Apis mellifera scutellata (africanized bee)

Pictures

Picture	Title	Caption	Copyright
Title Adults Caption An Africanized honey bee (dark specimen) and a European honey bee (pale specimen) on honeycomb. USA. Despite the obvious colour differences between these two individuals, they can't usually be identified by eye alone. Copyright ©Scott Bauer/USDA-ARS	Adults	An Africanized honey bee (dark specimen) and a European honey bee (pale specimen) on honeycomb. USA. Despite the obvious colour differences between these two individuals, they can't usually be identified by eye alone.	©Scott Bauer/USDA-ARS
Title Africanized honey bees Caption Close-up of Africanized honey bees (AHBs) surrounding a European queen honey bee (EHB), marked with a pink dot for identification. Since AHBs arrived in Texas in 1990, they've mated with EHBs and spread throughout the Southwest. But rather than commingling, AHBs tend to replace EHBs, partly because EHB queen bees mate disproportionately with African drones. Copyright ©Scott Bauer/USDA-ARS	Africanized honey bees	Close-up of Africanized honey bees (AHBs) surrounding a European queen honey bee (EHB), marked with a pink dot for identification. Since AHBs arrived in Texas in 1990, they've mated with EHBs and spread throughout the Southwest. But rather than commingling, AHBs tend to replace EHBs, partly because EHB queen bees mate disproportionately with African drones.	©Scott Bauer/USDA-ARS
Title Research and control Caption Entomologist David Gilley is part of the team investigating the usurpation of European honey bee colonies by swarms of Africanized honey bees. Because queenless colonies are particularly susceptible to usurpation, the team maintains a group of queenless colonies to lure usurpation swarms into their apiary to be studied. David Gilley is shown here requeening one of these 'bait colonies'. USA. Copyright ©Scott Bauer/USDA-ARS	Research and control	Entomologist David Gilley is part of the team investigating the usurpation of European honey bee colonies by swarms of Africanized honey bees. Because queenless colonies are particularly susceptible to usurpation, the team maintains a group of queenless colonies to lure usurpation swarms into their apiary to be studied. David Gilley is shown here requeening one of these 'bait colonies'. USA.	©Scott Bauer/USDA-ARS
Title ARS honey bee trap Caption Entomologist Justin Schmidt examines an ARS honey bee trap used to lure Africanized bee swarms and prevent their establishment in walls of buildings. Captured swarms are easily removed or destroyed with soapy water. USA. Copyright ©Scott Bauer/USDA-ARS	ARS honey bee trap	Entomologist Justin Schmidt examines an ARS honey bee trap used to lure Africanized bee swarms and prevent their establishment in walls of buildings. Captured swarms are easily removed or destroyed with soapy water. USA.	©Scott Bauer/USDA-ARS
Title Control measures Caption Northwest Fire District's Captain John Estes of Tucson, Arizona, uses a wide spray of water and chemical wetting agent as a means of subduing Africanized honey bees. Looking on is ARS entomologist Eric Erickson, who taught this control method to fire departments throughout Arizona. Copyright USDA-ARS	Control measures	Northwest Fire District's Captain John Estes of Tucson, Arizona, uses a wide spray of water and chemical wetting agent as a means of subduing Africanized honey bees. Looking on is ARS entomologist Eric Erickson, who taught this control method to fire departments throughout Arizona.	USDA-ARS

Identity

Preferred Scientific Name

• Apis mellifera scutellata Lepeletier

Preferred Common Name

• africanized bee

International Common Names

• English: African bee; african honey bee; African honeybee; Africanized bee; Africanized honeybee; Brazilian bee; killer bee
• Spanish: abeja africanizada; abeja de miel africana
• French: abeille africaine; abeille africanisée

Summary of Invasiveness

A. mellifera scutellata was imported from Africa to Brazil in 1956 to increase honey production, and 26 swarms accidentally escaped into the countryside where the queens mated with drones of the European honey bees. The subsequent poly-hybrid bees were named ‘Africanized honey bees’, and over a period of 50 years, they colonized most of South America and all of Central America. In 1990, they entered the USA through Texas, and they are now established throughout the south-western states and southern California.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Hymenoptera
•                         Family: Apidae
•                             Genus: Apis
•                                 Species: Apis mellifera scutellata

Notes on Taxonomy and Nomenclature

Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae) is one of a number of subspecies of A. mellifera (Western honey bee). The subspecies can be divided by their native continents as follows. Those originating in Europe include: carnica, caucasica, cecropia, cypria, iberiensis, ligustica, mellifera, remipes, ruttneri and sicula. Those originating in Africa include: adansonii, bandasii, capensis, intermissa, lamarcki, jemenitica, litoria, major, monticola, nubica, sahariensis, scutellata, unicolor and woyi-gambell. Those originating in the Middle East and Asia include: adamii, armeniaca, anatolica, macedonica, meda, pomonella and syriaca.

This datasheet includes data on pure Apis mellifera scutellata from its native range, but mostly on the invasive polyhybrid between A. mellifera scutellata and European honey bees (Apis mellifera) (Piereira and Chaud-Netto, 2005), commonly known as Africanized honey bees (AHB) or ‘killer bees’.

Description

A. mellifera scutellata is ca. 10-20 mm long, and brown with black stripes; workers are 10-15 mm long, drones 15-17 mm, and queens 18-20 mm.

Distribution

A. mellifera scutellata is native to eastern and southern Africa, from Ethiopia to South Africa. In Ethiopia, A. mellifera scutellata occupies the west, south and southwest humid midlands. Other morphoclusters in Ethiopia included A. mellifera jemenitica in the northwest and eastern arid and semi-arid lowlands; A. mellifera bandasii in the central moist highlands; A. mellifera monticola from the northern mountainous highlands; and A. mellifera woyi-gambell in south western semi-arid to sub-humid lowland parts of the country (Amssalu et al., 2004). Honey bees of Uganda represented an important biogeographical gap, defining the population structure of A. mellifera scutellata; however, morphometric analysis of worker honey bees has resolved this issue. At lower altitudes <200 m), honey bees formed one distinct morphocluster typical of A. mellifera scutellata throughout the continent. In comparison, those at higher altitudes (>2000 m) formed a separate distinct cluster of large, dark bees as mountain ecotypes (Radloff and Hepburn, 2001). The distribution of honey bee subspecies, including A. mellifera scutellata, in Taita Taveta District, Kenya, also appears to be related to altitude, with A. mellifera monticola found to be most widespread in the highlands and A. mellifera litorea was most common in lowland areas (Oden, 2001).

A. mellifera scutellata was introduced to Brazil in 1956 and from there it and its hybrids with European honey bees have spread to much of South America, to Central America and to the southern parts of North America.

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

A. mellifera scutellata was introduced into Brazil in 1956 by Warwick Estevam Kerr from South Africa and Tanzania, to assist the honey industry there because this species was better adapted to conditions in South America than various European subspecies of A. mellifera that he was using. The accidental release of queens and workers in 26 swarms into the Brazilian countryside was followed by subsequent hybridization with local European honey bee colonies (Piereira and Chaud-Netto, 2005). A. mellifera scutellata and the hybrids have since spread rapidly from the point of the original introduction near Rio Claro, Sao Paulo, Brazil, to as far south as mid-Argentina and to the north of Texas, Arizona, New Mexico, California and Nevada, USA (Kim and Oguro, 1999; Piereira and Chaud-Netto, 2005). The first record in the USA was from Hidalgo, Texas in 1993 and the bees were reported to advance at 100-300 miles (160-480 km) per year by colonizing existing hives or forming new hives in the wild (Kim and Oguro, 1999).

Introductions

Risk of Introduction

A. mellifera scutellata is an aggressive invader, and its spread is facilitated by a high adaptability to variable ecological conditions (Piereira and Chaud-Netto, 2005), indicating that further spread is likely. See Kaplan (2004) and Schneider et al. (2004) for a discussion of traits and behaviours that are responsible for making this species a successful invader.

Habitat

A. mellifera scutellata form colonies in tree hollows, rotted logs and man-made structures, such as wood and rock piles. A list of known nesting places of Africanized honey bees in the USA includes: trees and shrubs; wood pike or trash piles; flower pots; old tyres; ground holes; chimneys; storage sheds; wall cavities; attics and crawl spaces; roof overlaps and building eaves; underground utilities; water meters and sprinkler control boxes; old mine shafts or rock crevices; and evaporative coolers (UDAF, 2008). They can be found wherever such sites are found, most commonly in urban areas, agricultural land, forests (natural and managed), but also in riparian areas, coastal areas and occasionally in many other habitat types.

According to Oliveira and Cunha (2005), Africanized honey bees in the Americas are limited to regions of low altitude and cool winters, and principally occur in urban areas, and open or disturbed vegetation in Brazil, as opposed to the interior of dense forest, such as the Amazon. It was observed that Africanized honey bee workers did not visit baits in continuous forest or forest fragments, but were observed in deforested or open areas. This indicates that there is no possibility of source competition between Africanized and native bees within the Amazon forest, and also that large-scale beekeeping is unlikely to succeed in the region because the forest is not explored by Africanized bees.

Habitat List

Biology and Ecology

Genetics

Climate

Latitude/Altitude Ranges

Rainfall Regime

Natural enemies

Notes on Natural Enemies

Small hive beetles (Aethina tumida) are pests of honey bees and can damage combs, stored honey and pollen. A study by Neumann and Hartel (2004) showed that A. mellifera scutellata remove unprotected eggs and larvae of the beetles and this behaviour plays an important role in the apparent resistance of African honey bees towards infestations by small hive beetles. They are also affected by the mite Varroa destructor in the same way as other bee subspecies

A number of bacteria are associated with honey bees and in a study of those associated with A. mellifera capensis and A. mellifera scutellata by Jeyaprakash et al. (2003), Lactobacillus and Bifidobacterium were found. These have also been reported from other honey bee subspecies; however, other sequences were found associated with honey bees for the first time, e.g. Bartonella, Gluconacetobacter; Simonsiella/Neisseria; and Serratia. Another bacterium, the parasitic microbe Wolbachia, is found in workers and drones of A. mellifera scutellata and hybrid workers of A. mellifera capensis and A. mellifera scutellata (Hoy et al., 2003).

Gene sequencing of a microsporidium from honey bees in Zimbabwe has found Nosema apis, a fungal parasite of honey bees, causing nosemosis or nosema (Fries, 2002; Fries et al., 2003).

Means of Movement and Dispersal

Natural Dispersal (Non-Biotic)

Pathway Causes

Economic Impact

A. mellifera scutellata negatively affects the beekeeping industry because it competes with European honey bees (A. mellifera), invading nests and causing A. mellifera to produce less honey. Labour costs are high because harvesting of A. mellifera scutellata honey requires handling aggressive bees and frequently re-queening colonies. However, once these obstacles are overcome, beekeeping with A. mellifera scutellata is considered to be a good investment (Goncalves, 2004).

A possibly more significant negative economic impact is caused by livestock fatalities resulting from Africanized bee stings, of which there have been very many thousands, though the exact number and the actual economic cost have not been fully estimated.

Environmental Impact

Impact on Habitat

In pristine and fragmented Amazonian rainforest, tropical rain forest trees characteristically occur in low population densities and rely on animals for cross-pollination. However, Dick et al. (2003) studied pollen dispersal and found that in highly disturbed habitats, A. mellifera may expand genetic neighbourhood areas and so link fragmented and continuous forest populations. For example, Dinizia excelsa was found to thrive in pastures and forest fragments in Manaus, Brazil even in the absence of native pollinators, and A. mellifera scutellata was the predominant floral visitor in fragmented habitats, also replacing native insects in isolated pasture trees. Gene flow over 3.4 km was reported in pasture indicating the significance of this bee in possibly altering the genetic structure of remnant populations via frequent, long-distance gene flow (Dick, 2001).

Threatened Species

Social Impact

A. mellifera scutellata swarms were given the common name ‘killer bees’ after causing death in pets, livestock and humans. In Brazil, accidents caused by A. mellifera scutellata have been attributed to high swarming frequencies and the variety of shelters available to them in urban areas (Piereira and Chaud-Netto, 2005). Massive envenomations by honey bees can cause multiorgan dysfunction as a result of the direct toxic effects of the large venom load received, although the mechanisms behind this are not clearly understood (Betten et al., 2006). The Africanized honey bee is the most commonly implicated species in such attacks. Although reports of deaths vary, the first victim of Africanized honey bees was in 1991, with the first death in 1993, and to date they have apparently killed some 1000 humans in Brazil, with 175 deaths in Mexico since 1985, and seven deaths in the USA since 1993.

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Capable of securing and ingesting a wide range of food
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Has high reproductive potential
• Gregarious
• Reproduces asexually

• Changed gene pool/ selective loss of genotypes
• Conflict
• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Negatively impacts human health
• Negatively impacts livelihoods
• Reduced amenity values
• Reduced native biodiversity
• Threat to/ loss of native species

• Allelopathic
• Causes allergic responses
• Competition - monopolizing resources
• Pest and disease transmission
• Hybridization
• Induces hypersensitivity
• Interaction with other invasive species
• Poisoning

• Highly likely to be transported internationally accidentally
• Difficult to identify/detect in the field
• Difficult/costly to control

Uses List

Environmental

• Commercial pollinator

Human food and beverage

• Honey/honey flora

Medicinal, pharmaceutical

• Traditional/folklore

Detection and Inspection

Similarities to Other Species/Conditions

A. mellifera is similar to A. mellifera scutellata, and it is very difficult for the layperson to positively distinguish Africanized honeybees from common European honeybees, as apart from genetic analysis, a comparison of up to as 20 different body measurements is the only way to distinguish subspecies with any certainty (Kaplan, 2004). Also, the Africanized honey bee is more defensive around its nests than the European bee and has a tendency to sting in large numbers. A. mellifera can withstand colder temperatures compared to A. mellifera scutellata. A. mellifera scutellata is slightly smaller than A. mellifera and less selective in habitat selection.

Prevention and Control

Prevention

Early warning systems

Behavioural studies of A. mellifera scutellata have been undertaken with the aim of creating informed strategies for dealing with Africanized honey bees and educating the public (Mello et al., 2003). The removal of bee colonies and swarms in Sao Paulo, Brazil was positively correlated with average temperature and degree of insolation, and negatively correlated with relative humidity and rainfall, and it was shown to be likely that colonies nested in artificial constructions, whereas wandering swarms tended to nest in trees. Such results can be used to predict times of the year when people should be more alert for activity of the bees. State Agricultural Departments in the USA have been tracking and studying Africanized honey bees since their introduction, and in particular, the Utah Department of Agriculture and Food has been setting traps since 2003 as an early warning line (UDAF, 2008).

Public awareness

Public awareness facilitated by studies such as that by Mello et al. (2003) will hopefully lead to the removal of structures seen as potential nesting sites for the bees and avoidance of areas possibly harbouring swarms. Africanized bees are much more likely to attack to defend their colonies compared to European honey bees and attacks are provoked by vibration, noise or motion within 50 feet of the nest. Offending origins of such sounds include lawn mowers, leaf blowers and hedge trimmers, and the odour of freshly cut grass or citrus may also be a source of provocation. Animals and humans may be pursued for up to a quarter of a mile and bees remain agitated for up to 8 hours after disturbance (UDAF, 2008).

Control

Biological control

Chemical control

References

Akatsu IP, Pegoraro A, 2001. The thorax width of Apis mellifera scutellata Lepeletier 1836 (Hym. Apidae) workers, as a variable for pollen trap screen efficiency. (Largura do tórax de operárias de Apis mellifera scutellata Lepeletier, 1836 (Hym.,: Apidae) e eficiência da tela coletora de pólen.) Archives of Veterinary Science, 6(2):77-82.

Alcobia JJF, 1991. The horizontal moveable frame hive of Mozambique. In: Bees and beekeeping in southern Africa. Proceedings of the International Beekeepers' Symposium held at Stellenbosch, South Africa, 24-26 January, 1990 [ed. by Anderson, R. H.\Buys, B.]. Cape Town, South Africa: S. A. Federation of Beekeepers' Associations, 156.

Amssalu B, Nuru A, Radloff SE, Hepburn HR, 2004. Mulitvariate morphometric analysis of honey bees (Apis mellifera) in the Ethiopian region. Apidologie, 35(1):71-81.

Beekman M, Allsopp MH, Wossler TC, 2008. Factors affecting the dynamics of the honey bee (Apis mellifera) hybrid zone of South Africa. Heredity, 100:13-18.

Beekman M, Wossler TC, Martin SJ, Ratnieks FLW, 2002. Parasitic Cape honey bee workers (Apis mellifera capensis) are not given differential treatment by African guards (A. m. scutellata). Insectes Sociaux, 49(3):216-220.

Betten DP, Richardson WH, Tong TC, Clark RF, 2006. Massive honey bee envenomation-induced rhabdomyolysis in an adolescent. Pediatrics, 117(1):231-235. http://pediatrics.aappublications.org

Cabrera L, Ojeda de Rodríguez G, Céspedes E, Colina A, 2003. Antibacterial activity of honey of multifloral bees (Apis mellifera scutellata) from four apiarists zones in Zulia State, Venezuela. (Actividad antibacteriana de miel de abejas multiflorales (Apis mellifera scutellata) de cuatro zonas apícolas del estado Zulia, Venezuela.) Revista Cientifica, Facultad de Ciencias Veterinarias, Universidad del Zulia, 13(3):205-211.

Cairns CE, Villanueva-Gutiérrez R, Koptur S, Bray DB, 2005. Bee populations, forest disturbance, and Africanization in Mexico. Biotropica, 37(4):686-692. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1744-7429.2005.00087.x

Coulson RN, Pinto MA, Tchakerian MD, Baum KA, Rubink WL, Johnston JS, 2005. Feral honey bees in pine forest landscapes of east Texas. Forest Ecology and Management, 215(1/3):91-102.

Cox B, 1994. AHB in Puerto Rico. American Bee Journal, 134(10):668-669.

Delaplane KS, 2006. Africanized honey bees. The University of Georgia, Cooperative Extension. http://www.ent.uga.edu/bees/Publications/B1290.pdf

Dick CW, 2001. Genetic rescue of remnant tropical trees by an alien pollinator. Proceedings of the Royal Society of London. Series B, Biological Sciences, 268(1483):2391-2396.

Dick CW, Etchelecu G, Austerlitz F, 2003. Pollen dispersal of tropical trees (Dinizia excelsa: Fabaceae) by native insects and African honeybees in pristine and fragmented Amazonian rainforest. Molecular Ecology, 12(3):753-764.

Diniz NM, Soares AEE, Sheppard WS, Lama del MA, 2003. Genetic structure of honey bee populations from southern Brazil and Uruguay. Genetics and Molecular Biology, 26(1):47-52.

Francoy TM, Prado PRR, Gonçalves LS, Costa L da F, Jong D de, 2006. Morphometric differences in a single wing cell can discriminate Apis mellifera racial types. Apidologie, 37(1):91-97.

Fries I, 2002. African honey bees (Apis mellifera scutellata) and nosema (Nosema apis) infections. In: Bees without frontiers: Sixth European Bee Conference, Cardiff, UK, 1-5 July 2002 [ed. by Jones R] Cardiff, UK: International Bee Research Association, 50-55.

Fries I, Slemenda SB, Silva A da, Pieniazek NJ, 2003. African honey bees (Apis mellifera scutellata) and Nosema (Nosema apis) infections. Journal of Apicultural Research, 42(1/2):13-15.

Gervan NL, Winston ML, Higo HA, Hoover SER, 2005. The effects of honey bee (Apis mellifera) queen mandibular pheromone on colony defensive behaviour. Journal of Apicultural Research, 44(4):175-179.

Gonçalves LS, 2004. The big challenge: development of beekeeping with Africanized honey bees in Northeast Brazil. In: Proceedings of the 8th IBRA International Conference on Tropical Bees and VI Encontro sobre Abelhas, Ribeirão Preto, Brasil, 6-10 September, 2004 [ed. by Hartfelder KH, Jong D de, Pereira RA, Santos Cristino A dos, Morais MM, Tanaka ED, Lourenço AP, Silva JEB da, Almeida GF de, Nascimento AM do], 241-246.

Hepburn HR, 2001. The enigmatic Cape honey bee, Apis mellifera capensis. Bee World, 82(4):181-191.

Hepburn R, Radloff SE, 2002. Apis mellifera capensis: an essay on the subspecific classification of honeybees. Apidologie, 33(2):105-127.

Hicks RC, 1999. Africanized honey bees in Clark County, Nevada. In: Proceedings of the Annual Meeting of the Utah Mosquito Abatement Association [ed. by Dickson SL], 1-3.

Hoy MA, Jeyaprakash A, Alvarez JM, Allsopp MH, 2003. Wolbachia is present in Apis mellifera capensis, A. m. scutellata, and their hybrid in Southern Africa. Aphidologie, 34(1):53-60.

Human H, Nicholson SW, Dietemann V, 2006. Do honey bees, Apis mellifera scutellata, regulate humidity in their nest? Naturwissenschaften, 93(8):397-401.

Hunter LA, Jackman JA, Sugden EA, 1993. Detection records of Africanized honey bees in Texas during 1990, 1991 and 1992. Southwestern Entomologist, 18(2):79-89.

Härtel S, Neumann P, Kryger P, Heide CDvon, Moltzer GJ, Crewe RM, Praagh JPvan, Moritz RFA, 2006. Infestation levels of Apis mellifera scutellata swarms by socially parasitic Cape honeybee workers (Apis mellifera capensis). Apidologie, 37(4):462-470. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

ISSG (IUCN SSC Invasive Species Specialist Group), 2013. Global Invasive Species Database (GISD). IUCN SSC Invasive Species Specialist Group. http://www.issg.org/database/welcome/

ISSG, 2005. Global Invasive Species Database (GISD). Auckland, New Zealand: University of Auckland. http://www.issg.org/database

Jeyaprakash A, Hoy MA, Allsopp MH, 2003. Bacterial diversity in worker adults of Apis mellifera capensis and Apis mellifera scutellata (Insecta: Hymenoptera) assessed using 16S rRNA sequences. Journal of Invertebrate Pathology, 84(2):96-103.

Jones S, 2002. Know your stingers: subtle identification differences can help you predict behavior. Pest Control, 70(7):28-30.

Kairo M, Ali B, Cheesman O, Haysom K, Murphy S, 2003. Invasive species threats in the Caribbean region. Report to the Nature Conservancy. Curepe, Trinidad and Tobago: CAB International, 132 pp. http://www.issg.org/database/species/reference_files/Kairo%20et%20al,%202003.pdf

Kaplan JK, 2004. What's buzzing with Africanized honey bees? Agricultural Research (Washington), 52(3):4-8.

Kim KT, Oguro J, 1999. Update on the status of Africanized honey bees in the western states. Western Journal of Medicine, 170(4):220-222.

Lenzi M, Orth AI, 2004. Foraging activity of bees Apis mellifera scutellata on pink-pepper flowers. (Atividade forrageira da abelha Apis mellifera scutellata sobre as flores da aroeira-vermelha.) Agropecuária Catarinense, 17(3):61-63.

Martin S, Wossler T, Kryger P, 2002. Usurpation of African Apis mellifera scutellata colonies by parasitic Apis mellifera capensis workers. Apidologie, 33(2):215-232.

Martin SJ, Beekman M, Wossler TC, Ratnieks FLW, 2002. Parasitic Cape honey bee workers, Apis mellifera capensis, evade policing. Nature London, 415(6868):163-165.

Martin SJ, Kryger P, 2002. Reproduction of Varroa destructor in South African honey bees: does cell space influence Varroa male survivorship? Apidologie, 33(1):51-61.

Martin SJ, Medina LM, 2004. Africanized honey bees have unique tolerance to Varroa mites. Trends in Parasitology, 20(3):112-114.

McMillan D, 2005. Apicultural exotic disease surveillance report. Surveillance (Wellington), 32(4):9-10.

McNally LC, Schneider SS, 1996. Spatial distribution and nesting biology of colonies of the African honey bee Apis mellifera scutellata (Hymenoptera: Apidae) in Botswana, Africa. Environmental Entomology, 25(3):643-652.

Mello MHSH de, Silva EA da, Natal D, 2003. Africanized bees in a metropolitan area of Brazil: shelters and climatic influences. (Abelhas africanizadas em área metropolitana do Brasil: abrigos e influências climáticas.) Revista de Saúde Pública, 37(2):237-241.

Moritz RFA, 2002. Population dynamics of the Cape bee phenomenon: the impact of parasitic laying worker clones in apiaries and natural populations. Apidologie, 33(2):233-244.

Muli EM, Raina SK, Mueke JM, 2005. Royal jelly production in East Africa: performance potential of the honey bees, Apis mellifera scutellata and Apis mellifera monticola in Kenya. Journal of Apicultural Research, 44(4):137-140.

Neumann P, Hepburn HR, Radloff SE, 2000. Modes of worker reproduction, reproductive dominance and brood cell construction in queenless honeybee (Apis mellifera L.) colonies. Apidologie, 31(4):479-486.

Neumann P, Härtel S, 2004. Removal of small hive beetle (Aethina tumida) eggs and larvae by African honeybee colonies (Apis mellifera scutellata). Apidologie, 35(1):31-36.

Nielsen DI, Ebert PR, Page RE Jr, Hunt GJ, Guzmán-Novoa E, 2000. Improved polymerase chain reaction-based mitochondrial genotype assay for identification of the Africanized honey bee (Hymenoptera: Apidae). Annals of the Entomological Society of America, 93(1):1-6.

Nuru A, Amssalu B, Hepburn HR, Radloff SE, 2002. Swarming and migration in the honey bees (Apis mellifera) of Ethiopia. Journal of Apicultural Research, 41(1/2):35-41.

Oden AC, 2001. Honey bee Apis mellifera L. subspecies and apiclora in Taita Taveta District, Kenya. Minor Field Studies International Office, Swedish University of Agricultural Sciences, 172:45.

Ojar C, 2002. Introduced species summary project: Africanized honey bee (Apis mellifera scutellata). http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/Apis_mellifera_scutellata.htm#Threats

Oldroyd BP, 2002. The Cape honey bee: an example of a social cancer. Trends in Ecology and Evolution, 17(6):249-251.

Oliveira ML de, Cunha JA, 2005. Do Africanized honey bees explore resources in the Amazonian forest? Acta Amazonica, 35(3):389-394.

Outlaw WH Jr, Fleig M, Grüninger W, Magel E, Hampp R, 2000. Observations on honey plants and Africanized honey bees in the temperate-zone State of Rio Grande do Sul, Brazil. American Bee Journal, 140(5):401-404.

PaDIL, 2007. Apis mellifera scutellata. Pests and Diseases Image Library. http://www.padil.au/viewPestDiagnosticImages.aspx?id=370

Pegoraro A, Chaves Neto A, Dalla CMA, 2001. Production of eggs, larvae and pupae in Apis mellifera scutellata (Hym., Apidae) colonies. (Producão de ovos-larvas e pupas em colônias de Apis mellifera scutellata (Hym., Apidae).) Archives of Veterinary Science, 6(2):57-62.

Piereira AM, Chaud-Netto J, 2005. Africanized honey bees: biological characteristics, urban nesting behaviour and accidents caused in Brazilian cities (Hymenoptera: Apidae). Sociobiology, 46(3):535-550.

Pinto MA, Johnston JS, Rubink WL, Coulson RN, Patton JC, Sheppard WS, 2003. Identification of Africanized honey bee (Hymenoptera: Apidae) mitochondrial DNA: validation of a rapid polymerase chain reaction-based assay. Annals of the Entomological Society of America, 96(5):679-684.

Principal J, D'Aubeterre R, Graterol Z, 2004. Prevalance of Varroa destructor in Africanized honey bees (Apis mellifera scutellata Lepeletier) in Lara State, Venezuela. In: Proceedings of the 8th IBRA International Conference on Tropical Bees and VI Encontro sobre Abelhas, Ribeirao Preto, Brasil, 6-10 September, 347-350.

Quezada-Euan JJG, 2000. Hybridization between European and Africanized honey bees in tropical Yucatan, Mexico. Morphometric, allozymic and mitochondrial DNA variability in feral colonies. Apidologie, 31(3):443-453.

Rabe MJ, Rosenstock SS, Nielsen DI, 2005. Feral Africanized honey bees (Apis mellifera) in Sonoran Desert habitats of Southwestern Arizona. Southwestern Naturalist, 50(3):307-311. http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1894%2F0038-4909%282005%29050%5B0307%3AFAHBAM%5D2.0.CO%3B2

Radloff SE, Hepburn HR, 1999. Honey bees, Apis mellifera Linnaeus (Hymenoptera: Apidae), of the Drakensburg mountains in relation to neighbouring populations. African Entomology, 7(1):35-41.

Radloff SE, Hepburn HR, 2001. Population structure of Apis mellifera scutellata (Hymenoptera: Apidae): filling the Uganda gap. African Entomology, 9(2):131-135.

Radloff SE, Hepburn HR, Robertson MP, Hille Rvan, Davidson Z, Villet MH, 1996. Discriminant analysis of the honeybee populations of southwestern Africa. African Entomology, 4(1):1-6.

Roubik DW, Sakai S, Gattesco F, 2003. Canopy flowers and certainty: loose niches revisited. In: Arthropods of tropical forests: spatio-temporal dynamics and resource use in the canopy [ed. by Basset Y, Novotny V, Miller SE, Kitching RL] Cambridge, UK: Cambridge University Press, 360-368.

Sanchez Junior JLB, Malerbo-Souza DT, 2004. Pollinators frequency on cotton flowers and production. (Freqüência dos insetos na polinização e produção de algodão.) Acta Scientiarum - Agronomy, 26(4):461-465.

Sanford M, 1994. African Bee found in California. APIS Newsletter: Apicultural Information and Issues from IFAS/University of Florida. http://apis.ifas.ufl.edu

Schneider SS, Deeby T, Gilley DC, DeGrandi-Hoffman G, 2004. Seasonal nest usurpation of European colonies by African swarms in Arizona, USA. Insectes Sociaux, 51(4):359-364.

Schneider SS, DeGrandi-Hoffman G, Smith DR, 2004. The African honey bee: factors contributing to a successful biological invasion. Annual Review of Entomology, 49:351-376.

Schotman CYL, 1989. Plant pests of quarantine importance to the Caribbean. RLAC-PROVEG, No. 21:80 pp.

Sereno FTPS, Padilla FA, Message D, Vilela EF, 2004. Differentiation of Africanized, European and North African colonies of Apis mellifera using morphological characteristics. (Diferenciación de colonias Apis mellifera africanizadas, europeas y del norte de Âfrica por características morfológicas.) Archivos Latinoamericanos de Producción Animal, 12(Suplemento 1):22-25.

Sheppard WS, Smith DR, 2000. Identification of African-derived bees in the Americas: a survey of methods. Annals of the Entomological Society of America, 93(2):159-176.

Suazo A, Hall HG, 2002. Nuclear DNA PCR-RFLPs that distinguish African and European honey bee groups of subspecies. II: Conversion of long PCR markers to standard PCR. Biochemical Genetics, 40(7/8):241-261.

Suazo A, Lee MyeongLyeol, Hall HG, 2002. A locus with restriction fragment length plymorphisms characteristic of African and European honey bee (Hymenoptera: Apidae) groups of subspecies. Annals of the Entomological Society of America, 95(1):115-124.

Tarpy DR, undated. Africanized 'killer' bees: a problem for North Carolina? http://www.ncagr.gov/plantindustry/plant/apiary/ncahb_files/African_Bees_for_Bee_Keepers.pdf

Thimann R, Manrique AJ, 2002. Propolis harvest from hives of Africanized bees during the raining season in Guanare, Venezuela. (Recolección de propóleos en colonias de abejas africanizadas durante la temporada de lluvias en Guanare, Venezuela.) Zootecnia Tropical, 20(4):493-499.

Thoenes S, 2002. There's money in honey bees: honey bee control is an emerging, high-margin service. Pest Control, 70(7):20-21, 24, 26, 28.

UDAF, 2008. What are Africanized Honey Bees (AHBs)? State of Utah Department of Agriculture and Food: What are Africanized Honey Bees (AHBs)?. http://ag.utah.gov/pressrel/ahb_info.html

USDA, 2011. Africanized Honey Bees. http://www.ars.usda.gov/Research/docs.htm?docid=11059

Whitfield CW, Behura SK, Berlocher SH, Clark AG, Johnston JS, Sheppard WS, Smith DR, Suarez AV, Weaver D, Tsutsui ND, 2006. Thrice out of Africa: ancient and recent expansions of the honey bee, Apis mellifera. Science (Washington), 314(5799):642-645. http://www.sciencemag.org

Wossler TC, 2002. Pheromone mimicry by Apis mellifera capensis social parasites leads to reproductive anarchy in host Apis mellifera scutellata colonies. Apidologie, 33(2):139-163.

Links to Websites

Contributors

22/05/08 Original text by:

Claire Beverley, CABI, Nosworthy Way, Wallingford, Oxon OX10 8DE, UK

Distribution Maps

You can pan and zoom the map

Save map

Unsupported Web Browser:

One or more of the features that are needed to show you the maps functionality are not available in the web browser that you are using.

Please consider upgrading your browser to the latest version or installing a new browser.

More information about modern web browsers can be found at http://browsehappy.com/

Download KML file Download CSV file

This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Unported License.

Please click OK to ACCEPT or Cancel to REJECT

OK Cancel

This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Unported License.

Please click OK to ACCEPT or Cancel to REJECT

OK Cancel