Small hive beetle

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Small hive beetle
Small hive beetle.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Infraorder: Cucujiformia
Family: Nitidulidae
Genus: Aethina
Species:
A. tumida
Binomial name
Aethina tumida
Murray, 1867

The small hive beetle (Aethina tumida) is a beekeeping pest. [1] It is native to sub-Saharan Africa, but has spread to many other regions, including North America, Australia, and the Philippines.

Contents

The small hive beetle (SHB) can be a destructive pest of honey bee colonies, causing damage to honey comb, stored honey, and pollen. If a beetle infestation is abundant and concentrated, they may cause bees to abandon their hive. The presence of A. tumida can also be a marker in the diagnosis of colony collapse disorder for honey bees. The beetles can be a pest of stored combs, and honey (in the comb) awaiting extraction. Beetle larvae may tunnel through combs of honey, feeding and defecating, causing discoloration and fermentation of the honey.

Distribution

The small hive beetle was first discovered in the United States in 1996 and has now spread to many U.S. states including California, Connecticut, Florida, Georgia, Hawaii, Iowa, Illinois, Indiana, Kansas, Louisiana, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Nebraska, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas and Virginia.

In the summer of 2015, the discovery of a number of adult beetles as well as one infestation in all stages of brood in British Columbia's Fraser Valley triggered a temporary quarantine. [2] [3]

In Mexico, the small hive beetle has become established in at least eight states. Infestation levels are especially high in tropical areas such as the Yucatán. [4]

The small hive beetle was first detected in Belize in 2016 in the Corozal District. [5]

The small hive beetle has also spread to Australia, being first identified at Richmond, New South Wales, in 2002. Subsequently, it has affected many areas of Queensland and New South Wales. [6] It is possible that the import of package bees, honeybee or bumblebee colonies, queens, hive equipment or even soil constituted the potential invasion pathway of the small hive beetle; however, at the current state of evidence it is still unclear how small hive beetles actually reached Australia. [7]

The small hive beetle has now reached the southern Philippines in southern Mindanao, and there is great concern that it will spread through the country if hives and bees are moved from the southern Mindanao area where the beetle has already been identified. [8]

Small hive beetle was first detected in Calabria southern Italy in September 2014. It is currently confined to the provinces of Reggio Calabria and part of Vibo Valentia, where it has now become naturalized. In 2014 and 2019, outbreaks of infestation occurred in eastern Sicily. [9]

In Australia, the small hive beetle has been detected in every state and territory aside from the Northern Territory, however in Western Australia it is restricted to the Kimberly region [10] and the most recent outbreak in Tasmania was restricted to the Devonport region as of May 2023. [11] [12] Small hive beetle is no longer a notifiable pest in Victoria or South Australia. [13] [14]

Food resources

Small hive beetles are insectivores, and they can be fed on both animal and plant diets. Animal diets include the bee eggs, host body fluids and dead bees. Plant diets include fruit, nectar and pollen. Adults and larvae are fed by the same food. [15]

Description

Small hive beetles are categorized as small insects with length around 5-7mm and width around 2.5-3.5mm. Sexual dimorphism is observed since females are usually longer and heavier than male. The size of the beetles may vary based on environmental factors such as diet, temperature, and humidity. For their body structure, there are three pairs of walking legs, two antennae, and two pairs of wings. Their body is bilaterally symmetrical. They have a wide range of body coloration, from orange-brown to dark brown or black. This species is ectothermic, which means they rely on external heat sources to adjust body temperature. [15]

Younger small hive beetles have a very distinguishable appearance than adults. The beetle eggs are elongated with a white color. The beetle larvae are reduced white color with a segmented body shape. When the exoskeleton develops, the pupae turn to brown color like the adults. [15]

Parental care

Oviposition

Small hive beetles reproduce sexually on a year-round basis and they are polygynandrous, meaning that both males and females mate with multiple partners. The eggs are fertilized externally. Females can oviposit at least 1000 and up to 2000 eggs in her entire life. [1] Females prefer to lay eggs directly onto their food sources. They can also lay eggs in clusters in crevices within the beehive to improve the offspring survival rate. The beetles prefer strong hives for more resources to keep their body temperature during wintertime. On the contrary, they prefer weaker hives to invade and reproduce during the summertime. [15]

Social behavior

Reproduction seasonality

The peak of reproduction period varies in different places. Most small hive beetle populations reach the highest density during the summer and early autumn season with high temperature. Some populations show preference for the rainy season while others prefer the dry season. [16]

Function of pheromones

Pheromones are chemical substances that are released by organisms to impact the behavior of other members within the species. They are very important for group behavior and communication.

The small hive beetle (SHB) is a parasite and scavenger of bee colonies, especially for honeybee colonies, Apis mellifera . The honeybee’s alarm pheromone, especially of European honeybee (EHB), plays a significant role in colony defense against predators and parasites by recruiting more guard bees and attacking the invaders. However, it is shown that the volatiles of pheromone are attractive to SHBs. Therefore, there is a trade-off between recruiting more bees to defend the hive and to attract more attack from additional beetles. [17]

It has been proved that the antennae of SHBs has a lower threshold to sense the component of the volatiles released from the hive entrance, which highly improves the beetle’s ability to invade the colony since the bees are not able to sense and initiate attack at the invader under low pheromone concentration. [17]

The mutualistic relationship between yeast and SHB

It was found that nitidulid beetles can function as the vector for fungi [18] and yeast related to the SHB can release alarm pheromones, which has the same attractive effects as the pheromone produced by EHB. Therefore, yeast can facilitate the beetle’s invasion into the beehive. The bees themselves might also help attract the beetle if they get into contact with the yeast pores released by nitidulids. The volatiles released by bees and the yeast together suggest the availability of the food and attract more beetles, which might lead to the absconding or the complete collapse of the hive. [17]

Small hive beetles aggregation pheromone

Small hive beetles use pheromone-mediated aggregations to counteract the host defense response. [7] It is suggested that the aggregation pheromone is released by males because males infest before females. [7] The pheromone can signal both males and females.

Life history

Aethina tumida was previously known only from the sub-Saharan regions of Africa where it has been considered a minor pest of bees. [19] The life cycle information is known primarily from studies in South Africa.[ citation needed ]

Small hive beetles grow and develop via metamorphosis, which means the organism undergoes several distinct stages to become an adult. Females lay the eggs either in the gaps within the host hive or directly on the fruits or meat. The eggs hatch and become larvae after 1–6 days, but the time highly depends on the environmental humidity. The larvae feed on honey, pollen, bee brood cells and bee eggs. This stage creates the most severe damage toward the host hive as fermentation occurs when the larvae tunnel through the hive. The larval feeding stage generally lasts about 3–10 days and it may vary based on the amount of protein composition in the diet. The beetles then enter the wandering phase where they leave the hive and dig into the soil to pupate by creating a chamber about 5–20 cm beneath the soil. The larvae can aggregate and stay in the host colonies for a period up to 61 days before they pupate. Pupation takes about 2–20 weeks, depending on the temperature, texture, and humidity of the soil. When pupation terminates, the adult beetles emerge and start the mating process after one week. Adults may live up to 6 months and are often found on the rear portion of the bottom board of a hive. [15] [20] [21]

Parasitism

Host response and SHBs' counter-strategy

In response to the beetle’s invasion, the honeybee has developed some adaptive strategies. Firstly, they use guard bees to limit the beetles from entering the hive by attacking the invaders and minimizing the entrance size using propolis. [22]

Patrolling behavior around the nest and combs is well observed in the brood area within the hive. [23] The presence of bees within the colony can limit the number of beetles entering the hive, which is proposed to be due to the protective effects to the colony via the patrolling behavior of the host workers. The presence of bees within the colony can impact the number of beetles’ presence where more bees can limit the existence of beetles. [23]

The third strategy is to remove beetle eggs and larvae. SHB can hide from the patrolling behavior by ovipositing in the combs. To deal with this counter strategy, honeybee workers developed the adaptation to remove the beetle’s eggs and larvae. However, SHBs can still lay clutch of eggs in the crevices to escape from the host’s defense response. [7] Once SHBs get to this invasion stage, they can start the mass reproduction which leads to a completely structural collapse of the entire beehive. [22] Honeybees can either physically prevent beetles to lay clusters of eggs or to detect and remove eggs once laid. The strategy of “social encapsulation” is used to confine the beetles via propolis, which prisons the beetle and limit their activity. [24]

Host colony mobility

Under severe infestation situation, the host honeybee colonies might show an absconding and migration strategy where they would give up their nest and leave the left resources to the beetles. This strategy is usually followed by colony merging. [22]

Vector to the host

Small hive beetles help spread Paenibacillus larvae, which is a bacteria causing the American foulbrood. The infective P. larvae spores are transmitted to the honeybee as well as the honey if the host gets into contact with the infected small hive beetles. [25] Small hive beetle is also a vector for honeybee viruses, including deformed wing virus and sacbrood virus. They can bring the viruses to the honeybee host and cause widespread infection. [26]

Migration

Natural range expansion

Adult SHBs have strong flying ability up to 16 kilometers. [7] Adult beetles emerged immediately from pupation were found not to invade the honeybee hives that are close to their pupation location and they prefer the long distance flights. [7]

Human-associated dispersal

Human activity such as migratory beekeeping and movement of package bees and beekeeping equipment significantly facilitates the quick spread of small hive beetles around the world. It only took two years for the small hive beetles to move across the US. [7]

Enemies

Predators

Entomopathogenic fungi can function as a strong biopesticide for small hive beetles, such as Metarhizium and Beauveria. [27] The ant Pheidole megacephala is the predator specifically to small hive beetle larvae in Kenya. [16]

Competitor

There is no direct competitor with small hive beetles within honeybee colonies. Competition may appear when other organisms occasionally share the food left within the honeybee colonies and other resources with small hive beetles. [23]

Mating

Aggregation and copulation behavior

Copulation starts one week after the adults emerge from the soil and reaches the highest frequency after 18 days. Mating is much more common if beetles are aggregated than if they are in single pairs. [28]

Gender preference

Small hive beetles demonstrate an age-dependent gender preference during copulation. Within the window of 18 days after adults emerge, the male beetles show mating behavior with other males. After 18 days, both genders spend more time to interact and copulate with the opposite sex. [28]

Male-male interaction

The male beetles might interrupt the copulation process of other males. During the interruption, the males fight over each other using their tarsae, antennae and mandibles. [28]

Female-female interaction

The interruption caused by beetle’s pushing behavior during mating process is not only done by males. Instead, a majority of pushing behavior is demonstrated by females towards other members of the same sex during copulation. [28]

Female-male interaction

The first step for a male to approach a female is via thrilling and tapping the female to climb onto its body. Most of the time, the male eventually ceases his action and either demonstrates non-social behavior or leaves the female. If the male chooses to stay, he will thrill female’s pronotum, slide from female’s body and start copulation by inserting its aedeagus. The female can choose either to stay or to escape. If the female stays during the copulation, the male clings onto the female with its legs while rubbing its partner’s elytrae. Mostly, males can keep his position with the females during the interruption, but not during the escape. The male then continues to sit on the female’s back and repeats the entire copulation process. The complete mating behavior finishes when the male descends from the female and demonstrates non-social behavior. [28]

Damage to colonies and stored honey

Comb slimed by hive beetle larvae. Hives infested at this level will drive out bee colonies. Aethina tumida.jpg
Comb slimed by hive beetle larvae. Hives infested at this level will drive out bee colonies.

The primary damage to colonies and stored honey caused by the small hive beetle is through the feeding activity of the larvae. Larvae tunnel through comb with stored honey or pollen, damaging or destroying cappings and comb. Larvae defecate in honey, and the honey becomes discolored from the feces. Activity of the larvae causes fermentation and a frothiness in the honey; the honey develops a characteristic odor of decaying oranges. Damage and fermentation cause honey to run out of combs, creating a mess in hives or extracting rooms. Heavy infestations cause bees to abscond; some beekeepers have reported the rapid collapse of even strong colonies.[ citation needed ]

Control

The small hive beetle is considered a secondary pest in South Africa, and as such, has not been the subject of major control efforts. The beetle is most often found in weak or failing hives and rarely affects strong hives. However, differences in the housecleaning traits of the bees found in South Africa and the U.S. may mean very different responses to the beetles. Some early reports from Florida and South Carolina suggest the beetles may be more damaging there than in Africa. Para-dichlorobenzene (PDB) has been used for protecting empty stored combs. Coumaphos bee strips (Bayer Corporation) have been approved for use in hives for the control of small hive beetles in some states under an emergency registration.[ citation needed ]

Peppermint oil extract and even candy are used successfully. Beetles do not like peppermint, but honey bees savour this essential oil. [29]

Biological control through beneficial soil nematodes specific to the SHB is also effective. [30]

Beneficial nematodes are used by applying them to the soil while suspended in water. They may be applied as a pressurized spray or simply poured from a watering can. Once applied to the soil, nematodes burrow downward in search of insect pests. Once an insect is found, nematodes enter the body of the insect and release a powerful bacterium which quickly kills the pest. Released bacteria dissolve the internal tissues of the insect which becomes food for nematode growth and development. Mature nematodes then mate and lay eggs to produce more nematodes within the dead insect. Several such generations may occur over just a few days. After the inside of an insect is consumed, tiny infective-stage nematodes leave the dead insect shell and begin searching for more pests. As many as 350,000 nematodes may emerge from a single dead insect after only 10–15 days. Numbers depend on insect size.[ citation needed ]

There are also several traps currently on the market. The more effective ones are the Beetlejail Baitable, Hood Trap, the Freeman Beetle Trap, the West trap, the Australian, AJ's Beetle Eater, [31] and the Beetle Blaster. [32] All these traps use non-toxic oil to suffocate the beetles. This allows beekeepers to avoid having toxic chemicals in their beehives.[ citation needed ]

Fungal biocontrol may also be an effective method of control for the small hive beetle. [27] A study investigating the use of Metarhizium and Beuvaria fungi found could be used as control methods. It was found that if Metarhizium is added to soil containing the beetle, this would kill many of the larvae that entered it. It was also found that the fungi could negatively impact the fertility of the beetles. [27]

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  23. 1 2 3 Spiewok, Sebastian; Duncan, Michael; Spooner-Hart, Robert; Pettis, Jeff S.; Neumann, Peter (2008-11-01). "Small hive beetle, Aethina tumida, populations II: Dispersal of small hive beetles". Apidologie. 39 (6): 683–693. doi:10.1051/apido:2008054. ISSN   1297-9678. S2CID   10178081.
  24. Neumann, P.; Pirk, C.; Hepburn, H.; Solbrig, A.; Ratnieks, F.; Elzen, P.; Baxter, J. (2001-05-01). "Social encapsulation of beetle parasites by Cape honeybee colonies (Apis mellifera capensis Esch.)". Naturwissenschaften. 88 (5): 214–216. Bibcode:2001NW.....88..214N. doi:10.1007/s001140100224. ISSN   1432-1904. PMID   11482434. S2CID   1428187.
  25. Schäfer, Marc Oliver; Ritter, Wolfgang; Pettis, Jeff; Neumann, Peter (2010-01-01). "Small hive beetles, Aethina tumida, are vectors of Paenibacillus larvae". Apidologie. 41 (1): 14–20. doi:10.1051/apido/2009037. ISSN   1297-9678. S2CID   37530927.
  26. Eyer, Michael; Chen, Yan Ping; Schäfer, Marc O.; Pettis, Jeff S.; Neumann, Peter (January 2009). "Honey bee sacbrood virus infects adult small hive beetles, Aethina tumida (Coleoptera: Nitidulidae)". Journal of Apicultural Research. 48 (4): 296–297. Bibcode:2009JApiR..48..296E. doi:10.3896/IBRA.1.48.4.11. ISSN   0021-8839. S2CID   83566288.
  27. 1 2 3 Leemon, Diana (August 2012). "In-hive Fungal Biocontrol of Small Hive Beetle" (PDF). Rural Industries Research & Development Corporation. 12 (12).
  28. 1 2 3 4 5 Mustafa, Sandra G.; Spooner-Hart, Robert; Duncan, Michael; Pettis, Jeffery S.; Steidle, Johannes L. M.; Rosenkranz, Peter (2015-08-19). "Age and aggregation trigger mating behaviour in the small hive beetle, Aethina tumida (Nitidulidae)". The Science of Nature. 102 (9): 49. Bibcode:2015SciNa.102...49M. doi:10.1007/s00114-015-1300-9. ISSN   1432-1904. PMID   26286322. S2CID   10645938.
  29. NTALLI, Nikoletta G.; SPOCHACZ, Marta; ADAMSKI, Zbigniew (2022-05-25). "The role of botanical treatments used in apiculture to control arthropod pests". Apidologie. 53 (2): 27. doi:10.1007/s13592-022-00924-7. ISSN   1297-9678.
  30. Sanchez, WinDi; Shapiro, David; Williams, Geoff; Lawrence, Kathy (8 July 2021). "Entomopathogenic nematode management of small hive beetles (Aethina tumida) in three native Alabama soils under low moisture conditions". Journal of Nematology. 53: e2021–63. doi:10.21307/jofnem-2021-063. ISSN   0022-300X. PMC   8267404 . PMID   34286283.
  31. Ellis, Jamie. "Pests and Disease Videos". Dr. Jamie Ellis discusses the Small Hive Beetle. Honey Bee Research & Extension Laboratory University of Florida Entomology & Nematology Department. Archived from the original on 2011-07-18. Retrieved 2010-10-31. Another trap that has become increasingly popular is AJ's Beetle Eater...
  32. Hood, Michael Integrated Pest Management Archived 2016-07-12 at the Wayback Machine Dept. of Entomology, Soils, and Plant Sciences Clemson University, Clemson, South Carolina June 2010

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