JOURNAL OF

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AVIAN BIOLOGY
Article
Heavy and persistent rainfall leads to brood reduction and nest
failure in a passerine bird

Eva Maria Schöll and Sabine Marlene Hille

E. M. Schöll (https://orcid.org/0000-0003-3096-5885) ✉ (eva.schoell@boku.ac.at) and S. M. Hille (https://orcid.org/0000-0002-3680-7549), Inst. of

Wildlife Biology and Game Management, Univ. of Natural Resources and Life Sciences, Vienna, Vienna, Austria.

Journal of Avian Biology The process of reproduction is a sensitive period for bird offspring, since they are
00: 1–11, 2020 exposed to environmental conditions for several weeks. Within the long reproductive
2020: e02418
doi: 10.1111/jav.02418 period, adult birds do not only have to maintain their own body condition, they also
have to ensure nutritional needs of their nestlings. In the worst case, if weather condi-
Subject Editor: Christiaan Both tions are very harsh, breeding is not successful and fails. This suggests that weather
Editor-in-Chief: Thomas Alerstam conditions might be an important driver of breeding success. Here, we studied the
Accepted 23 April 2020 effect of weather conditions (temperature and precipitation) on the survival of 350
nests of great tits Parus major during nestling period in six years (2010–2015). We
used traditional generalized linear mixed model (binomial response variable) and a
dedicated survival-analysis program (MARK) to analyze the data. The two approaches
allowed us to highlight and compare the effect of weather conditions on a fine (nest-
ling survival) and coarse (nest fate) scale. Both methods showed that precipitation
explained most of the variation in individual nestling and overall nest survival during
the 15-day nestling period. Heavy and persistent rainfall did not only lead to brood
reduction, it ultimately also led to losses of the entire brood. Likely causes were the
negative effects of precipitation on thermoregulation, food availability and predation
risk. However, while reduced food availability most likely might have led to brood
reduction through selective individual nestling death, predation might have resulted
in total nest failures.

Keywords: MARK, nest survival, Parus major, precipitation, temperature

Introduction
Life history of an animal is explained by traits that on one hand affect reproduction,
like clutch size, number of broods per season, age at first reproduction and lifespan,
and on the other hand affect survival. Both reproduction and survival highly affect
population growth, but survival seems to be the strongest to model demographic pro-
cesses (Schmidt et al. 2005, Arlt et al. 2008, Rose et al. 2019), as animals can only
contribute to the local population if they survive until they start to reproduce (Stearns
1992). In birds, a physiologically demanding and critical timeframe of survival until
first reproduction is the period between hatching and fledging (Ricklefs 1969).
––––––––––––––––––––––––––––––––––––––––
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www.avianbiology.org Nordic Society Oikos
This is an open access article under the terms of the Creative Commons
Attribution-NonCommercial-NoDerivs License, which permits use and
distribution in any medium, provided the original work is properly cited, the 1
use is non-commercial and no modifications or adaptations are made

At this stage of development, several variables can severely
affect nestlings, but abiotic conditions (e.g. local weather con-
ditions) and their temporal variability are assumed to have a
major effect on nestling survival and therefore on reproduc-
tive success (O’Connor and Morgan 1982, Morrison and
Bolger 2002, Slobodník et al. 2013).
As climate change scenarios are forecasting changes in
mean temperature and precipitation (IPCC 2013), studies
assessing detailed information about the nature of climate and
its impact on animal populations are of high interest (Knape
and de Valpine 2011, Matter and Roland 2017). However,
while reference periods to calculate climatological averages
cover at least 30 years (World Meteorological Organization
2017), yearly changing weather conditions are assumed to
affect demographic parameters of local populations more
directly (Skagen and Adams 2012). Near- and long-term pro-
jections forecast that daily ambient temperature will become
more extreme and heavy precipitation events will become
more frequent in temperate regions (Stocker et al. 2013), but
little knowledge exists to which degree these extreme condi-
tions affect bird species (Martin et al. 2017).
While adult birds and offspring of precocial birds are usu-
ally well-suited and prepared for unfavorable temperature
and rainfall conditions, nestlings of altricial birds are most
vulnerable to weather conditions, because newly hatched
young lack highly insulating down feathers and relatively
waterproof plumage. Further, nestlings cannot regulate their
own body temperature in the first few days after hatching
(Kluijver 1951). Open nests are more affected by adverse
weather conditions than cavity nests (Mainwaring et al.
2014). Nevertheless, temperature and rainfall indirectly
affect survival of cavity-nesting birds (Bordjan and Tome
2014). Parental nest visitation rates decrease with increas-
ing amount and persistence of rainfall (Öberg et al. 2015);
thus, rainfall may negatively affect nestling growth (Keller
and van Noordwijk 1994). Foraging success might be driven
by weather conditions (Newton 1998), because prey activity
is reduced during cold and rainy periods (Avery and Krebs
1984). In addition, caterpillar availability is reduced dur-
ing periods with harsh weather (Schöll et al. 2016). Thus,
regional weather conditions can have a major effect on local
population dynamics, if survival of individual nestlings is
reduced or the entire brood is lost (Öberg et al. 2015).
Estimates of nest survival provide a sensitive metric to
study the effects of differences among environmental condi-
tions (Dinsmore et al. 2002, Rotella et al. 2007), whereas
estimates of individual nestlings in a nest are a useful measure-
ment of annual reproductive success (Armstrong et al. 2002).
Therefore, here, we analyzed firstly, daily nest survival rates of
great tit Parus major within the 15-day nestling period and
secondly, variation in individual nestling survival through-
out three consecutive 5-day intervals, using data on breeding
performance gathered within the 6-year study period. The
great tit is an excellent species for studying nest fate, since
it is breeding in nest boxes, which are easily monitored. We
used two methods to analyze variation in survival data by
incorporating variables and factors of ecological relevance
2
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(e.g. weather conditions): a dedicated survival-analysis pro-
gram (MARK, White and Burnham 1999) and traditional
generalized linear mixed model. Using a large sample size
of 350 nests, we analyzed nest fate across the 15-day nest-
ling period and divided the nestling period in three 5-day
intervals to study nestling survival. While nest survival analy-
ses only account for the effect of local weather conditions
on complete nest fates, the effect of temperature and rain-
fall could also be less dramatic, leading to brood reduction
through selective individual nestling death rather than to the
loss of the entire brood. Therefore, we assume that the results
of both survival analyses will be complementary rather than
contradictory.
Material and methods
Study area
Field work on great tits was carried out in a mixed deciduous
forest located in lower Austria (48°05′03″N, 15°55′24″E).
The vegetation is dominated by beech Fagus sylvatica and
acer Acer sp., along with some conifers (spruce Picea abies, fir
Abies alba, larch Larix decidua). Nest boxes for great tits were
provided within the study area (19.5 ha) along an altitudinal
gradient ranging from 491 to 887 m above sea level. Nest
boxes were monitored between March and July in the years
2010–2015. The number of boxes increased throughout the
six study years from 211 (in 2010) to 286 (in 2015). We
therefore operated with the occupancy rate of nest boxes by
great tits that varied between 28.0 and 48.4%.
Data collection – breeding data
Within our long-term study, nest boxes were frequently mon-
itored during nest building and egg-laying period every two
to three days. Once a clutch was completed and eggs in the
nest were warm, nests were not checked for the following 13
days to avoid disturbing the incubating female (mean incu-
bation period length in our study site: 12 days). Afterwards,
nests were daily revisited until the last nestling had hatched.
Once all nestlings had hatched, the brood was assumed to
be completed (day 1). In some rare cases, when nests were
checked 13 days after the assumed incubation start, the last
nestling had already hatched. In these cases, age of nestlings
could be readily and precisely determined (based on a com-
prehensive study made in that area in the year 2009, master
thesis project of J. Fingerlos).
We monitored a total of 408 great tit nests in the years
2010–2015. Since great tits are multiple brooded species and
nesting success differs in first and second broods (Kluijver
1951), we included only first broods and nests that started
within 30 days after the first egg was laid in each year in the
population in our analyses (Both et al. 2004, Both and Visser
2005). Thus, we had to exclude 51 nests because the first egg
in these nests was laid 30 days after the first egg had been laid
in the specific year. In addition, we had to exclude 7 nests,

because we did not know exact date of clutch initiation. The
fate of the nests was monitored on a regular basis every five
days (day 5, 10 and 15) after the last nestling had hatched,
but some nests failed before day 15. Therefore, sample sizes
for the two model approaches differ.
Weather data
Daily weather conditions were obtained from a weather
station, which was located within the study site (Office of
the Federal Government of Lower Austria, Hydrology and
Geoinformation, official webpage: <www.noe.gv.at>). Sum
of precipitation (mm) was measured every 15 min in the years
2011–2015, whereas in the year 2010, only data of daily sum
of precipitation were available. We extracted four precipita-
tion metrics to comprehensively study the effect of rainfall.
Daily sum of precipitation (mm) was measured (Bordjan and
Tome 2014) and number of days with rainfall (> 0 mm),
strong rainfall (> 10 mm, Skagen and Adams 2012) and
heavy rainfall (> 20 mm, Öberg et al. 2015) within the nest-
ling period were counted. To assess if the amount of precipi-
tation during the six study years was usual or extraordinary
for the study site, we calculated the 30-year average of precip-
itation measured at three meteorological stations surround-
ing the study area (distance 24–32 km) and compared these
data (Auer et al. 2012) to the data measured within the study
site in the years 2010–2015.
Ambient temperature (°C) was recorded every 15 min at
the weather station. To verify, if temperature data collected
at the meteorological station represent temperature within
the study site, ambient temperature was measured at 40
nest boxes along the altitudinal gradient in the year 2015.
Between 24-Apr and 29-May-2015, data on ambient tem-
perature was collected for periods between 10 and 16 consec-
utive days (using iButtons: every 3 min, resolution: 0.5°C).
Temperature buttons were fixed underneath nest boxes to
avoid direct solar irradiation at any daytime. Daily minimum
temperatures were extracted for a period of 24 h. These daily
minimum temperatures measured along the altitudinal gradi-
ent were then compared with daily minimum temperatures
measured at the meteorological station for the same periods
(Wilcoxon rank sum test with Bonferroni correction). We did
not find statistical differences between the temperature mea-
surements. Therefore, daily minimum temperature measured
at the weather station seemed to represent these temperature
measurements within the study site adequately.
Modeling of nest survival using RMark
During the last decades, nest survival has been usually esti-
mated based on the Mayfield method, which incorporates
exposure days to account for lacking data if parts of the stud-
ied nests were not monitored from the beginning of nest
initiation (Mayfield 1961, 1975). In addition, this method
accounts for irregular visits to nests, by assuming that nest fail-
ures occurred midway between the last two visits. Especially
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in natural systems, where nests are usually not checked on
a regular basis, this kind of analyses is highly advantageous.
While this advanced method has greatly improved the esti-
mation of nest survival itself, the approach assumes that
daily survival probability is constant within nesting stages
and among all nests, which is often biologically restrictive
or unrealistic (reviewed by of Jehle et al. 2004). Moreover,
the method does not allow incorporating covariates that may
explain variation in nest survival. Dinsmore et al. (2002)
therefore developed an extension of the Mayfield method
using the program MARK (White and Burnham 1999). The
method includes all the advancements of the Mayfield esti-
mation, but it also accounts for daily variation in survival
rates as a function of daily varying environmental conditions.
Therefore, daily survival of great tit nests during the 15-day
nesting period was modelled using the programs MARK
and R statistics (< www.r-project.org >). In addition, the R
package ‘RMark’ (Laake 2018) was used, which provides an
interface to use the software MARK. The terms ‘nest survival’
and ‘nest success’ are used in various ways, thus, we chose
the following definitions for the purpose of this paper. ‘Nest
survival’ is the probability that a nest will be successful within
the period between hatching and the following consecutive
15 days (nestling period). A nest was considered successful,
when a) at least one fully feathered young was observed 15
days after hatching in the nest (time of fledging), or b) the
nest was undisturbed at the time of fledging, but nestlings
had already fledged, or c) we heard fledglings nearby the nest
box 15 days after hatching. Nests were considered to have
failed, when a) we detected clear evidence of predation e.g.
destroyed nest material, injuries in dead nestlings or adults
indicating attacks by other animals, or b) nest were deserted
due to an unknown cause and chicks had starved. ‘Daily nest
survival (DNS) rate’ is the probability that a nest will survive
a single day within this period.
To model nest survival during the 15-day nestling period
four parameters were required: a) the first day the nest was
checked during the nestling period; b) the last day the nest
content was checked being alive; c) the last day the nest was
checked during the nestling period and d) the fate of the nest
(binary outcome: successful/failed) (Dinsmore et al. 2002).
These parameters allow the estimation of nest survival prob-
abilities, even if nests were not monitored from the beginning
of the nestling period or were controlled at various irregular
intervals. The program MARK uses this information to gen-
erate an appropriate encounter history (live–dead format) for
each nest covering the 15-day nestling period. Since we do
not know the exact day a nest failed, MARK models the fail-
ure of the nest during the interval from the last day the nest
content was checked being alive until the last day the nest was
checked during the nestling period (Dinsmore and Dinsmore
2007, Rotella 2018). We followed the approach described by
Dinsmore and Dinsmore (2007) and standardized the begin-
ning of the nestling period among years (22-Apr). We con-
verted all calendar dates to numerical dates and numbered
all nest-check dates sequentially thereafter. We defined the
3

maximal period great tit nests were active, which ranged from
22 April until 19 June (59 days), regardless of year. Since
nest age can affect daily survival rates (Grant et al. 2005), we
aimed to most precisely age the nests. When the last nestling
of a clutch had hatched, we aged this nest as nest age = 1. In
most cases, the nest was first controlled within the nestling
period, when only some of the nestlings in a nest had already
hatched. Thus, we checked these nests on a daily routine and
when the last nestling of a nest had finally hatched; we aged
this nest by back-calculating when the nest was checked the
first time during the nestling period using negative numbers
(nest age < 1). Only in some rare cases (n = 29), nests were
visited the first time within the nestling period when the last
nestling just had hatched some days before (nest age > 1).
To avoid any effect of these delayed nest checks on our nest
survival model, completion days for these 29 nests were put
in the analyses rather than actual check dates (Rotella 2018).
We included 58 individual covariates for each year,
accounting for daily weather conditions (minimum daily
temperature 24 h, daily sum of precipitation 24 h) and nest
age. These time-varying covariates corresponded to a unique
daily value for each nest and were included by building a
design matrix to model daily nest survival as a function of
weather conditions and nest age. Finally, DNS estimates of
the best model were calculated for each study year separately.
To compare apparent fledgling success of the six study years,
we raised the estimates of DNS to the power of 15 (DNS15),
since nests were monitored during the nestling period from
the hatching of the whole brood until day 15 after hatch. In
total, a series of 13 candidate models with different combina-
tions of the individual covariates was built (Supplementary
material Appendix 1 Table A1). Similar to the procedure
performed by Dinsmore et al. (2002) or Dinsmore and
Dinsmore (2007), we used a hierarchical-modeling approach
to keep the model set small, since the number of covariates
was large. First, we performed a set of six models containing
a constant, linear and quadratic time trend for daily survival
rate within and between years (Model 1–6). These models
should reflect time dependent survival rates (Grant et al.
2005, Berkunsky et al. 2016). Then, we used the model which
explained temporal variations within and between years best,
and added the individual covariates altitude (Slobodník et al.
2013), nest age (Grant et al. 2005) and number of hatchlings
(Mertens 1969) separately and additively (Model 7–10).
Finally, we included the weather covariates by adding each
one separately and also the interaction terms of temperature
and precipitation (Model 11–13) to the best model from the
set of models we had run before. Models were ranked using
Akaike’s information criterion corrected for small sample size
(AICC) to determine the best model (Burnham and Anderson
2002).
Modeling of nestling survival
To test if weather conditions affect not only nest sur-
vival during the nestling period, but also survival of
4
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individual nestlings, we used generalized linear mixed mod-
els (GLMMs) with a binomial distribution and a logit link
(package ‘lme4’, Bates et al. 2014). We created three sepa-
rate models, each lasting for five days during the nestling
period: day 1–5, day 6–10, day 11–15, where day 1 again
indicated the day, when a clutch was completely hatched.
We used the cbind() binomial approach for the response
variable, where we included the number of nestlings at the
start and end of each 5-day period (day 1/day 5, day 5/day
10, day 10/day 15). Note, that the sample size used for ana-
lysing nestling survival using GLMMs is lower than for the
nest survival analyses using RMark, because we could only
use nests, which were exactly visited on day 1, 5, 10 and
15. In addition, since some nests already failed within the
first days after complete hatching, the sample size decreased
throughout the nestling period.
We used a stepwise approach for model selection for each
5-day period. First, we identified, which precipitation param-
eter explained most variation in nestling survival (step 1;
Öberg et al. 2015). Each model included one of the precipi-
tation metrics, calculated for the respectively 5-day interval
within the nestling period (sum of precipitation (Bordjan and
Tome 2014), number of days with precipitation, number of
days with strong precipitation > 10 mm (Skagen and Adams
2012), number of days with heavy precipitation > 20 mm
(Öberg et al. 2015)), and a nested random effect (study year,
nestbox ID). The four preliminary models were ranked by
Akaike’s information criterion (AICc) to determine the model
with the best fit for the next step in the analyses.
In a second step, additional variables were included after
testing for multi-collinearity. Variance inflation factors
were below 2 and indicated no problems with collinearity
(Zuur et al. 2009). Thus, we added the variables number of
nestlings (on day 1, 5, 10 for the corresponding period), alti-
tude (of the nest box), Julian day (clutch completely hatched,
day 1 = 1 Apr) and average daily minimum ambient tem-
perature (for the same 5-day period as for the precipitation
variable) to the best model of step 1 (global model). We did
not include an interaction term between temperature and
precipitation, because Pearson’s correlation coefficient never
exceeded 0.3. Since predictor variables had very different
scales, we rescaled them by generating standardized scores
(z-score, subtracting the variable mean and dividing by the
variable standard deviation) to avoid problems with model
convergence. Then, this global model was used to generate a
set of models with all possible combinations of fixed effects
(R package ‘MuMIn’, Bartoń 2018). The models were graded
according to their AICc, and model-averaged coefficients for
a subset of models (ΔAICc < 2) were extracted. Since Akaike
weights of the best model was below 0.9 (Results, Table 3)
and therefore high model selection uncertainty existed, full-
model averaging was used (Grueber et al. 2011, Symonds and
Moussalli 2011). Upper and lower bounds of the 95% confi-
dence intervals were calculated for each parameter. Statistical
analyses were conducted using the software R 3.4.1 (< www.r-
project.org >).
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Table 1. Weather conditions measured at the meteorological station between 22 Apr and 19 June and the percentage of successful nests in
the six study years. 30-year average (1981–2010) of precipitation measured between April and June exhibited 231 ml.
Ambient temperature (°C) Daily precipitation (mm) Successful nests
Study year Min Max Mean (± SE) Min Max Mean (± SE) Sum (%)
2010 0.4 29.0 12.1 (0.08) 0 40.7 6.1 (1.27) 362.53 56.8
2011 −1.3 24.1 13.2 (0.07) 0 45.6 3.7 (1.21) 217.50 80.4
2012 0.7 27.0 13.4 (0.09) 0 21.4 2.7 (0.63) 160.50 84.5
2013 1.7 28.5 12.6 (0.08) 0 47.9 4.9 (1.28) 288.00 67.4
2014 0.3 29.3 12.7 (0.08) 0 98.6 5.3 (1.88) 313.10 63.6
2015 2.1 27.8 13.2 (0.08) 0 60.0 3.8 (1.33) 222.40 69.0

Results Modeling daily nest survival

Breeding data
Within the six study years, out of the 350 nests, 252 nests
were successful and 98 nests failed before nestlings were ready
to fledge. Nearly 42% of nest failure was due to abandon-
ment and 58% to nest predation. Compared to the other
study years, in 2012 a high number of the provided nest
boxes was occupied by great tits; almost twice as many nests
as monitored in the other five study years (Supplementary
material Appendix 1 Table A2). In total, 1640 nestlings sur-
vived until day 15 after hatch. Mean number of hatchlings
per nest was lowest in the year 2012 (Kruskal–Wallis rank
sum test, Chi-squared = 49.42, df = 5, p < 0.001, followed
by Wilcoxon rank sum tests with Bonferroni adjustment,
p < 0.05, see Supplementary material Appendix 1 Table A2),
whereas the number of hatchlings per nest did not differ in
the other years (Wilcoxon rank sum tests with Bonferroni
adjustment, p > 0.05). Although brood size (mean number
of nestlings) was low in the year 2012, the high occupancy
rate (almost 50% of nest boxes were occupied) and high over-
all nest survival (85% successful nests) led to a high num-
ber of fledglings in the year 2012 (Supplementary material
Appendix 1 Table A2).
Within the set of six preliminary candidate models, which
included constant, linear and quadratic time trends and
therefore allowed for temporal variation within and between
years, the model including a linear time trend and study year
received stronger support than the other models. Adding the
individual covariates altitude, nest age or number of nestlings
when the nest was first checked during the nestling period
did not substantially improve the model fit. However, the fit
of the model was improved when we added weather condi-
tions to this preselected model (Table 2).
The final best model including a linear time trend, study
year and daily sum of precipitation [confidence intervals
CI = −0.047, −0.023] carried the most support (wi = 65%,
Table 2). Daily sum of precipitation had a negative effect on
nest survival (βP = −0.04 [−0.047, −0.023], Fig. 1), whereas
the confidence interval for the effect of the linear time trend
included zero (βL = −0.02 [−0.057, 0.010]). We also ran a
post-hoc model including both a linear and a quadratic
effect of rainfall (βP = −0.03 [−0.077, −0.001], βPP = 0.00004
[−0.0004, 0.0005]), since weather conditions could have a
non-linear influence on nest survival (Mattsson and Cooper
2009, Post van der Burg et al. 2010), but it did not improve
the model fit (AICc = 688.80).

Table 2. Model selection table for great tit nest survival in the six
Weather conditions
study years ranked by ascending ΔAICc using the software MARK.
Variables included: linear time trend (L), quadratic time trend (Q),
Weather conditions in the nestling period between 22-Apr constant time trend (null model, .), nest age, altitude (A), number of
and 19-June varied within and between years (Table 1). hatchlings (NN), daily sum of precipitation (P), minimum daily tem-
Daily sum of precipitation varied between 0 and 98.6 ml perature (T).
(mean ± SE = 4.4 ± 0.54 ml) and ambient temperature
recorded every 15 min at the weather station through- Model Deviance K AICC ΔAICc wi
out the nestling period varied between −1.3 and 29.3°C SL +Y + P 670.80 8 686.83 0.00 0.65
(mean ± SE = 12.9 ± 0.03) in the study period. Average sum SL+Y+P×T 668.03 10 688.08 1.25 0.35
of precipitation measured between April and June in the SL+Y+T 685.24 8 701.27 14.44 0.00
SL+Y 689.63 7 703.66 16.83 0.00
30-year period between 1981 and 2010 exhibited 231 ml. SL+Y+NEST AGE 688.16 8 704.19 17.36 0.00
Compared to this prolonged period, we conducted our study SL+Y+A 689.26 8 705.29 18.46 0.00
during two wet springs (years 2010, 2014), one dry spring SL+Y+NN 689.46 8 705.49 18.66 0.00
(year 2012) and three intermediate springs (years 2011, SQ+Y 692.61 7 706.64 19.81 0.00
2013, 2015). The rainiest and coldest spring in our study S(.)+Y 695.42 6 707.44 20.61 0.00
period was spring 2010. In this year, only slightly more than SL 703.93 2 707.93 21.10 0.00
half of the nests were successful. In contrast, spring 2012 SL+Y+A+NEST AGE+NN 687.90 10 707.95 21.12 0.00
was the warmest and driest spring and nest survival reached SQ 706.26 2 710.26 23.43 0.00
S(.) 711.01 1 713.01 26.18 0.00
almost 85% (Table 1).

5

Figure 1. Effect of daily sum of precipitation (mm) on daily nest
survival rate (± 95% CI) of great tits in Austria (2010–2015). For
computing estimates of real parameters, the effect of the linear time
trend was fixed (Table 2). Each point in the figure refers to a single
great tit nest (n = 350).
One additional model shared a substantial weight of evi-
dence in the confidence set, a model including next to the
linear time trend and study year also an interaction between
precipitation and daily minimum ambient temperature mea-
sured over 24 h (wi = 35%). However, the confidence interval
for the effect of the interaction term of precipitation and tem-
perature included zero (model SL+Y+P×T: βP×T = 0.005 [−0.002,
0.012]). Annual average DNS during the nestling period
varied between 0.975 and 0.984 and apparent nestling sur-
vival within the 15-day period between hatching and fledging
ranged between 57.9 and 78.9% (Supplementary material
Appendix 1 Fig. A1).
Modeling nestling survival
For modeling early nestling survival (day 1–5, n = 348), the
subset of models with ΔAICc < 2 encompassed nine mod-
els (Table 3). The final model describing nestling survival
between day 1 and 5 included the variables number of heavy
rain days (> 20 mm), number of nestlings, average daily min-
imum temperature, altitude and Julian days. Nestling sur-
vival within the first five days after clutches were completely
hatched decreased with increasing number of days with heavy
rainfall (> 20 mm). Confidence intervals for the parameter
estimates of number of nestlings, temperature, altitude and
Julian days included zero, indicating little evidence that these
variables affected nestling survival within the first 5 days after
hatching (Table 4).
The subset of models explaining variation in nestling sur-
vival between day 6 and 10 (n = 317) encompassed six mod-
els (Table 3). The final models with ΔAICc < 2 included the
parameters number of heavy rain days (> 20 mm), Julian
days, average daily minimum temperature, altitude and
6
1600048x, 2020, 7, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/jav.02418 by Univ de Buenos Aires, Wiley Online Library on [03/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
number of nestlings. Increasing number of days with heavy
rainfall negatively affected nestling survival. Confidence
intervals for the parameters Julian days, temperature, altitude
and number of nestlings indicated little evidence of affecting
nestling survival (Table 4).
For modeling nestling survival between day 11 and 15
(n = 276) the subset encompassed five models (Table 3). All
final models describing variation in nestling survival included
the variables number of days with heavy rainfall (> 20 mm)
and Julian days. In addition, some models included the vari-
ables average daily minimum temperature, altitude and num-
ber of nestlings on day 10 (Table 4). Number of days with
heavy precipitation and Julian days (after clutches completely
hatched) negatively affected nestling survival. The variables
average daily minimum temperature, altitude and number of
nestlings had no effect on nestling survival between day 11
and 15 (confidence intervals included zero).
Discussion
Our results indicated that local weather conditions explained
most variation in daily survival rate of great tit nestlings; in
particular precipitation was a dominant driver of reproduc-
tive success. Precipitation led to brood reduction through
selective individual nestling death, but it ultimately also had
a negative effect on the entire brood. Further variables, which
were included in the analyses of nest survival (altitude, nest
age, number of nestlings) did not explain variation in nest
fate. Clear evidence for the effect of ambient temperature
on overall nest fates or survival of individual nestlings could
not be found (Bordjan and Tome 2014), but birds seem to
respond to year-to-year changes in climatic conditions in an
ecological time frame (Skagen and Adams 2012); and as a
consequence the probability that nestlings survive the period
between hatching and fledging varied in the six study years.
Analyses of nest fates using the program MARK and tradi-
tional approaches analysing nestling survival using GLMMs
showed similar results, thereby showing that both analyses
can complement and support each other. Thus, analysing sur-
vival on a coarse level can already provide good insights in
the complex interactions and processes of how environmen-
tal conditions affect nest survival. The nest survival analysis
using MARK is highly advantageous especially for studies in
natural systems, where survival is not checked on a regular
basis. We assume that precipitation caused variation in indi-
vidual nestling survival and nest fate: 1) directly, as it might
have affected thermoregulation and 2) indirectly, through its
effect on food availability (Schöll et al. 2016) and predation
risk. However, while reduced food availability most likely
might have led to brood reduction through selective individ-
ual nestling death (Siikamäki 1996), predation might have
caused the loss of the entire brood leading to nest failures.
Nestlings cannot maintain their own body temperature
during the first days after hatch (Mertens 1977, Whittow
2000). Feathers are not yet fully developed and nestlings
easily suffer immediate physical damage during this period
 1600048x, 2020, 7, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/jav.02418 by Univ de Buenos Aires, Wiley Online Library on [03/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Table 3. Subsets of generalized linear mixed effect models (ΔAICc < 2) explaining the variation in nestling survival in great tits based on a
multi-model inference approach. Variables: number of heavy rain days (n heavy rain), number of nestlings (n nestlings), average daily mini-
mum temperature (temp), Julian days clutch completely hatched (J days), altitude.
Response Model subsets AICc ΔAICc AICc weight
Nestling survival day 1–5 = Intercept + n heavy rain + n nestlings 614.4 0.0 0.197
= Intercept + n heavy rain 614.8 0.4 0.167
= Intercept + n heavy rain + n nestlings + temp 615.2 0.8 0.133
= Intercept + n heavy rain + n nestlings + J days 615.5 1.1 0.117
= Intercept + n heavy rain + temp 616.2 1.8 0.080
= Intercept + n heavy rain + n nestlings + altitude 616.3 1.9 0.079
= Intercept 616.4 2.0 0.076
= Intercept + n heavy rain + n nestlings + temp + J days 616.4 2.0 0.075
= Intercept + n heavy rain + J days 616.4 2.0 0.075
Nestling survival day 6–10 = Intercept + n heavy rain + J days 669.6 0.0 0.304
= Intercept + n heavy rain + J days + temp 670.7 1.1 0.180
= Intercept + n heavy rain 671.1 1.5 0.147
= Intercept + n heavy rain + J days + altitude 671.3 1.7 0.130
= Intercept + n heavy rain + J days + n nestlings 671.5 1.9 0.121
= Intercept + n heavy rain + temp 671.5 1.9 0.119
Nestling survival day 11–15 = Intercept + n heavy rain + J days 742.3 0.0 0.318
= Intercept + n heavy rain + J days + temp 743.0 0.7 0.220
= Intercept + n heavy rain + J days + altitude 743.1 0.8 0.214
= Intercept + n heavy rain + J days + n nestlings 744.1 1.8 0.130
= Intercept + n heavy rain + J days + temp + altitude 744.2 1.9 0.119

(Newton 1998). To reduce heat loss during cold and rainy
periods, nestlings can clump and roost together, thus, brood
size can affect energy expenditure for heat production and
therefore nestling survival (Mertens 1969). However, survival
of individual nestlings was not related to the number of sib-
lings in our study site. Temporal patterns (e.g. nestling age)
seem to affect nest survival (Grant et al. 2005, Boulton et al.
2008), because shortly after hatching young nestlings are
incapable of thermoregulation (Whittow 2000) and rely on
females to keep them warm (Kluijver 1951). If females spent
more time within nest boxes, they have to reduce their food
provisioning behavior (Radford et al. 2001, Öberg et al. 2015)
and might face a trade-off between brooding nestlings and
feeding themselves (Bordjan and Tome 2014). Ultimately,
adult birds might be forced to abandon nests (Bordjan and
Tome 2014) to cover their own energy demands, if meta-
bolic costs of searching for food for nestlings are too high.
Within our long-term study, 11.7% of all nests were aban-
doned between 2010 and 2015. When nestlings grow older,
gain body weight and change to an endothermic metabolism
(Mertens 1977), females can slowly reduce their brooding
behavior and increase time spent searching for food (Kluijver

Table 4. Model-averaged summary statistics of generalized linear mixed effect models: estimates of coefficients and unconditional standard
error (SE), lower and upper limits of the 95% confidence intervals (CI 95%) and relative importance values of the model parameters.
Important parameters affecting response variables are highlighted in bold. Variables: number of heavy rain days (n heavy rain), number of
nestlings (n nestlings), average daily minimum temperature (temp), Julian days clutch completely hatched (J days), altitude.
Response Variables Estimate SE CI 95% Relative importance
Nestling survival day 1–5 (Intercept) 5.536 0.607 4.347; 6.725
n heavy rain −0.432 0.219 −0.861; −0.004 1.00
n nestlings 0.373 0.223 −0.065; 0.811 0.60
temp 0.232 0.229 −0.218; 0.681 0.29
altitude −0.106 0.214 −0.526; 0.314 0.27
J days −0.207 0.232 −0.662; 0.248 0.08
Nestling survival day 6–10 (Intercept) 4.768 0.466 3.855; 5.681
n heavy rain −0.616 0.205 −1.017; −0.214 1.00
J days −0.393 0.218 −0.820; 0.033 0.73
temp −0.244 0.220 −0.676; 0.188 0.30
altitude 0.132 0.214 −0.288; 0.552 0.13
n nestlings −0.109 0.224 −0.549; 0.330 0.12
Nestling survival day 11–15 (Intercept) 2.943 0.196 2.558; 3.327
n heavy rain −0.535 0.136 −0.801; −0.268 1.00
J days −0.359 0.155 −0.663; −0.055 1.00
temp 0.179 0.165 −0.145; 0.502 0.34
altitude 0.155 0.147 −0.134; 0.443 0.33
n nestlings 0.079 0.145 −0.204; 0.363 0.13

7

1951). Adult birds and nestlings also suffer large heat loss if
they are exposed to rainfall or get wet. Short periods and low
intensity of rain might be less problematic, since plumage of
adult birds is at least temporarily rainproof (Kennedy 1970)
and nest boxes can shield drizzle. However, heavy precipita-
tion events might have negatively affected nestling survival,
if the nesting material got soaked (Wesolowski et al. 2002).
In addition, if adult birds released moisture within nest boxes
during food provisioning, nestlings can rapidly lose body
mass (Whitehouse et al. 2013) during rainy periods, because
energy demands to maintain body temperature constant
increase if birds’ plumage is wet (reviewed by Kennedy 1970,
Wilson et al. 2004).
Nestlings of many bird species and great tits are mainly
fed with invertebrate prey (Naef-Daenzer and Keller 1999,
Naef-Daenzer et al. 2000). Caterpillars, e.g. of winter moth
Operophtera brumata, are one of the main food sources of
great tit nestlings, especially in study sites, where they occur
only during a short annual peak (Naef-Daenzer and Keller
1999). In our study site, caterpillars are available throughout
the great tit breeding season (Schöll et al. 2016), but nestling
survival between day 11 and 15 after hatch was negatively
associated with the Julian day of hatching (clutch completely
hatched, day 1 = 1 Apr).
Since insect development itself is highly dependent on
temperature and rainfall (Price 1997), weather conditions
should also have an indirect effect on individual nestling
survival. During rainy periods, caterpillars are less active
(Tamburini et al. 2013) and might be difficult to detect by
great tits. Caterpillar availability is reduced during harsh
weather conditions (Schöll et al. 2016), because heavy and
persistent rainfall increases mortality of caterpillars (Dennis
and Sparks 2007). As food availability is one factor limit-
ing nestling survival (Boulton et al. 2008), optimal weather
conditions in the year 2012 might have been advantageous
for caterpillar survival and development (Newton 1998).
In contrast, low food availability during years with harsh
weather conditions might have negatively affected nestling
survival, because parents could not provide enough food for
the whole brood, or might have been forced to abandon nests
to increase their own chances of survival. Moreover, adult
birds seek shelter during strong rainfall events and reduce
feeding rates (Radford et al. 2001, Öberg et al. 2015, but see
Geiser et al. 2008). As adult birds might not completely com-
pensate for periods of rainfall and lack of food by increasing
feeding frequency after rainfall has stopped (Radford et al.
2001), individual nestlings can either die from hypothermia
or starvation, if they do not receive enough food to meet the
demands of heat production, which increase during low tem-
perature periods (Newton 1998).
Bordjan and Tome (2014) have also studied the effects of
weather conditions on nest survival of great tits. They found
indications that rainfall might cause nest abandonment, and
reasoned that food-shortages during cold and wet periods had
affected nest fate. However, in many passerine species, nest
predation is one of the most important reasons for complete
8
1600048x, 2020, 7, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/jav.02418 by Univ de Buenos Aires, Wiley Online Library on [03/03/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
nest failure and therefore, analyses of nest survival should also
include predated nests (but see Bordjan and Tome 2014).
Since food availability is affected by weather conditions and
reduced during periods with harsh weather (Schöll et al.
2016), an increased begging behavior of hungry nestlings
(Kölliker et al. 1998) might have attracted predators (Leech
and Leonard 1997, Fogarty et al. 2017). In addition, as nest-
ing season advances and nestlings mature, nestlings require
more food (García-Navas and Sanz 2011). Since parental
activity at the nest increases, because more frequent nest
visits are necessary to provide food for nestlings, the prob-
ability of predation due to diurnal predators increases as well
(Martin et al. 2000). Indeed, predation occurred in 16.3%
of all studied nests (n = 350) throughout the 15-day nestling
period in our study site. In contrast to studies from Lithuania
(Juškaitis 2006), Germany (Koppmann-Rumpf et al. 2003)
or Czech Republic (Adamík and Král 2008), where only
9.6%, 8.4% and respectively 3.2% of great tit nests were
destroyed in the prolonged period between egg-laying and
fledging, we showed that predation had a huge effect on nest
survival. However, compared to a study of great tits nesting in
natural tree cavities in Poland, where predation accounted for
66.7% of nest failures (Maziarz et al. 2015), predation risk
was lower for our great tit population breeding in nest boxes
(accounted for 58% of nest failures in our study site, see also
Purcell et al. 1997, but see Mitrus 2003).
One of the most important predators in our study site,
the edible dormouse Glis glis, is mainly active between sunset
and sunrise (Rodolfi 1994). It does not only depredate eggs
and nestlings; it can also predate adult birds resting within
nest boxes (Adamík and Král 2008). Until now, it is not clear,
whether dormice find occupied nest boxes just by chance due
to a overlap in timing of breeding seasons, or whether they
actively search for occupied nest boxes (Adamík and Král
2008). Although great tits might be able to defend cavities
against intruders (Adamík and Král 2008), they might prob-
ably abandon nests after a dormouse tried to enter, because
of high predation risk. It is assumed, that local abundance
of predators increases as the breeding season progresses and
therefore, nest survival declines throughout the nesting sea-
son (Grant et al. 2005). However, during the last decades,
activity seasons of the edible dormouse might have shifted
into early spring and is assumed to overlap with the nesting
season of great tits (Koppmann-Rumpf et al. 2003, but see
Bieber et al. 2018).
Our study showed that local weather conditions in general,
and heavy and persistent rainfall in particular had a strong
negative effect on nestling survival. Among all years, mean
number of hatchlings per nest box was the lowest in spring
2012. In this year, almost 50% of the available nest boxes were
occupied, and the high nest box breeding density might have
led to high intraspecific competition. Thus, a strong density-
dependent effect might have resulted in the low average num-
ber of hatchlings in the year 2012 (Van Balen 1973, Dhondt
2010). However, in contrast to the other study years, the
high overall nest survival (85% successful nests) during the

nestling period finally resulted in a high number of fledglings
in this year. Spring 2012 was the driest spring in our study
site and these conditions seemed to increase the probability
of nestling survival until fledging, compared to wet years,
where number of hatchlings was higher, but nest survival was
reduced and finally fewer nestlings had fledged. In contrast
to our study, extremely harsh weather conditions occurred
during spring 2012 in eastern England and affected breeding
success negatively (Whitehouse et al. 2013). Thus, studying
geographic variation of weather patterns seems important to
model animal population trends and will be of high ecological
and conservation interest, since the Intergovernmental Panel
on Climate Change (IPCC) forecasts an increasing number
of heavy rain events in future (IPCC 2013). Ambient tem-
perature and precipitation do not only affect survival of nest-
lings, they can have a complex interacting impact on different
age classes, e.g. recruitment of fledglings (Öberg et al. 2015,
Gyurácz et al. 2016) or parental survival (Öberg et al. 2015).
Acknowledgements – Thanks to M. Polo Aparisi, M. de los Milagros
Alvarez Martinez and Raimund Barth for help with fieldwork.
Further, we thank two anonymous reviewers for their helpful
comments on the original draft of the manuscript.
Funding – This study was funded by the Federal Ministry for
Sustainability and Tourism, Austria; Wolfgang Pauli Fellowship,
Univ. of Vienna and the Austrian Federal Forests.
Author contributions – The first author collected data in the field,
analyzed the data, performed statistical analysis and wrote the
manuscript. The second author originally developed methodology
and supervised the project.
Conflicts of interest – There is no conflict of interest.
Permit(s) – Permission to handle our study animals was given by the
Office of the Provincial Government of Lower Austria. The Austrian
Federal Forests gave their permission to work on their land.
Transparent Peer Review
The peer review history for this article is available at https://
publons.com/publon/10.1111/jav.02418
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