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Potentially Misidentified Species:
It is exceedingly difficult for a non-entomologist to positively distinguish
Africanized honeybees from common European honeybees. Aside from genetic
analysis, comparison of as many as twenty different body measurements remains
the only way to distinguish the strains with certainty (Kaplan 2004).
Apis mellifera scutellata are most easily recognized for their behavioral
dissimilarities to European honeybees. They respond quickly and aggressively
to disturbance, defend hives more intensely than European honeybees and will
chase the source of the disturbance for a mile or more (Sanford and Hall 2005).
II. HABITAT AND DISTRIBUTION
Regional Occurrence:
The native range of Apis mellifera scutellata is the eastern and southern
portions of Africa. Introduction to the New World began with the intentional
importation and subsequent accidental release in Brazil followed, in turn, by
spread of the strain through Central- and South America and into the southern
U.S. (Sanford and Hall 2005).
The known introduced U.S. range now includes
portions of California, Nevada, Arizon, New Mexico, Texas, Oklahoma, Arkansas, Louisiana, and Florida. The strain is considered to be officially established in Florida.
IRL Distribution:
As of March 2006, Africanizede honeybees had been reported from 14 Florida counties, including the
southern three counties (St. Lucie, Martin, Palm Beach) of the IRL watershed.
III. LIFE HISTORY AND POPULATION BIOLOGY
Age, Size, Lifespan:
Adult Apis mellifera scutellata workers attain a body length of around 19 mm,
which is only slightly smaller than the average European honeybee length.
Female worker bees live for around a month while male drones survive for 5-10
weeks. Queens typically live for 1-3 years (ISSG 2005).
Abundance:
Africanized honeybees have been found in 14 Florida counties and are now
considered to be established in the state.
Reproduction:
Reproduction is fascinating, complex, and an extensively studied facet of
honeybee life history. A cursory overview is presented here:
When queen honeybees mate with male drones, the fertilized eggs that are
produced are destined to hatch into sterile female workers unless, as larvae,
they are fed large quantities of royal jelly
secreted by hive workers. The royal jelly diet causes larvae to mature into
young reproductive queens. Queens can also produce unfertilized eggs, which
mature into reproductive male drones (Breed 2003).
The actual mating between honeybee queens and drones occurs during swarming
events when the old queen, young queens, drones, and about half of an
established colony's workers leave the nest to establish new ones. Seasonal
synchrony in reproductive swarming ensures that drones from many different
colonies are available to mate with emerging queens, thus minimizing the
effects of inbreeding. Queens are capable of mating with multiple drones whose
sperm they then store and use to fertilize eggs for their entire lifetime
(Breed 2003, Collins 2006).
European honeybees and Africanized honeybees exhibit important differences in
swarming behavior that are key to the successful spread of the Africanized
strain. Where both strains co-occur, European honeybee queens mate
disproportionately with Africanized drones. This is believed to happen both
because Africanized bees produce more drones per colony than European bees and
also because queens inseminated by both Africanized and European drones will
preferentially use the African semen first in the production of the next
generation of offspring. Additionally, Africanized colonies mature at a faster
rate allowing more swarms to split off from a hive in a given amount of time,
again conferring an advantage upon the Africanized reproductive individuals
(Kaplan 2004). Other reproductive differences conferring a competitive
advantage on Africanized honeybees are described below.
If young queens are isolated from Africanized colonies, they will have only
resident European drones with which to mate. Such mating results in offspring
with diluted African tendencies toward aggression (Sanford and Hall 2005). As
the numbers of Africanized colonies increase in an area, however, the incidence
of mating between Africanized queens and drones will also increase, resulting
in more aggressive progeny.
Embryology:
Africanized honeybee queens are capable of producing 1,500 eggs a day. Most of these are destined
to become workers, some will become drones, and a few will become new queens.
Larvae hatch out is just three days, as opposed to more than a week in European
bees. Initial growth is also extremely rapid. Fed nectar and pollen by adult
workers, larvae attain 900 times the initial weight of the egg in under one
week (Winston 1992).
IV. PHYSICAL TOLERANCES
Temperature:
Africanized honeybees are descended from stocks that evolved in the tropics and,
as such, are ill-equipped to withstand prolonged cold winters. They are
believed to be limited to tropical and subtropical habitats.
A study by Abramson et al. (1997) supports the belief that Africanized
honeybees are behaviorally and physiologically unsuited to low temperatures.
Bees abandoned a hive experimentally maintained at 9°C after 14 days, while
survivorship of individual bees exposed to 0°C for 4 hours was low.
Water Availability:
Apis mellifera scutellata evolved in the arid environment of eastern and
southern Africa. Until quite recently (ca. 2005, but see below) migration of A. m.
scutellata eastward from Texas had been largely suppressed. Villa et al.
(2002) suggest that the relatively high precipitation (greater than 125
cm/year) in the southeastern United States was a key reason why Africanized
bees failed to establish populations east of Texas.
V. COMMUNITY ECOLOGY
Trophic Mode:
Like all honeybees, Apis mellifera scutellata feeds on nectar and pollen.
Pollen is the chief source of dietary protein and nectar (and honey once it is
made through partial-digestion of nectar by workers), is the chief source of
carbohydrate. Honey is stored in honeycombs in the hive and the honey stores
are relied on heavily during the winter or when foraging conditions become
poor. However, the tropical A. m. scutellata does not stockpile honey
to the degree that European bees do (ISSG 2005).
Africanized honeybees are more solitary in their foraging than European bees
and they often forage in adverse conditions where European bees would cease
foraging (Camazine and Morse 1988, Winston 1992). They are also more likely to
abscond from (abandon) their nests during periods of low food availability to
establish new nests elsewhere. This behavior is characteristic of tropical
honeybee strains evolved in areas experiencing frequent prolonged adverse
environmental conditions (Camazine and Morse 1988, Schneider 1990).
Associated Species:
Apis mellifera scutellata co-occurs with and typically hybridizes or outcompetes
the European honeybee strain in Florida.
VI. INVASION INFORMATION
Invasion History:
The history of the introduction of Apis mellifera scutellata to the Americas is
well-documented. The strain was first domesticated in the scrub desert
environment of central South Africa and became known as an aggressive but
productive, disease-resistant strain that produced quality honey.
Growth of the honey industry in Brazil In the 1950s was tempered by relatively
poor performance of imported European honeybees. In 1956 the Brazilian
government sent a geneticist named Warwick Kerr to collect African honeybee
queens to bring back for experimental breeding. Interbreeding
African queens and European drones (via artificial insemination) produced the
hybrid strain that came to be known as Africanized honeybees (Kerr 1967,
Winston 1992).
The accidental release by a Brazilian beekeeper of more than two-dozen AHB
queens in 1957 led to widespread hybridization with naturalized European
honeybee populations. The reasonable assumption that outcrossing would quickly
dilute the aggressive tendencies of the Africanized strain proved to be
incorrect as reports of feral bee swarms attacking livestock began to multiply.
Africanized honeybees spread rapidly through continued outcrossing with
naturalized honeybee populations, from Brazil up though South America and into
Central America and Mexico. The strain first arrived in the United States in
1990, when they were identified near Hidalgo, TX (Kaplan 2004).
From 1995-2005, A. m. scutellata were captured in bait hives maintained
by Florida Department of Agriculture and Consumer Services staff at shipping
ports such as Jacksonville, Miami and Tampa. Throughout this period,
Africanized bees appeared unable to gain a foothold in the state (Woods 2005).
As of 2005, however, established A. m. scutellata populations have been
documented in more than a dozen counties in peninsular Florida. The arrival in
Florida appears to be disjunct to the spread of A. m. scutellata from
Texas. More likely, Florida introductions were human-facilitated as shipping,
trucking, and/or rail transport may have accidentally carried undetected
Africanized bees into the state.
Potential to Compete With Natives:
The rapid spread of Apis mellifera scutellata into new areas is largely
accomplished through hybridization with naturalized European honeybees. The
reproductive advantages enjoyed by A. m. scutellata (see above)
typically allow it to eventually dominate areas formerly occupied by
naturalized European strain colonies (Kaplan 2004).
European honeybee colonies can also be directly taken over by Africanized bees.
Takeovers are perpetrated by swarming or absconding queens and a contingent of
workers and appear to primarily be a means to gain access to favorable nest
sites, food resources, and/or large worker populations (Vergara et al. 1993).
Queenless colonies and small or failing colonies are most likely to be taken
over but large, strong colonies can be infiltrated as well (Rinderer and
Helmich 1991) The actual process of colony takeover is complex and not
entirely understood, but it involves the initial acquisition of the resident
colony odor by invading Africanized bees prior to hive invasion and killing of
resident queens if present (Vergara et al. 1993).
Possible Economic Consequences of Invasion:
Honeybees may be the most economically important insects in the world. The
first European honeybees were brought to the New World perhaps three centuries
ago, and the economic benefit of this historic introduction here and elsewhere
throughout the world is incalculable. In addition to producing honey and
beeswax, honeybees also pollinate a great many agricultural crops. A 2007 New York Times article cited a
Cornell University study estimating the annual value of honeybee crop
pollination in the U.S. at more than $14 billion.
The notion that African honeybees are "killer bees" is largely hyperbole,
although a handful of U.S. fatalities (and at least 175 from Mexico since 1985)
indicate that Africanize hives should be treated with caution. Although the
sting of a single Africanized bee is no worse than that of a single European
bee, Africanized bees do swarm more readily and defend the hive more
aggressively than European varieties.
If the number of Africanized bee colonies continues to expand, takeovers of
cultivated European hives by Africanized queens during periods of high swarming
could cause considerable problems for Florida beekeepers. Potential economic
problems for Florida beekeepers if their stocks became Africanized are probably
those relating to the tendencies of A. m. scutellata for producing less
stored honey and for excessive swarming and nest abandonment (absconding).
None of those attributes are desirable in cultivated honeybee hives.
VII.
REFERENCES
Abramson C.I., Aquino I.S., Azeredo G.A., and J.M. Price. 1997. Some
preliminary studies on the ability of Africanized honey bees (Apis
mellifera L.) to tolerate cold temperatures when placed inside a
refrigerator. Psychological Reports 81:707-718.
APHIS. 1993. Africanized honey bee fact sheet. Document FACTS-05. Available
online.
Breed M.D. 2003. Animal Behavior. Online textbook available online.
Camazine S. and R. Morse. 1988. The Africanized Honeybee. American Scientist
76:465-471.
Collins A.M. 2006. Sperm storage in Apis mellifera, proteomics, genomics
and technology. Invited Symposia Presentation, International Union for the
Study of Social Insects (IUSSI) 2006 Congress. Abstract available online.
Kaplan J.K. 2004. What's buzzing with Africanized honey bees? USDA Agricultural
Research Magazine 52:4-8. Available online.
Kerr W.E. 1967. The history of the introduction of Africanized honey bees to
Brazil. South African Bee Journal. 39:3-5.
Rinderer T.E. 1986. Africanized bees: An overview. American Bee Journal.
126:98-100.
Sanford M.T. and H.G. Hall. 2005. African honey bee: What you need to know.
UF/IFAS Fact Sheet ENY-114. Reviewed: March 1995. Revised: September 2005.
Available online.
Vergara C., Dietz A., and A. Perez-de-Leon. 1993. Female Parasitism of European
Honey Bees by Africanized Honey Bee Swarms in Mexico. Scientific American 269:
84-90.
Villa J.D., Rinderer T.E., and J.A. Stelzer. 2002. Answers to the puzzling
distribution of Africanized bees in the United States or "Why are those bees
not moving east of Texas?" American Bee Journal 142:480-483.
Winston M.L. 1992. Killer Bees. The Africanized Honey Bee in the Americas.
Harvard University Press, Cambridge, Mass. 176p.
Report by:
J. Masterson, Smithsonian Marine Station
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