BIRD STUDY

https://doi.org/10.1080/00063657.2022.2026876

Modelling population dynamics and trends in migratory birds from non-

standardized multi-species ringing data: the potential of multi-model selection
Tina Petrasa and Al Vrezec b,c

a
Institute of the Republic of Slovenia for Nature Conservation, Izola, Slovenia; bNational Institute of Biology, Ljubljana, Slovenia; cSlovenian
Museum of Natural History, Ljubljana, Slovenia

ABSTRACT ARTICLE HISTORY

Capsule: Long-term and non-standardized migratory bird ringing data can be used in models Received 17 August 2020
controlling variation in bird ringing methodology for reliable population trend estimations. Accepted 25 November 2021
Aims: Bird ringing data usually cover long periods and might reflect long-term population
changes. However, they are mainly derived during non-standardized multi-species catching at
numerous sites during the autumn migration period. We searched for the best modelling
approach to determine reliable species population dynamics and trend estimation models
based on annual multi-species bird ringing data.
Methods: We used ringing data from the Slovenian Bird Ringing Scheme and selected data in
three steps according to temporal, quantitative, and qualitative data selection. Annual indices
were constructed based on two types of denominators, ringing days, and ringing totals, vs. a
robust model without a denominator. We ran 20 candidate-generalized additive models
describing migrating population dynamics for 15 bird species by combining different data
selection approaches and denominators.
Results: We found that the models were species-specific, although the universal model could also
be applied to most species. We propose a general model construction approach for population
trend assessments from non-standardized bird ringing data. The estimates obtained by this
approach were comparable to the overall European population trends derived from breeding
survey data.
Conclusions: Bird ringing data from the autumn migration period are a valuable resource for
assessing continental scale population trends taking into account the whole population (non-
breeders and juveniles included) and even some rare and endangered species, but should be
conducted according to standard protocols to ensure reliable statistical inference of population
trends.

Population monitoring should constitute an essential
part of population ecology studies and species
conservation management, since change in abundance
is one of the most common responses to changes in
ecosystems (Acevedo-Whitehouse & Duffus 2009).
The duration of the monitoring period is critical for
distinguishing long-term trends from short-term
fluctuations (Primack 2006). To detect significant
population trends in birds, continuous monitoring
should be carried out for 10 years or more (White 2019).
In Europe, for example, bird population trends are
currently assessed using data from a well-established
network of national breeding bird counts, which
provide reliable regional and continental trend
estimates (Gregory et al. 2007, Heldbjerg et al. 2019).
However, alongside breeding bird count data, bird
ringing databases are another source of long-term
time series data, which could be used to estimate
trends (Busse 1990, Bibby et al. 1992, Peach et al.
1999, Rintala et al. 2003, Saurola & Francis 2004,
Briedis & Keišs 2016). Compared to count data,
ringing data offer a number of potential benefits. First,
there are longer runs of historical data from ringing
schemes, which give the opportunity to analyse species
population dynamics over longer time periods. Most
breeding bird count schemes refer to the last 30 years,
with the first scheme started in 1966 (Heldbjerg et al.
2019), while most of the ringing scheme datasets cover
periods of 100 or more years, with the first scheme
started in 1899 (Preuss 2001, Greenwood 2009, Møller
& Fiedler 2010, Saurola et al. 2013, Bairlein et al.
2014). Second, ringing data cover bird populations on

CONTACT Al Vrezec al.vrezec@nib.si, al.vrezec@gmail.com

Supplemental data for this article can be accessed at https://doi.org/10.1080/00063657.2022.2026876.
© 2022 British Trust for Ornithology
2 T. PETRAS AND A. VREZEC
a larger, continental scale, while breeding bird count
data are limited to local or regional populations.
During autumn migration, when most of the ringing
is conducted, birds aggregate from larger areas thus
enabling more comprehensive insight into continental
scale population trends, including birds from remote
areas with otherwise scarce ornithological activity
(Spina & Volponi 2008). Third, different aspects of
population data are captured in ringing data. While
breeding bird count data consider breeding pairs or
territorial males only, ringing data provide
information on the whole population, including
breeders, non-breeders, and immature individuals, an
aspect that is already well implemented in the
Constant Effort Site ringing programme (Robinson
et al. 2009, Kampichler & van der Jeugd 2011).
Finally, data on ringed migratory birds also include
those for some rare and endangered species (e.g.
Bluethroat Luscinia svecica and Moustached Warbler
Acrocephalus melanopogon), which are not included in
common bird censuses but are regularly ringed during
autumn migration (Spina & Volponi 2008, Kralj et al.
2013, Valkama et al. 2014, European Bird Census
Council 2017).
The initial aim of bird ringing was to track migration
patterns and routes of birds and not to monitor
population changes, which make some ringing data
overall less standardized and potentially more biased.
The main assumption when using bird ringing data in
population trend analyses is that there is a close link
between the number of birds ringed annually and
ringing effort (Hjort & Lindholm 1978). Thus, reliable
and straightforward estimations of population trends
are possible when the methodology is standardized.
Such standardizations are advancing as a part of
ringing schemes in Europe, Asia, and North America
(Baillie et al. 1986, DeSante et al. 1995, Sokolov et al.
2000, Robinson et al. 2009). However, many bird
ringing schemes in Europe have been operating
continuously for 100 years or more, and much of the
long-term datasets were obtained in a non-
standardized way, which hampers analyses of
population trends. The source of the variability in the
data lies in variation in the number of ringers,
locations, catching methods, number, and type of nets
and/or traps, the use of playback, weather conditions,
and habitats. These variables have their own trends
and this hinders the straightforward estimation of
relevant population trends for the species concerned
(Busse 1990, Sutherland 1996, Rintala et al. 2003).
Since annual changes in ringing effort are almost
impossible to estimate at a long-term historical level
(Hjort & Lindholm 1978, Rintala et al. 2003), indirect
analyses that increase the reliability of statistical
inference from these data are of key importance in the
evaluation of population trends (Mason & Hussey
1984, Rintala et al. 2003, Briedis & Keišs 2016).
Various methods have been suggested for the
standardization of heterogeneous bird ringing data;
for example, the comparison of single-species totals
with other species or with other parameters that can
be estimated from available data (Ginn 1969), or the
identification of indicator species with similar
ecological traits, to control for variation in bird
ringing methodology (Perrins 2003, Rintala et al.
2003, Keišs et al. 2007, Briedis & Keišs 2016).
In this paper, our aim was to combine different
approaches to account for bird ringing data variability
and thus determine the best species population
dynamics and trend estimation models based on
annual multi-species bird ringing datasets. The bird
ringing datasets were mainly derived during non-
selective and non-standardized multi-species catching
with mist-nets at numerous sites during the autumn
migration period. For trend estimation, therefore, it
was necessary to distinguish individual species ringing
data from non-specific multi-species ringing data. We
attempted to explore whether one general model could
be used to assess population trends, or whether the
modelling of bird ringing data must be species-
specific. The main aims of our study were: (1) to
reduce bias to the greatest extent possible in the
assessment of ringing effort, (2) to estimate the long-
term trends of the migrating populations of the target
species, (3) to compare the best models for assessing
population trends, and (4) to assess model trend
estimates against known trends derived from breeding
bird count data.
Methods
Target species and selection of bird ringing data
We used bird ringing data from the Slovenian Bird
Ringing Scheme and selected 15 bird species that
sufficiently describe the variability in the bird ringing
dataset according to several ecological and
methodological criteria, as well as differences in
sample sizes (Table 1). The Slovenian Bird Ringing
Scheme is run by the Slovenian Museum of Natural
History, and the ringing database encompasses data
on ringed bird species that have been continuously
collected since 1927 (Božič 2009). In the current
analysis, we included data from the 17-year period
between 2000 and 2016, during which several field
ringing stations were operating on a more or less
 BIRD STUDY 3

Table 1. Ecological and methodological traits for 15 migratory bird species that describe the variability in the bird ringing dataset,
with the mean number of annually ringed birds at autumn migration in the period 2000–2016 (±sd). W: woodland, F: farmland,
M: marshland, U: urban. Status – RB: regular breeder, OB: rare occasional breeder, PM: passage migrant, RW: rare wintering,
S: habitat specialist at stopover sites (wetlands), G: habitat generalist at stopover sites, NA: no data available. Data references:
Cramp & Perrins (1983), Mukhin et al. (2005), Mukhin et al. (2008), European Bird Census Council (2017), and BirdLife International
(2020).
Number of birds Annual
ringed annually at Habitat Responding to European
autumn migration in specificity at playback of the Included in population
Slovenia (2000- Breeding Breeding Status in migration Garden Warbler night playback trend estimate
Species 2016) habitat period Slovenia stopover sites song vocalisations (2000–2016)
Tree Pipit Anthus 112 ± 41 W/F IV – VIII RB, PM G No No −0.6%
trivialis
European Pied 122 ± 49 W late IV - OB, PM G No No −1.4%
Flycatcher late VI
Ficedula
hypoleuca
Icterine Warbler 345 ± 175 W late V - VII OB, PM G No No −1.2%
Hippolais
icterina
Eurasian Wryneck 252 ± 72 W/F V – VI RB, PM G No No +0.2%
Jynx torquilla
Red-backed 224 ± 85 F V - VII RB, PM G No No −0.7%
Shrike Lanius
collurio
River Warbler 17 ± 10 W late V - RB, PM S NA No −2.7%
Locustella mid-VII
fluviatilis
Common 145 ± 65 F/M late IV - RB, PM S Yes No −0.2%
Grasshopper mid-VII
Warbler
Locustella
naevia
Common 181 ± 41 W/F late IV - RB, PM G NA No +0.3%
Nightingale mid VII
Luscinia
megarhynchos
Spotted 81 ± 27 W/U mid-V - RB, PM G No No −0.6%
Flycatcher mid-VIII
Muscicapa
striata
Common Redstart 62 ± 24 W/F late IV - RB, PM G No No +2.2%
Phoenicurus mid-VII
phoenicurus
Common 2207 ± 484 W IV - early RB, PM, G NA Yes +3.1%
Chiffchaff VIII RW (background)
Phylloscopus
collybita
Eurasian Blackcap 20728 ± 5608 W/F mid-IV - RB, PM, G Yes Yes +4.6%
Sylvia atricapilla VIII RW (background)
Garden Warbler 6972 ± 2131 W/F IV - VII RB, PM G Yes Yes (main voice) −1.2%
Sylvia borin
Common 472 ± 147 F IV - VII RB, PM G No No −1.2%
Whitethroat
Sylvia
communis
Song Thrush 322 ± 71 W mid-III (IV) RB, PM, G NA No +1.6%
Turdus - mid- RW
philomelos VIII

regular annual basis during the autumn migration
period (Vrezec et al. 2014; Figure 1). There were 450
active ringing locations during the study period, 86%
of which were active for less than 75% of the study
period (1–12 years) and 14% of which were active for
more than 75% of the study period (13–17 years).
Among the latter group, 17 ringing stations were
active every year and were evenly distributed
throughout Slovenia (Figure 1). During autumn
migration, birds were caught, mostly with mist-nets,
and information on ring number, species, age, sex,
wing length, mass, date and place of ringing, and
name of ringer was collected. Since our study focused
on the population dynamics of migrating birds in the
autumn, we excluded from the dataset all ringed
nestlings and data from the spring period (January to
June). The use of playback of a recording of birdsong
significantly increases the catch rate (Mukhin et al.
4 T. PETRAS AND A. VREZEC

Figure 1. Ringing stations that were operating during the autumn migration periods between 2000 and 2016 in Slovenia. Locations
that were active at least 75% of the time during the 17-year period are marked with black spots, while locations that were active less
than 75% of the time during the 17-year period are marked with white spots.

2008). In Slovenia, this method has been used since 1995
(Šere 2001). Playback was usually used during mist-
netting in the autumn migration period and broadcast
during the night from about 01:00 to one hour after
sunrise (B. Lapanja, pers. comm.). A recording of the
song of the Garden Warbler Sylvia borin was used
with at least six additional species vocalizing in the
background, predominantly Common Chiffchaff
Phylloscopus collybita and Blackcap Sylvia atricapilla
(Table 1). The recording was taken at the Ljubljana
Marsh in central Slovenia during the breeding season
by Tomi Trilar (Slovenian Museum of Natural

Table 2. Set of a priori models for assessing the population dynamics of the autumn migratory populations of 15 bird species
considering different combinations of data selection approaches and denominators for calculating indices.
No. Model Autumn migration period Ringing effort Indicator species Denominator
Species- Selected days with
General specific defined minimum No Selected days No. of
autumn autumn No ringing of ringed birds or indicator with at least ringed birds No. of
migration migration effort ringed bird species one indicator without ringing
period period selection species selection species No nestlings days
1 GEN_NO_NO_N x x x x
2 GEN_NO_NO_B x x x x
3 GEN_NO_NO_D x x x x
4 GEN_NO_IND_B x x x x
5 GEN_NO_IND_D x x x x
6 GEN_SEL_NO_N x x x x
7 GEN_SEL_NO_B x x x x
8 GEN_SEL_NO_D x x x x
9 GEN_SEL_IND_B x x x x
10 GEN_SEL_IND_D x x x x
11 SPE_NO_NO_N x x x x
12 SPE_NO_NO_B x x x x
13 SPE_NO_NO_D x x x x
14 SPE_NO_IND_B x x x x
15 SPE_NO_IND_D x x x x
16 SPE_SEL_NO_N x x x x
17 SPE_SEL_NO_B x x x x
18 SPE_SEL_NO_D x x x x
19 SPE_SEL_IND_B x x x x
20 SPE_SEL_IND_D x x x x

History). The playback was used consistently
throughout the study period, but bias occurred due to
its non-systematic use with respect to the duration of
playback, combination with other recordings during
the daytime and uneven use between ringing stations.
There were approximately 110 ringing stations (24%)
that were using night playback, while the playback
broadcast during the daytime was applied regularly at
all ringing stations.
As ringing data were collected in a non-standardized
way, without clear reference to the protocol used (e.g.
the number of mist nets, duration of ringing, weather
data, playback used or not), we selected the data in
three steps according to the following criteria before
conducting analyses (Table 2):
(1) Temporal data selection. As our study focuses on
the autumn migration period, we defined two
time periods: general and species-specific. The
general autumn migration period encompasses the
whole autumn migration period and was
arbitrarily defined as the second half of the year,
from 15 July to 31 December. The species-specific
time period for the target species was determined
from ringing data. In the analysis, only the
months in which ringed individuals of the target
species constituted 5% or more of the total
number of ringed birds in the autumn migration
period (across all years from 2000 to 2016) were
included in the core migration period for the
target species in Slovenia.
(2) Quantitative data selection (ringing effort selection).
Multi-species ringing data are composed of data
obtained by intensive mist-netting in the
migration period as well as data which were not
relevant for our analysis, such as those on
occasional ringing of nestlings or found birds and
species-specific ringing. Therefore, we selected
ringing days according to the number of
individual birds and the number of species ringed
per day. To define thresholds for ringing days
relevant for migration monitoring, we plotted the
number of ringed birds and ringed species per
day per location and then visually defined the
point on the curve delineating intensive vs. non-
intensive ringing days. We used the geom_vline
function from the package ggplot2 in the
statistical program R (R Development Core Team
2018), to add a vertical line representing the
threshold between intensive vs. non-intensive
ringing days (Teetor 2011).
(3) Qualitative data selection (target species sample
selection). One of the shortcomings of using
BIRD STUDY 5
ringing data collected in a non-standard way is
that there are no reliable data on target species
samples vs. samples in which the target species
was not likely to be caught and ringed. In
previous studies, species totals as a proportion of
totals of ecological indicator species have been
proposed to overcome this problem (Ginn 1969,
Perrins 2003, Rintala et al. 2003, Keišs et al.
2007). Our approach to account for unreliable
data was to first select all species that occurred in
the days when the target species were ringed from
2000 to 2016 during the species-specific autumn
migration period in Slovenia. Among the selected
species, we identified potential indicators as
species that co-occurred with the target species in
75% or more of the ringing days during the
autumn migration period. The Barn Swallow
Hirundo rustica was excluded from the potential
indicator species since it was known to be ringed
in large numbers at night roosts within the
framework of the EURING Swallow Project which
proceeded in Slovenia until 2013 (Vrezec et al.
2014) and therefore could potentially obscure the
trend direction of the target species. Further, we
modelled the presence/absence of the target
species (response variable) with the presence/
absence of the potential indicators (predictors) for
each day. When selecting indicator species, we
assumed that they behaved in a similar way to the
target species during migration and were thus
more likely to be caught together with the target
species (Bairlein 1983, Rintala et al. 2003, Briedis
& Keišs 2016). This approach also acknowledged
the inconsistent use of playback, which might
attract target species in a similar way as indicator
species. We identified such species co-occurrence
patterns and possible species groupings with a
dendrogram created in the PAST software
program (Hammer 2019), applying Ward’s
method, which combines clusters by minimizing
variance, measured by the sum of squares
(Murtagh & Legendre 2014, https://www.
statisticshowto.com/wards-method/). Logistic and
stepwise regression (Thompson 1989, Royle et al.
2012, Smith 2018) were performed in the R
software program using the package MASS (R
Development Core Team 2018). Data analyses
were based on the assumption that birds were
trapped and ringed randomly. The result was a
guild of species which best indicated the
probability of the presence of the target species.
To obtain a finer selection of the indicator species
guild, we set the significance level at P < 0.01. A
6 T. PETRAS AND A. VREZEC
sampling day for the target species was defined as a
day in which the target species had the potential to
be caught when ringing was carried out, i.e. a day in
which the target species or at least one of the
indicator species was caught.
To standardize data characterized by variable ringing
effort, we constructed annual indices based on two types
of denominators: (i) ringing days and (ii) ringing totals
for all birds without nestlings. In addition, we also
applied a robust model without a denominator, using
only (iii) absolute numbers of ringed target species per
year. We ran 20 candidate-generalized additive models
(GAMs) describing migrating population dynamics for
15 bird species by combining different data selection
approaches and denominators (Table 2).
Population dynamics modelling
The deviance explained (DE), which is a statistic that is
identical to pseudo-R 2, was used for model comparison
and selection and was calculated with the following
equation (Wiersma & Skinner 2011):
DE = 1–(residual deviance/null deviance)
Among the candidate models, the model with the
greatest percentage of explained deviance for each
species was considered the best model. The selected
model was chosen for further analyses of population
dynamics. Non-significant models and those with
explained deviance under 50% were rejected and were
not used in further trend analyses. We fitted our data
using a GAM, which results in non-linear, more
complex trends (Simpson 2018). In addition, the fitted
curve of the GAM allows for comparison of the
dynamics between different models (species). As a
type of smoothing, we used a thin plate spline, which
is more general and especially useful for noisier data
(Craven & Wahba 1979, Hastie & Tibshirani 1986).
For model validation, we used standard errors and
confidence intervals. We applied the bootstrap
technique, a statistical tool that can be used to
quantify the uncertainty associated with a given
estimator (Varian 2005, Banjanovic & Osborne 2016,
James et al. 2017). Here, we resampled the indices
from the best models, using 95% confidence intervals
with the recommended number (399) of bootstrap
replicates for data similar to ours (Fewster et al. 2000).
We used normal bootstrap confidence intervals, where
the standard error (se) is computed as the standard
deviation of the bootstrap distribution. Subsequently,
the 95% confidence intervals are computed as the
sample mean ± 1.96 ∗ se (Banjanovic & Osborne 2016).
Additionally, we determined the universal model, i.e.
the model which fitted best across the widest range of
species, to be served as the best overall model for
assessing population trends from multi-species bird
ringing data on autumn migrants. We thereby selected
all species for which the best models were not
rejected. For each model in each species, we calculated
the proportion of DE by the best model, giving the
value 1.0 to the best model. Furthermore, we
calculated the mean proportion of DE for each model
across all species and defined the universal model as
the model with the highest mean proportion of DE,
which indicates the overall best fit in the greatest
number of species.
Since the GAM does not provide population trend
estimates, we applied fitted values from the GAM in a
general linear model (GLM) for estimating the
population growth rate, as the model coefficient gives
the predicted change in indices for one-unit increase
per year (Carey 2013).
Results
Examined populations
According to data from the literature and recoveries
stored in the Slovenian Bird Ringing Centre database,
the migratory directions of the target species followed
routes from Scandinavia southwards, and from Russia
and Finland towards the south-west (Figure 2). Some
populations of Grasshopper Warbler Locustella naevia,
Common Chiffchaff, and Song Thrush Turdus
philomelos migrate in a more eastern direction, from
Russia and Belarus, while only the Red-backed Shrike
Lanius collurio has a generally south-eastern migratory
direction (Cramp & Perrins 1983, Andreotti et al.
1999, Wernham et al. 2002, Cepák et al. 2008, Spina &
Volponi 2008, Božič 2009, Korner-Nievergelt et al.
2012, Tøttrup et al. 2012, Kralj et al. 2013, Bairlein
et al. 2014, Busse et al. 2014, Ouwehand et al. 2015,
Pedersen et al. 2018, Belgian Ringing Scheme 2019).
Thresholds for data selection
In the temporal data selection step, we defined the
species-specific migration period for the target species,
which was used in further analyses (online Table S1).
Furthermore, ringing effort selection was set to
locations with a daily minimum of 50 ringed birds
and five ringed species (online Figure S1). The
number of indicator species that showed the best co-
 BIRD STUDY 7

Figure 2. General migration routes of the target species through Slovenia. Abbreviations: ANTTRI: Tre Pipit Anthus trivialis, FICHYP:
European Pied Flycatcher Ficedula hypoleuca, HIPICT: Icterine Warbler Hippolais icterina, JYNTOR: Wryneck Jynx torquilla, LANCOL: Red-
backed Shrike Lanius collurio, LOCFLU: River Warbler Locustella fluviatilis, LOCNAE: Grasshopper Warbler Locustella naevia, LUSMEG:
Common Nightingale Luscinia megarhynchos, MUSSTR: Spotted Flycatcher Muscicapa striata, PHOPHO: Common Redstart
Phoenicurus phoenicurus, PHYCOL: Common Chffchaff Phylloscopus collybita, SYLATR: Blackcap Sylvia atricapilla, SYLBOR: Garden
Warbler Sylvia borin, SYLCOM: Common Whitethroat Sylvia communis, TURPHI: Song Thrush Turdus philomelos.

occurrence with the target species after performing
logistic and stepwise regression varied between one
(for River Warbler Locustella fluviatilis) and eight (for
Icterine Warbler Hippolais icterina) with a median of
five, and were specified as indicators (Table S1).
Hierarchical clustering distinctly separated four
groups of species according to similarity of indicator
species (Figure 3): (1) Grasshopper Warbler, Wryneck
Jynx torquilla, and Common Nightingale Luscinia
megarhynchos; (2) Pied Flycatcher Ficedula hypoleuca,
Red-backed Shrike, Garden Warbler, Icterine Warbler,
and Spotted Flycatcher Muscicapa striata; (3)
Common Chiffchaff and Song Thrush; and (4) Tree
Pipit Anthus trivialis, River Warbler, Common
Redstart Phoenicurus phoenicurus, Blackcap, and
Common Whitethroat Sylvia communis. The first level
of selection was done according to migration period
and the second-level selection according to
preferences of habitat and migratory direction.

Population dynamics model selection

Figure 3. Ward’s hierarchical clustering of analysed species
(guilds) according to common indicator species. See Figure 2
for species codes.
For each species, we selected the best GAM with the
greatest percentage of explained deviance and thus the
highest explanatory power (Table 3; online Table S2).
However, not all of the best models were actually
significant or explained at least 50% of the deviance.
Quantitative data selection and different denominators
primarily contributed to the division between accepted
and rejected best models, while the other types of data
selection (temporal and qualitative data selection) did
not have such power.
The best models for the Tree Pipit and Blackcap did
not have denominators and were non-significant (Table
8 T. PETRAS AND A. VREZEC

Table 3. Best and universal general additive models of population dynamics for the 15 target bird species. Estimated annual growth
rate (trend) was assessed in a GLM. Significant codes: ∗∗∗ P < 0.001, ∗∗ P < 0.01, ∗ P < 0.05. Abbreviations: UM: universal model (the
general autumn migration period from 15 July to 31 December, selected days with ≥50 ringed birds and ≥5 ringed species, no
indicator species selection, number of ringing days as denominator), CI: confidence intervals, DE: deviance explained.
Parametric coefficients
Approximate Estimated
Best GAM model/ Lower Upper significance of DE Model annual growth
Species UM Estimate CI CI smooth terms (F) (%) acceptance rate (GLM) Trend
Tree Pipit GEN_SEL_NO_N 80.765 63.940 96.920 2.8 15.9 Rejected Uncertain
European Pied Flycatcher GEN_SEL_IND_D 0.427 0.361 0.492 3.6∗ 77.6 Accepted −0.012∗∗∗ Decline
UM 0.216 0.179 0.257 3.5∗ 66.3 Accepted −0.022∗∗∗ Decline
Icterine Warbler SPE_SEL_NO_B 0.006 0.005 0.008 3.4∗ 75.0 Accepted −0.042∗∗∗ Decline
UM 0.669 0.518 0.822 8.1∗∗ 47.5 Rejected Uncertain
Eurasian Wryneck SPE_SEL_NO_B 0.005 0.004 0.005 2.1 70.1 Rejected Uncertain
∗ ∗∗∗
Red-backed Shrike SPE_SEL_NO_B 0.003 0.003 0.004 4.5 84.9 Accepted −0.019 Decline
UM 0.368 0.307 0.429 7.9∗∗ 74.7 Accepted −0.052∗∗∗ Decline
River Warbler GEN_NO_NO_D 0.094 0.068 0.119 10.8∗∗ 62.0 Accepted −0.090∗∗∗ Decline
UM 0.034 0.025 0.043 7.4∗∗ 54.9 Accepted −0.082∗∗∗ Decline
Common Grasshopper Warbler GEN_SEL_NO_D 0.301 0.247 0.359 8.9∗∗ 56.5 Accepted −0.059∗∗∗ Decline
UM 0.301 0.247 0.359 8.9∗∗ 56.5 Accepted −0.059∗∗∗ Decline
Common Nightingale GEN_SEL_NO_B 0.002 0.002 0.003 9.8∗∗∗ 77.4 Accepted +0.054∗∗∗ Increase
UM 0.351 0.312 0.389 5.0∗ 64.6 Accepted +0.028∗∗∗ Increase
Spotted Flycatcher GEN_NO_IND_D 0.934 0.807 1.054 12.5∗∗∗ 80.3 Accepted −0.013∗∗∗ Decline
UM 0.146 0.124 0.166 6.5∗∗ 70.2 Accepted −0.023∗∗∗ Decline
Common Redstart SPE_SEL_IND_B 0.108 0.085 0.131 3.6∗ 56.6 Accepted +0.041∗∗∗ Increase
UM 0.110 0.088 0.132 0.5 3.1 Rejected Uncertain
Common Chiffchaff SPE_NO_NO_B 0.030 0.025 0.035 6.2∗∗ 78.2 Accepted +0.041∗∗∗ Increase
UM 3.912 3.538 4.270 3.7∗ 55.3 Accepted +0.009 Stable
Eurasian Blackcap SPE_SEL_NO_N 17711 15106 20337 2.7 33.3 Rejected Uncertain
∗∗∗
Garden Warbler SPE_NO_IND_B 0.904 0.890 0.917 24.5 96.6 Accepted −0.001 Stable
UM 14.39 12.50 16.36 9.1∗∗ 91.0 Accepted −0.039∗∗∗ Decline
Common Whitethroat SPE_SEL_IND_B 0.042 0.037 0.048 9.0∗∗∗ 90.2 Accepted −0.003 Stable
UM 0.930 0.783 1.067 17.1 ∗∗∗ 80.4 Accepted −0.050∗∗∗ Decline
Song Thrush SPE_NO_NO_B 0.004 0.003 0.005 5.2∗ 25.7 Rejected Uncertain

3). The best model for the Wryneck was also non-
significant, while the model for the Song Thrush was
significant but had low explained deviance (26%). Due
to low explanatory power, all four of these species
were excluded from further analyses since the current
dataset, or our set of a priori models, did not
sufficiently account for sampling biases in these four
species.
The best estimated models were in most cases those
in which the denominators were the number of ringed
birds and in which migration time was specified by
the species-specific period. All of these models were
characterized by high explained deviance, which
mostly varied between 75% and 97% (Table 3).
Models with ringing days as denominators were
determined by the general migration period and had
56–80% of explained deviance, but only in species
with lower mean annual ringing totals (less than 150
ringed birds per year; Table 3). Data selection
according to qualitative (indicator species) and
quantitative criteria (ringing effort) also contributed to
model improvement.
Model 8 (the general autumn migration period from
15 July to 31 December, selected days with ≥50 ringed
birds and ≥5 ringed species, no indicator species
selection and with number of ringing days as
denominator) fitted best across the widest range of
species with an average proportion of DE compared to
the best model being 0.78, which was determined as a
universal model. The second (model 10) and third
(model 18) best candidate universal models had 0.74
and 0.72 of DE, respectively.
Estimation of population trends
Modelling of population dynamics was conducted only
for nine out of 15 species with accepted models, i.e.
significant models with adequate explanatory power
 BIRD STUDY 9

(˃50% explained deviance) and for which it was possible Model selection
to determine acceptable universal models (Table 3,
We have demonstrated that the accepted best models
Figure 4). In all of these species the universal model
of population dynamics in the studied migratory birds
showed the same direction of population trend, but in
derived from non-standardized long-term bird ringing
three species (Common Chiffchaff, Garden Warbler,
data significantly improve sampling effort control.
Common Whitethroat) the significance of the trend
Most of the species analysed are long-distance
was changed (Table 3). The universal models for the
migrants, with a migratory peak in August.
Common Nightingale showed an increasing trend
Presumably, the night playback of calls influenced
with estimated annual growth rate of 2.8%. In the
the capture rates of certain species, especially Sylvia
majority of species (European Pied Flycatcher, Red-
warblers and the Common Chiffchaff (Mukhin et al.
backed Shrike, River Warbler, Grasshopper Warbler,
2008; Table 1), and attracted species even to
Spotted Flycatcher, Garden Warbler, and Common
suboptimal habitats (Herremans 1990, Mukhin et al.
Whitethroat) there was a declining trend, with an
2008). Nevertheless, playback is often used during
annual rate of change ranging from −2.2% to −8.2%.
bird ringing sessions in autumn, as it allows larger
The trend for the Common Chiffchaff was stable
sample sizes to be obtained, but such data might be
(Table 3). With respect to population dynamics, the
biased in several ways (Mukhin et al. 2008, Arizaga
Pied Flycatcher, Spotted Flycatcher, and Garden
et al. 2015, De La Hera et al. 2017). Therefore,
Warbler showed patterns with greater fluctuations
models of species that respond to playback are
(Figure 4).
expected to have more error and presumably
In comparison with the general estimated population
reduced explanatory power.
trends in Europe based on breeding survey data for the
Four of the models were rejected (Tree Pipit,
same time period (Table 1), the estimated trends from
Blackcap, Wryneck, and Song Thrush). Within these
ringing data matched the direction in all accepted
models, only one species that responds to acoustic
models, when considering best models as well as a
signals was included (Blackcap), so response to the
universal model. However, the magnitude of trend
playback per se was not the key bias for statistical
estimates was found to be generally higher in ringing
inference. The rejected models were primarily those
than in count data.
with no denominators. They showed low explained
deviance and insufficient model fitting. Unlike the
Discussion other non-significant models, the model for the
Wryneck had ringed birds in the denominators and a
We have shown that non-standardized multi-species
high proportion of explained deviance. Further
bird ringing can provide useful long-term data for
modelling, taking into account other aspects of
statistically reliable population trend estimation. The
variability in the data, would therefore be required in
best models were found to be species-specific, but
future studies to search for other possibilities in
we have found that an universal model with the
controlling biases in non-standardized ringing data.
best fit in the greatest number of studied species
Most of the models were accepted and showed that
provided similar and well supported trend estimates
the selections proposed in the study resulted in good
for most of the species. Generally, data derived from
model fits. The most powerful models were those that
variable ringing activity are too heterogenous to
considered selected data according to quantitative
allow for a uniform modelling procedure; therefore,
(ringing effort) data selection. Through quantitative
our approach was to construct 20 a priori models
selection we essentially removed incidental findings or
for model selection, which allowed us to derive
occasional ringing sessions and thus gained more
reliable population trend estimates comparable to
homogenous samples, with mist-netting as the
overall European population trends derived from
prevailing method. This type of selection provided the
breeding survey data (European Bird Census Council
best fit in 64% of the accepted models.
2017). Although abundance estimates based on non-
The best estimated models with qualitative data
standardized methods have largely been avoided in
selection alone were found only in two species.
the past (Pollock et al. 2002), the recent
Qualitative data selection, alone or together with other
development of more sophisticated statistical
selections, provided the best fit in 45% of the accepted
modelling helps to control for different sources of
models and hence contributed to the power of models.
variability in data (Walker & Taylor 2017) and thus
The use of indicators has already been shown to be a
provides more powerful and reliable statistical
beneficial approach (Rintala et al. 2003). Our study is
inference.
10 T. PETRAS AND A. VREZEC
Figure 4. Population dynamics of nine bird species with standard errors and partial deviance residuals according to the universal
model.
based on the assumption that particular species
congregate in specific habitats at stopover sites, which
is a well-known migratory pattern that is associated
with food requirements (Bairlein 1983). Considering
such species guilds, we defined potential sampling
(ringing) days for the target species according to the
presence of indicator species.
Differentiation between groups of indicator species
was based mainly on the migration period. In two
groups at the first level of differentiation (Figure 3),
the main separation parameter was the migration
period, with the first group formed by species
migrating from July to September and the second
group formed by species migrating from August to
October. Such separation was expected, as we chose
indicator species in the species-specific migration
periods. In each first level species group, there were an
additional two groups representing a second level of

differentiation (altogether four) separating species by
habitat preferences and migratory direction. One was
comprised of species with a south-western autumn
migratory direction and occurring predominantly in
farmland habitats: the Grasshopper Warbler, Wryneck,
and Common Nightingale (Cramp & Perrins 1983,
Cepák et al. 2008, Spina & Volponi 2008). The second
group included two species, the Common Chiffchaff
and Song Thrush, which are woodland species with a
western autumn migratory direction (Cramp & Perrins
1983, Andreotti et al. 1999, Busse et al. 2014).
The models in which the numbers of ringed birds were
considered as the denominator typically showed higher
explained deviance. Such models were more powerful,
as they were better at controlling for sampling effort
variability than those denominated by ringing days. For
instance, differences in the time and duration of the
catch, the use of a different number of nets, the use of
playback or methods other than mist-netting might
result in different numbers of birds caught per ringing
day. In such cases, ringing days do not adequately
control for the variable sampling effort, especially for
more abundant species. In contrast, for species which
were ringed in lower daily numbers, ringing days better
account for changes in ringing activity since ringing
days remained more stable throughout the period than
the number of ringed birds. On the contrary, the large
number of species, i.e. nine of 11 species with accepted
best models, could be modelled by a universal model
with ringing days as the denominator. The trends,
obtained by the best and universal models, showed the
same direction, with slight differences in estimated
annual growth rate between them.
All models with denominators provided good
improvement in model fit for the majority of species.
This is in contrast to some previous studies, where
data obtained in a non-standardized way with
multiple sources of bias were better suited to robust
modelling without denominators (Walker & Taylor
2017, Boersch-Supan et al. 2019). In such instances,
the sample size may play an important role. The large
dataset obtained from multiple locations could
obscure the biases and heterogeneity in the samples
(Kaplan et al. 2014) and may increase the precision in
trend estimates (Gregory et al. 2004, Fink et al. 2020).
With respect to the rejected models, further
improvement of the models should be made to obtain
more powerful species-specific models of population
trends from long-term bird ringing data. It is
important to note that it is very likely that estimating
population trends from past bird ringing data are not
possible for all species due to a species-specific catch
rate, which might be too accidental and unrelated to
BIRD STUDY 11
the actual population dynamics of the species. In such
cases, modelling is not possible.
Population dynamics and trends
Our results of long-term trend analyses using bird
ringing data were in agreement with the overall
directions of European trends derived from breeding
surveys (European Bird Census Council 2017).
However, the magnitude of the observed deviations
from European trends was most probably derived from
the fact that ringing data addressed different aspects of
a population than did breeding count data. The latter
consider only breeding pairs or territorial males, while
ringing data refer to the whole population, including
immature individuals, and thus also reflect reproductive
information for the population from the current year.
Additionally, the differences in trends could be
related to the low accuracy of overall European trend
estimates of some species, resulting from the rough
methodology of combining the results of existing
national schemes (van Strien et al. 2001). Both
datasets obtained from the large scale pan-European
breeding survey and from autumn bird ringing
activity include many bird populations. In this regard,
the trend estimates based on breeding surveys are
assessed artificially, by a compilation of trends from
different countries (van Strien et al. 2001). In contrast,
bird populations from bird ringing data are combined
naturally, resulting in assessment of the actual overall
trend of the European population, especially if the
breeding range of autumn migrating populations is
known for a certain area. Further studies would be
needed to compare trends assessed from autumn bird
ringing data for selected species from several countries
to verify the coherence of the ringing data and their
consistency with European census data.
When comparing trends obtained from breeding and
migratory population surveys we should consider
important distinctions between them: (i) data on
migratory populations are gathered from a wider
geographical area and (ii) in the autumn migratory
population, breeders, and non-breeders are combined,
each of which might show its own trend. The
generally accepted consensus that trend estimates
using bird ringing data coincide with trends obtained
by other methods (Berthold 2004) is especially true
when comparing trends from the autumn migration
period. At that time of the year, birds show more
flexible behaviour in selecting habitats (Mason 1979,
Moore et al. 1995, Parrish 2000) and are therefore
more likely to be caught. Moreover, such positive
correlations in trends obtained by different bird
12 T. PETRAS AND A. VREZEC
censuses and survey techniques are especially common
in those species that exhibit the same population
trends over a large area (Berthold 2004). We argue
that ringing data on migrants can provide more
accurate and representative trend estimates at the
continental scale, but have low power in assessing
population change at the local or regional scale
compared to the breeding population censuses.
In conclusion, when variability in ringing effort is
taken into account, long-term bird ringing data are a
valuable source of information for describing patterns
of long-term population dynamics. Firstly, there is great
potential for performing historical analyses when other
sources of data on birds are not available. Secondly, the
approach could serve as a beneficial supplement to
other bird monitoring programmes (Rintala et al. 2003).
For example, the monitoring of local breeding birds
could assess population dynamics on a limited number
of sites at a local or regional scale, while monitoring in
the migration period could provide population data
over extensive geographical areas, especially if ringing
recoveries reveal the geographic origin of the migrating
population. In future studies we should combine trend
estimates of autumn migratory populations across
Europe to fully understand the dynamics of migratory
birds and their cross-continental connectivity.
The main issue in trend assessments derived from
ringing data is variable ringing effort, which produces
data samples that are difficult to compare. Within
ringing sites, ringing effort, and sampling
methodology should be as consistent as possible,
particularly in regard to the length of mist-nets used,
their position, time of netting, and the use of playback
calls. The use of playback can significantly increase the
catching rate of some target species (Mukhin et al.
2008) and might greatly influence the unexplained
variability in ringing data if not used consistently,
despite it resulting in larger samples. When playback
is used, for example in monitoring of specific target
species, a strict protocol should be developed,
including type of recording, broadcast time, duration,
and loudness, to assure its consistent use and to avoid
data biases. On the other hand, for assessing trends
from long-term historical multi-species ringing data,
we recommend using a universal model approach
with data obtained from the whole autumn migration
period by selecting the most intensive ringing days
(i.e. ≥50 ringed birds and ≥5 ringed species) and by
using ringing days as denominator. However, attention
should be given to the significance of the trend
estimates, since in some species biases in the data
cannot be accounted for with this approach, as shown
in this study. Our models were derived from multi-
species ringing using mist-nets only, but in the future
trend assessments from multi-species historical data
based on other catching methods should be tested in
order to obtain patterns of population dynamics over
longer time periods that might reveal population
statuses of bird species over 100 or more years.
Acknowledgements
We are particularly grateful to the volunteer ringers from the
Slovenian Bird Ringing Centre (Slovenian Museum of
Natural History) who provided data on birds in the period
2000–2016: Dušan Belingar, Franc Bolta, Darjo Bon, Ivo
Božič, Franc Bračko, Igor Brajnik, Jože Bricelj, ml., Jože
Bricelj, st., Alfonz Colnar, Stane Černalogar, Marjan Debelič,
Dušan Dimnik, Jože Dolinšek, Stanko Drašček, Dare Fekonja,
Jernej Figelj, Jože Geiser, Marjan Gobec, Jože Gračner, Dejan
Grohar, Peter Grošelj, Vojko Havliček, Ludvik Jakopin,
Marko Jankovič, Tone Jankovič, Leon Kebe, Milovan Keber,
Brane Koren, Stane Kos, Jelko Kozjak, Brane Lapanja, Ivan
Lipar, Anton Lisec, Zvonko Lončarevič, Tone Macele, Tomaž
Mihelič, Jurij Mikuletič, Jože Nered, Žan Pečar, Miro
Perušek, Dušan Petkovšek, Rajko Piciga, Zdravko Podhraški,
Dušan Pogačar, Milan Pustoslemšek, Aljaž Rijavec, Miran
Romšak, Luka Simončič, Branko Slabanja, Andrej Sovinc,
Željko Šalamun, Dare Šere, Iztok Škornik, Pavle Štirn, Vlado
Štolfa, Polde Štricelj, Rudolf Tekavčič, Tomi Trilar, Andrej
Trontelj, Miro Vamberger, Lojza Vesel, Bogdan Vidic, Milan
Vogrin, Iztok Vreš, Davorin Vrhovnik, Eva Vukelič, and Ivan
Zlobko. Special thanks go to Dare Šere, who coordinated bird
ringing in Slovenia from 1987 to 2011, and to Dare Fekonja,
who has coordinated bird ringing in Slovenia since 2012. We
are very grateful to Przemysław Busse for helpful discussions
about the evaluation of bird ringing data. Dare Šere, Tomi
Trilar, and Brane Lapanja kindly provided useful information
on bird ringing methodology. We would like to thank Andrej
Kapla for map preparation. We acknowledge using data
source on European trends of common birds by EBCC/
BirdLife/RSPB/CSO.
Funding
Bird ringing in Slovenia is financially supported by the
Ministry of Culture of the Republic of Slovenia. The study is
a part of TP’s PhD work, which is financially supported by
the Ministry of Education, Science and Sport. AV was
supported by the Ministry of Culture of the Republic of
Slovenia and research core funding No. P1-0255 by the
Slovenian Research Agency.
ORCID
Al Vrezec http://orcid.org/0000-0002-4699-6451
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