Agriculture, Ecosystems and Environment 216 (2016) 44–50

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Agriculture, Ecosystems and Environment

journal homepage: www.elsevier.com/locate/agee

Lost colonies found in a data mine: Global honey trade but not pests or
pesticides as a major cause of regional honeybee colony declines
Robin F.A. Moritza,b,c,* , Silvio Erlera
a
Institut für Biologie, Molekulare Ökologie, Martin-Luther-Universität Halle-Wittenberg, Hoher Weg 4, 06099 Halle (Saale), Germany
b
Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa
c
German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany

A R T I C L E I N F O A B S T R A C T

Article history: Recent losses of honeybee (Apis mellifera) colonies have been linked to several non-exclusive factors; such
Received 12 June 2015 as pests, parasites, pesticides (e.g., neonicotinoids) and other toxins. Whereas these losses pose a threat
Received in revised form 23 September 2015 to apiculture, the number of globally managed colonies appeared to be less affected because beekeepers
Accepted 25 September 2015
replace lost colonies. From a socioeconomic and ecological perspective the number of managed colonies
Available online xxx
is arguably more relevant when addressing the issue of apiculture and pollination services provided by
honeybees. We here use the FAO data base to dissect the interactions between the global honey market
Keywords:
and the number of colonies. Global scale analyses do not show a general colony decline. Whereas Western
Socioeconomics
Honey
Europe and the US show suffer colony declines, other regions show considerable increase. We could not
Global trade find any link between the colony dynamics and the occurrence of specific pathogens or the use of
Colony losses pesticides. In contrast, changes in the political and socioeconomic system show strong effects on
Beekeeping apiculture. In addition, many countries show a tight negative correlation between honey import and the
number of managed colonies. For some countries, the amount of honey produced per colony is highly
positively correlated with the amount of honey imports, and we cannot exclude large scale relabeling of
imported to nationally produced honey. It is very clear that honey trade is a dominating factor for the
number of managed colonies since countries with a strong import and export ratio are those suffering
most strongly from colony declines.
ã 2015 Elsevier B.V. All rights reserved.

1. Introduction climate change, landscape alteration, agricultural intensification

The past decade has seen a multitude of dramatic reports on
honeybee (Apis mellifera) declines that raised great public and
societal concern. Indeed the value of honeybees for human society
has been estimated to exceed 153 billion (Gallai et al., 2008)
primarily as key pollinators of many crops. The function as
generalist pollinators also puts them into a key position for any
ecosystem functioning that arguably exceeds any conceivable
pecuniary value. A global decline of honeybee colonies would be of
particular concern, because of large scale declines of wild
pollinators (Biesmeijer et al., 2006) and a global loss of pollinators
would clearly have profound and devastating impact on life on
earth, a sinister scenario that sparked a huge debate in the public
media. Pathogens, pesticides, and their interactions, but also
* Corresponding author at: Institut für Biologie, Molekulare Ökologie, Martin-
Luther-Universität Halle-Wittenberg, Hoher Weg 4, 06099 Halle (Saale), Germany.
Fax: +49 345 5527264.
E-mail address: robin.moritz@zoologie.uni-halle.de (R.F.A. Moritz).
and non-native species (González-Varo et al., 2013) have been
attributed to have substantially contributed to the losses of
honeybee colonies (Kluser et al., 2010; Le Conte et al., 2012; Martin
et al., 2012; Potts et al., 2010a). However, historically reports about
regional large scale losses of honeybee colonies are both recurrent
and frequent, with dramatic events dating back to medieval times
(Fleming, 1871). Today mass losses can be typically linked to
diseases or regional poisoning (Pistorius et al., 2009). A prominent
exception has been the so called colony collapse disorder (CCD)
which killed millions of colonies in the US (vanEngelsdorp et al.,
2009). In spite of considerable research efforts (Cox-Foster et al.,
2007; Stokstad 2007) CCD could not be clearly associated with a
specific pathogen (Anderson and East 2008) or poisoning. As a
consequence interactions among pests, pathogens and pesticides
were suspected to have caused these massive colony deaths
(vanEngelsdorp and Meixner, 2010; Nguyen et al., 2010; Potts et al.,
2010a). Recently, neonicotinoid pesticides have been suggested to
play a potential role in these colony losses (Johnson et al., 2010;
Johnson, 2015) which led to a ban of these compounds by the
European Commission (2013). However, several recent reviews

http://dx.doi.org/10.1016/j.agee.2015.09.027
0167-8809/ã 2015 Elsevier B.V. All rights reserved.
 R.F.A. Moritz, S. Erler / Agriculture, Ecosystems and Environment 216 (2016) 44–50
show that the documentation of colony losses can be regional
extremely variable. Potts et al. (2010b) showed in their pan-
European study, including data between 1965 and 2005, the losses
rarely exceeded 30% at the national scale. Although this may be
high from an apicultural perspective, one should keep in mind that
this may well be in the range of sustainable mortality rates
expected in the wild if a queen lives for three years and issues a
reproductive swarm every year.
In general however, although the loss of colonies can be
disastrous for the individual apicultural operation, the number of
lost colonies is much less relevant than the number of existing
colonies from a societal or ecological perspective. For Europe the
densities of managed honeybee colonies have recently been
reported to have fallen below one colony per square kilometre
(Chauzat et al., 2013) which is more than an order of magnitude
less that the density of wild honeybees in the African Kalahari
desert in regions without any beekeeping and rather unfavourable
environmental conditions for honeybees (Jaffé et al., 2010). For
pollination services the number of bees, both wild and managed,
available in the blooming season are essential, not those that died
after or before the season. The number of honeybee colonies lost
are of course most important for apiculture because the bee-
keepers need to take measures to compensate for the losses.
Honeybee colonies are typically lost over winter (Genersch et al.,
2010), and this places the beekeeper into a pivotal position for
honeybee conservation. It is apicultural routine to replace lost
colonies by splitting one colony into two for swarm prevention and
to compensate for the winter losses (Ambrose, 1992). This also
allows for a vivid market for honeybee colonies in many countries
that is contributing towards the compensation of regional and local
colony losses. Yet whereas the number of colony losses are often
poorly documented and require the combination of diverse sources
of different quality (e.g., Potts et al., 2010b), the numbers of
managed colonies have been officially documented for 100
countries since 1961 in the global database managed by the FAO
(Food and Agriculture Organization of the United Nations).
Whereas, the absolute data may be difficult to compare among
countries due to differences in national statistical procedures, and
some obvious inconsistencies are rather apparent as we will show
later, the relative changes of the long term trends may be
nevertheless be informative. This data has been used before to
document the decline of honeybee colonies in Europe and the US
over the past between 1961 and 2009 in spite of a global increase of
colony numbers (Aizen and Harder, 2009a; vanEngelsdorp and
Meixner, 2010; Moritz et al., 2010). However, the data also showed
that the global increase of colonies is insufficient to compensate for
the even higher demand for pollination which by far exceeded the
availability of honeybee colonies (Aizen and Harder 2009a; Breeze
et al., 2014). In addition to colony numbers, the data gathered by
the FAO also includes the national production, import, and export
of marketed honey of the various nations. The combination of
these various sources of information related to honeybees and
their products might not only reveal imprecisions in the data sets
but also allow for the detection of the major patterns and
eventually allow for interpretations in search of the causal reasons
of a potential colony decline.
2. Material and methods
We used the data sets for colony numbers (1961–2013),
national honey production (tonnes, 1961–2013), import and export
of marketed honey (tonnes, 1961–2011) from the FAO database
(FAOSTAT (http://faostat.fao.org). For some countries data sets
were fused to have a complete data set thru 2013: (1) former Soviet
Union (URS = Armenia, Azerbaijan, Belarus, Estonia, Georgia,
Kazakhstan, Kyrgyzstan, Latvia, Lithuania, Republic of Moldova,
45
Russian Federation, Tajikistan, Turkmenistan, Ukraine, Uzbeki-
stan), (2) former Yugoslavia (YUG = Bosnia and Herzegovina,
Croatia, Montenegro, Serbia, Serbia and Montenegro, Slovenia,
the former Yugoslav Republic of Macedonia), (3) former Czecho-
slovakia (TCH = Czech Republic and Slovakia), and (4) Belgium-
Luxembourg (BEL). Data analysis and statistics (comprising
correlation and GLM) were done using simple spreadsheet
software and STATISTICA 8.0 (StatSoft, Tulsa, OK). Values are
given as means and standard errors. We use the data set to test if
colony numbers change in response the arrival of the major
honeybee pest, the parasitic mite Varroa destructor, and the
introduction of neonicotinoid pesticides. We compare those
changes with the effects of socioeconomic transformations known
to have had major impact on national economies. We test if the
amount of honey produced per colony is plausible, given the
climatic conditions of the various countries in question. We finally
test if honey trade may have any impact on the number of managed
colonies in those countries for which all four metrics are available:
colony number, national honey production, honey import and
honey export.
3. Results and discussion
3.1. The FAO database—colony numbers
More than 100 countries provided data to the FAO-database and
globally there is a significant trend of colony increase of more than
60% over the past 50 years from 1961–2013 (Fig. 1). Clearly, the
globally collected data does not support the notion of a global
colony decline. However, the variability among the reporting
countries is huge, ranging from declines down to one forth to more
than the four fold increase of the colony number as of 1961. At a
continental scale the most dramatic decline occurred in Europe
between 1989 and 1995 where almost seven million colonies
disappeared within only five years (Fig. 1). This loss in managed
colony numbers coincided with the collapse of the socialistic
regimes in the Eastern European countries after 1989 (Aizen and
Harder 2009a; Moritz et al., 2010; Smith et al., 2013). The
association of the colony decline with the political changes in East
and Central Europe is very robust since the change in the number
of colonies among the various countries shows a highly significant
correlation (mean r = 0.79  0.04, n = 7; TCH and HUN excluded,
as no significant colony decline was observed during this period) in
the five years before and after the societal transitions in Eastern
Europe and the Balkan (Fig. 2). Comparing the impact of each
country and the years of colony decline in a general linear model in
the Eastern European and Balkan data set, the factor ‘year’ was
much more important than ‘country’, with both significantly
influencing colony declines (GLM for normalized data (as of 1984
for each country), with country as random and year as fixed
factors; country: F = 8.96, dF = 6, P = 0.000001; year: F = 14.68,
dF = 10, P < 0.000001). Thus although there was considerable
national variability among the different Eastern European and
Balkan countries, the temporal dynamics over time were highly
similar. This also shows that the FAO data base is in principle
suitable to detect major impacts on the number of colonies. In
contrast, the arrival of the parasitic mite V. destructor in the early
eighties (Potts et al., 2010b; Rosenkranz et al., 2010) had no
detectable effect on the number of managed colonies in the full
European data set. On the contrary, the number of colonies in
Europe increased from 21.4 Mio to 22.4 Mio between 1980 and
1990 (Fig. 1). So, although V. destructor has been clearly identified
as the major pest and foremost factor in colony losses (Genersch
et al., 2010; Potts et al., 2010a), it did not affect the number of
managed colonies reported to the FAO at all. Thus to be explicit:
this does not show that Varroa does not kill honeybee colonies, it
46 R.F.A. Moritz, S. Erler / Agriculture, Ecosystems and Environment 216 (2016) 44–50

Fig. 1. Relative changes in total colony numbers, as compared to 1961, summarized for Europe (empty circles), Southern Europe (filled circles) and whole world (grey circles).

shows how beekeepers efficiently adopt their operations to comply
with the challenges set by pests even as lethal as V. destructor. It
also makes it unlikely that other less lethal pests and pathogens
ever had a major impact on the number of managed colonies at the
global scale.
In addition to the dramatic declines in colony numbers after the
societal transition in Eastern Europe, Western Europe has seen a
less dramatic but constant and highly significant decline of
colonies in the past three decades. At the average there was a 1%
decline per year (b = 0.012, r2 = 0.95, P < 0.001, n = 20 years 1993–
2013; countries: Austria, Belgium, Switzerland, Germany, France,
Liechtenstein, and Luxemburg). This is nearly five times higher as
the decline observed in the US in the same period which suffers
from a constant decline of about 0.21% per year in the past two
decades (b = 0.002, r2 = 0.26, P < 0.001, n = 20 years 1993–2013).
Colony numbers in other regions of the world show a
fundamentally different development. The number of managed
colonies more than doubled over the past 50 years at a rate of
2.5% per year in Southern Europe (b = 0.02, r2 = 0.96, P < 0.001)
(Fig. 1). An even steeper increase can be observed for South
America (5.2  0.20), Africa (3.3  0.11) and Asia (4.4  0.07).
Extreme examples of colony increase in the past five decades
include Myanmar (157-fold), Pakistan (7-fold), Senegal (35-fold),
Syria (11-fold), Uruguay (9-fold) and Vietnam (17-fold).
The few atypical declines were again associated with grave
socioeconomic changes. These include:
1) A dramatic decline of 66% of the colonies in Madagascar after
1977, subsequent to the political coup from which apiculture
never recovered.
2) A 73% decline in Burundi during the civil war, a loss which has
however rapidly overcompensated during the past decade with
three times as many colonies today as in 1961.
3) A similar effect can be seen in Zambia, where the country
recovered after the implementation of multi-party democracy

Fig. 2. Average numbers of managed colonies as derived from the GLM analysis (black filled circles) before and after the collapse of the socialistic regimes in East and Central
Europe. Grey circles and lines show the values for the individual countries. (ALB—Albania, BLG—Bulgaria, DEU—Germany, POL—Poland, ROM—Romania, URS—Union of Soviet
Socialist Republics, YUG—Yugoslavia).
 R.F.A. Moritz, S. Erler / Agriculture, Ecosystems and Environment 216 (2016) 44–50
from more than two decades of decline of about 66% during the
rule of the one-party politics.
4) After the oil boom in Venezuela in the late 1970s–early 1980s
the number of managed colonies dropped to only 11% of that of
1961. The colony numbers recovered to 86% but only to crash
again after the economic crisis in Venezuela in 1994. Apiculture
never recovered during the last two decades and remained at
28% of the colony number reported 50 years ago.
5) The start of the military rule in Bolivia in 1964 was also followed
by an extreme decline in managed colonies and the FAO records
end in 1973 with a reported zero colonies.
In spite of potential ambiguities in the FAO data base, none of
the colony number dynamics of the past 50 years, neither increase
nor decrease, show any relation to the arrival of novel pests or the
use of novel pesticides or toxins in the respective countries. Even
the use of the controversially discussed and now partially banned
neonicotinoid agrochemicals did not cause any abrupt decrease in
colony numbers in any of the national data provided nor did they
impact on the long term changes in colony numbers which went up
or down irrespective of the use or ban of pesticides in the data
providing countries (Eisenstein 2015; Staveley et al., 2014). Yet it
appears the long term declines, as shown for Western Europe and
the US, are those of largest concern because they have been almost
linear, consistent and stable over more than two decades. These
declines cannot be explained by pests, pathogens, pesticides or
societal collapses and hence it may be helpful to use additional
information to extract potential causes.
3.2. The honey data base
Some additional relevant data on apiculture can be obtained by
extracting the FAO honey data base. It reflects honey production
and trade (import and export) (Fig. 3). Globally the production has
almost doubled over the past five decades and since it are
honeybee colonies that produce the honey the data set may allow
for the interpretation of the colony number dynamics from a
different perspective. In particular, it can serve to conduct
plausibility checks on the colony number data. For example the
Fig. 3. Global honey production (empty circles), import (black circles) and export (grey circles) between 1961 and 2013. Import and export data were only available till 2011.
The proportion of honey traded increased in relation to the global production over time from 11% in 1961 to 31% in 2011. Nevertheless, the majority of honey is still marketed at
the national level.
47
dramatic decline in Italy in 2007–08 with a reported 500,000
colonies disappearing was not visible in the national honey
production neither in the following nor in the five subsequent
years. Actually the average honey production was even slightly
(but by no means significant, P = 0.9) less in the five years before
(9600.0 t) compared to the five years after the reported decline
(9764.4 t). It seems highly unlikely that the surviving colonies
worked more than twice as hard to compensate these upsetting
losses. It is much more likely that colony numbers were incorrect
estimates either before or after 2007. Similarly there was an
increase of 300,000 colonies in Uruguay in 1999, which was highly
welcomed but also well buffered in the national honey production
and actually fell to only half of that level the year after (typically
the honey is marketed the year after the harvest). So again the data
sets of colony number and honey production lack consistency.
Nevertheless, in spite of these specific national inconsistencies,
at the overall global scale there is a highly significant correlation
(mean r = 0.70  0.03, n = 103) between the number of colonies and
the honey produced as reported by Aizen and Harder (2009b).
Although this seems trivial, a more regional analysis reveals
surprising variance. The data fall into two distinctly different
groups: those countries with a colony decline and those with a
colony increase. Countries with an increase in managed colonies
show a positive slope between the change in honey production per
colony and the number of managed colonies (b = 0.29  0.64).
Countries with a colony decline surprisingly have steep negative
slope (b = 1.32  0.64). Hence the reported honey production per
colony increased in response to the colony decline (Fig. 4). In some
countries this increase in reported colony productivity seems to be
biologically impossible given the regional environmental and
climatic conditions in these countries. There are two possible
explanations for these implausibilities: either the actual numbers
of colonies are in reality much higher than reported or the amount
of honey reported as nationally produced is much less than
reported. When looking at the honey trade it becomes clear that
countries with a positive correlation between production per
colony and colony number are the main honey exporters whereas
the countries with a negative correlation are those importing
honey. Equally interesting is that these countries show a positive
48 R.F.A. Moritz, S. Erler / Agriculture, Ecosystems and Environment 216 (2016) 44–50

Fig. 4. Changes in colony number and honey production (kg) per colony for 20 European countries. The x-axis shows the difference in % between the mean number of colonies
of the first 5 years (1961–1965) with those of the last five years (2009–2013) in the data base. Countries with colony declines (filled circles, negative x-values) show a negative
slope between colony number and honey production per colony. Countries with a colony increase (open circles, positive x-values) show a slight positive slope with honey
production per colony. Both data sets are plotted in a log10-scale to better visualize the different relationships between colony decline and productivity per colony. (ALB—
Albania, AUT—Austria, BEL—Belgium-Luxemburg, BGR—Bulgaria, CHE—Switzerland, CYP—Cyprus, DEU—Germany, GRC—Greece, FIN—Finland, FRA—France, HUN—Hungary,
ITA—Italia, POL—Poland, PRT—Portugal, ROM—Romania, SWE—Sweden, TCH—Czechoslovakia, TUR—Turkey, URS—Union of Soviet Socialist Republics, YUG—Yugoslavia).

correlation between import and export (mean r = 0.41  0.08,
n = 20), hence the more is imported also export increases showing
that commercial honey trade takes an increasing share of the
market. The negative correlation between colony number and
production per colony implies that the yield per managed colony
should have increased, in some of these countries more than
doubled (e.g., BEL and SWE produced in the last 5 years 51 kg/
colony and 54 kg/colony, respectively, more compared to the first 5
years of data collection). Given the reports that foraging conditions
for bees have decreased rather than increased (Potts et al., 2010a),
it seems unlikely that the enhanced colony productivity can be
explained by bees working harder. If we assume that the biological
capacity of a colony to produce honey has remained constant over
time, and the foraging conditions have not become doubly
rewarding, it is more likely that the honey marketing and/or
colony management has changed. An extreme case may be that of
the honey produced in Belgium. If one divides the honey produced
by the number of managed colonies, Belgian honeybee colonies
seem to globally excel with an average production of 83 kg per
colony per year (average over the years 2009–2013). Such numbers

Fig. 5. Negative correlation between honey trade and the decline in colony numbers in Europe [mean of (2007–2011) as percent of mean (1961–1965)]. Countries with strong
colony declines trade more honey than they produce. In contrast, countries with a colony increase not only produce but also consume more honey than before (2007–2011
compared to 1961–1965). (Country codes as in Fig. 4).
 R.F.A. Moritz, S. Erler / Agriculture, Ecosystems and Environment 216 (2016) 44–50
Fig. 6. Change in honey production (kg/year) per colony (mean  SE) for the 89 countries with consistent data on colony numbers and national honey production between
1961 and 2013. There is an increase in productivity per colony (b = 0.162, r2 = 0.95, P < 0.001, n = 89) but also a significant increase in the variance over time (b = 0.012, r2 = 0.55,
P < 0.001, n = 89).
are unheard of in Western Europe, where average honey yields per
colony of 40 kg reflect excellent honey yield seasons (Genersch
et al., 2010). At the same time the national Belgian honey
production shows globally the highest correlation with the amount
of yearly imported honey (r = 0.98, P < 0.001), one cannot exclude
recurrent irregular reporting of imported honey as nationally
produced to the FAO. Whereas it is plausible that honey export can
correlate with the number of colonies, it seems hard to believe that
Belgian honeybees adjust their productivity according to the
imports.
The example from Belgium, as well as Sweden, may be special
cases. It seems clear that marketing of honey has changed over the
past five decades as apiculture became more professionalized. It
may well be that slowly changing societal values, from hobby to
business beekeeping, are important drivers of colony declines.
Clearly, the increase of honey trade in relation to the national
honey production in Europe goes hand in hand with the colony
decline over the five decades listed in the FAO data base (Fig. 5). We
found a highly significant negative correlation between honey
trade and the corresponding change in the number of managed
colonies (r = 0.55; P = 0.01, n = 20). Countries suffering the
strongest colony declines were those where honey trade became
much more important than production.
4. Conclusions
In spite of the manifold ambiguities of the FAO data base it is
certainly possible to identify grave and sustainable colony declines
in many industrialized countries. Indeed the density of managed
honeybee colonies lies orders of magnitudes below those of wild
honeybee populations with little or no beekeeping as shown for
the Kalahari desert, arguably not a preferred habitat for honeybees
(Jaffé et al., 2010). Perhaps disappointing for some media, the
declines detectable in the FAO data set are not due to pathogens,
pests, pesticides, climate change or any other factor of timely
public interest. It is the decline in beekeeping activity and the
increase of honey trade (Fig. 5) that is of concern. A global honey
market with low honey prices in exporting countries may make it
less attractive for professional beekeepers in importing countries
to produce honey with their own colonies. This is well supported
49
by the negative correlation between honey production and the
number of colonies in countries with a colony decline (Fig. 4).
Although, globally the honey production per colony increased by
16% per year (b = 0.162, r2 = 0.98, P < 0.001, n = 89) over the past
decades (1961–2013), with an average of 18.38  0.35 kg (Fig. 6),
the reported steep increases in colony productivity in countries
with colony declines are not likely to be real. Although improved
management may have improved the efficiency of honey produc-
tion and harvesting in many exporting countries, it is much more
likely that the global increase in colony numbers is the primary
driver of the increase of global honey produced. This is also
supported by the significant increase in the variance of colony
productivity among the various countries over the past decades
(b = 0.012, r2 = 0.55, P < 0.001, n = 89) (Fig. 6) which is strongly
driven by the massive (and not necessarily plausible) increase in
productivity per colony in the high honey trade countries.
Whereas in many countries the global markets are in support of
further developing apiculture, in may now be time in the
industrialized countries to reconsider policies on supporting
apiculture. In these countries it used to be the large number of
hobbyist beekeepers that kept a few hives in the backyard that
contributed the largest share in the number of colonies by far. To
them the market value of honey may not be the prime driver for
their operation. If the linear colony declines of the past 50 years in
countries like the US, Germany, Austria and Switzerland continue
at the current pace, in the following decades it may well fall below
levels where we can only hope that wild bee pollinators, other than
honeybees, can provide sufficient pollination services (Garibaldi
et al., 2013). Given the reports on pathogens, negative impacts of
pesticides and habitat loss of wild bee pollinators, this is probably
an overly optimistic expectation (Fürst et al., 2014). In a few
countries in Western Europe there are now first reports on
increasing numbers of beekeepers. Recent data from UK (Vanber-
gen et al., 2014) show how tightly the increase colony number is
correlated by the number of beekeepers. The path back to the old
colony densities in Europe and North America may have been
reopened in response to the medial attention of global honeybee
declines. This is a very first positive signal that should be fostered
by receiving more support from governmental institutions. If this
increase in the interest for apiculture has been partly due to the
50 R.F.A. Moritz, S. Erler / Agriculture, Ecosystems and Environment 216 (2016) 44–50

media hype on colony losses this is clearly to be welcomed, even Genersch, E., von der Ohe, W., Kaatz, H., Schroeder, A., Otten, C., Büchler, R., Berg, S.,
though it may very likely be for different reasons than many media Ritter, W., Mühlen, W., Gisder, S., Meixner, M., Liebig, G., Rosenkranz, P., 2010.
The German bee monitoring project: a long term study to understand
wanted the general public to believe. periodically high winter losses of honey bee colonies. Apidologie 41 (3), 332–
352.
Acknowledgement González-Varo, J.P., Biesmeijer, J.C., Bommarco, R., Potts, S.G., Schweiger, O., Smith,
H.G., Steffan-Dewenter, I., Szentgyörgyi, H., Woyciechowski, M., Vilà, M., 2013.
Combined effects of global change pressures on animal-mediated pollination.
Financial support was granted by the Federal Ministry of Food, Trends Ecol. Evol. 28 (9), 524–530.
Agriculture and Consumer Protection (Germany): FIT-BEE project Jaffé, R., Dietemann, V., Allsopp, M.H., Costa, C., Crewe, R.M., Dall’Olio, R., De La Rúa,
P., El-Niweiri, M.A.A., Fries, I., Kezic, N., Meusel, M.S., Paxton, R.J., Shaibi, T.,
(Grant no. 511-06.01-28-1-71.007-10). Stolle, E., Moritz, R.F.A., 2010. Estimating the density of honeybee colonies
across their natural range to fill the gap in pollinator decline censuses. Conserv.
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