Varroa mite | |
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Varroa destructor adult female in dorsal (top) and ventral (lower) views | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Chelicerata |
Class: | Arachnida |
Order: | Mesostigmata |
Family: | Varroidae |
Genus: | Varroa |
Species: | V. destructor |
Binomial name | |
Varroa destructor Anderson & Trueman, 2000 [1] | |
Varroa destructor, the Varroa mite, is an external parasitic mite that attacks and feeds on honey bees and is one of the most damaging honey bee pests in the world. [2] [3] A significant mite infestation leads to the death of a honey bee colony, usually in the late autumn through early spring. Without management for Varroa mite, honey bee colonies typically collapse within 2 to 3 years in temperate climates. [4] These mites can infest Apis mellifera , the western honey bee, and Apis cerana , the Asian honey bee. Due to very similar physical characteristics, this species was thought to be the closely related Varroa jacobsoni prior to 2000, but they were found to be two separate species after DNA analysis.
Parasitism of bees by mites in the genus Varroa is called varroosis. The Varroa mite can reproduce only in a honey bee colony. It attaches to the body of the bee and weakens the bee. [5] The species is a vector for at least five debilitating bee viruses, [5] including RNA viruses such as the deformed wing virus (DWV). The Varroa mite is the parasite with possibly the most pronounced economic impact on the beekeeping industry and is one of multiple stress factors contributing to the higher levels of bee losses around the world. [6] Varroa mite has also been implicated as one of the multiple causes of colony collapse disorder.
Management of this pest focuses on reducing mite numbers through monitoring to avoid significant hive losses or death. 3% of bees infested in a hive is considered an economic threshold where damage is high enough to warrant additional management. Miticides are available, though some are difficult to time correctly while avoiding harm to the hive, and resistance has occurred for others. Screened bottom boards on hives can be used for both monitoring and mite removal, and drone comb that mites prefer can be used as a trap to remove mites from the hive. Honey bee lines in breeding programs also show partial resistance to Varroa mite through increased hygienic behavior that is being incorporated as an additional management strategy.
The adult female mite is reddish-brown in color, while the male is white. Varroa mites are flat, having a button shape. They are 1–1.8 mm long and 1.5–2 mm wide, and have eight legs. [7] Varroa mites lack eyes. [8] These mites have curved bodies that allow them to fit between the abdominal segments of adult bees. [9]
Host bee species can help differentiate mite species in the genus Varroa ; both V. destructor and Varroa jacobsoni parasitize Apis cerana , the Asian honey bee, but the closely related mite species originally described as V. jacobsoni by Anthonie Cornelis Oudemans in 1904 does not attack Apis mellifera , the western honey bee, unlike V. destructor. Until 2000, V. destructor was thought to be V. jacobsoni and resulted in some mislabeling in the scientific literature. [1] [10] The two species cannot be easily distinguished with physical traits and have 99.7% similar genomes, [11] so DNA analysis is required instead. [1] [12] Because the more virulent and damaging species V. destructor could not be distinguished at the time, most pre-2000 research on western honey bees that refers to V. jacobsoni was actually research on V. destructor. [4]
Other Varroa species V. underwoodi and V. rindereri can also parasitize honey bee species and can be distinguished from V. destructor and V. jacobsoni with slight differences in body size and setae characteristics, though each of the four species within the Varroa genus have similar physical characteristics. [13] [14] If a Varroa species is found on a western honey bee, it will typically be V. destructor except where V. underwoodi is present, such as in Papua New Guinea. [14]
The name "Varroa mite" is typically used as the common name for V. destructor after the species was considered separate from V. jacobsoni. [9]
Mite species | Bee host |
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Varroa destructor | western honey bee, Asian honey bee |
Varroa jacobsoni | Asian honey bee |
Varroa rindereri | Apis koschevnikovi |
Varroa underwoodi | western honey bee, Asian honey bee, Apis nigrocincta , Apis nuluensis |
Varroa mite has two distinct genetic strains from when it switched hosts from the Asian honey bee to the western honey bee: Korean and Japanese. The Korean strain that occurred in 1952 is now found worldwide in high frequencies, while the Japanese strain that started around 1957 occurs in similar areas at much lower frequencies. [11] Varroa mite has low genetic diversity, which is typical for an invasive species undergoing a range or host expansion. [15]
Varroa mites originally only occurred in Asia on the Asian honey bee, but this species has been introduced to many other countries on several continents, resulting in disastrous infestations of European honey bees. [16]
Introduction data prior to 2000 is unclear due to confusion with V. jacobsoni. By 2020, V. destructor was confirmed to be present throughout North America excluding Greenland, South America, most of Europe and Asia, and portions of Africa. The species was not present in Australia as well as Oman, Congo, Democratic Republic of Congo, and Malawi. It was suspected to not be present in Sudan and Somalia. [17] [18] Mites were found in 2022 in New South Wales in Australia. [19]
Female mites enter brood cells to lay eggs on the comb wall after the cell is capped. Eggs are approximately 0.2 to 0.3 mm in diameter and cannot be seen without magnification. These eggs hatch into male and female protonymphs that are both transparent white. Immature mites can only feed on capped brood, so the life cycle cannot be completed during broodless periods. Protonymphs molt into deuteronymphs that more closely resemble the curved body of adults before they molt into adults. Development time from egg to adult is 6–7 days. Males will not leave brood cells and only mate with females present in the brood cell. [9]
Adult females can be found feeding both on brood and adult bees. After reaching the adult stage, females will leave the brood cell and enter a phoretic stage where mites attach to adult bees in order to disperse. Mites will feed on adult bees at this time and can be transmitted from bee to bee during this stage. Nurse bees are preferred hosts in order to be moved to new brood cells. Because the nurse bee spends more time around the drone brood (i.e., male bees) rather than the worker brood, many more drones are infected with the mites. [4] These phoretic females can also be transmitted to other hives through bee contact or hive equipment transfer. The phoretic stage can last for 4.5–11 days during brood production periods or up to five to six months when no brood is present in winter months. Female mites have a life expectancy of 27 days when brood is present. [9]
After the phoretic stage, female mites leave the adult bee and enter brood cells with bee larvae. Drone cells are preferred over workers. These females are called foundress mites, and they bury themselves in brood food provided by worker bees before the cell is capped. Brood cell capping begins egg cell activation for a foundress mite while she emerges to feed on the larva. [11] She will lay a single unfertilized egg after feeding to produce a male mite. After laying this egg, fertilized eggs to produce females are laid approximately once a day. Both the mother and nymphs will feed on the developing pupa. Unless multiple foundress mites are present in a cell, mating occurs between siblings when they reach the adult stage. Once females mate, they are unable to receive additional sperm. [9] Varroa mite's genetic bottleneck is also likely due to its habit of sibling mating. [11]
Adult mites feed on both adult bees and bee larvae by sucking on the fat body, an insect organ that stores glycogen and triglycerides with tissue abundant under epidermis and the surrounding internal body cavity. [5] [20] [21] As the fat body is crucial for many bodily functions such as hormone and energy regulation, immunity, and pesticide detoxification, the mite's consumption of the fat body weakens both the adult bee and the larva. Feeding on fat body cells significantly decreases the weight of both the immature and adult bee. Infested adult worker bees have a shorter lifespan than ordinary worker bees, and they furthermore tend to be absent from the colony far more than ordinary bees, which could be due to their reduced ability to navigate or regulate their energy for flight. [4] [22] [23] Infested bees are more likely to wander into other hives and further increase spread. Bees will occasionally drift into other nearby hives, but this rate is higher for Varroa infested bees. [9] [24]
Adult mites live and feed under the abdominal plates of adult bees primarily on the underside of the abdominal region on the left side of the bee. Adult mites are more often identified as present in the hive when on top of the adult bee on the thorax, but mites in this location are likely not feeding, but rather attempting to transfer to another bee. [5]
Varroa mites have been found on flowers visited by worker bees, which may be a means by which phoretic mites spread short distances when other bees, including from other hives, visit. [25] [26] They have also been found on larvae of some wasp species, such as Vespula vulgaris , and flower-feeding insects such as the bumblebee, Bombus pensylvanicus , the scarab beetle, Phanaeus vindex , and the flower-fly, Palpada vinetorum . There have not been any indications Varroa mites are able to complete their life cycle on these insects, but instead they become distributed to other areas while a mite is still alive on these insects. [27]
Open wounds left by the feeding become sites for disease and virus infections. The mites are vectors for at least five and possibly up to 18 debilitating bee viruses, [5] including RNA viruses such as the deformed wing virus.
Prior to the widespread introduction of Varroa mite, honey bee viruses were typically considered a minor issue. Virus particles are directly injected into the bee's body cavity and mites can also cause immunosuppression that increases infection in host bees. Varroa mites can transmit the following viruses: [4]
Deformed wing virus is one of the most prominent and damaging honey bee viruses transmitted by Varroa mites. It causes crumpled deformed wings that resemble sticks and also causes shortened abdomens. [4] [28]
There is some evidence that harm from both Varroa mite and associated viruses they transmit may be a contributing factor that leads to colony collapse disorder (CCD). [2] While the exact causes of CCD are not known, infection of colonies from multiple pathogens and interaction of those pathogens with environmental stresses is considered by entomologists to be one of the likely causes of CCD. [29] [30] Most scientists agree there is not a single cause of CCD. [31]
Mite populations undergo exponential growth when bee broods are available, and exponential decline when no brood is available. In 12 weeks, the number of mites in a western honey bee hive can multiply by roughly 12. Mites often invade colonies in the summer, leading to high mite populations in autumn. [32] High mite populations in the autumn can cause a crisis when drone rearing ceases and the mites switch to worker larvae, causing a quick population crash and often hive death. [33] Various management methods are used for Varroa mite integrated pest management to monitor and manage damage to hives.
Beekeepers use several methods for monitoring levels of Varroa mites in a colony. [34] They involve either estimating the total number of mites in a hive by using a sticky board under a screen bottom board to capture mites falling from the hive or estimating the number of mites per bee with powdered sugar or an ethanol wash. [35]
Monitoring for mites with a sticky board can be used to estimate the total number of mites in a colony over 72 hours using the equation:
where b is the number of mites found on the sticky board and c is the number of estimated mites in the colony. However, the bee population in a colony also needs to be known to determine what population of mites is tolerable with this method. [35]
Mite counts from a known quantity of bees (i.e., 300 bees) collected from brood comb are instead often used to determine mite severity. Mites are dislodged from a sample of bees using non-lethal or lethal means. The bees are shaken in a container of either powdered sugar, alcohol, or soapy water to dislodge and count mites. Powdered sugar is generally considered non-lethal to honey bees, but lethal methods such as alcohol can be more effective at dislodging mites. [36] [35] 3% of the colony being infested is considered an economic threshold damaging enough to warrant further management such as miticides, though beekeepers may use other management tactics in the 0–2% infestation range to keep mite populations low. [35]
Varroa mites can be treated with commercially available acaricides that must be timed carefully to minimize the contamination of honey that might be consumed by humans. The four most common synthetic pesticides used for mite treatments with formulations specific for honey bee colony use are amitraz, coumaphos, and two pyrethroids, flumethrin and tau-fluvalinate, while naturally occurring compounds include formic acid, oxalic acid, essential oils such as thymol and beta acids from hops resin (e.g. lupulone). Many of these products whether synthetic or naturally produced can negatively affect honey bee brood or queens. These products often are applied through impregnated plastic strips or as powders spread between brood frames. [35]
Synthetic compounds often have high efficacy against Varroa mites, but resistance has occurred for all of these products in different areas of the world. Pyrethroids are used because a concentration that will kill mites has relatively low toxicity to honey bees. [35] Compounds derived from plants have also been assessed for mite management. Thymol is one essential oil with efficacy against mites, but can be harmful to bees at high temperatures. Other essential oils such as garlic, oregano, and neem oil have had some efficacy in field trials, though most essential oils that have been tested have little to no effect. Essential oil use is widespread in hives with many of those uses being off-label or in violation of pesticide regulations in various countries. Hop beta acids are lupulones obtained from hop plants and have been used in products marketed for mite control. [35]
Chemical | Efficacy | Notes |
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Amitraz | High (75–90%) | Not very affordable; slower occurrence of resistance |
Coumaphos | Low | Use decreased over time due to low efficacy and resistance |
Flumethrin | High (73–97%) | Negative effects to honey bees less severe than tau-fluvalinate |
Tau-fluvalinate | Low | Efficacy lost due to resistance; reduced brood survival and queen size |
Formic acid | High (35–75%) | Efficacy varies based on temperature, brood population, and proximity to the chemical within the hive; can cause brood or queen mortality |
Oxalic acid | High (near 100%) | Used during broodless periods only and increases grooming behavior; no known cases of resistance |
Thymol | Moderate (50–80%) | Similar temperature-based issues to formic acid, not effective under 15 °C |
Hop beta acids | Moderate (43–88%) | Low toxicity to humans and bees |
Resistance to pyrethroids has occurred in the Czech Republic and the UK due to a single amino acid substitution on Varroa mite's genome. Underlying mechanisms for resistance in other pesticides, such as coumaphos, are still unknown. [37]
Varroa mites can also be controlled through nonchemical means. Most of these controls are intended to reduce the mite population to a manageable level, not to eliminate the mites completely. [35]
Screened bottom boards are used both for monitoring and can modestly reduce mite populations by 11–14%. Mites which fall from the comb or bees can land outside the hive instead landing on a solid bottom board that would allow them to easily return to the nest. [35]
Varroa infest drone cells at a higher rate than worker brood cells, so drone cells can be used as a trap for mite removal. Beekeepers can also introduce a frame with drone foundation cells that encourage bees to construct more drone cells. When the drone cells are capped, the frame can be removed to freeze out mites. This labor-intensive process can reduce mite levels by about 50–93%, but if trap cells are not removed early enough before mites emerge, mite populations can spike. This method is only viable in spring and early summer when drones are produced. [35]
Heat is also sometimes used as a control method. The mites cannot survive temperatures near 40 °C (104 °F), but brief exposure to these temperatures do not harm honey bees. Devices are marketed intended to heat brood to these temperatures, though the efficacy of many of these products has not been reviewed. [35] [38]
Powdered sugar used for estimating mite counts in hives has also been considered for mite management as it or other inert dusts were believed to initiate grooming responses. Long-term studies do not show any efficacy for reducing mite populations. [35]
The Asian honey bee, is more hygienic with respect to Varroa mite than western honey bees, which is in part why mite infestations are more pronounced in western honey bee colonies. Efforts also have been made to breed hygienic honey bees heritable behavior traits, such as those with resistance to Varroa mites. Honey bee lines with resistance include Minnesota Hygienic Bees, Russian Honey Bees, and Varroa sensitive hygiene. [39] [40] [41]
Hygienic behaviors include [39] [42] workers removing pupae heavily infested with mites, which kills both the developing bee and immature mites, and grooming or removal from the brood cell, which increases adult mite mortality. Mites removed from host pupae are at an incorrect life stage to re-infest another pupa. An extended phoretic period in adult female mites has also been noticed.
Hygienic behavior is effective against diseases such as American foulbrood or chalkbrood, but the efficacy of this behavior against mites is not well-quantified; colonies with this behavior alone do not necessarily result in Varroa mite resistant colonies that can survive without miticide treatments. The efficacy of this behavior can vary between bee lines in comparison studies with Minnesota hygienic bees removing 66% of infested pupae, while Varroa sensitive hygiene bees removed 85% of infested pupae. There are minimal trade-off costs to hives that have this hygienic behavior, so it is being actively pursued in bee breeding programs. [39]
Researchers have been able to use RNA interference by feeding honey bees mixtures of double-stranded RNA that target expression of several Varroa mite genes, such as cytoskeleton arrangement, transfer of energy, and transcription. This can reduce infestation to 50% without harm to honey bees and is being pursued as an additional control method for Varroa mite. [43] [44]
A honey bee is a eusocial flying insect within the genus Apis of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmopolitan distribution of honey bees, introducing multiple subspecies into South America, North America, and Australia.
The Apis mellifera mellifera is a subspecies of the western honey bee, evolving in central Asia, with a proposed origin of the Tien Shan Mountains and later migrating into eastern and then northern Europe after the last ice age from 9,000BC onwards. Its original range included the southern Urals in Russia and stretched through northern Europe and down to the Pyrenees. They are one of the two members of the 'M' lineage of Apis mellifera, the other being in western China. Traditionally they were called the Black German Bee, although they are now considered endangered in Germany. However today they are more likely to be called after the geographic / political region in which they live such as the British Black Bee, the Native Irish Honey Bee, the Cornish Black Bee and the Nordic Brown Bee, even though they are all the same subspecies, with the word "native" often inserted by local beekeepers, even in places where the bee is an introduced foreign species. It was domesticated in Europe and hives were brought to North America in the colonial era in 1622 where they were referred to as the English Fly by the Native Americans.
THYMOVAR is a product to control the Varroa mite on bees and contains the essential oil thymol.
Nosema apis is a microsporidian, a small, unicellular parasite recently reclassified as a fungus that mainly affects honey bees. It causes nosemosis, also called nosema, which is the most common and widespread of adult honey bee diseases. The dormant stage of N. apis is a long-lived spore which is resistant to temperature extremes and dehydration, and cannot be killed by freezing the contaminated comb. Nosemosis is a listed disease with the Office International des Epizooties (OIE).
The Russian honeybee refers to honey bees that originate in the Primorsky Krai region of Russia. This strain of bee was imported into the United States in 1997 by the USDA Agricultural Research Service's Honeybee Breeding, Genetics & Physiology Laboratory in Baton Rouge, Louisiana, in response to severe declines in bee populations caused by infestations of parasitic mites, and has been used in breeding programs to improve existing stocks. Many Russian queens openly mate with drones from various stock, creating colonies that are genetically hybrid. Some of these 'uncontrolled' hybrids may exhibit "increased aggressiveness, reduced honey production and a decrease in their ability to withstand mites and detrimental expressions of other traits as well."
Varroa is a genus of parasitic mesostigmatan mites associated with honey bees, placed in its own family, Varroidae. The genus was named for Marcus Terentius Varro, a Roman scholar and beekeeper. The condition of a honeybee colony being infested with Varroa mites is called varroosis.
Varroa jacobsoni is a species of mite that parasitises Apis cerana. The more damaging Varroa destructor was previously included under the name V. jacobsoni, but the two species can be separated on the basis of the DNA sequence of the cytochrome oxidase I gene in the mitochondrial DNA.
Aethina tumida,commonly known as small hive beetle (SHB), is a beekeeping pest. It is native to sub-Saharan Africa, but has spread to many other regions, including North America, Australia, and the Philippines.
Deformed wing virus (DWV) is an RNA virus, one of 22 known viruses affecting honey bees. While most commonly infecting the honey bee, Apis mellifera, it has also been documented in other bee species, like Bombus terrestris, thus, indicating it may have a wider host specificity than previously anticipated. The virus was first isolated from a sample of symptomatic honeybees from Japan in the early 1980s and is currently distributed worldwide. It is found also in pollen baskets and commercially reared bumblebees. Its main vector in A. mellifera is the Varroa mite. It is named after what is usually the most obvious deformity it induces in the development of a honeybee pupa, which is shrunken and deformed wings, but other developmental deformities are often present.
The western honey bee or European honey bee is the most common of the 7–12 species of honey bees worldwide. The genus name Apis is Latin for 'bee', and mellifera is the Latin for 'honey-bearing' or 'honey-carrying', referring to the species' production of honey.
Colony collapse disorder (CCD) is an abnormal phenomenon that occurs when the majority of worker bees in a honey bee colony disappear, leaving behind a queen, plenty of food, and a few nurse bees to care for the remaining immature bees. While such disappearances have occurred sporadically throughout the history of apiculture, and have been known by various names, the syndrome was renamed colony collapse disorder in early 2007 in conjunction with a drastic rise in reports of disappearances of western honey bee colonies in North America. Beekeepers in most European countries had observed a similar phenomenon since 1998, especially in Southern and Western Europe; the Northern Ireland Assembly received reports of a decline greater than 50%. The phenomenon became more global when it affected some Asian and African countries as well. From 1990 to 2021, the United Nation’s FAO calculated that the worldwide number of honeybee colonies increased 47%, reaching 102 million.
Apis cerana, the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of Apis koschevnikovi and both are in the same subgenus as the western (European) honey bee, Apis mellifera. A. cerana is known to live sympatrically along with Apis koschevnikovi within the same geographic location. Apis cerana colonies are known for building nests consisting of multiple combs in cavities containing a small entrance, presumably for defense against invasion by individuals of another nest. The diet of this honey bee species consists mostly of pollen and nectar, or honey. Moreover, Apis cerana is known for its highly social behavior, reflective of its classification as a type of honey bee.
Varroa sensitive hygiene (VSH) is a behavioral trait of honey bees (Apis mellifera) in which bees detect and remove bee pupae that are infested by the parasitic mite Varroa destructor. V. destructor is considered to be the most dangerous pest problem for honey bees worldwide. VSH activity results in significant resistance to the mites.
Beekeeping is first recorded in Ireland in the seventh century. It has seen a surge in popularity in modern times, with the membership of beekeeping associations exceeding 4,500. The median average number of hives per beekeeper is three hives, while the average honey output per hive is 11.4 kg. The growth in the practice has occurred despite increased pressures on bees and beekeepers due to parasites, diseases and habitat loss.
Chronic bee paralysis virus (CBPV) commonly affects adult Apis mellifera honey bees and causes a chronic paralysis that can easily spread to other members of a colony. Bees infected with CBPV begin to show symptoms after 5 days and die a few days after. Chronic bee paralysis virus infection is a factor that can contribute to or cause the sudden collapse of honeybee colonies. Since honeybees serve a vital role in ecological resilience, it is important to understand factors and diseases that threaten them.
Slow bee paralysis virus (SBPV) is a virus discovered in England in 1974 that infects honeybees, bumblebees, and silkworms through Varroa destructor mite infestations. The virus causes paralysis in the front two pairs of legs of adult bees eventually killing its hosts. The virus is in the iflaviridae family of viruses. Infection by iflaviridae viruses is among the leading cause of death of honeybee colonies. As bees and silkworms are of great economic and biological importance, the virus is the subject of ongoing research.
Mite Biting Is one of the behavioral mechanisms of honey bees used to fight off the ectoparasitic mites Varroa destructor. This behavior has been studied since the late 1990s for honey bee breeding and improvement of honeybee stocks towards mite resistance. Krispn Given and Dr. Greg Hunt at Purdue University started a hierarchical selective breeding program in 1997–present for increased mite-biting and grooming behavior of European honey bee. A group of Midwest bee breeders visiting the Purdue bee lab were inspired to start the Heartland Honey Bee Breeders Cooperative as a result of their pioneering work.
Apis mellifera adansonii(Western African bee) is a subspecies of the Western honey bee with probably the largest range of Apis mellifera in Africa, belonging to the A (Africa) Lineage of honey bees. Originally identified by Michael Adansonin in his Histoire naturelle du Seneegal in 1757. Initially the name adsansonii was misapplied to A. m. scutelleta and in particular to the Africanised bees of South America.
Apis mellifera syriaca is known by the common name of the Syrian honey bee, sometimes also called the Palestine honey bee.
The fat body is a relatively large organ distributed throughout the insect body, preferentially underneath the integument and surrounding the gut and reproductive organs
In the autumn though, brood rearing declines and the number of multiple-infested cells increases.