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

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Contributors

22/05/08 Original text by:

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

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