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The ectoparasitic mite, Varroa destructor , and the viruses that it transmits, kill the colonies of European honey bees Apis mellifera kept by beekeepers unless the bees are treated with miticides.
Nevertheless, there exist populations of wild colonies of European honey bees that are persisting without being treated with miticides.
We hypothesized that the persistence of these wild colonies is due in part to their habits of nesting in small cavities and swarming frequently.
We tested this hypothesis by establishing two groups of colonies living either in small hives 42 L without swarm-control treatments or in large hives up to L with swarm-control treatments.
We followed the colonies for two years and compared the two groups with respect to swarming frequency, Varroa infesttion rate, disease incidence, and colony survival.
Colonies in small hives swarmed more often, had lower Varroa infestation rates, had less disease, and had higher survival compared to colonies in large hives.
These results indicate that the smaller nest cavities and more frequent swarming of wild colonies contribute to their persistence without mite treatments.
Citation: Loftus JC, Smith ML, Seeley TD How Honey Bee Colonies Survive in the Wild: Testing the Importance of Small Nests and Frequent Swarming.
PLoS ONE 11 3 : e Received: November 9, ; Accepted: February 12, ; Published: March 11, This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Data Availability: Data hes been placed on Cornell University's digital repository "eCommons". Funding: This study was supported by the Eastern Apiculture Society of North America, the National Institute of Food and Agriculture Project No.
NYC , and a National Science Foundation Graduate Research Fellowship DGE to MLS. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist. In recent decades, beekeepers worldwide have faced numerous challenges in maintaining healthy honey bee Apis mellifera colonies [ 1 ].
A variety of factors have contributed to the elevated rate of colony mortality, but perhaps the most significant is the introduction from Asia of the ectoparasitic mite, Varroa destructor , which acts as an efficient vector of the viruses of honey bees [ 3 — 5 ].
The rate of colony loss in Europe and North America nearly tripled after the arrival of Varroa in the s and s [ 6 ].
These mites have introduced a new viral transmission route that has altered the viral landscape and caused a massive loss of diversity in Deformed Wing Virus DWV [ 7 ], the pathogen that is linked with the demise of honey bee colonies [ 8 ].
Without treatments for Varroa , managed honey bee colonies almost always die within two or three years [ 9 , 10 ]. Even though Varroa infestations lead to the deaths of honey bee colonies managed by beekeepers unless they are given mite-control treatments, several investigators have reported populations of European honey bee colonies living in the wild that have persisted without mite-control treatments, despite being infested with Varroa Brazil [ 11 ], Russia [ 12 ], Sweden [ 10 ], France [ 13 ], and United States [ 14 ].
In all of these populations, selective pressures by the mites and viruses have probably produced genetic changes in the bees that give them intrinsic resistance to these parasites and pathogens.
We know, for example, that the population of wild colonies in the Arnot Forest in the U. However, there may also be environmental factors that are making it possible for wild colonies to survive mite infestations without mite treatments, when managed colonies cannot.
We hypothesized that the relatively small nest cavities of wild colonies might partially explain their greater ability to survive Varroa infestations without treatments.
In North America, wild honey bees occupy tree cavities with volumes of 30 to 60 L [ 16 ], whereas managed colonies are usually housed in hives with volumes of to L so that they have sufficient room to create large honey stores for beekeepers to harvest.
Because wild colonies live in small nest cavities, which are conducive to swarming [ 17 ], and because they are not subject to beekeeping practices for swarm control, wild colonies probably swarm more often than managed colonies.
We also hypothesized that more frequent swarming by wild honey bee colonies, together with their reduced brood rearing because they have smaller nests , hinders Varroa reproduction and so makes these wild colonies less vulnerable to the mites and to the diseases they spread.
Varroa depends on honey bee brood for reproduction, so this broodless period may help further shrink the Varroa population in a colony that has swarmed.
To test the hypothesis that small nest cavities contribute to the ability of wild colonies to persist without Varroa treatments, we performed an experiment that compared two groups of colonies.
In one group, the colonies lived in small 42 L hives and were left alone. These were our "small-hive colonies," which simulated wild colonies of honey bees.
In the other group, the colonies lived in large hives up to L and were managed in ways that reduced their swarming and maximized their honey production: queen cells were removed periodically and colonies were given two deep hive bodies for a brood chamber plus another two deep hive bodies "honey supers" for honey storage.
These were our "large-hive colonies", which simulated typical managed colonies of honey bees. We monitored the brood and adult bee populations, mite infestation rates, incidences of disease, occurrences of swarming, honey production, and survival of the colonies in both groups over a two-year period May —April We predicted that the small-hive colonies would experience more frequent swarming, lower Varroa infestation rates, lower incidences of disease, lower honey production, and higher colony survival than the large-hive colonies.
The site consisted of a field with two mowed areas for two apiaries: one for the small-hive colonies and one for the large-hive colonies.
The two apiaries were spaced 60 m apart, center to center. Each apiary had open land to the east, south, and west, and thus received good sun exposure.
And each had a windbreak to the north, either a storage building or a grove of spruce trees, and thus was well sheltered.
Also, each apiary contained six hive stands for pairs of hives, with each pair separated from its neighboring pair by 4 m. On 22 May , we installed in both apiaries 12 nucleus colonies in 5-frame hives.
Each nucleus colony's hive contained 5 full-depth Langstroth frames 48 x 23 cm : 2 frames of comb—one filled with brood, one partially filled with pollen and honey—covered with adult bees, 1 frame of comb filled with honey but without bees, 1 frame of empty comb, and 1 frame of beeswax comb foundation.
We obtained the frames of bees and brood for the 24 nucleus colonies from 12 source colonies living in an apiary 4. We took 4 frames of bees and brood from each source colony, so each source colony provided the bees and brood for one colony in both the small-hive and the large-hive treatment groups.
This ensured that the two treatment groups started out with the same average Varroa infestation rate of adult bees. Each nucleus colony was given an open-mated Italian queen bee purchased from Olivarez Honey Bees, Inc Chico, California and all 24 queens were accepted.
All were then marked with a dot of yellow paint on the thorax. On 5 June , we transferred all the colonies in both apiaries from their 5-frame hives into frame Langstroth hives, each of which had a volume of 42 L.
Adjacent colonies were given different colored hives, to minimize drifting of workers and drones between colonies. We also installed an entrance reducer in each hive so each one had a small, cm 2 -entrance opening.
On 5 July , we inspected the colonies and gave additional bees or brood, or both, to the three smallest colonies in each group, to bring all the colonies up to the same strength.
Specifically, we gave one colony in each treatment group 2 frames that were filled with capped brood but were not covered with worker bees, and we gave 2 colonies in each group 2 frames that were both filled with capped brood and covered with worker bees.
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