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Plenary Address - 8:15-8:55 AM
Quantifying variation in migratory strategies using ring-recoveries
G.M. Siriwardena, C.V. Wernham & S.R. Baillie
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Bird populations have traditionally been labelled as “migrant” or “resident” on the
basis of field observations and qualitative interpretations of patterns of
ring-recoveries. However, even such a non-systematic approach has identified many
intermediate species where only part of the population migrates (partial migrants) or
where different components of the population migrate to different extents (differential
migrants). A method that would allow a quantitative definition of migratory tendency to
be derived for many species would facilitate investigations into the ecological causes
and life-history consequences of migratory behaviour. Species or populations could then
be placed objectively into the continuum between true residency and an obligate,
long-distance migratory habit.
We present a novel method for the analysis of ring-recovery data sets that produces just
such a quantitative index of migratory tendency for British birds, developed as part of
the BTO's Migration Atlas project. The method uses distributions of ringing-to-recovery
distances to classify individual species' patterns of movement relative to those of
other species. The areas between species' cumulative distance distributions are treated
as inter-species dissimilarities and a one-dimensional map is then constructed using
multi-dimensional scaling. We have used the method in example analyses to show how it
could be used to investigate the factors that affect the migratory strategies that
species adopt, such as diet, territoriality and distribution, and in studies of their
consequences for demographic parameters such as annual survival and the timing of
breeding. We have also conducted initial analyses to show how temporal changes in the
indices could reveal otherwise unmeasured population consequences of environmental
change and thus have an important application in conservation science. Finally, we
discuss how our approach to producing indices of migratory tendency could be enhanced to
reduce the bias that follows from spatial or temporal variation in reporting rates and
how they could be made more broadly valuable by incorporating other data sets and
recovery data from other countries.
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Individual Papers
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8:55-9:20 AM
How do geometric constraints influence migration patterns?
- K. Thorup & C. Rahbek
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Null models exclusively invoking geometric constraints have recently been
demonstrated to provide new insight as to what explains geographic patterns
of species richness and range size distribution. Analyses of migration
patterns have traditionally been conducted in the absence of appropriate
simulations and analytical models. Here we present a null model exclusively
invoking geometric constraints and a more advanced analytical model
incorporating spread along a migration direction that allow investigation of
the influence of physiographical and physiological boundaries for
terrestrial taxa, with ocean and sea as geometric constraints, in relation
to observed patterns of migration. Our models take into account the low
recovery probability of terrestrial taxa over sea. The null model was not
found to explain any of the directional variation in the ring-recoveries,
but when comparing the distribution of data modeled using a simple
clock-and-compass model with distributions of ring-recoveries, geometric
constraints were found to explain up to 22% of the variation in
ring-recoveries. However, the assumed directional concentrations per step
used in the model were much higher than expected, and the qualitative fit of
the model was rather poor even when non-terrestrial sites of recoveries were
excluded.
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9:20-9:45 AM
Estimating dispersal in birds: taming the unknown.
A.J. van Noordwijk
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A fundamental problem in measuring dispersal is that the observations made are
restricted by the observations that could possibly be made. This quickly leads
to a situation where the distribution of observers in time and space
determines the observations more than the behaviour of the species studied.
Although we will never know where individuals lost out of sight went to, or
where immigrants came from, it is possible to obtain estimates of dispersal
distances that have been corrected for distance specific observation
probabilities (Thomson, van Noordwijk & Hagemeijer, J. Appl Stat in press).
Here I want to explore a complementary approach, we can use the frequency of
immigrants of unknown origin as an estimate of the unknown part of a
'dispersal as a function of distance' curve. Getting a number for how much we
don't know allows us to get a better perspective on what we do know. In nearly
all data sets, we will have a short range of distances in which nearly all
potential recruits were marked, a further range of distances in which a
proportion of all potential recruits were marked and a furthest range of
distances in which no potential recruits were marked. Where we can get an
estimate of the proportion marked in the middle range, from population
densities or distribution of suitable habitat, we can derive estimates for the
numbers of immigrants coming from these distances. Although we cannot know
where the rest of the immigrants came from, we do have an estimate of "the
total area under the curve" of the furthest distances. I will apply these
ideas to data on natal dispersal in Swallows, Blue Tits and Great Tits. This
approach will allow us to compare dispersal curves between groups (e.g.
sexes), but also curves for the same species among areas differing in the
distribution of suitable habitat (differing in fragmentation).
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9:45-10:10 AM
Multistate modeling of movement and philopatry of breeding ross's geese
- K. Drake & R. Alisauskas
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We estimated rates of breeding philopatry and dispersal within the Queen Maud
Gulf metapopulation of Ross's Geese (Chen rossii) using multistate
modeling of
neckband observations at five breeding colonies, 1999-2002. Philopatry was
female-biased, but probability of philopatry varied among colonies. Despite
probabilities of =0.62 for breeding philopatry to a given colony by either
sex, this study demonstrates that annual movement among breeding colonies was
substantial, and greater than previously assumed. We provide behavioral
evidence (consistent with genetic studies which suggested little or no
phylogeographic structure in the closely-related lesser snow goose, Chen
caerulescens) for extensive interconnectedness among breeding populations.
High likelihood of annual movements among colonies emphasize (1) the influence
of dispersal on changes in breeding distribution, and (2) provides insight
about the contributions of emigration and immigration to colony-specific
population growth rates. Although movement was the primary parameter of
interest, estimates of male survival from multistate modeling differed from
survival estimates from band recovery models, whereas female survival
estimates were consistent with those from band recovery models. We suspect the
difference between sexes in survival estimated from multistate models was due
to higher rates of neckband loss in males compared to females, as has been
found in numerous other studies, and/or higher rates of permanent emigration
by males from sampled areas. The importance of violating model assumptions
depends upon the biological parameter of interest, and whether inference is
restricted to the sample population or extended to the entire metapopulation.
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10:25-10:45 AM
Modelling Dolphin Behaviour using Multi-State Models with Time-Varying
Covariates
- S. Brooks & R. King
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Effective management is the key to the protection of many endangered species.
Identification of the primary factors affecting survival will often lead to
the introduction of strategies to improve survival rates. In this talk, we
consider a small population of Hector's dolphins located of the coast of New
Zealand and the impact that the establishment of a seasonal sanctuary has on
their survival and migration rates. We use a multi-state model to describe the
movement of the dolphins around the habitat and adopt a Bayesian approach
using reversible jump Markov chain Monte to distinguish between a wide range
of competing models. We also examine the impact of the inclusion of
time-varying covariates such as catch-effort information and demonstrate the
added value that these data provide in terms of both model discrimination and
parameter estimation. In particular, we find a whole class of models that
provide a far better fit to the data (and therefore better prediction and
ultimately better management) than those adopted in previous analyses.
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10:45-11:05 AM
Spatial distribution of breeding Pied Flycatchers Ficedula hypoleuca in
respect to their natal sites
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L. Sokolov, N. Chernetsov, V. Kosarev, D. Leoke, M. Markovets, A. Tsvey, A.
Shapoval
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Research project on the study of philopatry and dispersal of Pied Flycatchers
Ficedula hypoleuca has been underway on the Courish Spit on the Baltic
since
1980. A total of 8006 nestlings were ringed in the Russian part of the Courish
Spit. In subsequent years, 512 individuals (6,4%) were recaptured in the whole
study plot, 300 males and 212 females. The aim of this study was to test two
alternative hypotheses proposed to explain the distribution of breeding Pied
Flycatchers in respect to their natal site. The breeding area imprinting
hypothesis (Löhrl 1959, Berndt & Winkel 1979, Sokolov et al. 1984) assumes
that in spring, migrants try to return to a relatively small area, with a
radius of one to several km. The other hypothesis forwarded by Vysotsky (2001)
suggests that all first-year birds and many adults in spring arrive randomly
to an area much larger than the initial study area. In 2000, some 800
nest-boxes were added so that their overall number reached 1340 and the length
of the study plot reached 44 km. To study natal dispersal, we compared the
observed distribution of natal dispersal distances with the distribution
derived from the model assuming random arrival to the study area.
Significantly more Pied Flycatchers were recaptured within several km from the
natal site that at larger distance, and this was not a function of the control
effort. Therefore, it is concluded that the home area imprinted by juvenile
Pied Flycatchers has a diameter of several kilometres and not several dozens
of kilometres.
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11:05-11:25 AM
Timing and routes of migration based on ring recoveries and randomization
methods - H. Lokki & P. Saurola
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In the first EURING Technical Meeting we recommended randomization tests for
comparing wintering areas of different species or cohorts of birds on the
basis of ring recoveries. In this contribution we will use randomization
methods when trying to answer the following important questions of migration
studies: Do the timing and/or routes of migration of different species, sexes
or age classes differ from each other? Can anything statistically sound be
inferred from ring recoveries? As a rule, the data consists of non-normally
distributed recovery records (ti, x(ti), y(ti)) where x(ti) are longitudes,
y(ti) latitudes of birds recorded in N dates ti, i = 1, 2, … , N. We assume
that these records constitute a representative sample of places where the
birds have been found during their lives. At first, in order to sum up the
information included in the recovery locations in an "average route", we
create a time-window and slide it through the set of recoveries. In each
time-window we sample the records with replacement and compute the average of
the records in the sample. By repeating this procedure many times we obtain an
empirical distribution of the average. Then we combine these distributions
computed for each time-window to get the average route of the birds. For
comparing two routes, we compute the difference of the averages of the records
of the two groups in each time-window. Then we compute the sum of the
differences. In order to find out whether or not the observed sum of
differences is statistically significant we compute the empirical distribution
of the sum of differences as follows. In each time-window we randomize many
times the inclusion of records into the two groups. Then we compute
differences of the averages of the randomized groups and accordingly the sums
of the differences. The ranking of the observed sum in the empirical
distribution of the sums of differences determines whether or not the two
routes are similar geographically and in respect of timing. Finally we
illustrate the methods with data sets extracted from the Finnish recovery
database.
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11:25-11:45 AM
Multistate modeling of brood amalgamation in white-winged scoters (melanitta
fusca
deglandi)
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Joshua J. Traylor, Ray T. Alisauskas, and F. Patrick Kehoe
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There is a proclivity for female North American waterfowl to lose or abandon
offspring after hatch. Often these abandoned ducklings are subsequently brooded by
foster hens giving rise to the phenomena of post-hatch brood amalgamation (PHBA).
The potential fitness implications that arise from this behavior has brought about
considerable debate on physiological or ecological motivations for PHBA of young,
and potential costs and benefits of ducklings brooded in amalgamated broods. Few
studies have estimated the effects of PHBA on offspring survival and have not
followed individually marked offspring: most studies have been restricted to
studying PHBA based on maternal characteristics because only broods have been
individually color marked. Here, we explore relationships between probabilities of
movements from maternal to amalgamated broods by using a population of
individually marked white-winged scoters (i.e., females and offspring) (Melanitta
fusca deglandi) and pertinent ecological covariates utilizing multistate modeling.
We tested hypotheses about movement probabilities and 1) hatch date, 2) brood
size, 3) female size and condition, 4) duckling size and condition, and 5)
weather. We defined ducklings as belonging to one of 3 social strata: (A) natal
mother with no foster ducklings, (B) foster mother and conspecific non-siblings,
(C) natal mother with conspecific non-siblings and limited our study of movement
probabilities to two weeks after hatch because this period is when most mortality
and movement occurred. Thus, we estimated probabilities of survival, recapture,
and movement for each stratum focusing on covariates for movement. We evaluate
hypotheses suggested to motivate PHBA (e.g., accidental mixing, brood size and
brood success, and energetic stress hypotheses) and evaluate the influence on
duckling survival. Because the importance of predation in motivating abandonment
or adoption of ducklings remains equivocal, we discuss how duckling traits, in a
population located near high gull densities, may provide insights into the
relative importance of predation to PHBA.
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