify the role of floaters in sexual selection (table 5). Although floater individuals have recently been incorporated into the picture of population processes and conservation biology, their identity has seldom been established in field studies. Establishing the existence of “surplus” individuals depends on verifying the identity of non-territorial birds, a difficult proposition requiring specific programmes for marking individuals and so identifying each and every territorial bird in the study area. It should become a priority in avian studies to identify the floater fraction. Moreover, the implications of floaters in EPCs and egg dumping should be further clarified through detailed observational studies. Are most floaters really NR? It is insufficient to genotype nestlings; we need to genotype the floaters themselves. Another question raised is whether all floaters are involved in these alternative reproductive strategies. Genotyping is unable to detect the real NR fraction, for which observational studies of marked birds are necessary. The identification of floaters should be conducted across years to estimate the duration of exclusion status for individuals. How frequently do floaters change their status during their lifetimes? Is it a once-in-a-lifetime transition or can territorial status be lost after being acquired? Lethal or injurious exclusion should be quantified if possible to try to estimate the survival implications of floater status. If a considerable fraction of natural populations suffers permanent exclusion from the breeding pool through competitive processes, the implications would be similar to those of the typical forces underlying natural selection, such as disease, predation, exposure and starvation.

Darwin (1871), the originator of the idea of competition for breeding resources and of sexual selection, wrote that selection processes unrelated to intraspecific competition -which he called natural selection- were probably stronger than those acting upon competition between individuals for participation in the breeding pool. In his view, natural selection was continuously removing a certain fraction of individuals from the breeding population by not allowing them to survive, while sexual selection just established the relative number of offspring of those able to breed. He was thereby underestimating the force that he was first to detect and understand. Sexual selection is also generating non-breeders, although they may be alive and healthy. In this way, by contributing not only to reduce but to prevent reproduction by a fraction of the population, it is shaping phenotypes for competition in both sexes. The levels of aggression and signalling exhibited by many birds may not be fully understood without considering that many individuals in natural populations never get a chance to breed. Existing phenotypes should express in their morphology, physiology and behaviour the relentless drive through evolutionary time to avoid this fate.


A. Velando invited me to contribute a review and commented on several preliminary versions. L. M. Bautista helped with the graph. An anonymous reviewer contributed to improving a previous version. I was supported while writing by project CGL2013-48193-C3-3-P of Dirección General de Investigación Científica y Técnica.



Aebischer, N. J. and Wanless, S. 1992. Relationships between colony size, adult non-breeding and environmental conditions for shags Phalacrocorax aristotelis on the Isle of May, Scotland. Bird Study , 39:43–52. Google Scholar


Alatalo, R. V. , Lundberg, A. and Stahlbrandt, K. 1983. Do pied flycatcher males adopt broods of widowed females? Oikos , 41:91–93. Google Scholar


Alisauskas, R. T. 1987. Morphometric correlates of age and breeding status in American coots. Auk , 104:640–646. Google Scholar


Andersson, M. 1982. Female choice selects for extreme tail length in a widowbird. Nature , 299:818–820. Google Scholar


Andersson, M. 1994. Sexual Selection. Princeton University Press. Princeton. Google Scholar


Andersson, M. 2001. Relatedness and the evolution of conspecific brood parasitism. American Naturalist , 158:599–614. Google Scholar


Antor, R. J. , Margalida, A. , Frey, H. , Heredia, R. , Lorente, L. and Sese, J. A. 2007. First breeding age in captive and wild bearded vultures Gypaetus barbatus. Acta Ornithologica , 42:114–118. Google Scholar


Arcese, P. 1987. Age, intrusion pressure and defence against floaters by territorial male song sparrows. Animal Behaviour , 35:773–784. Google Scholar


Barraquand, F. , Hoye, T. T. , Henden, J.-A. , Yoccoz, N. G. , Gilg, O. , Schmidt, N. M. , Sittler, B. and Ims, R. A. 2014. Demographic responses of a site-faithful and territorial predator to its fluctuating prey: long-tailed skuas and arctic lemmings. Journal of Animal Ecology , 83:375–387. Google Scholar


Bayne, E. M. and Hobson, K. A. 2001. Effects of habitat fragmentation on pairing success of ovenbirds: Importance of male age and floater behavior. Auk , 118:380–388. Google Scholar


Berg, E. C. 2005. Parentage and reproductive success in the white-throated magpie-jay, Calocitta formosa a cooperative breeder with female helpers. Animal Behaviour , 70:375–385. Google Scholar


Blanco, G. , Pais, J. L. , Fargallo, J. A. , Potti, J. , Lemus, J. A. and Davila, J. A. 2009. High proportion of non-breeding individuals in an isolated red-billed chough population on an oceanic island (La Palma, Canary Islands). Ardeola , 56:229–239. Google Scholar


Blas, J. , Cabezas, S. , Figuerola, J. , López, L. , Tanferna, A. and Hiraldo, F. 2013. Carotenoids and skin coloration in a social raptor. Journal of Raptor Research , 47:174–184. Google Scholar


Blas, J. and Hiraldo, F. 2010. Proximate and ultimate factors explaining floating behavior in long-lived birds. Hormones and Behavior , 57:169–176. Google Scholar


Blas, J. , Sergio, F. , Wingfield, J. C. and Hiraldo, F. 2011. Experimental tests of endocrine function in breeding and nonbreeding raptors. Physiological and Biochemical Zoology , 84:406–416. Google Scholar


Bornschein, M. R. , Pizo, M. A. , Sobotka, D. D. , Belmonte-Lopes, R. , Golec, C. , Machadode-Souza, T. , Pie, M. R. and Reinert, B. L. 2015. Longevity records and signs of aging in marsh antwren Formicivora acutirostris (Thamnophilidae). Wilson Journal of Ornithology , 127:98–102. Google Scholar


Bortolotti, G. R. 1994. Effect of nest-box size on nest-site preference and reproduction in American kestrels. Journal of Raptor Research , 28:127–133. Google Scholar


Bowman, R. and Bird, D. M. 1986. Ecological correlates of mate replacement in the American kestrel. Condor , 88:440–445. Google Scholar


Bradley, J. S. , Wooller, R D. and Skira, I. J. 2000. Intermittent breeding in the short-tailed shearwater Puffinus tenuirostris. Journal of Animal Ecology , 69:639–650. Google Scholar


Breitwisch, R. 1989. Mortality patterns, sex ratios, and parental investment in monogamous birds. In, Current ornithology, Vol. 6 ( Power, D. , ed.). Plenum Press , New York , pp 1–50. Google Scholar


Bretagnolle, V. , Mougeot, F. and Thibault, J. C. 2008. Density dependence in a recovering osprey population: demographic and behavioural processes. Journal of Animal Ecology , 77:998–1007. Google Scholar


Brown, D. R. and Long, J. A. 2007. What is a winter floater? Causes, consequences, and implications for habitat selection. Condor , 109:548–565. Google Scholar


Brown, J. L. 1969. Territorial behaviour and population regulation in birds. Wilson Bulletin , 81:293–329. Google Scholar


Bruinzeel, L. W. and Van De Pol, M. 2004. Site attachment of floaters predicts success in territory acquisition. Behavioral Ecology , 15:290–296. Google Scholar


Bruinzeel, L. W. , Van De Pol, M. and Trierweiler, C. 2006. Competitive abilities of oystercatchers (Haematopus ostralegus) occupying territories of different quality. Journal of Ornithology , 147:457–463. Google Scholar


Butchart, S. H. M. 2000. Population structure and breeding system of the sex-role reversed, polyanduous bronze-winged jacana Metopidius indicus. Ibis , 142:93–102. Google Scholar


Butchart, S. H. M. , Seddon, N. and Ekstrom, J. M. M. 1999. Polyandry and competition for territories in bronze-winged jacanas. Journal of Animal Ecology , 68:928–939. Google Scholar


Cain, K. E. and Rosvall, K. A. 2014. Next steps for understanding the selective relevance of female-female competition. Frontiers in Ecology and Evolution , 2:32. Google Scholar


Cam, E. , Hines, J. E. , Monnat, J. Y. , Nichols, J. D. and Danchin, E. 1998. Are adult nonbreeders prudent parents? The kittiwake model. Ecology , 79:2917–2930. Google Scholar


Campioni, L. , Delgado, M. M. and Penteriani, V. 2010. Social status influences microhabitat selection: breeder and floater eagle owls Bubo bubo use different post sites. Ibis , 152:569–579. Google Scholar


Campioni, L. , Lourenco, R. , Delgado, M. M. and Penteriani, V. 2012. Breeders and floaters use different habitat cover: should habitat use be a social status-dependent strategy? Journal of Ornithology , 153:1215–1223. Google Scholar


Carmen, W. J. 2004. Noncooperative breeding in the California scrub-jay. Studies in Avian Biology , 28:1–100. Google Scholar


Caro, J. , Ontiveros, D. and Pleguezuelos, J. M. 2011. The feeding ecology of Bonelli's eagle (Aquila fasciata) floaters in southern Spain: implications for conservation. European Journal of Wildlife Research , 57:729–736. Google Scholar


Carrete, M. , Donázar, J. A. and Margalida, A. 2006a. Density-dependent productivity depression in Pyrenean bearded vultures: Implications for conservation. Ecological Applications , 16:1674–1682. Google Scholar


Carrete, M. , Donázar, J. A. , Margalida, A. and Bertran, J. 2006b. Linking ecology, behaviour and conservation: does habitat saturation change the mating system of bearded vultures? Biology Letters , 2:624–627. Google Scholar


Carrick, R. 1963. Ecological significance of territory in the Australian magpie, Gymnorrhina tibicen. Proceedings International Ornithological Congress , 13:740–753. Google Scholar


Coady, C. D. and Dawson, R. D. 2013. Subadult plumage color of female tree swallows (Tachycineta bicolor) reduces conspecific aggression during the breeding season. Wilson Journal of Ornithology , 125:348–357. Google Scholar


Conrad, K. F. , Johnston, P. V. , Crossman, C. , Kempenaers, B. , Robertson, R. J. , Wheelwright, N. T. and Boag, T. 2001. High levels of extra-pair paternity in an isolated, low-density, island population of tree swallows (Tachycineta bicolor). Molecular Ecology , 10:1301–1308. Google Scholar


Cooper, N. W. , Murphy, M. T. , Redmond, L. J. and Dolan, A. C. 2009. Density-dependent age at first reproduction in the eastern kingbird. Oikos , 118:413–419. Google Scholar


Darwin, C. R. 1871. The Descent of Man, and Selection in Relation to Sex. John Murray. London. Google Scholar


Delgado, M. M. , Penteriani, V. , Nams, V. O. and Campioni, L. 2009. Changes of movement patterns from early dispersal to settlement. Behavioral Ecology and Sociobiology , 64:35–43. Google Scholar


Delius, J. D. 1965. A population study of skylarks Alauda arvensis. Ibis , 167:466–492. Google Scholar


Driscoll, D. E. , Jackman, R. E. , Hunt, W. G. , Beatty, G. L. , Driscoll, J. T. , Glinski, R. L. , Gatz, T. A. and Mesta, R. I. 1999. Status of nesting bald eagles in Arizona. Journal of Raptor Research , 33:218–226. Google Scholar


Dwyer, J. F. , Fraser, J. D. and Morrison, J. L. 2012. Range sizes and habitat use of nonbreeding crested caracaras in Florida. Journal of Field Ornithology , 84:223–233. Google Scholar


Dwyer, J. F. , Fraser, J. D. and Morrison, J. L. 2013. Within-year survival of nonbreeding crested caracaras. Condor , 114:295–301. Google Scholar


Eguchi, K. , Yamaguchi, N. , Ueda, K. , Nagata, H. , Takagi, M. and Noske, R. 2007. Social structure and helping behaviour of the greycrowned babbler Pomatostomus temporalis. Journal of Ornithology , 148:S203–S210. Google Scholar


Emlen, S. T. and Wrege, P. H. 2004. Size dimorphism, intrasexual competition, and sexual selection in wattled jacana (Jacana jacana), a sex-role-reversed shorebird in Panama. Auk , 121:391–403. Google Scholar


Ewen, J. G. , Armstrong, D. P. and Lambert, D. M. 1999. Floater males gain reproductive success through extrapair fertilizations in the stitchbird. Animal Behaviour , 58:321–328. Google Scholar


Fedy, B. C. and Stutchbury, B. J. M. 2004. Territory switching and floating in white-bellied antbird (Myrmeciza longipes), a resident tropical passerine in Panama. Auk , 121:486–496. Google Scholar


Fischer, C. A. and Keith, L.B. 1974. Population responses of central Alberta ruffed grouse to hunting. Journal of Wildlife Management , 38:585–600. Google Scholar


Garant, D. , Sheldon, B. C. and Gustafsson, L. 2004. Climatic and temporal effects on the expression of secondary sexual characters: Genetic and environmental components. Evolution , 58:634–644. Google Scholar


Githiru, M. , Lens, L. , Bennun, L. A. and Perrins, C. 2006. Experimental evidence of ‘floaters’ in two isolated populations of an Afrotropical forest bird. Ostrich , 77:28–35. Google Scholar


Gómez De Segura, A. , Martínez, J. M. and Alcántara, M. 2012. Population size of the endangered bearded vulture Gypaetus barbatus in Aragon (Spain): an approximation to the Pyrenean population. Ardeola , 59:43–55. Google Scholar


Gray, E. M. 1996. Female red-winged blackbirds accrue material benefits from copulating with extra-pair males. Animal Behaviour , 53:625–639. Google Scholar


Gruell, A. , Gross, J. and Steiner, J. 2007. Singing activity, territoriality and polygyny in the hoopoe Upupa epops in the Lake Neusiedl area, Austria. Vogelwelt , 128:67–78. Google Scholar


Harris, M. P. 1970. Territory limiting the size of the breeding population of the oystercatcher (Haematopus ostralegus) - a removal experiment. Journal of Animal Ecology , 39:707–713. Google Scholar


Harris, M. P. and Wanless, S. 1995. Survival and non-breeding of adult common guillemots Uria aalge. Ibis, 137:192–197. Google Scholar


Hegyi, G. , Török, J. , Garamszegi, L. Z. and Rosivall, B. 2006. Rapid temporal change in the expression and age-related information content of a sexually selected trait. Journal of Evolutionary Biology , 19:228–238. Google Scholar


Heusmann, H. W. and Bellville, R. 1978. Effects of nest removal on starling populations. Wilson Bulletin , 90:287–290. Google Scholar


Hockey, P. A. R. , Wanless, R. M. and Von Brandis, R. Demographic resilience of territorial island birds to extinction: the flightless Aldabra rail Dryolimnas (cuvieri) aldabranus as an example. Ostrich , 82:1–9. Google Scholar


Holt, R. F. and Martin, K. 1997. Landscape modification and patch selection: The demography of two secondary cavity nesters colonizing clearcuts. Auk , 114:443–455. Google Scholar


Johnson, K. , Du Val, E. , Kielt, M. and Hughes, C. 2000. Male mating strategies and the mating system of great-tailed grackles. Behavioral Ecology , 11:132–141. Google Scholar


Katzner, T. E. , Ivy, J. A. R. , Bragin, E. A. , Milner-Gulland, E. J. and DeWoody, J. A. 2011. Conservation implications of inaccurate estimation of cryptic population size. Animal Conservation , 14:328–332. Google Scholar


Kazama, K. , Niizuma, Y. and Watanuki, Y. 2012. Intraspecific kleptoparasitism, attacks on chicks and chick adoption in black-tailed gulls (Larus crassirostris). Waterbirds , 35:599–607. Google Scholar


Kempenaers, B. , Everding, S. , Bishop, C. , Boag, P. and Robertson, R. J. 2001. Extrapair paternity and the reproductive role of male floaters in the tree swallow (Tachycineta bicolor). Behavioral Ecology and Sociobiology , 49:251–259. Google Scholar


Kendeigh, S. C. 1941. Territorial and mating behaviour of the house wren. Illinois Biological Monographs , 18:1–120. Google Scholar


Knapton, R. W. and Krebs, J. R. 1974. Settlement patterns, territory size and breeding density in the song sparrow (Melospiza melodia). Canadian Journal of Zoology , 52:1413–1420. Google Scholar


Kokko, H. , Harris, M. P. and Wanless, S. 2004. Competition for breeding sites and site-dependent population regulation in a highly colonial seabird, the common guillemot Uria aalge. Journal of Animal Ecology , 73:367–376. Google Scholar


Komdeur, J. 1996. Breeding of the Seychelles magpie robin Copsychus sechellarum and implications for its conservation. Ibis , 138:485–498. Google Scholar


Komdeur, J. and Edelaar, P. 2001. Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment. Behavioral Ecology , 12:706–715. Google Scholar


Krebs, J. R. 1982. Territorial defence in the great tit (Parus major): do residents always win? Behavioral Ecology and Sociobiology , 11:185–194. Google Scholar


Leisler, B. , Beier, J. , Staudter, H. and Wink, M. 2000. Variation in extra-pair paternity in the polygynous great reed warbler (Acrocephalus arundinaceus). Journal für Ornithologie , 141:77–84. Google Scholar


Lenda, M. , Maciusik, B. and Skorka, P. 2012. The evolutionary, ecological and behavioural consequences of the presence of floaters in bird populations. Northwestern Journal of Zoology , 8:394–408. Google Scholar


Lewis, R.A. and Zwickel, F.C. 1980. Removal and replacement of male blue grouse on persistent and transient territorial sites. Canadian Journal of Zoology , 58:1417–1423. Google Scholar


Linz, G. M. , Sawin, R. S. , Lutman, M. W. and Bleier, W. J. 2011. Modeling parental provisioning by red-winged blackbirds in North Dakota. Prairie Naturalist , 43:92–99. Google Scholar


Loewenthal, D. , Paijmans, D. M. and Hockey, P. A. R. 2015. How do African black oystercatchers Haematopus moquini recruit into high-density populations? Ostrich , 86:1–8. Google Scholar


Lyon, B. E. and Montgomerie, R. 2012. Sexual selection is a form of social selection. Philosophical Transactions of the Royal Society B, 367:2266–2273. Google Scholar


Marra, P. P. and Holmes, R. T. 1997. Avian removal experiments: Do they test for habitat saturation or female availability? Ecology , 78:947–952. Google Scholar


Martínez, J. A. and Zuberogoitia, I. 2002. Factors affecting the vocal behaviour of eagle owls Bubo bubo: effects of sex and territorial status. Ardeola , 49:1–9. Google Scholar


Manuwal, D. A. 1974. Effects of territoriality on breeding in a population of Cassin's auklet. Ecology , 55:1399–1406. Google Scholar


McNabb, E. G. , Kavanagh, R. P. and Craig, S. A. 2007. Further observations on the breeding biology of the powerful owl, Ninox strenua in south-eastern Australia. Corella , 31:6–9. Google Scholar


Møller, A. P. 1988. Female choice selects for male sexual tail ornaments in the monogamous swallow. Nature , 332:640–642. Google Scholar


Møller, A. P. and Szep, T. 2005. Rapid evolutionary change in a secondary sexual character linked to climatic change. Journal of Evolutionary Biology , 18:481–495. Google Scholar


Morales, J. , Gordo, O. , Lobato, E. , Ippi, S. , Martínez-De La Puente, J. , Tomás, G. , Merino, S. and Moreno, J. 2014. Femalefemale competition is influenced by forehead patch expression in pied flycatcher females. Behavioral Ecology and Sociobiology , 68:1195–1204. Google Scholar


Moreno, J. 2015. The incidence of clutch replacements in the pied flycatcher Ficedula hypoleuca is related to nest-box availability: evidence of female-female competition? Ardeola , 62:67:80. Google Scholar


Moreno, J. , Velando, A. , González-Braojos, S. , Ruíz-De-Castañeda, R. and Cantarero, A. 2013. Females paired with more attractive males show reduced oxidative damage: possible direct benefits of mate choice in pied flycatchers. Ethology , 119:1–11. Google Scholar


Moreno, J. , Gil, D. , Cantarero, A. and López-Arrabé, J. 2014. Extent of a white plumaje patch covaries with testosterone levels in female pied flycatchers Ficedula hypoleuca. Journal of Ornithology , 155:639–648. Google Scholar


Morton, E. S. , Derrickson, K. C. and Stutchbury, B. J. M. 2000. Territory switching behavior in a sedentary tropical passerine, the dusky antbird (Cercomacra tyrannina). Behavioral Ecology , 11:648–653. Google Scholar


Moulton, L. L. , Linz, G. M. and Bleier, W. J. 2013. Responses of territorial and floater male red-winged blackbirds to models of receptive females. Journal of Field Ornithology , 84:160–170. Google Scholar


Mumme, R. L. 2015. Demography of slatethroated redstarts (Myioborus miniatus): a non-migratory Neotropical warbler. Journal of Field Ornithology , 86:89–102. Google Scholar


Newton, I. 1979. Population Ecology of Raptors. Poyser. Berkhamsted. Google Scholar


Newton, I. 1994. Experiments on the limitation of bird breeding densities: a review. Ibis , 136:397–411. Google Scholar


Newton, I. 1998. Population Limitation in Birds. Academic Press. San Diego. Google Scholar


Newton, I. and Marquiss, M. 1991. Removal experiments and the limitation of breeding density in sparrowhawks. Journal of Animal Ecology , 60:535–544. Google Scholar


Newton, I. and Rothery, P. 2001. Estimation and limitation of numbers of floaters in a Eurasian sparrowhawk population. Ibis , 143:442–449. Google Scholar


Peer, K. , Robertson, R. J. and Kempenaers, B. 2000. Reproductive anatomy and indices of quality in male tree swallows: The potential reproductive role of floaters. Auk , 117:74–81. Google Scholar


Penteriani, V. and Delgado, M. M. 2012. There is a limbo under the moon: what social interactions tell us about the floaters' underworld. Behavioral Ecology and Sociobiology , 66:317–327. Google Scholar


Penteriani, V. and Delgado, M. M. 2008. Owls may use faeces and prey feathers to signal current reproduction. Plos One , 3:e3014. Google Scholar


Penteriani, V. , Delgado, M. M. and Campioni, L. 2009. Quantifying space use of breeders and floaters of a long-lived species using individual movement data. Science of Nature , 102:21. Google Scholar


Penteriani, V. , Ferrer, M. and Delgado, M. M. 2011. Floater strategies and dynamics in birds, and their importance in conservation biology: towards an understanding of nonbreeders in avian populations. Animal Conservation , 14:233–241. Google Scholar


Pierotti, R. 1980. Spite and altruism in gulls. American Naturalist , 115:290–300. Google Scholar


Pilz, K. M. and Smith, H. G. 2004. Egg yolk androgen levels increase with breeding density in the European starling, Sturnus vulgaris. Functional Ecology , 18:58–66. Google Scholar


Piper, S. E. 2001. Elucidating population structure in the long-tailed wagtail Motacilla clara: The use of the space-time diagram. Ardea , 89:113–121. Google Scholar


Piper, W. H. , Tischler, K. B. and Klich, M. 2000. Territory acquisition in loons: the importance of take-over. Animal Behaviour , 59:385–394. Google Scholar


Piper, W. H. , Walcott, C. , Mager, J. N. , Perala, M. , Tischler, K. B. , Harrington, E. , Turcotte, A. J. , Schwabenlander, M. and Banfield, N. 2006. Prospecting in a solitary breeder: chick production elicits territorial intrusions in common loons. Behavioral Ecology , 17:881–888. Google Scholar


Porter, J. M. and Coulson, J. C. 1987. Longterm changes in recruitment to the breeding group, and the quality of recruits at a kittiwake Rissa tridactyla colony. Journal of Animal Ecology , 56:675–689. Google Scholar


Pryke, S. R. and Andersson, S. 2003. Carotenoidbased epaulettes reveal male competitive ability: experiments with resident and floater redshouldered widowbirds. Animal Behaviour , 66:217–224. Google Scholar


Richner, H. 1989. Habitat specific growth and fitness in carrion crows (Corvus corone corone). Journal of Animal Ecology , 58:427–440. Google Scholar


Ridley, A. R. , Raihani, N. J. and Nelsonflower, M. J. 2008. The cost of being alone: the fate of floaters in a population of cooperatively breeding pied babblers Turdoides bicolor. Journal of Avian Biology , 39:389–392. Google Scholar


Rivera, J. L. , Vargas, F. H. and Parker, P. G. 2011. Natal dispersal and sociality of young Galapagos hawks on Santiago Island. Open Ornithology Journal , 4:12–16. Google Scholar


Rohner, C. 1995. Great horned owls and snowshoe hares - what causes the time-lag in the numerical response of predators to cyclic prey. Oikos , 74:61–68. Google Scholar


Rohner, C. 1997a. Non-territorial ‘floaters’ in great horned owls: Space use during a cyclic peak of snowshoe hares. Animal Behaviour , 53:901–912. Google Scholar


Rosenfield, R. N. , Sonsthagen, S. A. , Stout, W. E. and Talbot, S. L. 2015. High frequency of extra-pair paternity in an urban population of Cooper's hawks. Journal of Field Ornithology , 86:144–152. Google Scholar


Rudnick, J. A. , Katzner, T. E. , Bragin, E. A. and DeWoody, J. A. 2008. A non-invasive genetic evaluation of population size, natal philopatry, and roosting behavior of non-breeding eastern imperial eagles (Aquila heliaca) in central Asia. Conservation Genetics , 9:667–676. Google Scholar


Ryder, T. B. , Horton, B. M. and Moore, I. T. 2011. Understanding testosterone variation in a tropical lek-breeding bird. Biology Letters , 7:506–509. Google Scholar


Saitou, T. 2001. Floaters as intraspecific brood parasites in the grey starling Sturnus cineraceus. Ecological Research , 16:221–231. Google Scholar


Sandell, M. I. and Diemer, M. 1999. Intraspecific brood parasitism: a strategy for floating females in the European starling. Animal Behaviour , 57:197–202. Google Scholar


Sarah, E. A. , Dit Durell, L. V. and Clarke, R. T. 2004. The buffer effect of non-breeding birds and the timing of farmland bird declines. Biological Conservation , 120:375–382. Google Scholar


Schmutz, J. A. , Wright, K. G. , DeSorbo, C. R. , Fair, J. , Evers, D. C. , Uher-Koch, B. D. and Mulcahy, D. M. 2014. Size and retention of breeding territories of yellow-billed loons (Gavia adamsii) in Alaska and Canada. Waterbirds , 37:53–63. Google Scholar


Schweizer, C. L. and Whitmore, R. C. Movements of the mangrove warbler in Baja California Sur. Western Birds , 44:262–272. Google Scholar


Sergio, F. , Blas, J. and Hiraldo, F. 2009. Predictors of floater status in a long-lived bird: a cross-sectional and longitudinal test of hypotheses. Journal of Animal Ecology , 78:109–118. Google Scholar


Shutler, D. and Weatherhead, P. J. 1991. Owner and floater red-winged blackbirds: determinants of status. Behavioral Ecology and Sociobiology , 28:235–241. Google Scholar


Shutler, D. and Weatherhead, P. J. 1994. Movement patterns and territory acquisition by floater red-winged blackbirds. Canadian Journal of Zoology , 72:712–720. Google Scholar


Smith, J. N. M. and Arcese, P. 1989. How fit are floaters? Consequences of alternative territorial behaviours in a non-migratory sparrow. American Naturalist , 133:830–845. Google Scholar


Smith, S. M. 1976. Ecological aspects of dominance hierarchies in black-capped chickadess. Auk , 93:95–107. Google Scholar


Smith, S. M. 1978. The ‘underworld’ in a territorial sparrow: adaptive strategy for floaters. American Naturalist , 112:571–582. Google Scholar


Snetsinger, T. J. , Herrmann, C. M. , Holmes, D. E. , Hayward, C. D. and Fancy, S. G. 2005. Breeding ecology of the puaiohi (Myadestes palmeri). Wilson Bulletin , 117:72–84. Google Scholar


Stacey, P. B. and Koenig, W. D. 1990. Cooperative Breeding in Birds: Long-term Studies of Ecology and Behaviour. Cambridge University Press. Cambridge. Google Scholar


Stutchbury, B. J. 1992. Experimental evidence that bright coloration is not important for territory defense in purple martins. Behavioral Ecology and Sociobiology , 31:27–33. Google Scholar


Stutchbury, B. J. and Robertson, R. J. 1985. Floating populations of female tree swallows. Auk , 102:651–654. Google Scholar


Stutchbury, B. J. and Robertson, R. J. 1987. Behavioural tactics of sub-adult female floaters in the tree swallow. Behavioral Ecology and Sociobiology , 20:413–419. Google Scholar


Szuba, K. J. and Bendell, N. F. 1988. Nonterritorial males in populations of spruce grouse. Condor , 90:492–496. Google Scholar


Taborsky, M. 1994. Sneakers, satellites, and helpers - Parasitic and cooperative behavior in fish reproduction. In, P. J. B. Slaters , J. S. Rosenblatt , C. T. Snowdon and M. Milinski (Eds): Advances in the Study of Behavior Vol. 23, pp. 1–100. Elsevier. San Diego. Google Scholar


Tanferna, A. , López-Jiménez, L. , Blas, J. , Hiraldo, F. and Sergio, F. 2013. Habitat selection by black kite breeders and floaters: Implications for conservation management of raptor floaters. Biological Conservation , 160:1–9. Google Scholar


Tobias, J. A. , Montgomerie, R. and Lyon, B. E. 2012. The evolution of female ornaments and weaponry: social selection, sexual selection and ecological competition. Philosophical Transactions of the Royal Society B , 367:2274–2293. Google Scholar


Tobler, M. and Smith, H. G. 2004. Specific floater home ranges and prospective behaviour in the European starling, Sturnus vulgaris. Naturwissenschaften , 91:85–89. Google Scholar


Turrin, C. and Watts, B. D. 2014. Intraspecific intrusion at bald eagle nests. Ardea , 102:71–78. Google Scholar


Veiga, J. P. , Polo, V. , Arenas, M. and Sánchez, S. 2012. Intruders in nests of the spotless starling: Prospecting for public information or for immediate nesting resources? Ethology , 118:917–924. Google Scholar


Veiga, J. P. , Polo, V. , Arenas, M. and Sánchez, S. 2013. Nest intrusions in relation to breeding status in the spotless starling. Behaviour , 150:1553–1566. Google Scholar


Velando, A. , Lessells, C. M. and Márquez, J. C. 2001. The function of female and male ornaments in the inca tern: evidence for links between ornament expression and both adult condition and reproductive performance. Journal of Avian Biology , 32:311–318. Google Scholar


Vili, N. , Szabo, K. , Kovacs, S. , Kabai, P. , Kalmar, L. and Horvath, M. 2013. High turnover rate revealed by non-invasive genetic analyses in an expanding eastern imperial eagle population. Acta Zoologica Academiae Scientiarum Hungaricae , 59:279–295. Google Scholar


Village, A. 1990. The Kestrel. Poyser. Calton. Google Scholar


Villavicencio, C. P. , Apfelbeck, B. and Goymann, W. 2013. Experimental induction of social instability during early breeding does not alter testosterone levels in male black redstarts, a socially monogamous songbird. Hormones and Behavior , 64:461–467. Google Scholar


Voegeli, M. , Laiolo, P. , Serrano, D. and Tella, J. L. 2008. Who are we sampling? Apparent survival differs between methods in a secretive species. Oikos , 117:1816–1823. Google Scholar


Watson, A. 1985. Social class, socially-induced loss, recruitment and breeding of red grouse. Oecologia , 67:493–498. Google Scholar


Watson, A. and Jenkins, D. 1968. Experiments on population control by territorial behaviour in Red Grouse. Journal of Animal Ecology , 37:595–614. Google Scholar


Weatherhead, P. J. and Boag, P. T. 1995. Pair and extrapair mating success relative to male quality in red-winged blackbirds. Behavioral Ecology and Sociobiology , 37:81–91. Google Scholar


Westcott, D. A. and Smith, J. N. M. 1994. Behavior and social-organization during the breeding-season in Mionectes oleagineus, a lekking flycatcher. Condor , 96:672–683. Google Scholar


Westneat, D. F. and Stewart, I. R. K. 2003. Extra-pair paternity in birds: causes, correlates and conflict. Annual Review of Ecology, Evolution and Systematics , 34:365–396. Google Scholar


West-Eberhard, M. J. 1983. Sexual selection, social competition and speciation. Quarterly Review of Biology , 58:155–183. Google Scholar


Wiebe, K. L. 2011. Nest sites as limiting resources for cavity-nesting birds in mature forest ecosystems: a review of the evidence. Journal of Field Ornithology , 82:239–248. Google Scholar


Wischhoff, U. , Marques-Santos, F. , Ardia, D. R. and Roper, J. J. 2015. White-rumped swallows prospect while they are actively nesting. Journal of Ethology , 33:145–150. Google Scholar


Wynne-Edwards, V. C. 1962. Animal Dispersion in Relation to Social Behaviour. Oliver and Boyd. Edinburgh. Google Scholar


Yasukawa, K. , Enstrom, D. A. , Parker, P. G. and Jones, T. C. 2009. Epaulet colour and sexual selection in the red-winged blackbird: a field experiment. Condor , 111:740–751. Google Scholar


Young, E. C. 1972. Territory establishment and stability in McCormick's skua. Ibis , 114:234–244. Google Scholar


Zhang, W.-W. , Ma, J.-Z. and Li, J.-B. 2011. Preliminary study on conspecific brood parasitism and defense mechanism of Fulica atra. Chinese Journal of Zoology , 46:19–23. Google Scholar


Zilberman, R. , Moav, B. and Yom-Tov, Y. 1999. Extra-pair paternity in the socially monogamous orange-tufted sunbird (Nectarinia osea osea). Israel Journal of Zoology , 45:407–421. Google Scholar


Zilberman, R. , Moav, B. and Yom-Tov, Y. 2001. Territoriality and mate guarding in the orangetufted sunbird (Nectarinia osea). Israel Journal of Zoology , 47:275–286. Google Scholar
Juan Moreno "The Unknown Life of Floaters: The Hidden Face of Sexual Selection," Ardeola 63(1), 49-77, (1 June 2016).
Published: 1 June 2016

densidad de población
extra-pair paternity
paternidad extrapareja
population density
recursos reproductivos
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