Ibis (2007), 149, 396–404
© 2007 The Authors
Journal compilation © 2007 British Ornithologists’ Union
Blackwell Publishing Ltd
Resident and transient dynamics, site fidelity and
survival in wintering Blackcaps Sylvia atricapilla:
evidence from capture–recapture analyses
EDUARDO J. BELDA,
1
* EMILIO BARBA
2
& JUAN S. MONRÓS
2
1
Universidad Politécnica de Valencia, Crta. Nazaret-Oliva, s/n, 46730 Gandía, Spain
2
‘Cavanilles’ Institute of Biodiversity and Evolutionary Biology, University of Valencia, Apartado Oficial 2085,
E-46071 Valencia, Spain
In their winter quarters, migrant birds may either remain within a small area (resident
strategy) or move frequently over a large area looking for locally abundant food (transient
strategy). It has been suggested that both strategies could simultaneously occur in the same
population. We used time-since-marking capture–recapture models to infer the coexistence
of these two behavioural strategies (transient and resident) among wintering Blackcaps
Sylvia atricapilla using weekly recapture data over a 7-year period. A related question is
whether Blackcaps, if surviving to the next winter, always return to the same wintering area,
so we also used this approach to analyse winter site fidelity and to estimate annual survival
probabilities. Model selection supported the existence of heterogeneity in survival estimates
for both the within-season and the interannual survival probabilities, i.e. there was evidence
for the existence of transients. It was estimated that 26% of the Blackcaps were resident
during the winter. Mean apparent annual survival probability was 0.46 (se = ±0.11).
However, there was some evidence suggesting that not all individuals showed winter site
fidelity. The estimated proportion of individuals that, if alive, returned to the wintering area
was 28%. This is the first study to show the existence of these two behavioural strategies
(residence and transience) among wintering Blackcaps, and the first confirming this pattern
using capture–recapture models. These models considering transient and resident dynamics
may become an important tool with which to analyse wintering strategies.
Survival is a crucial element of life history, especially
in short-lived organisms (e.g. Stearns 1992, Newton
2001). Migration can be viewed as a strategy to survive
during harsh or extreme environmental conditions,
which is well illustrated in many bird species that
undertake long migrations (Alerstam 1991, Berthold
2001, Senar & Borrás 2004). However, in spite of
survival being the ultimate objective of migration,
there are, to our knowledge, few studies analysing
the probabilities of survival in wintering areas (e.g.
Senar & Borrás 2004).
There are many factors that may affect winter
survival (for a recent review see Senar & Borrás 2004).
Two factors that have an important influence on sur-
vival are the search for and defence of food resources
(Senar & Borrás 2004), and the previous knowledge
of the wintering area, i.e. winter site fidelity (Newton
1972, Cuadrado et al. 1995). With regard to the first
factor, two main strategies of searching for food have
been described: the ‘resident’ and the ‘transient’
strategies. In frugivorous birds, wandering in search
of fruits may improve winter survival by preventing
food shortages that eventually occur in discrete
territories (Herrera 1985, Levey & Stiles 1992, Rey
1995). This is known as a transient strategy, in which
individuals locate and exploit alternative food
supplies over a large area (e.g. Newton 1972, Senar
et al . 1992, Pérez-Tris & Tellería 2002a, Newton 2004).
Alternatively, when resources are predictable and
plentiful, wintering birds should behave as residents,
with movements limited to a relatively small area,
and possibly showing territorial behaviour (Cuadrado
1995). Senar et al. (1992) (see also Senar & Metcalfe
*Corresponding author.
Email: ebelda@dca.upv.es