Popul Ecol
DOI 10.1007/s10144-016-0563-0
ORIGINAL ARTICLE
Numerical responses of an avian predator to prey fluctuations
in a temperate latitude: breeders vs. entire population
Marek Panek1
Received: 18 February 2016 / Accepted: 21 October 2016
Ó The Society of Population Ecology and Springer Japan 2016
Abstract Distinct numerical responses of predators to
fluctuations in the abundance of their prey are often
observed in northern regions but occur more rarely in
temperate latitudes. This statement is, however, mostly
based on observations of breeding populations, while in
some predators, for example in raptors, numerous non-
breeding floaters can occur. I estimated the breeding den-
sity and reproductive performance (nest survey) as well as
the density of entire population (transects with distance
sampling) of the common buzzard Buteo buteo in western
Poland (52°N) in the years 2005–2014 to test the
hypotheses that in temperate latitudes: (1) the breeding
population of these birds does not show any numerical
response, understood as annual changes in their abundance;
(2) its reproductive success, however, changes with the
abundance of main prey, the common vole Microtus
arvalis; and (3) hence the entire buzzard population (in-
cluding potential immature floaters) present in a given area
during the nesting period responds numerically with some
time delay. The reproductive success of buzzards was
positively correlated with their prey abundance. Contrary
to my predictions, however, the breeding population of
buzzards showed a slight numerical response with a 3-year
lag and the entire population tracked vole fluctuations
without any time delay. The immediate numerical response
of the entire buzzard population was probably caused by
large-scale movements of floaters. Such rapid numerical
responses of some predators may contribute to the relative
& Marek Panek
m.panek@pzlow.pl
1
Polish Hunting Association, Research Station, Sokolnicza 12,
64-020 Czempiń, Poland
stability of prey populations in temperate latitudes com-
pared to northern regions.
Keywords Common buzzard  Common vole  Non-
breeding floaters  Poland  Population dynamics
Introduction
Predators can respond functionally and numerically to fluc-
tuations in the availability of their food resources (Solomon
1949; Angelstam et al. 1984). The numerical response often
consists of changes in reproduction or mortality rates in
accordance with fluctuations in the abundance of main prey,
which means that the number of adult predators present in an
area at a given time can change in relation to prey abundance
but with some time delay. Such delayed numerical responses
have been described most often in resident mammalian
predators (e.g., Korpimäki et al. 1991; O’Donoghue et al.
1997; Angerbjörn et al. 1999), but were also found among
raptors, for example in goshawks (Tornberg et al. 2005) and
gyrfalcons (Nielsen 1999). In these species high reproduc-
tive success was observed in years with peak abundances of
their fluctuating prey, but as their young reach reproductive
maturity only after a few years, the breeding populations of
these raptors track the changes in prey abundance with 2- or
3-year lags, respectively. In nomadic predators, thus in some
other raptors, numerical responses without a time delay can
occur, resulting not from variable reproduction or mortality
rates, and consisting of large-scale movements to areas
abundant in prey (e.g., Korpimäki and Norrdahl 1991; Kor-
pimäki 1994; Salamolard et al. 2000).
Distinct fluctuations in prey abundance and related
numerical responses of predators often occur in the
northern regions of Eurasia and North America (Hanski
123

et al. 1991, 2001; Korpimäki et al. 2005). In contrast, such
phenomena have been observed less often or on a smaller
scale in temperate latitudes, where mainly generalist
predators live and which have at their disposal numerous
alternative prey species. They therefore respond mainly
functionally (i.e., by switching between prey) and thus their
populations are relatively stable (Korpimäki 1986; Hanski
et al. 1991; Turchin and Hanski 1997; Korpimäki et al.
2005). Clear numerical responses, however, have some-
times been observed in temperate regions, for example in
the Montagu’s harrier in France (Salamolard et al. 2000).
On the other hand, the numerical responses of raptors
are typically described on the basis of data about their
breeding populations. At the same time the local densities
of these birds can also depend on the presence of non-
breeding ‘‘floaters’’ (Newton 1979; Rohner 1996; Kenward
et al. 2000). The fraction of floaters potentially includes
both immature young individuals and adults without mates
or breeding territories (Penteriani et al. 2011). In the case
of the common buzzard (Buteo buteo Linnaeus, 1758), i.e.,
the most abundant raptor species in Europe (Newton 1979),
a clear numerical response of the nesting fraction was
noted in Fennoscandia (Reif et al. 2004). Although an
effect of food richness on the reproductive success of
buzzards has sometimes been found in temperate regions in
the continent of Europe, clear and regular fluctuations in
their breeding populations were not usually observed there
(Kostrzewa and Kostrzewa 1990; Je˛drzejewski et al. 1994;
Goszczyński 1997; Selås 2001; Sim et al. 2001). On the
other hand, the presence of numerous floaters has been
detected in this species (Kenward et al. 2000). In this paper
I therefore test hypotheses resulting from the knowledge
described above, predicting that in temperate latitudes: (1)
the breeding populations of common buzzards does not
reveal any numerical response, understood as annual
changes in their abundance; (2) however, these birds show
some demographic response, i.e., their reproductive suc-
cess changes with the availability of their main prey; (3)
hence the entire buzzard population present in a given area
during the nesting period (i.e., territorial birds and imma-
ture floaters together) responds numerically with some time
delay.
Methods
I carried out the study in a 75 km2 area located near
Czempiń, south of Poznań, in western Poland (52°090 N,
16°450 E), in a region dominated by farmland with frag-
mented forests covering 20% of its area. The study site
consisted mainly of arable land (80%). Several small for-
ests (35–270 ha) and patches of woodland (\1–15 ha)
covered only 10% of this area. Individual crop fields were
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Popul Ecol
relatively large in some places as they amounted to
10–100 ha (60% of agricultural land), but in other places
they usually did not exceed 5 ha (40%). The main crops
were cereals (58%); other cultivated plants included oil-
seed rape, maize, sugar beets, potatoes and alfalfa. Con-
siderable annual fluctuations in the abundance of common
voles (Microtus arvalis Pallas, 1779) took place in the
study area, and high vole densities occurred especially in
large plantations of alfalfa and oilseed rape (Ryszkowski
1982; Panek and Hušek 2014). According to an ornitho-
logical review from the end of the 20th century, the com-
mon buzzard was a widespread and fairly common raptor
in this region of Poland, and its breeding density ranged
there from 0.5 to 5 pairs per 10 km2 depending on location
(Bednorz et al. 2000).
The occurrence of breeding pairs of common buzzards
in the study area and their reproductive success was esti-
mated in the years 2005–2014 using the methodology
commonly utilised in research on birds of prey (e.g.,
Newton 1979; Je˛drzejewska and Je˛drzejewski 1998; Panek
and Hušek 2014). In March and April, forests and patches
of woodland were searched for buzzard nests. Located
nests were checked once every 2 weeks from a distance of
several dozen metres using binoculars. It was assumed that
in a given nest a breeding attempt occurred if in April or
May (i.e., during the incubation period) a buzzard sitting
on the nest was observed at least once. A breeding attempt
was considered as successful when at least one fully
feathered offspring was observed on the nest before the
anticipated fledging time (typically in the second half of
June). The number of fledglings was also estimated in this
period. Annual values of several demographic variables
were calculated on the basis of the above field data: the
density of breeding pairs, the proportion of pairs with
reproductive success (annual sample: 25–31 pairs), the
mean number of fledglings per successful nest (annual
sample: 17–27 nests), the production of fledglings per
nesting pair and the production of fledglings in the whole
study area.
The index of the entire buzzard population present in
the study area during the nesting season was estimated in
the middle of June. By this time both members of nesting
pairs have already finished watching offspring at nests and
they hunted in their territories, but young birds still stayed
in nests. Adult buzzards were counted on line transects
with distance sampling (Buckland et al. 1993). This
method has previously been used successfully in the case
of common buzzards (Kenward et al. 2000). Six perma-
nent transect routes totalling 86 km ran along field and
forest roads, and were evenly distributed over the study
area. The routes were traversed by foot during the
morning hours and all sites of buzzard observations were
marked on maps (1:10,000) based on available landmarks.
Popul Ecol

Their perpendicular distances were then calculated from Results

these maps (to the nearest 10 m). Annual samples of
recorded individuals ranged from 28 to 70. The index of The density of the buzzard breeding population in the years
buzzard density (with confidence intervals, CI) was esti- 2005–2014 changed from 3.3 to 4.1 pairs per 10 km2
mated using the Distance 6.0 software (Thomas et al. (mean 3.8, n = 10, SD = 0.3; Fig. 1a). Reproductive
2009). success was achieved by 63–89% of pairs attempting
In central Europe the common vole is the main prey of nesting in a given year (mean 75.9, n = 10, SD = 10.0);
common buzzards in the areas dominated by agriculture successful pairs raised 1.4–1.9 fledglings (mean 1.6,
(Mebs 1964; Goszczyński and Piłatowski 1986; Je˛drze- n = 10, SD = 0.2); annual production per nesting pair
jewska and Je˛drzejewski 1998; Šotnár and Obuch 2009). In amounted to 0.9–1.7 fledglings (mean 1.2, n = 10,
the study area, common voles constituted 72% of prey SD = 0.3); and annual production per 10 km2 was equal to
items in the spring-summer buzzard diet evaluated during 3.4–6.9 fledglings (mean 4.6, n = 10, SD = 1.1). The
the period of this study (M. Panek, unpublished data). I index of entire population density in individual years
estimated the abundance of common voles by counting changed from 4.6 to 14.1 individuals per 10 km2 (mean
their burrow entrances (Mackin-Rogalska et al. 1986). The 8.0, n = 10, SD = 3.0; Fig. 1b). No correlation was found
counts were carried out in March each year between 2002 between the density of breeders and the density of the
and 2014. Six permanent transect routes totalling 66 km in entire buzzard population (r = –0.024, P = 0.9, n = 10).
length were evenly distributed throughout the agricultural The mean number of vole burrow entrances in the years
land. Only the entrances with clear signs of use (fresh 2002–2014 ranged from 1.1 (n = 6, SE = 0.2) to 25.6
digging, droppings, pieces of food) located within 3 m of (n = 6, SE = 3.2) per kilometre of transect routes and the
both sides of the transect routes were recorded. Due to a
curvilinear relationship between the density of burrow
a
entrances and the population density of common voles
(Mackin-Rogalska et al. 1986), the annual mean numbers
of burrow entrances per 1 km of transect routes were
logarithmically transformed and then used as an index of
vole abundance.
The relationship between the demographic variables of
buzzards and the index of vole abundance was examined
by Pearson’s correlation analysis. To check whether the
density of breeders and the density of the entire buzzard
population was dependent on the abundance of voles with
or without a time lag, cross-correlations were performed
with vole data both from the same year and from 1 to
3 years ago. The effect of breeding performance in pre-
vious years on the two buzzard density estimates was b
analysed using the annual number of fledglings in the
whole study area. The initiation of breeding in these birds
occurs most often in the third year of their life, but can be
earlier (Newton 1979; Davis and Davis 1992; Kenward
et al. 2000). I therefore calculated correlations between
the two density estimates and the fledgling production
observed 1, 2 and 3 years ago as well as the accumulated
number of fledglings in the three previous years. During
my analyses, several correlations were sometimes per-
formed on the same data set. Therefore, the Bonferroni
correction was used, i.e., when a given correlation coef-
ficient was found to be statistically significant taking into
account the standard significance level (a = 0.05), I
checked whether its P value was equal to or lower than
Fig. 1 The various density estimates of common buzzards: a breeding
the corrected level (0.05/n, where n is the number of
pairs (nest survey), b entire population (an index estimated by line
calculated correlations). Positive results are marked with transects, thus including potential non-breeders; with ±95% CI) in
‘‘*’’ after P. western Poland in the years 2005–2014

123
 Popul Ecol

b
Fig. 2 The index of vole abundance in early spring (the mean number
of common vole burrow entrances per kilometre of transects,
transformed logarithmically) in western Poland in the years 2002–2014

logarithmic index of vole abundance ranged from 0.09 to

3.24. Three peak periods and two lows occurred in the
study years (Fig. 2).
The index of vole abundance was positively related to
the proportion of buzzard pairs with reproductive success
in a given year (r = 0.690, P = 0.03, n = 10), the number
of fledglings per successful pair (r = 0.741, P = 0.01*,
n = 10), and the annual production of fledglings per
nesting pair (r = 0.759, P = 0.01*, n = 10).
When the vole abundance index was correlated with the
two buzzard density estimates (i.e., breeders and entire
population) in the same year and with 1–3 years delay, no Fig. 3 Relationships between the index of vole abundance and the
two density estimates of common buzzards: a breeding pairs (nest
significant relationship was found in the case of current survey) in the same year and with a 3-year delay, b entire population
breeding density, but some response of the breeding frac- in the same year (an index estimated by line transects, thus including
tion (insignificant with the Bonferroni correction) was potential non-breeders) in western Poland in the years 2005–2014.
found for 3-year delay (Table 1; Fig. 3a). However, the Regression lines for significant relationships are showed, but in a the
regression line is only for breeding pairs with a 3-year delay
index of entire population density was significantly affec-
ted by vole availability in the same year (Table 1; Fig. 3b).
The density of the buzzard breeding population was not study area in the previous year (r = -0.065, P = 0.9, n = 9),
significantly correlated with the fledgling production in the 2 years ago (r = -0.420, P = 0.3, n = 8), 3 years ago
(r = 0.042, P = 0.9, n = 7), or the total production in the
Table 1 Correlations (coefficients r and their P values) between the
previous 3 years (r = 0.212, P = 0.6, n = 7). Similarly, the
two density estimates of common buzzards, i.e., breeding pairs (nest
survey) and entire population (an index estimated by line transects, index of the entire buzzard population density was not sig-
thus including potential non-breeders), and the index of vole abun- nificantly related to the fledging production in the previous
dance in the same year (0) and with 1–3 years delay in western year (r = -0.140, P = 0.7, n = 9), 2 years ago (r = 0.352,
Poland (n = 10 in all cases)
P = 0.4, n = 8), 3 years ago (r = 0.193, P = 0.7, n = 7), or
Time delay Buzzard density estimate the accumulated production in the previous 3 years
Breeders Entire population (r = 0.207, P = 0.7, n = 7).

Year 0 r = -0.206, P = 0.6 r = 0.745, P = 0.01* Discussion

Year -1 r = -0.036, P = 0.9 r = 0.414, P = 0.2
Year -2 r = 0.295, P = 0.4 r = 0.286, P = 0.4 In accordance with my predictions, common buzzards from
Year -3 r = 0.641, P = 0.046 r = -0.147, P = 0.7 western Poland responded demographically to fluctuations
* Significant also after the Bonferroni correction for multiple in the abundance of their main prey in terms of changes in
comparisons their reproductive success. Similar relationships were

123
Popul Ecol
found during a number of previous studies on this species
(e.g., Newton 1979; Je˛drzejewski et al. 1994; Goszczyński
1997; Selås 2001; Sim et al. 2001), although on occasion
its breeding performance did not change with fluctuations
in the availability of main prey (Krüger 2004; Reif et al.
2004). In this study I measured the food abundance in the
early spring, i.e., 2–3 months before the period of feeding
nestlings by buzzards. The used index of vole density,
however, apparently also described sufficiently well the
availability of this prey in the late spring. Newton (1979)
stated that food abundance can affect the course and results
of breeding in raptors to a large extent by the effects on the
condition of females at the beginning of the breeding
season, and thus on the size of clutches.
Contrary to my predictions, the breeding population of
buzzards most probably showed a slight numerical
response with a 3-year lag in relation to the fluctuations of
their main prey in the study area located in a temperate
latitude. This delay corresponds to the typical age of
maturation and the first instance of breeding in the common
buzzard (Newton 1979; Davis and Davis 1992; Kenward
et al. 2000). It suggests that the observed numerical
response of the breeding population resulted from the
changes of reproductive success in the preceding period. In
fact, the breeding density was correlated neither with the
fledgling production 3 years earlier nor with the total
production during the three previous years, but this may be
associated with the smaller sample size (the number of
years with available data) than in the case of other
analyses.
My estimations of breeding buzzard densities and entire-
population density indices were based on different methods
and could contain some biases. For example, in the years
with limited vole availability the density of the entire
population was often lower than the density of nesting
buzzards (i.e., doubled pair density; Fig. 1). This might
potentially mean that in the years with scant food unsuc-
cessful birds left their breeding territories, and thus the
study area, in search of more favourable feeding condi-
tions. On the other hand, the observed relationships
between the above mentioned density values in individual
years may be only accidental. It therefore seems inappro-
priate to directly compare the density estimations obtained
by the two different methods. However, it is unlikely that
the observed changes in vole abundance affected the
accuracy of these two buzzard density estimations for two
reasons. First, because the breeding density did not vary
with the fluctuations in vole abundance, and second,
because the distance sampling method used to estimate the
index of entire population density is based on functions
which model the probability of detecting animals (Buck-
land et al. 1993). This means that potential effects of vole
fluctuations on the detection of buzzards in the field (e.g.,
by changes in used habitats or secretive behaviour) are
already taken into account. Therefore, the comparison of
relationships between the two density estimates and the
index of vole abundance seems appropriate.
However, contrary to the breeding population, the den-
sity of entire buzzard population present in the study area
during the nesting period clearly tracked the fluctuations of
vole abundance without any time-lag. Thus, this finding
was also inconsistent with my predictions. Such rapid
numerical response occurs in some raptor species but
consists of long-range movements to regions with peak
numbers of their fluctuating prey to perform reproduction
in good feeding conditions (Korpimäki 1985, 1994; Kor-
pimäki and Norrdahl 1991; Salamolard et al. 2000; Millon
and Bretagnolle 2008). In western Poland, the fluctuations
of vole availability undoubtedly affected the density of
non-breeding buzzards, as the density of nesting birds did
not show significant relationships with the current level of
main prey abundance. The fraction of non-breeders might
potentially include both territorial birds without clutches
(and even pairs after early nest losses or desertions, and
thus omitted during the survey of nesting pairs), local
young birds from previous years and alien floaters. How-
ever, it is unlikely that the number of non-breeders among
territorial buzzards (if such birds were present in the study
area) was positively related to the abundance of voles, and
furthermore, the density of entire buzzard population was
not dependent on the production of fledglings in this pop-
ulation in the previous years. This suggests that the
immediate numerical response of the entire buzzard pop-
ulation consisted mainly of the movements of floaters.
The fidelity of breeding buzzards to their territories in
the study area remains unknown. In the temperate regions
of Europe these birds tend to stay in their territories
throughout the year and typically do not move further than
2 km (Weir and Picozzi 1975; Kenward et al. 2001;
Schindler et al. 2012), although as close as in north-eastern
Poland a substantial part of nesting buzzards leave their
territories at the beginning of winter and migrate different
distances (Je˛drzejewska and Je˛drzejewski 1998). On the
other hand, young common buzzards typically take natal
dispersal and usually move a distance of a few to several
tens of kilometres (Davis and Davis 1992; Walls and
Kenward 1998; Kenward et al. 2001). After some period of
movements they usually settle in a range and only make
short excursions, but do not continue to wander; however,
sometimes they can move again, typically in spring, to
settle in a new area (Walls and Kenward 1998; Kenward
et al. 2001). Juvenile and non-territorial individuals, both in
the common buzzard (Kenward et al. 2001; Schindler et al.
2012) and in other bird species (Penteriani et al. 2011),
typically stay in areas providing rich food (but not suit-
able nesting sites as in the case of breeders). Such habits of
123

non-breeding buzzards could result in the immediate
numerical response observed in western Poland.
The oscillations of small rodent populations occurring in
northern latitudes are characterised by considerable ampli-
tudes and synchronization over large areas, while the fluctu-
ations of these animals observed in temperate latitudes have
relatively slight amplitudes and are synchronized over small
areas (Korpimäki et al. 2005). This local scale of the changes
in prey availability in temperate latitudes may mean that even
less mobile species are able to track these changes directly by
movements. In the same area in western Poland red foxes also
responded numerically without any time lag to the changes of
vole abundance (M. Panek and R. Kamieniarz, unpublished
data). Such rapid tracking prey fluctuations show a stabilizing
effect on the populations of this prey in contrast to the delayed
response (Korpimäki 1985; Hanski et al. 1991, 2001). The
factors causing geographic gradient in rodent oscillations, i.e.,
their disappearance from north to south, include the increase
in the occurrence of generalist predators, the availability of
alternative prey species and higher habitat heterogeneity in the
more southern regions (Korpimäki 1986; Hanski et al.
1991, 2001; Turchin and Hanski 1997). Immediate numerical
responses of some predators to local changes in the abundance
of small rodents, such as those found in western Poland, can be
another phenomenon associated with the relative stability of
prey populations in temperate latitudes. Moreover, this study
confirmed the importance of non-breeders in population
dynamics of some raptors and other bird species (Kenward
et al. 2000; Penteriani et al. 2011) because taking into account
this fraction completely changed the observed pattern of
numerical response showed by the buzzard population.
Acknowledgements I am grateful to my colleagues from the
Research Station PHA in Czempiń for their help in data collection,
and to Javier Balbontı́n, Robert Kamieniarz, Hannu Pietiäinen and
anonymous reviewer for their valuable comments to the manuscript. I
also thank Ian Hatcher for language improvements.
Compliance with ethical standards
Conflict of interest The author declares that he has no conflict of
interest.
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