Apis mellifera scutellata (africanized bee)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Rainfall Regime
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Economic Impact
- Environmental Impact
- Threatened Species
- Social Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
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 InvasivenessTop of page
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 TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hymenoptera
- Family: Apidae
- Genus: Apis
- Species: Apis mellifera scutellata
Notes on Taxonomy and NomenclatureTop of page
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’.
DescriptionTop of page
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.
DistributionTop of page
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 TableTop of page
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.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Botswana||Present||Native||McNally and Schneider (1996)|
|Ethiopia||Present||Native||Amssalu et al. (2004)||West, south & southwest humid midlands|
|Kenya||Present||Native||Oden (2001); Muli et al. (2005)||Taita Taveta District|
|Namibia||Present||Radloff et al. (1996)|
|South Africa||Present||Native||Radloff and Hepburn (1999); Beekman et al. (2002)|
|Tanzania||Present||Native||Piereira and Chaud-Netto (2005)|
|Uganda||Present||Native||Radloff and Hepburn (2001)|
|Zimbabwe||Present||Native||Fries et al. (2003)|
|Costa Rica||Present||Introduced||1983||CABI (Undated)||Original citation: Tarpy (undated)|
|El Salvador||Present||Introduced||1985||CABI (Undated)||Original citation: Tarpy (undated)|
|Mexico||Present||Introduced||1986||Invasive||Cairns et al. (2005)||Yucatan (Quintana Roo)|
|Panama||Present||Introduced||1982||Invasive||Roubik et al. (2003)||forests|
|Puerto Rico||Present||Introduced||1994||Invasive||Cox (1994); Kairo et al. (2003)|
|Trinidad and Tobago||Present||Introduced||1979||CABI (Undated); Schotman (1989)||Not Tobago; Original citation: Tarpy (undated)|
|United States||Present||CABI (Undated a)||Present based on regional distribution.|
|-Arizona||Present||Introduced||Piereira and Chaud-Netto (2005); CABI (Undated)|
|-Arkansas||Present, Localized||Introduced||Invasive||USDA (2011)|
|-California||Present||Introduced||1994||Sanford (1994); Kim and Oguro (1999); Piereira and Chaud-Netto (2005)|
|-Louisiana||Present, Localized||Introduced||Invasive||USDA (2011)|
|-Nevada||Present||Introduced||1998||Hicks (1999); Piereira and Chaud-Netto (2005)||Clark County. Swarms in Las Vegas in July 1998.|
|-New Mexico||Present||Introduced||Piereira and Chaud-Netto (2005)|
|-Texas||Present||Introduced||Invasive||Hunter et al. (1993); Kim and Oguro (1999); Coulson et al. (2005); USDA (2011)|
|-Utah||Present, Localized||Introduced||Invasive||USDA (2011)|
|Australia||Present||CABI (Undated a)||Present based on regional distribution.|
|-Victoria||Present||Introduced||Invasive||CABI (Undated)||Original citation: ISSG (IUCN SSC Invasive Species Specialist Group) (2013)|
|New Zealand||Present||Introduced||Invasive||McMillan (2005)||Rangitoto Island|
|Argentina||Present||Introduced||1969||CABI (Undated); Diniz et al. (2003); Piereira and Chaud-Netto (2005)||Original citation: Tarpy (undated)|
|Bolivia||Present||Introduced||1967||CABI (Undated)||Original citation: Tarpy (undated)|
|Brazil||Present||CABI (Undated a)||Present based on regional distribution.|
|-Parana||Present||Introduced||Akatsu and Pegoraro (2001)||Mandirituba|
|-Rio Grande do Sul||Present||Introduced||Outlaw et al. (2000)||temperate|
|-Sao Paulo||Present||Introduced||1956||Invasive||Gonçalves (2004); Sanchez Junior and Malerbo-Souza (2004); Piereira and Chaud-Netto (2005)||Ribeirao Preto; Near Rio Claro|
|Colombia||Present||Introduced||1979||CABI (Undated)||Original citation: Tarpy (undated)|
|Ecuador||Present||Introduced||1981||CABI (Undated)||Original citation: Tarpy (undated)|
|French Guiana||Present||Introduced||1974||CABI (Undated)||Original citation: Tarpy (undated)|
|Guyana||Present||Introduced||1975||CABI (Undated)||Original citation: Tarpy (undated)|
|Paraguay||Present||Introduced||1965||CABI (Undated)||Original citation: Tarpy (undated)|
|Peru||Present||Introduced||1977||CABI (Undated)||Original citation: Tarpy (undated)|
|Uruguay||Present||Introduced||1971||Diniz et al. (2003)|
|Venezuela||Present||Introduced||1977||CABI (Undated); Thimann and Manrique (2002); Cabrera et al. (2003)||Original citation: Tarpy (undated)|
History of Introduction and SpreadTop of page
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).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Sao Paulo||South Africa||1956||Yes||No|
Risk of IntroductionTop of page
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.
HabitatTop of page
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 ListTop of page
|Terrestrial – Managed||Cultivated / agricultural land||Principal habitat||Harmful (pest or invasive)|
|Cultivated / agricultural land||Principal habitat||Natural|
|Cultivated / agricultural land||Principal habitat||Productive/non-natural|
|Protected agriculture (e.g. glasshouse production)||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Protected agriculture (e.g. glasshouse production)||Secondary/tolerated habitat||Productive/non-natural|
|Managed forests, plantations and orchards||Principal habitat||Harmful (pest or invasive)|
|Managed grasslands (grazing systems)||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Managed grasslands (grazing systems)||Secondary/tolerated habitat||Natural|
|Disturbed areas||Principal habitat||Harmful (pest or invasive)|
|Rail / roadsides||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Urban / peri-urban areas||Principal habitat||Harmful (pest or invasive)|
|Buildings||Principal habitat||Harmful (pest or invasive)|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Principal habitat||Harmful (pest or invasive)|
|Natural forests||Principal habitat||Natural|
|Natural forests||Principal habitat||Productive/non-natural|
|Natural grasslands||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Natural grasslands||Secondary/tolerated habitat||Natural|
|Natural grasslands||Secondary/tolerated habitat||Productive/non-natural|
|Riverbanks||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Scrub / shrublands||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Scrub / shrublands||Secondary/tolerated habitat||Natural|
|Coastal dunes||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Coastal dunes||Secondary/tolerated habitat||Natural|
Biology and EcologyTop of page
The period from egg to adult is approximately 18.5 days for worker bees and 16 days for queens, and adult longevity is approximately 30 days for workers, 5-10 weeks for drones, and 1-3 years for queens. For more details on egg, larvae and pupae production in A. mellifera scutellata colonies in Mandirituba, Parana, Brazil, see Pegoraro et al. (2001). One queen can produce 1500 eggs a day, and when a queen mates with a drone, the fertilized egg becomes a female worker bee, whereas unfertilized eggs become male drones, and larvae develop into queens if they are fed large quantities of royal jelly, a very nutritious creamy white liquid consisting of hypopharyngeal and mandibular gland secretions. Colonies reproduce by frequent swarming, and one colony can result in 17 other colonies in a year.
Africanized honey bees are limited to hot tropical and warm sub-tropical habitats with separate wet and dry seasons, in comparison to hot and cold seasons in cooler temperate areas. Temperate climatic restrictions seem to be a natural limit to the expansion of Africanized honey bees at latitudes of approximately 35-40°. Radloff and Hepburn (1999) reported on A. mellifera worker bees in South Africa and described three morphoclusters: an unnamed population at >1500 m altitude; bees considered to be A. mellifera scutellata x A. m. capensis hybrids; and A. m. scutellata surrounding the mountains at <1500 m altitude. High humidity is necessary for honey bee brood development and honey bees are very efficient at regulating the biophysical parameters of their hive according to the needs of the colony. However, Human et al. (2006), studying A. mellifera scutellata, suggested that regulation of humidity is adjusted within sub-optimal limits.
ClimateTop of page
|A - Tropical/Megathermal climate||Preferred||Average temp. of coolest month > 18°C, > 1500mm precipitation annually|
|B - Dry (arid and semi-arid)||Preferred||< 860mm precipitation annually|
|C - Temperate/Mesothermal climate||Tolerated||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Rainfall RegimeTop of page Bimodal
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Varroa destructor||Parasite||Adult Female/Adult Male||to genus|
Notes on Natural EnemiesTop of page
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 DispersalTop of page
Natural Dispersal (Non-Biotic)
Pathway CausesTop of page
|Breeding and propagation||From South Africa & Tanzania to Brazil||Yes||Yes||Piereira and Chaud-Netto, 2005|
|Escape from confinement or garden escape||From South Africa & Tanzania to Brazil, then escaped into the Brazilian countryside||Yes||Piereira and Chaud-Netto, 2005|
|Intentional release||From South Africa & Tanzania to Brazil ,to aid honey bee production in Brazil||Yes||Piereira and Chaud-Netto, 2005|
|People sharing resources||Yes||Piereira and Chaud-Netto, 2005|
Economic ImpactTop of page
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 ImpactTop of page
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).
A. mellifera scutellata is spreading throughout tropical and subtropical America, hybridizing with and for the most part replacing European honey bees (mainly Apis mellifera mellifera and Apis mellifera ligustica).
Threatened SpeciesTop of page
Social ImpactTop of page
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 FactorsTop of page Invasiveness
- 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
- Reproduces asexually
- Changed gene pool/ selective loss of genotypes
- 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
- Causes allergic responses
- Competition - monopolizing resources
- Pest and disease transmission
- Induces hypersensitivity
- Interaction with other invasive species
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect in the field
- Difficult/costly to control
Uses ListTop of page
- Commercial pollinator
Human food and beverage
- Honey/honey flora
Detection and InspectionTop of page
Similarities to Other Species/ConditionsTop of page
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 ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
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 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).
ReferencesTop of page
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.
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.
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
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.
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.
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.
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/
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.
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
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.
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.
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.
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.
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Piereira A M, 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.
Roubik D W, 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 S E, Kitching R L]. Cambridge, UK: Cambridge University Press. 360-368.
Sanchez Junior J L B, Malerbo-Souza D T, 2004. Pollinators frequency on cotton flowers and production. (Freqüência dos insetos na polinização e produção de algodão.). Acta Scientiarum - Agronomy. 26 (4), 461-465.
Thimann R, Manrique A J, 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.
USDA, 2011. Africanized Honey Bees., http://www.ars.usda.gov/Research/docs.htm?docid=11059
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22/05/08 Original text by:
Claire Beverley, CABI, Nosworthy Way, Wallingford, Oxon OX10 8DE, UK
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