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Looking for “the Best Bee”. An experiment about interactions between origin and environment of honey bee strains in Europe

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The honey bee Apis mellifera, of which there are currently 28 identified subspecies and numerous ecotypes, have been evolving and adapting to a wide range of environments for hundreds of thousands of years within their native range of Europe, Africa and Asia. Honey bees have been widely dispersed over the past several hundred years and are now also established in the Americas and Australia. Today, the high loss of colonies worldwide is attributed to a combination of factors, including parasitic mites, pathogens, pesticides and malnutrition. The COLOSS network of European scientists asks the questions: Does beekeeper selection for productivity lead to genetic deficiency, and are locally adapted populations being displaced by the movement of various honey bee types to locations beyond their native range? A major research effort explores these questions, looking at numerous types of honey bees that are endemic to specific areas of Europe or have become adapted after several decades of breeding. Beekeepers in the U.S. also consider these questions through interest in locally adapted bees and “survivor” feral bees, although the situation is very different. Our honey bees are not native and were derived from relatively small founder populations, thus we lack the evolutionary diversity of subspecies and ecotypes that exist in Europe. We also lack the strong support of institutions and beekeeper organizations devoted to the selection and maintenance of specific subspecies, as established in many European countries. Feral populations in the U.S., previously considered a mixed source of raw genetic variation, have been devastated by the impact of Varroa mites. Through semen collections from Old World sources, Washington State University has been involved in the importation and distribution of additional honey bee genetic diversity in the U.S. Associated with the importations, cryopreserved germplasm from “pure” Old World subspecies has been deposited in the WSU Germplasm Repository for future breeding and conservation needs.Through such measures, we hope to enhance domestic bee breeding programs by providing additional genetic diversity to improve bee health in the U.S.
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(Originally published in German in “die
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Introduction by Susan Cobey
The honey bee Apis mellifera, of
which there are currently 28 iden-
tified subspecies and numerous
ecotypes, have been evolving and adap-
ting to a wide range of environments for
hundreds of thousands of years within
their native range of Europe, Africa and
Asia. Honey bees have been widely di-
spersed over the past several hundred
years and are now also established in the
Americas and Australia. Today, the high
loss of colonies worldwide is attributed to
a combination of factors, including pa-
rasitic mites, pathogens, pesticides and
The COLOSS network of European
scientists asks the questions: Does bee-
keeper selection for productivity lead to
genetic deciency, and are locally adap-
ted populations being displaced by the
movement of various honey bee types
to locations beyond their native range?
A major research effort explores these
questions, looking at numerous types of
honey bees that are endemic to specic
areas of Europe or have become adapted
after several decades of breeding.
Beekeepers in the U.S. also consider
these questions through interest in locally
adapted bees and “survivor” feral bees,
although the situation is very different.
Our honey bees are not native and were
derived from relatively small founder po-
pulations, thus we lack the evolutionary
diversity of subspecies and ecotypes that
exist in Europe. We also lack the strong
support of institutions and beekeeper or-
ganizations devoted to the selection and
maintenance of specic subspecies, as
established in many European countries.
Feral populations in the U.S., previously
considered a mixed source of raw genetic
variation, have been devastated by the
impact of Varroa mites.
Through semen collections from Old
World sources, Washington State Uni-
versity has been involved in the impor-
tation and distribution of additional
honey bee genetic diversity in the U.S.
Associated with the importations, cryo-
preserved germplasm from “pure” Old
World subspecies has been deposited
in the WSU Germplasm Repository for
future breeding and conservation needs.
Through such measures, we hope to en-
hance domestic bee breeding programs
by providing additional genetic diversity
to improve bee health in the U.S.
The international research network CO-
LOSS (Prevention of COlony LOSSes, was founded in 2008 and
received funding from the EU COST pro-
gram until 2012. The network aims to pro-
mote international collaboration on research
about colony losses. Within COLOSS, the
working group “Diversity and Vitality”
(now Research Network for Sustainable Bee
Breeding, investi-
gated the survival of honey bee colonies in
relation to their genetic origin and their ad-
aptation to environmental factors such as cli-
mate, diseases and beekeeping management.
Europe-wide comparison
To study the complex interactions be-
tween honey bee colonies and their en-
vironment, we conducted a very large
experiment involving colleagues from 11
countries. In this experiment, we compared
16 different strains of honey bees in differ-
ent environments for two and a half years,
with respect to characters such as honey
yield, survivability and susceptibility to
diseases. The experimental apiaries were
distributed across Europe, reaching from
Finland in the North to Sicily and Greece in
the South (gure 1). The different strains in
the experiment consisted of breeding lines
maintained at the institutes involved, local
breeding stock, regional bees that had not
been subjected to breeding efforts or lines
from conservation programs. The strains be-
longed to the ve subspecies Apis mellifera
1 LLH, Bee Institute, Erlenstrasse 9, 35274
Kirchhain, Germany
2 Consiglio per la Ricerca e la sperimenta-
zione in agricoltura – Unità di ricerca di
apicoltura e bachicoltura (CRA-API), Via
di Saliceto 80, 40128 Bologna, Italy
3 Faculty for Agricultural Science and Food,
bul. Aleksandar Makedonski b.b., 1000
Skopje, Republic of Macedonia
4 Research Institute of Horticulture, Apicul-
ture Division, 24-100 Pulawy, Poland
5 Agricultural University of Athens, Labora-
tory of Agricultural Zoology and Entomol-
ogy, 75 Iera Odos St., Athens 11855 Greece
6 The University of Applied Sciences Marko
Marulic in Knin, Croatia
7 Hellenic Institute of Apiculture –Hellenic
Agr. Org. ‘DEMETER’ , Nea Moudania,
8 Agricultural University of Plovdiv, 12,
Mendeleev Str, Plovdiv 4000, Bulgaria
9 Faculty of Agriculture, University of Za-
greb, Svetosimunska 25, 10000 Zagreb,
10 University of Aarhus, DJF, Research Cen-
tre Flakkebjerg, 4200 Slagelse, Denmark
11 INRA, UR 406 Abeilles et Environne-
ment, Laboratoire Biologie et Protection
de l’abeille, Site Agroparc, 84914 Avignon,
12 MTT, Agrifood research Finland, 31600
Jokioinen, Finland
13 Apiculture Division, Warmia and Mazury
University, Sloneczna 48, 10-710 Olsztyn,
American Bee Journal664
Cecelia Costa and Marina Meixner in Croaa
mellifera, A. m. carnica, A. m. ligustica, A.
m. macedonica and A. m. siciliana.
Each strain was present with at least ten
colonies in at least three of the 21 apiaries.
In every apiary, the local strain was com-
pared to at least two “foreign” strains.
Uniform starting conditions
The colonies were uniformly built in the
summer of 2009, either from shook swarms
or from splits, and the experimental queens
were introduced. The experiment started
on October 1, 2009, when all colonies con-
sisted of offspring of the new queens, and
ended on March 31, 2012.
All colonies were evaluated in regular
intervals. Colony development, amount of
brood and all other characters were assessed
according to international recommendations
(Büchler et al., 2013). These were based on
the traditional Apimondia guidelines, but
were expanded to include characters such
as brood hygiene. Thus, they were adapted
to the challenges of selection of vital and
resistant bees. In addition, at several times
bee samples were taken from each colony
and examined for bee diseases.
A colony was considered as lost when it
had either collapsed or the colony strength
was considered insufcient for further sur-
vival. Queenlessness or the presence of a
drone-laying queen was also regarded as
colony loss.
No medication was used during the en-
tire experiment; however, it was possible to
perform a total brood removal for control of
Varroa mites (per apiary). To prevent spill-
over of mites from collapsing colonies, the
Varroa infestation of each colony was moni-
tored continuously, and colonies in danger
of collapsing were treated. At the same time,
they were counted as lost and excluded from
further analyses (the complete test protocol
is described in Costa et al., 2012).
Hybridization reduces gentleness
Although we observed noticeable differ-
ences in behavior and performance between
strains that originated from breeding pro-
grams and strains that had received little
selective effort in the past, no single strain
showed superior performance at all loca-
tions. However, we most noticeably ob-
served that strains showing strong signs of
hybridization in the genetic analysis (Fran-
cis et al., 2014a) scored signicantly lower
in the assessment for gentleness (Uzunov et
al., 2014).
Local strains survive longer
Of the 597 colonies we could analyze, 94
(15.7%) survived until the end of the exper-
iment. We observed drastic differences in
survival time and disease load, both between
locations and between the genetic strains.
At some locations, for instance in Lunz
(Austria) or Schenkenturn (Germany), all
colonies had already collapsed by the sec-
ond winter (2010/2011), while colonies in
Avignon (France) survived longest with an
average of almost two years. Survival time
between the strains also differed noticeably.
Here, we observed a signicant difference
in survival time between local strains and
foreign strains. While in any given location
a colony of a foreign strain survived on av-
Figure 1: Map of Europe showing the 21 test locaons covering 11 countries.
Each locaon is indicated by a black dot, with its name shown in the white
box. The genec lines maintained at each locaon are indicated as leers be-
low each name. The legend at top right corner links the leers to the genec
lines. The abbreviaons mean: CarB = Carnica Bann (Germany), CarC = Carnica
Croaa, CarG = Carnica Kunki (Poland ), CarK = Carnica Kirchhain (Germany),
CarP = Carnica Gasiory (Poland ), CarL = Carnica Lunz (Austria), CarV = Carnica
Veitshöchheim (Germany), LigI = Ligusca Italy, LigF = Ligusca Finland, MacB
= Macedonica Bulgaria, MacG = Macedonica Greece, MacM = Macedonica
Macedonia, MelF = Mellifera France, MelL = Mellifera Læsø (Denmark), MelP
= Mellifera Poland, Sic = Siciliana. The leer in the circle next to each locaon
indicates the respecve local strain. Example: In Kirchhain, the strains D, E,
and N were tested, with CarK (D) being the local strain. In addion, CarP (E)
and MelF (N) were tested. Copyright Internaonal Bee Research Associaon.
Reprinted from Francis et al. (2014) with permission of the editors of Journal of
Apicultural Research.
June 2015 665
erage 470 days, the mean survival time of a
local colony was 553 days. Local bees thus
survived on average 83 days longer than for-
eign ones (Büchler et al., 2014).
Reasons for losses
The most frequent evident reasons for
colony loss were Varroa (38%), problems
with the queen (loss, drone layer, etc., 17%)
and Nosema (8%). All other reasons (star-
vation, robbing, unspecied winter loss,
other diseases, unknown reason) were less
frequent, but together accounted for 37% of
the losses.
Varroa infestation inuenced by location
The infestation with Varroa mites was
signicantly more strongly inuenced by
the apiary location compared to the genetic
origin of the colony (Meixner et al., 2014).
The Varroa infestation rates differed greatly
over the individual apiary locations. In some
places we observed a fast buildup of mite
populations, while in other locations infesta-
tion rates increased much more slowly. The
differences between experimental stations
were often much higher than the differences
between surviving and collapsing colonies
at one single station. In the autumn of 2010,
for instance, we observed extremely high
infestation rates between 30% and 40% at
the experimental locations of Unije (Croa-
tia) and Dimovci (Bulgaria). In spite of
these high infestations, many colonies at
these two locations survived the following
winter. In contrast, mite infestation rates at
stations in Poland and Italy increased more
slowly and remained below 10%, even after
two years without medication. In Kirchhain
(Germany), the autumn 2010 mean infesta-
tion rate of surviving colonies was 9.1%,
while in collapsing colonies it was 24.3%
(Büchler et al., 2014).
The differing length of season and the
resulting differences in colony development
certainly were among the main reasons for
the differences in mite population devel-
opment across the experimental colonies
(Hatjina et al. 2014). Our results indicate
that there is substantial variation of Varroa
damage thresholds across different regions
of Europe. To determine these thresholds,
comprehensive investigations involving
Field workshop pin test in Germany
(l) COLOSS WG4 Workshop | Palermo, Italy | 05 - 09 Nov 2012 (r) COLOSS WG4 Workshop | Forssa, Finland | 23 - 27
Jan 2012
sufciently large numbers of colonies are
Nosema not among major causes for losses
The gut parasite Nosema was present in
almost all locations, but colony losses as-
cribed to Nosema were low and in the ma-
jority (25 of 37 cases) occurred in a single
location (Le Bine, Italy) at the beginning
of the experiment. The Nosema spore load
across the experimental colonies was over-
all rather low; only at the locations in Po-
land and Italy were higher spore numbers
occasionally observed. In most apiaries we
only observed the “new” Nosema species
Nosema ceranae, while Nosema apis was
restricted to few locations and mostly oc-
curred in mixed infections with N. ceranae.
Pure N. apis infections were sporadically
found only in Finland and Poland. Thus,
our data do not support Nosema ceranae as
a major cause for substantial colony losses
(Meixner et al., 2014).
The frequency of virus infections (Acute
Bee Paralysis Virus and Deformed Wing
Virus) was also strongly inuenced by the
apiary location. For instance, in autumn
2010 in samples from Finland, no viruses
at all were found, while both viruses were
present in all analyzed samples from Bul-
garia. Overall, we could not determine an
effect of genetic origin on the frequency of
virus infections. However, an in-depth study
preformed on samples from the Greek loca-
tion (one of the largest, containing 4 geno-
types) showed that local colonies tended to
have lower levels of pathogens. In this case
study the seasonal trends of the viruses were
conrmed (lower levels in spring, higher in
autumn), together with the signicant cor-
relation between varroa and DWV (Francis
et al., 2014 b).
Local bees may have an advantage
Thus, our results clearly demonstrate that
location effects play a predominant role in
the occurrence of bee diseases. Both local
and foreign bees suffered from parasites and
other pathogens. Yet, the mean survival du-
ration of local bee origins was signicantly
longer than that of foreign ones. Possibly,
this ostensible contradiction indicates that
local bees may command more resources
for keeping parasites and pathogens in
check, due to their better adaptation to the
local environment, climate and vegetation,
but also to the locally prevailing manage-
ment methods. In addition, newer research
demonstrated that especially viruses exhibit
substantial genetic variation across regions
that may inuence their virulence (Cornman
et al., 2013). It could be possible that local
bees are better adapted to “their” strains of
viruses and are therefore better able to cope
with them.
The best bee does not exist!
In conclusion, our experiment demon-
strated that “the best bee” showing excellent
performance and superior disease tolerance
across all environments does not exist. In-
stead, the local bees were not only the most
long-lived, but in many cases also received
better scores for gentleness and honey yield.
Therefore, we suggest devoting more
attention to the preservation of the variety
of genetic resources of honey bees across
Europe. One way to achieve this could be
the establishment of conservation areas to
protect endangered populations from uncon-
American Bee Journal666
trolled introgression of imported strains. In
particular, however, we would like to em-
phasize the necessity of regional selection
and breeding efforts. Such efforts would
contribute to an improvement of local bees
and, in consequence, increase their accep-
tance with local beekeepers. Special at-
tention within such programs should be
devoted to traits like disease tolerance and
The uncontrolled importation of bee
strains from different areas endangers well-
adapted local bee populations and is often
not even to the advantage of the beekeeper,
as our experimental results show. For the
common beekeeper, our recommendation
would be to purchase queens from local
breeders whose material has been selected
after long-term comparative testing in their
own region.
The results of this experiment have been
published with open access in a series of sci-
entic articles in a Special Issue (May 2014)
of Journal of Apicultural Research (www. and are listed in the refer-
ences. This article provides an overview on
the most signicant results.
Büchler R; Andonov S; Bienefeld K;
Costa C.; Hatjina F; Kezic N; Kryger
P; Spivak M; Uzunov A; Wilde J
(2013). Queen rearing and selection.
In: Dietemann V; Ellis J D; Neumann P
(Eds) The COLOSS BEEBOOK: stan-
dard methods for Apis mellifera research.
Journal of Apicultural Research, 52(1):
DOI 10.3896/IBRA.
Büchler R; Costa C, Hatjina F, Andonov
S, Meixner MD, Le Conte Y, Uzunov A,
Berg S, Bienkowska M, Bouga M, Dra-
zic M, Dyrba W, Kryger P, Panasiuk B,
Pechhacker H, Petrov P, Kezic N, Ko-
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environmental effects on the survival of
Apis mellifera L. colonies in Europe.
Journal of Apicultural Research, 53(2):
205- 214.
Cornman RS; Boncristiani H; Dainat B;
Chen Y P; Vanengelsdorp D; Weaver
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the western honey bee, Apis mellifera,
inferred from deep sequencing. BMC
Genomics: 14
Costa C, Büchler R, Berg S, Bienkowska
M, Bouga M, Bubalo D, Charistos L,
Le Conte Y, Drazic M, Dyrba W, Fil-
lipi J, Hatjina F, Ivanova E, Kezic N,
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... Z ostatnich badań akcji COLOSS (2014-2016) wynika, że pszczoły lokalnych populacji np. Apis m. mellifera są najbardziej przystosowane do warunków środowiskowych i lepiej radzą sobie z chorobami, pasożytami oraz innymi patogenami (17,44,54,90). ...
... Średni czas przeżywalności pszczół miejscowych był jednak znacznie dłuższy niż w przypadku pszczół nielokalnych. Lokalne pszczoły mogą dysponować większymi zasobami, aby powstrzymać pasożyty i patogeny dzięki ich lepszemu przystosowaniu do lokalnego środowiska, klimatu i roślinności (54). ...
... pszczół środkowoeuropejskich w całej Europie. Jednym ze sposobów osiągnięcia tego celu może być ustanowienie obszarów chronionych zagrożonych populacji (17,44,54,90). ...
In Poland, as in the whole world, there is a growing risk of extinction of the honeybee, especially the subspecies of the native middle-European bee. The main factors for the disappearance of native bee lines are environmental degradation, diseases and pathogens, as well as the introduction of imported queen bees of other breeds into domestic breeding. In this situation, it is particularly important to protect the genetic resources of native bees, which currently live in small areas covered by protection programs. The aim of this work is to review the possibilities offered by morphological and genetic examinations in the conservation breeding of native honey bee lines. It was found that the implementation of programs for the protection of native middle-European bees should be continued because of the growing risk of losing or diluting the valuable gene pool of native bees. Only the combination of phenotypic analysis and analysis based on DNA markers can effectively contribute to the protection of the native middle-European bee..
... According to the recent COLOSS research, local bees, such as Apis m. mellifera, live longest and they often perform best in terms of quietness and honey yield. Also, local bees are more immune to parasites and other pathogens (12,28,37,62). It has been established that the effect of genotypes on the location plays a predominant role in the occurrence of diseases, such as the deformed wing virus associated with the epidemic of varroosis. ...
... The mean time of survival of sick local bees, such as Apis m. mellifera, is much longer than that of nonlocal bees. Local bees can have larger resources to contain parasites and pathogens owing to better accommodation to local climate and environment (37). Subsequent research confirmed that bee families with local queens (including middle-European bees) live longer -for instance by 83 days -than non-local ones (12). ...
... Accordingly, more attention should be paid to the preservation of diversified genetic resources of honey bees, including of middle-European bees, throughout Europe. The establishment of areas of protection for endangered populations is one of the means of achieving this goal (12,28,37,62). Today, middle-European bees are protected mainly in areas where this species evolved. ...
The population of the honey bee, Apis mellifera, continues to shrink. The middle-European bee, Apis m. mellifera, is particularly at risk in Europe. The drop in the number of middle-European bees is so huge that the insect is under the threat of extinction. Today, they live on small areas covered by the protection of genetic resources. Apis m. mellifera is protected mainly in areas where this species evolved: for instance, in Switzerland, Latvia, Norway, Sweden, Finland, Denmark, France, Germany, Poland or Russia. This paper presents methods used to preserve and protect Apis m. mellifera in Europe and research on the descent and original extent of the species. It also reviews opportunities created by the implementation of various types of programs for the protection of genetic resources of Apis m. mellifera and ways of employing morphological and genetic studies for the conservative breeding of middle-European bees. The paper demonstrates that the protection of Apis m. mellifera in Europe is necessary, considering the decreasing size, and the threat of hybridization, of this population. The use of the morphometric evaluation and DNA analysis methods have made it possible to track and compare likely directions of propagation of genes in the long history of evolution of bees. Moreover, these methods have given us better insight into the ongoing processes. The current use of these methods for reliable identification of bee breeds helps to protect Apis m. mellifera more effectively. European programs for the protection of genetic resources of bees are based on the following two main paradigms: the breeding of local isolated populations on islands and establishment of protected inland areas for the conservative breeding of contained swarms. All these programs share and are successful in achieving the goal that consists in the preservation of the characteristics of Apis m. mellifera as unchanged as possible, with retention of the maximum genetic diversity of the species.
... К сожалению, лечение пчел от клеща приводит к загрязнению продуктов пчеловодства. Поскольку приобретенная устойчивость пчел к этому паразиту может передаваться по наследству, то для решения проблемы необходимо использовать селекцию [3][4][5]. В 2017 г. на базе Института пчеловодства в г. Кирхгайне (Германия) Европейская комиссия создала международный консорциум по исследованию пчел EurBeST (European Bee Selection Team) ( ...
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Бюхлер Р., Узунов А., Ильясов Р. А., Коста С., Мейкснер М., Ле Конте И., Мондет Ф., Ковачич М., Андонов С., Каррек Н. Л., Димитров Л., Бассо, Б., Биенковска М., Далл’Олио Р., Хатджина Ф., Вирц У. Проект EURBEST: тестирование пчел на устойчивость к клещу варроа // Пчеловодство. ‒ 2022. № 2. ‒ C. 62-64. (Büchler R., Uzunov A., Ilyasov R. A., Costa C., Meixner M., Le Conte Y., Mondet F., Kovacic M., Andonov S., Carreck N. L., Dimitrov L., Basso B., Bienkowska M., Dall’Olio R., Hatjina F., Wirtz U. EURBEST project: testing bees for resistance to the mite Varroa // Russian Journal of Beekeeping "Pchelovodstvo". ‒ 2022. № 2. ‒ P. 62-64.) В 2017 году Европейская комиссия создала международный консорциум по исследованию пчел EurBeST (European Bee Selection Team) на базе Института пчеловодства в Кирхгайне, Германия. В рамках основной части проекта EurBeST было проведено пять крупномасштабных исследований, включающих семь стран Европейского Союза и 130 пчеловодов. Команда EurBeST определила и отобрала 23 линии пчел, принадлежащие к шести подвидам пчел, а также пчел гибридного происхождения. Пчелы были тестированы на общие признаки и признаки устойчивости к клещу Varroa. Тестирование пчел было проведено на двух различных уровнях: [1] сравнительное тестирование исследователями нескольких тестовых линий пчел на одной пасеке, [2] сравнительное тестирование коммерческими пчеловодами нескольких тестовых линий пчел вместе со своими линиями пчел в обычных полевых условиях. Проект EurBeST испытал более 3500 семей в течение одного сезона и считается одним из крупнейших исследований по оценке медоносных пчел на устойчивость к клещу Varroa в Европе. В результате проведенных исследований EurBeST были получены сведения о линиях медоносных пчел, обладающих устойчивостью к клещу Varroa и способные подавлять размножение клещей Varroa в семье. Селекции пчел является важным и единственно возможным инструментом для перехода к экологически чистому пчеловодству без лечения от болезней. Было доказано, что селекция пчел на устойчивость к клещу Varroa работает, но она дорогостоящая. Гигиеническое поведение против клеща Varroa и подавление его развития являются полезным критериями для отбора пчел, устойчивых к клещу Varroa. Однако затраты на тестирование пчел для селекционеров высоки и должны быть компенсированы государством. In 2017, the European Commission set up the international bee research consortium EurBeST (European Bee Selection Team) based at the Institute of Beekeeping in Kirchhain, Germany. Five large-scale surveys involving seven European Union countries and 130 beekeepers have been carried out in the core part of the EurBeST project. The EurBeST team identified and selected 23 honey bee lines belonging to six bee subspecies as well as bees of hybrid origin. They were tested for common traits and traits of resistance to the Varroa mite. Honey bee testing has been performed at two different levels: [1] comparative testing by researchers of several test honey bee lines in a single apiary; and [2] comparative testing by commercial beekeepers of several test honey bee lines together with their honey bee lines under normal field conditions. The EurBeST project has evaluated more than 3500 colonies in a single season and is considered one of the biggest studies in Europe for the evaluation of the resistance of honey bees to the Varroa mite. The results of the EurBeST surveys have shown that Varroa mite-resistant honey bee lines can suppress Varroa mite reproduction in honey bee colonies. Honey bee selection is an important and the only possible tool for the transition to environmentally friendly beekeeping without disease treatment. The selection of bees for resistance to the Varroa mite has been proven to work, but it is expensive. Hygienic behavior of honey bees against the Varroa mite and suppression of its development are useful criteria for selecting Varroa mite-resistant honey bees. However, the cost of testing bees for breeders is high and must be compensated by the government.
... However, the advancement in digital technology has resulted in a rapidly growing number of Citizen Science projects, as this facilitated the communication between the Citizen Scientists and the researchers. Citizen Science is not new within COLOSS group either, as several studies did anticipate on beekeepers' help and contribution (see GEI experiment, CSI Pollen), the size of which and the results were astonishing (Brodschneider et al., 2019;Meixner et al., 2015). Druschke and Carrie (2012), name the Citizen Scientists 'ambassadors for science' as they interact directly with other members of society conveying knowledge and experience, especially because they do go beyond the simple data collection. ...
... The necessity Variability of morphological characteristics of middle-European bees of the 'Northern M' line of preserving native bees as a general environmental requirement that should be met by establishment of sanctuaries, i.e. areas taking account of the reproductive biology of bees, was realized as early as in the 1960s and 1970s (21). Recent COLOSS action studies (2014)(2015)(2016) suggest that bees from local populations, such as Apis m. mellifera, are the most adapted to environmental conditions and are better at fighting diseases, parasites and other pathogens, and one of the ways to achieve this, could be to establishing of protected areas of endangered populations (8,25,32,53). Today, implementation of updated programs for the protection of genetic resources of middle-European bees (four lines: 'Augustów M', 'Kampinos M', 'Northern M' and 'Asta M') is monitored by the Ministry of Agriculture and Rural Development acting through the Animal Husbandry Institute of the State Research Institute and through the National Center for Animal Breeding. The Olecko Apiary owned by the KRIR Breeding Apiary Ltd. based in Parzniew (former Animal Breeding and Insemination Institute Ltd. in Bydgoszcz) has been one of the direct contractors for such programs since January 1, 2014. ...
The aim of the study was to evaluate the variability of morphological characteristics of native middle-European bees (Apis m. mellifera) of the ‘Northern M’ line. The research covered characteristics of breed (the length of proboscis, the cubital index), body size (the width of tergite 4 and the sum of widths of tergites 3 and 4) and wing size (length and width). The study compared bees harvested from a leading apiary and from collaborating apiaries participating in a program for the protection of genetic resources of bees of this line. The material for the study was harvested in 10 consecutive years. The samples were collected by the “cluster drawing” method (the multi-stage method of clustering described by Zee et al. in 2013). Each sample consisted of 25 to 30 bees. The frames were loaded in an instrument for the morphological measurement of bees (Apimeter). Seven measurements were taken on prepared body parts of each bee. The length and width of the wing and the length of the cubital vein were measured on the right front wing (hereinafter referred to as the “wing”). In addition, the width of abdominal tergites 3 and 4 and the length of proboscis were measured in each instance. In total, 4 291 bees were harvested and 30 037 measurements were taken. The conclusion is that the program for the protection of genetic resources of bees of the ‘Northern M’ line can be implemented in Poland based on the leading apiary and on the collaborating apiaries, and bees of this line display characteristics of middle-European bees. Moreover, the study demonstrated a consistency of values of the studied characteristics of the ‘Northern M’ line with the applicable references of morphological characteristics for Apis m. mellifera. In addition, based on a review of results of the author’s research and based on collected literature originating from the 1960s, the study proves that a dwarfing trend has emerged among middle-European bees.
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The loss of honey bees has drawn a large amount of attention in various countries. Therefore, the development of efficient methods for recovering honey bee populations has been a priority for beekeepers. Here we present an extended literature review and report on personal communications relating to the characterization of the local and bred stock of honey bees in the Russian Federation. New types have been bred from local colonies (A. mellifera L., A. m. carpatica Avet., A. m. caucasia Gorb.). The main selection traits consist of a strong ability for overwintering, disease resistance and different aptitudes for nectar collection in low and high blooming seasons. These honey bees were certified by several methods: behavioral, morphometric and genetic analysis. We illustrate the practical experience of scientists, beekeepers and breeders in breeding A. mellifera Far East honey bees with Varroa and tracheal mite resistance, which were the initial reasons for breeding the A. mellifera Far Eastern breed by Russian breeders, Russian honey bee in America, the hybrid honey bee in Canada by American breeders, and in China by Chinese beekeepers. The recent achievements of Russian beekeepers may lead to the recovery of beekeeping areas suffering from crossbreeding and losses of honey bee colonies.
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The aim of this study was to investigate the diversification of morphological features of the Dark European honey bee of the Augustow M line. The authors studied the proboscis length and cubital index, as features determining the affiliation to the species; the width of tergite 4 and the sum of widths of tergites 3 + 4, as indicators of the bee body size; and the length and width of the right forewing. They compared bees sampled from (1) the “lead apiary”, (2) “associate apiaries” and (3) “conservation area apiaries”—apiaries situated in the conservation area established by the national program for the conservation of genetic resources of this bee line. The conclusion was that it is possible to protect bees of the Augustow M line under the existing program, based on resources available to the lead, associate and conservation area apiaries. The bees studied have the essential features of the Dark European honey bee and the values of parameters tested are consistent with the morphological feature references valid for Apis m. mellifera. On the other hand, based on the authors’ research and on other studies described in literature of 1960s, there is a dwarfing trend in the Dark European honey bee of the Augustow M line.
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Dar, S.A., Dukku, U.H., Ilyasov, R.A., Kandemir, I., Lee, M.L., Özkan Koca, A. 2019. Chapter 4. The classic taxonomy of Asian and European honey bees. 107-137 pp. In: R.A. Ilyasov and H.W. Kwon. [eds.]. Phylogenetics of Bees. CRC Press, Taylor and Francis Group, Boca Raton, London, New-York, USA. 290 pp. ISBN 9781138504233. Abstract. Honey bees of the genus Apis, belonging to the family Apidae and the superfamily Apoidea in the order of insects Hymenoptera. The number of Apis species and their identification methods are discussed. According to different authors, the number of species of the genus varied from 6 to 24. While Apis mellifera inhabits West Asia, Africa and Europe, the ranges of all other species, including Apis cerana, are limited to Asia. A. mellifera and A. cerana are two species widely used in agriculture for the pollination, the production of honey and other products. They have adapted to wide climatic conditions. Intraspecific taxonomy for both species is incomplete and contradictory. In this review, all available studies of A. mellifera and A. cerana are analyzed to ordering the modern taxonomy of honey bees. We found that there are 27 subspecies for A. mellifera and 7 subspecies for A. cerana. However, these data are not ultimate, since some subspecies of A. mellifera and A. cerana remain unexplored.
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Diseases are known to be one of the major contributors to colony losses. Within a Europe-wide experiment on genotype - environment interactions, an initial 621 colonies were set up and maintained from 2009 to 2012. The colonies were monitored to investigate the occurrence and levels of key pathogens. These included the mite Varroa destructor (mites per 10 g bees), Nosema spp. (spore loads and species determination), and viruses (presence/absence of acute bee paralysis virus (ABPV) and deformed wing virus (DWV)). Data from 2010 to the spring of 2011 are analysed in relation to the parameters: genotype, environment, and origin (local vs. non-local) of the colonies in the experiment. The relative importance of different pathogens as indicators of colony death within the experiment is compared. In addition, pathogen occurrence rates across the geographic locations are described.
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Honey bee colonies exhibit a wide range of variation in their behaviour, depending on their genetic origin and environmental factors. The COLOSS Genotype-Environment Interactions Experiment gave us the opportunity to investigate the phenotypic expression of the swarming, defensive and hygienic behaviour of 16 genotypes from five different honey bee subspecies in various environmental conditions. In 2010 and 2011, a total of 621 colonies were monitored and tested according to a standard protocol for estimation of expression of these three behavioural traits. The factors: year, genotype, location, origin (local vs. non-local) and season (only for hygienic behaviour) were considered in statistical analyses to estimate their effect on expression of these behaviours. The general outcome of our study is that genotype and location have a significant effect on the analysed traits. For all characters, the variability among locations was higher than the variability among genotypes. We also detected significant variability between the genotypes from different subspecies, generally confirming their known characteristics, although great variability within subspecies was noticed. Defensive and swarming behaviour were each positively correlated across the two years, confirming genetic control of these characters. Defensive behaviour was lower in colonies of local origin, and was negatively correlated with hygienic behaviour. Hygienic behaviour was strongly influenced by the season in which the test was performed. The results from our study demonstrate that there is great behavioural variation among different subspecies and strains. Sustainable protection of local genotypes can be promoted by combining conservation efforts with selection and breeding to improve the appreciation by beekeepers of native stock.
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Adaptation of honey bees to their environment is expressed by the annual development pattern of the colony, the balance with food sources and the host -parasite balance, all of which interact among each other with changes in the environment. In the present study, we analyse the development patterns over a period of two years in colonies belonging to 16 different genotypes and placed in areas grouped within six environmental clusters across Europe. The colonies were maintained with no chemical treatment against varroa mites. The aim of the study was to investigate the presence of genotype -environment interactions and their effects on colony development, which we use in this study as a measure of their vitality. We found that colonies placed in Southern Europe tend to have lower adult bee populations compared to colonies placed in colder conditions, while the brood population tends to be smaller in the North, thus reflecting the shorter longevity of bees in warmer climates and the shorter brood rearing period in the North. We found that both genotype and environment significantly affect colony development, and that specific adaptations exist, especially in terms of adult bee population and overwintering ability. 234 Hatjina and Costa et al.
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An international collaborative experiment was run from 2009 to 2012 (Costa et al., 2012) with the aim of understanding genotype-environment effects on survival and health status of honey bee colonies headed by queens of different European origins that were tested in various locations under differing environmental conditions. No chemical treatment against bee diseases was performed on any of the test colonies. The colonies were managed according to a common protocol, which included an assessment of their health status (for details, see Meixner et al., 2014). One apiary located in Chalkidiki, Greece, was selected for a detailed case study analysis of parasites and pathogens, including an absolute quantification of viral titres. The colonies were sampled regularly between autumn 2009 and autumn 2011 and analysed for Varroa destructor and Nosema spp. as described in Meixner et al. (2014). For virus analysis, approximately 30 worker bees from each colony were sampled in March and September of 2010, and April and October of 2011, and analysed for deformed wing virus (DWV), black queen cell virus (BQCV), acute bee paralysis virus (ABPV), Kashmir bee virus (KBV) and Israeli paralysis virus (IAPV). Extraction of RNA, qPCR and quantification of virus titers followed the methods described in Francis et al. (2013), with ABPV, KBV and IAPV assessed together in a single assay (AKI). Viral titres (acute paralysis complex (AKI), DWV and BQCV), Nosema spp. infection levels and V. destructor mite infestation levels during the seven sampling times were compared to the survival status of the colony at the end of the experiment. The data were analysed for patterns, trends, correlations and significant differences. Of the 39 starting colonies, 31 could be sampled for quantitative virus analysis in the spring of 2010. They belonged to the genotypes
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The survival and performance of 597 honey bee colonies, representing five subspecies and 16 different genotypes, were comparatively studied in 20 apiaries across Europe. Started in October 2009, 15.7% of the colonies survived without any therapeutic treatment against diseases until spring 2012. The survival duration was strongly affected by environmental factors (apiary effects) and, to a lesser degree, by the genotypes and origin of queens. Varroa was identified as a main cause of losses (38.4%), followed by queen problems (16.9%) and Nosema infection (7.3%). On average, colonies with queens from local origin survived 83 days longer compared to non-local origins (p < 0.001). This result demonstrates strong genotype by environment interactions. Consequently, the conservation of bee diversity and the support of local breeding activities must be prioritised in order to prevent colony losses, to optimize a sustainable productivity and to enable a continuous adaptation to environmental changes. La influencia del origen genético y su interacción con los efectos del medio ambiente en la supervivencia de las colonias de Apis mellifera L. en Europa Resumen La supervivencia y el rendimiento de 597 colonias de abejas, representando cinco subespecies y 16 genotipos distintos, se estudiaron comparativamente en 20 apiarios en Europa. Iniciado en Octubre de 2009, el 15.7% de las colonia sobrevivieron sin ningún tratamiento 206 Büchler et al.
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The COLOSS GEI (Genotype-Environment Interactions) Experiment was setup to further our understanding of recent honey bee colony losses. The main objective of the GEI experiment was to understand the effects of environmental factors on the vitality of European honey bee genotypes. This paper aims to describe the genetic background and population allocation of the bees used in this experiment. Two wing morphometric and two genetic methods were employed to discriminate bee populations. Classical morphometry of 11 angles on the wings were carried out on 350 bees. Geometric morphometry on 19 wing landmarks was carried out on 381 individuals. DNA microsatellite analysis was carried out on 315 individuals using 24 loci. Allozyme analysis was performed on 90 individuals using six enzyme systems. DNA microsatellite markers produced the best discrimination between the subspecies (Apis mellifera carnica, A. m. ligustica, A. m. macedonica, A. m. mellifera and A. m. siciliana) used in the experiment. Morphometric methods generally showed an intermediate level of discrimination, usually best separating A. m. siciliana and A. m. ligustica from the remaining populations. Allozyme markers lack power to discriminate at the level of individual bees, and given our sample size, also fail to differentiate subspecies. Based on DNA microsatellites, about 69% of the individuals were assigned to the same subspecies as originally declared, and 17% were found to belong to a different subspecies. Fourteen percent of the samples were found to be of mixed origin and could not be assigned to any subspecies with certainty. We further discuss the caveats of the methods and details of the sampled bees, their origins and breeding programmes in their respective locations.
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Here we cover a wide range of methods currently in use and recommended in modern queen rearing, selection and breeding. The recommendations are meant to equally serve as standards for both scientific and practical beekeeping purposes. The basic conditions and different management techniques for queen rearing are described, including recommendations for suitable technical equipment. As the success of breeding programmes strongly depends on the selective mating of queens, a subchapter is dedicated to the management and quality control of mating stations. Recommendations for the handling and quality control of queens complete the queen rearing section. The improvement of colony traits usually depends on a comparative testing of colonies. Standardized recommendations for the organization of performance tests and the measurement of the most common selection characters are presented. Statistical methods and data preconditions for the estimation of breeding values which integrate pedigree and performance data from as many colonies as possible are described as the most efficient selection method for large populations. Alternative breeding programmes for small populations or certain scientific questions are briefly mentioned, including also an overview of the young and fast developing field of molecular selection tools. Because the subject of queen rearing and selection is too large to be covered within this paper, plenty of references are given to facilitate comprehensive studies
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Deep sequencing of viruses isolated from infected hosts is an efficient way to measure population-genetic variation and can reveal patterns of dispersal and natural selection. In this study, we mined existing Illumina sequence reads to investigate single-nucleotide polymorphisms (SNPs) within two RNA viruses of the Western honey bee (Apis mellifera), deformed wing virus (DWV) and Israel acute paralysis virus (IAPV). All viral RNA was extracted from North American samples of honey bees or, in one case, the ectoparasitic mite Varroa destructor. Coverage depth was generally lower for IAPV than DWV, and marked gaps in coverage occurred in several narrow regions (< 50 bp) of IAPV. These coverage gaps occurred across sequencing runs and were virtually unchanged when reads were re-mapped with greater permissiveness (up to 8% divergence), suggesting a recurrent sequencing artifact rather than strain divergence. Consensus sequences of DWV for each sample showed little phylogenetic divergence, low nucleotide diversity, and strongly negative values of Fu and Li's D statistic, suggesting a recent population bottleneck and/or purifying selection. The Kakugo strain of DWV fell outside of all other DWV sequences at 100% bootstrap support. IAPV consensus sequences supported the existence of multiple clades as had been previously reported, and Fu and Li's D was closer to neutral expectation overall, although a sliding-window analysis identified a significantly positive D within the protease region, suggesting selection maintains diversity in that region. Within-sample mean diversity was comparable between the two viruses on average, although for both viruses there was substantial variation among samples in mean diversity at third codon positions and in the number of high-diversity sites. FST values were bimodal for DWV, likely reflecting neutral divergence in two low-diversity populations, whereas IAPV had several sites that were strong outliers with very low FST. This initial survey of genetic variation within honey bee RNA viruses suggests future directions for studies examining the underlying causes of population-genetic structure in these economically important pathogens.
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An international experiment to estimate the importance of genotype-environment interactions on vitality and performance of honey bees and on colony losses was run between July 2009 and March 2012. Altogether 621 bee colonies, involving 16 different genetic origins of European honey bees, were tested in 21 locations spread in 11 countries. The genetic strains belonged to the subspecies A. m. carnica, A. m. ligustica, A. m. macedonica, A. m. mellifera, A. m. siciliana. At each location, the local strain of bees was tested together with at least two “foreign” origins, with a minimum starting number of 10 colonies per origin. The common test protocol for all the colonies took into account colony survival, bee population in spring, summer and autumn, honey production, pollen collection, swarming, gentleness, hygienic behaviour, Varroa destructor infestation, Nosema spp.infection and viruses. Data collection was performed according to uniform methods. No chemical treatments against Varroa or other diseases were applied during the experiment. This article describes the details of the experiment set-up and the work protocol.