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Summary
1. Few studies have been capable of monitoring the nonterritorial sector of a population because
of the typically secretive behaviour of floating individuals, despite the existing consensus over the
demographic importance of floating. Furthermore, there is almost no information on floating
behaviour for migratory species.
2. The factors that determine whether an individual will be a floater or a territory owner have been
framed into five, non-mutually exclusive hypotheses: (i) territory holders are morphologically
superior to floaters (resource-holding potential hypothesis); (ii) age confers skills and fighting
motivation which lead to social dominance and territory ownership (age hypothesis); (iii) occu-
pancy time of a site determines asymmetries in its knowledge, familiarity and value for potential
contenders (site-dominance hypothesis); (iv) contenders use an arbitrary rule to settle contests
leading to pre-defined cut-off points for a biologically meaningful trait (e.g. age, body size) separ-
ating floaters from territory holders (arbitrary convention hypothesis); and (v) floaters set up a
‘war of attrition’ at arbitrarily chosen territories (arbitrary attrition hypothesis).
3. We tested these hypotheses using long-term data on a long-lived, migratory raptor, the black kite
Milvus migrans Boddaert.
4. Floating status was best explained by the concerted action of mechanisms consistent with the
age and site-dominance hypotheses.
5. In both cross-sectional and longitudinal analyses, acquisition of a territory was determined by
a complex interaction between age and early arrival from migration, suggesting: (i) a progressive
incorporation of early arriving individuals in the territorial contingent of the population, and (ii)
the existence of an alternative restraint strategy of delayed territoriality mediated by long-term
acquisition of social dominance.
6. Such results suggested that territory acquisition was mediated by the establishment of site
dominance through pre-emption and, secondarily, despotism. In this population, age and arrival
date aligned individuals along a demographic continuum ranging from successful breeders mono-
polizing high-quality resources to floaters with no resources, consistent with the notion of floating as
an extreme form of breeding failure.
Key-words: dominance, floating, nonbreeders, territoriality, territory acquisition
Table 1. Hypotheses explaining the determinants of the status of individuals as floaters or territory holders. Predictions have been adapted to
the model system under analysis (a migratory, long-lived bird)
Mechanism determining
Hypothesis territorial status Predictions Supported†
†Symbol legend: *the prediction was supported by the Results of this study; ‡this prediction compares floaters with any type of territory holder;
§this prediction compares floaters with territory holders at their first year of successful territory acquisition (first-time ‘breeders’), that is,
between floaters that succeeded or not in acquiring a territory that year. ¶For individuals repeatedly sampled in consecutive years, longitudinal
improvements in arrival dates are more pronounced for individuals in transition between floater and territory holder status in successive years
than for individuals that remain floaters in consecutive years. ††This hypothesis predicts no clear separation between territory holders and
floaters in terms of quality traits such as age or body size, because floaters can eventually obtain a territory even if they lose all fights with owners
(Stutchbury 1991).
(review in Zach & Stutchbury 1992), to the point that they characteristics will lead to higher success during contests over
have been referred to as a ‘shadow’ population living in a territories. (ii) The age (or social dominance) hypothesis
secretive ‘underworld’ (Smith 1978; Rohner 1997). Further- proposes that age (and its associated decline in residual
more, there is almost no information on floating behaviour reproductive value) confers skills and fighting motivation
for migratory species (Porter 1988; Stutchbury 1991). which lead to social dominance and thus to higher likelihood
The factors that determine whether an individual will be a of successfully acquiring and defending a territory (Rohwer,
floater or a territory owner have been usually framed into the Ewald & Rohwer 1981; Grafen 1987; Shutler & Weatherhead
following five, non-mutually-exclusive hypotheses (summa- 1991). This hypothesis predicts that younger individuals
rized in Table 1): (i) the resource-holding potential (RHP) will be more likely to be floaters and to loose contests over
hypothesis suggests that, compared to floaters, territory territories. Some authors consider this hypothesis as a special
holders will have characteristics (larger size, strength etc.) that case of RHP (Shutler & Weatherhead 1991). (iii) The site
promote higher capabilities to acquire and defend resources dominance (or value asymmetry) hypothesis states that the
(Mönkkönen 1990; Lozano 1994; Pryke & Andersson 2003). occupancy time of a site determines asymmetries in its know-
This hypothesis predicts that territory holders will be larger ledge, familiarity and value for potential contenders (Krebs
than floaters or morphologically different, and that such 1982; Beletsky & Orians 1989; Tobias 1997). It predicts that,
© 2008 The Authors. Journal compilation © 2008 British Ecological Society, Journal of Animal Ecology, 78, 109–118
Predictors of floater status 111
all else being equal, territory holders will have spent more 1997), which confounds direct comparisons between floater
time at the site than floating individuals which did not succeed and holder characteristics (e.g. differences in body condition
in acquiring it. (iv) The arbitrary convention (or uncorrelated caused by habitat rather than territorial status). Therefore, as
asymmetry) hypothesis postulates that contenders use an noted by other authors (Newton 1992; Zach & Stutchbury
arbitrary rule to settle contests, such ‘the resident always 1992; Danchin & Cam 2002), there is a great need for empirical
wins’, or ‘the older always wins (Davies 1978; Rohwer 1982). studies where floaters co-exist with breeders and can be
It predicts the existence of some pre-defined cut-off point for observed before they become territorial and without removing
a biologically meaningful trait (e.g. age, body size), below territory holders. This would represent a much needed com-
which all individuals will be either all floaters or all territory plement to previous experimental or indirect analyses.
holders. (v) The arbitrary attrition hypothesis claims that Here we use a long-term data set on the population of a
floating individuals choose an arbitrary number of territories long-lived migratory raptor, the black kite Milvus migrans
where they regularly intrude and challenge the owner in a Boddaert, in which floaters co-exist with territory holders and
‘war of attrition’ won by who is able to persist the longest can be easily identified. Previous studies on floaters in raptors
(Stutchbury 1991). It predicts no clear separation between have been extremely few, most of them focused on small-sized
territory holders and floaters in terms of quality traits such as short-lived species, none of them was conducted on migrants,
age or body size, because floaters can eventually obtain a and they were mostly based on indirect methods (removal
territory even if they lose all fights with owners (Stutchbury experiments, or estimation of floater numbers based on breeders
1991). It is important to note that the above hypotheses are demographic rates) (Village 1990; Rohner 1996; Ferrer &
not mutually exclusive. Harte 1997; Hunt 1998; Kenward et al. 2000; Newton & Rothery
Overall, the relative importance of these hypotheses in 2001). None of them was framed as a test of the above hypotheses.
predicting territorial status remains unclear, as inconsistent
results have even been obtained for different populations of
the same species (e.g. Eckert & Weatherhead 1987; Pryke & Methods
Andersson 2003). Therefore, there is a great need for further
STUDY AREA
empirical tests in order to reach a consensus over a potential
general pattern. To date, studies on floaters have mainly Kites were studied in a 430-km2 plot located in Doñana National
included: (i) model species which are small-sized, short-lived Park (south-western Spain). The landscape was characterized by
and year-round residents (reviews in Newton 1992; and seasonally flooded marshland, scrublands, grasslands, and mobile
Zach & Stutchbury 1992); (ii) theoretical models examining sand dunes along the sea shore.
the consequences of floating for individual fitness and
population regulation (e.g. Zach & Stutchbury 1992; Kokko
STUDY SPECIES
& Sutherland 1998); (iii) estimates of floater abundance
(Rohner 1996; Kenward et al. 2000; Newton & Rothery 2001); The black kite is a medium-sized, migratory raptor. In our popula-
(iv) removal experiments (Newton 1992); and (v) analyses tion, the age of first territorial establishment ranges between 1 and 7
years and the maximum recorded life span is 23 years (Blas, Sergio
of territory acquisition where floater characteristics are
& Hiraldo 2009). On return from migration, individuals settle on
indirectly inferred by examining individuals at their first
breeding territories, which usually include a nest site and an area of
incorporation into the territorial sector of the population (e.g. 20–200 m around it, while foraging occurs over wider, undefended
Porter 1988; Arcese 1989). Given the secretiveness of floating, (communal) areas. The population was stable during the study
unavoidably some of these methods yield an incomplete period at around 500 breeding pairs plus 400–500 nonbreeding indi-
picture of floater characteristics because of inherent biases viduals congregated in six communal roosts (Blas 2002). The roosts
and assumptions. Commonly reported biases include: (i) the are regularly distributed within the matrix of breeding territories
difficulty to detect and monitor all floater age classes in and the floaters use hunting areas that overlap widely with those
observational studies (e.g. Smith & Arcese 1989; Stutchbury of breeders (i.e. floaters and breeders fully co-exist). Breeding
1991; Kenward et al. 2000); (ii) the incapability to distinguish and natal dispersal distances are short (median 302 m and 4800 m
between nonbreeding territorial holders and true floaters respectively) and extensive surveys suggest the absence of emigration
to other populations (Forero et al. 1999; Forero, Donázar & Hiraldo
(e.g. Eckert & Weatherhead 1987; Kenward et al. 2000); (iii)
2002).
the younger age classes of floaters never reclaim territories and
are thus missed in some removal experiments (e.g. Shutler &
Weatherhead 1994); (iv) studies on the age of first territorial FIELD PROCEDURES
acquisition unavoidably miss a large portion of floaters which
In all analyses, we employ reproductive data collected between 1997
die without ever being territorial; (v) in removal experiments,
and 2000. However, ringing activities started earlier in the following
the contests between the removed owner trying to regain its
staggered manner: since 1964, as many nestlings as possible have
territory against its artificially induced replacement yield been marked every year with metal rings. Since 1986, the nestlings
unclear information because both contenders may believe to were also marked with a white plastic ring with a black, three-
be the owner, an unrealistic condition in a natural setting (e.g. character alphanumeric code, which can be read by telescope
Shutler & Weatherhead 1992); (vi) floaters occupy areas and without disturbing the birds. In addition, since 1986, we used can-
habitats disjunct from those of breeders (e.g. Ferrer & Harte non nets and padded leg-hold traps throughout the study area to
© 2008 The Authors. Journal compilation © 2008 British Ecological Society, Journal of Animal Ecology, 78, 109– 118
112 F. Sergio, J. Blas & F. Hiraldo
capture and mark adults with metal and plastic rings. Many of these Table 3). Because body size and condition were never significant
had been metal-banded as nestlings before 1986, which subsequently and were available for a sub-sample of trapped birds, we re-ran the
allowed us to sample all the age classes in the population. For each analysis without fitting body measurements. This allowed to increase
trapped adult, we measured body mass to the nearest 5 g, tarsus the sample size to 714 individuals.
length to the nearest 0·1 mm, and wing length and tail length to the To further test whether body size, age or arrival date afforded a
nearest 1·0 mm. Overall, up to 2000, 2367 nestlings were marked higher likelihood of territory acquisition for floaters (prediction 1b,
only with metal bands, and 4257 nestlings plus 1076 adults were 2b, 3b), we built the same GLMM as above, but restricted the anal-
marked with both metal and plastic bands. Individuals were sexed by ysis to territory holders at their first year of successful territory
molecular analysis of a blood sample (Ellegren 1996) or by multiple acquisition (first-time ‘breeders’, n = 111 individuals when including
observations of copulation behaviour. the effect of body measures, n = 280 individuals without including
Marked territory holders were searched intensively by visiting the effect of body measures, details in Table 3). While the previous
breeding territories every 4–5 days. A marked individual was assigned models compared floaters with all territory owners, this analysis dis-
to a specific territory when it was observed there in more than criminated cross-sectionally between floaters that succeeded or not
three separate visits. In our population, floaters regularly gather to in acquiring a territory that year.
sleep at communal roosts throughout the spring–summer (authors’ To gain a further understanding of the link between arrival date
unpublished telemetry data on > 40 floaters). Roost sites were and territorial status, we conducted a longitudinal analysis of the
checked in the evening from hides every 4–5 days throughout the improvement in arrival dates in relation to age and territorial status
spring–summer and we classified as floaters all individuals that for individuals sampled in consecutive years (prediction 3c). For
were (i) detected at roosts in ≥ two visits separated by ≥ 15 days each marked individual whose arrival date was known in consecutive
in the period 15 April–15 June, and (ii) never observed holding a years, we: (i) calculated whether its arrival improved (became earlier
breeding territory in that year. Telemetry data on > 40 confirmed or not) in consecutive years; and (ii) tested by means of a binomial
floaters indicated that such procedure could reliably identify floater test whether there was a preponderance of individuals improving
status. their arrival for each age class category (e.g. transition from age 1 to
Arrival date from migration was expressed as the date of first 2 years old, 2 to 3 years old, etc) and for each of three territorial
observation of a marked individual, which is known to be a reliable status categories: (i) floater, (ii) territory holder, and (iii) individuals
estimate of true settlement date (details in Sergio et al. 2007a). in transition between the status of floater and territory holder in
The extremely frequent visits to territories allowed detection of consecutive years.
numerous cases in which a marked bird was observed to hold a To test whether age, arrival date or body measures predicted
territory and to be replaced by another individual at a later date. In successful territory acquisition during a takeover (predictions 1c, 2c,
all the cases in which this was directly observed, it was accompanied 3d, 4a, 5c), we built a GLM with takeover success as the dependent
by fighting, which sometimes escalated into talon-locking, with variable and sex, age, arrival date, body size and body condition as
individuals grasping each other’s talons in flight and falling to the the explanatory variables. To increase sample size, for this analysis
ground, where fighting continued until one of the contestants we employed all the takeover episodes recorded between 1992 and
escaped. For this reason, we assume that most of the observed 2006. Because winners of takeovers included both individuals
replacements were aggressive evictions. Hereafter, we refer to such that were previously floaters and individuals that previously held a
contests as ‘takeovers’, to the expelled bird as the ‘evicted’ and to the territory elsewhere (individuals switching to another territory),
new occupant as the ‘winner’. we conducted two separate analyses. In the first, we compared floaters
that succeeded in usurping a territory (i.e. which abandoned floater
status) with floaters that failed to evict the previous territory owner
HYPOTHESIS TESTING AND STATISTICAL ANALYSES
during a takeover attempt (n = 39). In the second, we compared
Because univariate metrics have been criticized as measures of floaters that failed to acquire a territory during a takeover attempt
body size (Freeman & Jackson 1990), we estimated size by means with experienced territory holders that succeeded in switching to
of the first axis (PC1, hereafter ‘body size’) of a principal components another territory and evicting its previous owner (n = 50).
analysis built using tarsus, wing and tail length. The PC1 explained All models were built through a backward stepwise procedure
62% of the variation in size and had high positive loadings for wing where the least significant terms or interactions were sequentially
length (r = 0·87), tarsus length (0·52) and tail length (0·68). We used removed until obtaining a minimal adequate model that only retained
body mass corrected for skeletal size as an index of condition. In significant effects at the 5% probability level (Crawley 1993). All
particular, as mass varied with year and body size, we standardized tests are two-tailed, statistical significance was set at α < 0·05, and
it by using the residuals of such relationship as an index of body condi- all means are given ± 1 SE.
tion (see Sergio et al. 2007b, unpublished data and references therein).
The five testable hypotheses and their main predictions are sum-
marized in Table 1. Throughout, we assume that early arrival from Results
migration allowed individuals more time to acquire general familiarity
with the area and to establish site dominance through pre-emption PRELIMINARY, DESCRIPTIVE TESTS
(Sergio et al. 2007a; predictions 3a–d).
Compared to territory holders, floaters of both sexes were
To test the effect of age, arrival date and body measures on terri-
smaller, younger and arrived later from migration (Table 2;
torial status (predictions 1a, 2a, 3a, 5a), we built a generalized linear
mixed model (GLMM, Littel et al. 1996) with year (as a random Figs 1 and 2). Body condition did not vary significantly
factor), individual identity (as a random factor), sex, age, arrival between floaters and territory holders (Table 2), but its
date, body size, body condition and their first-order interactions seasonal variations differed between the two status categories
as explanatory variables and territorial status (floater vs. territory (Fig. 3): the body condition of territory holders declined
holder) as the dependent variable (n = 170 individuals, details in rather constantly through the breeding cycle. In contrast, the
© 2008 The Authors. Journal compilation © 2008 British Ecological Society, Journal of Animal Ecology, 78, 109–118
Predictors of floater status 113
Table 2. Generalized linear mixed model logistic regressions (with binomial errors and a logit link function) testing the effect of sex, age, arrival
date from migration, body size and body condition on the territorial status of black kites in Doñana National Park (Spain). (a) comparison
between floaters and territory holders for 72 males and 98 females for which body measurements were available; (b) comparison between floaters
and territory holders for an extended sample of 361 males and 353 females for which body measurements were not available‡; (c) comparison
between floaters and individuals acquiring a territory for the first time in their life (n = 146 males and 134 females); (d) comparison between
floaters that succeeded in usurping a territory and floaters that failed to evict the previous territory owner during a takeover attempt (n = 17
males and 22 females); (e) comparison between floaters that failed to acquire a territory and experienced territory holders that succeeded in
switching to another territory and evicting its previous owner during a takeover attempt (n = 19 males and 31 females). Random factors (year
and individual identity) are only shown when their effect was significant
Percentage of
Variable Parameter estimate ± SE F P deviance explained
†1, territory holder; 2, floater. This analysis employs only individuals for which body measurements were available. ‡1, territory holder;
2, floater. Because the effect of body measures was not significant in Model a, the model was repeated on a sample of individuals for which body
measurements were or not available. This allowed a marked increase in sample size. §1, individual holding a territory for the first time in its life;
2, floater. ¶1, floater which failed to usurp a territory; 2, floater that succeeded to evict a previous owner. ††1, floater which failed to usurp a
territory; 2, experienced territory holders that succeeded in switching to another territory and evicting its previous owner.
Table 3. Mean (± SE) age, arrival date from migration, body size and body condition of floaters and territory holders of both sexes in a black
kite population of Doñana National Park (Spain)
Age 2·39 ± 0·06 (400) (167†) 6·76 ± 0·13 (637) (63†) 24·93 < 0·0001
Arrival date 123·9 ± 1·6 (300) (134‡) 86·9 ± 0·6 (697) (146‡) 27·94 < 0·0001
Body size −0·49 ± 0·09 (128) (33§) 0·28 ± 0·06 (228) (141§) 7·49 < 0·0001
Body condition −11·85 ± 8·18 (127) (33 §) 6·72 ± 5·82 (224) (138§) 1·88 0·06
†Individuals of unknown sex included in the sample; ‡individuals of unknown sex or age included in the sample; §individuals of unknown age
included in the sample.
Table 4. Percentage of cases in which the arrival date from migration improved from 1 year to the next for a given individual. For territory
holders, only data up to 5 years of age were considered, in order to make them comparable to the data on floaters. Improvements above 75% are
highlighted in bold
†Number of individuals, each one sampled in two successive years; ‡tested by means of a binomial test; §individuals sampled in the year of their
first territorial establishment and in the previous year (their last year as floaters); ¶not available, or sample size too low for any meaningful
estimate or test.
In contrast, there was support for the age hypothesis and, suggesting that the effect of site dominance can be overcome
partly, for the site-dominance hypothesis. Age can lead to a by direct aggression coupled with social dominance. This
higher likelihood of territory acquisition in five non-exclusive implies a concerted action of the mechanisms proposed by
ways: (i) age can directly confer or be associated with higher the site-dominance and age hypotheses. In agreement with
dominance status, as shown in many studies (e.g. Smith this, territorial status was consistently predicted by the
et al. 1980; Rohwer et al. 1981; Cronin & Field 2007); (ii) interaction between age and arrival date.
older individuals have lower residual reproductive value and In particular, the comparison of Figs 2 and 4 suggested
are thus expected to be ready to fight harder for a territory a complex interplay between age and arrival in determining
(Grafen 1987; Shutler & Weatherhead 1994). This is con- territorial status. In the cross-sectional analysis shown in
sistent with the age effect that we observed in takeover fights. Fig. 2a, territory holders constantly improved their arrival
(iii) Conversely, in long-lived species with dangerous weap- with increasing age, particularly so in the youngest age
onry, like raptors, younger individuals are more likely to classes. However, in the longitudinal analysis of Fig. 4,
refrain from physical confrontations given the risk of lethal territory holders did not show such pronounced improve-
injuries and their likelihood of surviving to the next breeding ment in arrival dates. By exclusion, this implies a process of
season. (iv) In long-lived species, the cost of reproduction is progressive incorporation of early arriving birds into the
often high (e.g. Tavecchia et al. 2001), selecting for delayed territorial pool of the population, confirmed by the longitu-
territoriality as a voluntary restraint in order to enhance dinal analyses in which the transition from floater to terri-
longevity and the quality of the territory eventually acquired torial status coincided with pronounced improvements in
(Zach & Stutchbury 1992; Hunt 1998; Kokko & Sutherland arrival dates (Table 4, Fig. 4). In turn, such progressive
1998). Consistent with this idea, in our population delayed removal of early arriving birds from the floater sector of the
breeding led to higher longevity and older birds monopolized population may explain why floaters did not show any further
the best territories (Sergio et al. 2007a, unpublished; Blas et al. cross-sectional improvement in arrival date beyond the age
2009). Such strategy may contribute to accumulate older of 3 years (Fig. 2a). Therefore, only late-arriving birds seemed
birds in the territorial sector of the population. (v) Finally, to accumulate in the older floater age classes. Such ‘late, old’
even assuming that vacancies are filled randomly by the first floaters could be either low-quality individuals incapable
individuals encountering them, older individuals will have to advance their arrival, or individuals with an alternative,
accumulated more time and thus a higher probability to delayed breeding strategy. The latter explanation seems more
chance upon a vacancy than younger ones, contributing to likely, because such delayed breeders had a better nutritional
the older age-structure of territory holders (Shutler & Weath- status (Blas 2002) and a higher longevity (Blas et al. 2009),
erhead 1994). However, this mechanism is unlikely in our which is usually a reliable predictor of lifetime reproductive
model-system, because it assumes that territory holders do success (Newton 1989).
not suffer takeovers (Eckert & Weatherhead 1987) and it The above reasoning is consistent with two alternative
cannot explain why the winners of takeovers were older than ‘career-decisions’ co-existing in the population: (i) an invest-
the evicted individuals. ment in early arrival and early incorporation into the ter-
In agreement with the above ideas, most previous studies ritorial sector of the population; and (ii) an investment in
have shown that floaters are usually younger than territory long-term survival and in acquisition of social dominance.
holders (Porter 1985; Smith & Arcese 1989; Shutler & Weath- The first strategy entails a route to territoriality mainly medi-
erhead 1991; Sol et al. 2005; Blanco et al. 2007; but see also ated by the establishment of site dominance, while the second
Stutchbury 1991). However, in our population age alone may rely more on the dominance status conferred by age.
could not be the sole predictor of territorial status because However, considering that 84% of the individuals become
there was (i) wide overlap in the age structure of floaters and territorial within 4 years of age, the site-dominance route to
holders (Fig. 1), and (ii) pronounced variation among indi- territoriality seems to be the predominant one in the popula-
viduals in the age of first territorial establishment (range: 1–7 tion. All the above reinforces the idea of previous analyses,
years old). Therefore, two individuals of the same age could and extends it to the floater contingent, that access to breed-
still belong to different categories of territorial status. ing territories in this population is mediated by pre-emption
Variation in arrival dates may help to explain such differences and, secondarily, despotism (see Sergio et al. 2007a).
in territorial status within age classes (Fig. 2, see below). To date, we are aware of only one previous study which
Independent of age, early arriving birds had a higher focused on the potential link between arrival date and terri-
likelihood of acquiring a territory (Table 4, Fig. 2). Early torial status (Porter 1985). Its results were remarkably similar
arrival from migration may allow floaters to: (i) find more to ours: in Kittiwakes Rissa tridactyla, improvements in
numerous vacant territories; (ii) have more time to encounter arrival dates were particularly pronounced in the younger age
vacancies and gain familiarity with the general area; (iii) classes, and, within each age class, floaters arrived later than
pre-empt territories (sensu Rodenhouse, Sherry & Holmes established breeders. Longitudinal improvements in arrival
1997) for long enough to develop subsequent site dominance. were particularly marked for individuals passing from floater
All this is consistent with the site-dominance hypothesis. On to territorial status. Although more studies will be needed,
the other hand, in our system, occupying a territory too early the above observations suggest that the patterns found for our
could lead to subsequent eviction by an older individual, population may be common to many other migratory species.
© 2008 The Authors. Journal compilation © 2008 British Ecological Society, Journal of Animal Ecology, 78, 109–118
Predictors of floater status 117
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L. Marchesi, K. Norris, an anonymous associate editor and two anonymous in a Eurasian Sparrowhawk population. Ibis, 143, 442–449.
referees for comments on a previous draft of the manuscript. Part of this study Peer, K., Robertson, R.J. & Kempenaers, B. (2000) Reproductive anatomy and
was funded by the research projects PB96-0834 of the Dirección General de indices of quality in male tree swallows: the potential reproductive role of
Investigación Científica y Tecnológica, JA-58 of the Consejería de Medio floaters. Auk, 117, 74–81.
Ambiente de la Junta de Andalucía and by the Excellence Project RNM 1790 Penteriani, V., Otalora, F., Sergio, F. & Ferrer, M. (2005) Environmental
of the Junta de Andalucía. stochasticity in dispersal areas can explain the ‘mysterious’ disappearance
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