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TOMAS MOZER, APICULTURAL CONSULTANT/APIARY
TECHNICIAN, UNIVERSITY OF FLORIDA.
The incomplete historical record documenting the origins of Apis
mellifera in the Americas is open to interpretation. Crane, E.
(1999) cites: "We do not know when honey bees first reached
New Spain (Mexico)... As early as 1513, Herrera's Obra de Agricultura
mentioned the difficulty of transporting bees to the newly discovered
Indies... Brand (1970, pblshd. 1988) concluded...that the Spanish
probably introduced honey bees in the 1520's or 1530's... Perkins
(1926) quoted verbatim from a report written in the 1600's: 'a
swarm must have settled on a ship about to sail from Spain...and
on reaching what is now Veracruz the bees flew ashore and...in[to]
a cask which a priest provided as a hive'." [The World History
of Beekeeping and Honey Hunting; Routledge, N.Y.: (p. 361...)]
Nelson, E. V. (1967) in "History
of Beekeeping in the United States" comments: "The
actual date of importation of the first colonies of honey bees...to
North America is unknown... In 1763, the English introduced colonies
of bees into Florida, although they may have been brought earlier
to St. Augustine by the Spanish." [Beekeeping in the United
States; USDA/ARS Agriculture Handbook no. 335:(p.2...)]
Oertel, E. (1976) writes in
"Bicentennial bees: early records of honey bees in the eastern
United States" that "Bartram (1792)...noted...bee tree[s]
on the banks of the [lower] St. John's river in 1765 and... considerable
honey [harvests]... In 1765 De Brahm...began an official land
survey in East Florida...[and reported]...bees were frequently
seen... Barton (1802) stated that the honey bees in Florida,
after having been introduced by the Spaniards, had by 1785 increased
into innumerable swarms... We can speculate...about...the feasible
routes for swarms to take, whether natural swarms or box hives
transported by man...[including] along the Gulf coast."
[American Bee Journal 116:70...]
Attempts to elucidate the nature
of "New World" feral honey bees need to incorporate
observations by Daly, H. V. et al. (1991): "Feral honey
bees in California are mongrel populations, partially differentiated
in
morphometrics from managed colonies as well as from European
subspecies...The geographic variation is presumably adaptive...[and]
points to an underlying genetic differentiation among feral populations
in California." ["Clinal geographic variation in feral
honey bees in
California, USA"; Apidologie 22:591-609. (abstract):
http://danrrec.ucdavis.edu/sierra_foothill/
research_publications7.html#g_8]
A preliminary analysis by Mozer, T. (2002) of Florida "ferals"
found in the vicinity of ports-of-entry points to similarities
in reports of the survival of feral bees with "non-commercial"
lineage, suggesting naturalized "New World" ecotypes
that are conceivably descendants of the earliest introductions.
["Observations on feral honey bees in Florida, USA";
http://www.flareal.com/fsbanews.htm]
The complexities of understanding
honey bee ancestry are illustrated in a study by Schiff, N. M.
& Sheppard, W. S. (1993) on genetic variation of feral populations
in the southern U.S. in which "422 feral bee colonies [were]
sampled...from nine states (Florida was not included)... [and]
results are as follows: [~22% were of] European honey bee races
considered 'western' races (A. mellifera mellifera/A. mellifera
iberica)...[~77% were of] 'eastern' races (A. mellifera carnica/A.
mellifera ligustica)...thought to make up the majority of the
present U.S. commercial honey bee stock...[~1%] African mtDNA
was from the Egyptian honey bee (A. mellifera lamarckii), not
A. mellifera scutellata [from sub-Saharan Africa]." ["Mitochondrial
DNA evidence for the 19th century introduction of African honey
bees into the United States"; Experientia 49:350-352. (cited
in APIS newsletter):
http://www.ifas.ufl.edu/~mts/apishtm/apis94/apjul94.htm#2]
In a later publication, Schiff
& Sheppard (1995) report that "commercial
bees...[are] significantly different than the feral population
of the
southern United States, where 36.7% of 692 feral colonies had
the A.m. mellifera/iberica haplotype (Schiff et al. 1994). The
lack of A.m. mellifera[/iberica] haplotypes in the commercial
population is indicative of restricted gene flow between feral
and commercial populations. Until A.m. ligustica was introduced
by bee breeders in 1859, A.m. mellifera[/iberica] was the only
subspecies present in the United States. Gene flow between commercial
and feral populations likely has occured through swarming and
open matings since that time. However, the maternally inherited
mtDNA of A.m. mellifera[/iberica] has made little intrusion into
commercial populations, demonstrated by the low frequency of
A.m. mellifera[/iberica] mtDNA haplotypes in this group (3%).
Perhaps, through selection or other breeding practices, bee breeders
have contributed to this asymmetry in mtDNA haplotype frequencies.
A [third] haplotype, A.m. lamarckii, which was present in 2%
of feral colonies (Schiff et al. 1994) was not found in the sampled
breeder queen population." ["Genetic Analysis of Commercial
Honey Bees from the Southeastern United States". J. Econ.
Entomol.
88(5):1216-1220. http://www.beesource.com/pov/ahb/jee1995.htm]
Coincidentally, Hall, H. G.
& McMichael, A. M. (2001) found that "Frequencies of
Restriction Fragment-Length Polymorphisms Indicate That Neotropical
Honey Bee Populations Have African and West European Origins:
In this study, ancestry in New World bees was inferred from allele
identities and frequencies at a highly polymorphic nuclear locus
in Old and New World honey bee populations...in combination with
mitochondrial DNA types. In bees from the United States, collected
before the invasion of African bees, east and west European alleles
were found at frequencies of 83 and 17%, respectively, which
is consistent with previously identified nuclear and mitochondrial
DNA markers. Colonies from two neotropical countries, Mexico
and Honduras, had African mitochondrial DNA and high frequencies
of African nuclear DNA alleles. Consistent with previous findings,
east European alleles were absent or detected at low frequencies
in these colonies. However, west European alleles were found
at frequencies from 26 to 31%. These results suggest that queen
offspring of the African queens first introduced into Brazil
mated with west European drones, incorporating neutral markers
that have since remained in the expanding population of feral
African bees. The results point to little paternal introgression
from managed east European colonies encountered by the African
bees spreading through the neotropics."
[Ann. Entomol. Soc. Am. 94(5): 670-676.
http://esa.edoc.com/annals/v94n5/v94n5p670.pdf]
It is interesting to note an
apparent parallel in an "Old World" setting, as communicated
by Franck, P. et al. (1998): "Apis mellifera is composed
of [at least] three evolutionary branches including mainly African
(branch A), West and North European (branch M) and South-East
European (branch C) populations. The existence of morphological
clines extending from the equator to the polar circle through
Morocco and Spain raised the hypothesis that the branch M originated
in Africa. Mitochondrial DNA analysis revealed that branches
A and M were characterized by highly diverged lineages implying
very remote links between both branches. It also revealed that
mtDNA haplotypes from lineages A coexisted with haplotypes M
in the Iberian peninsula and formed a South-North frequency cline,
suggesting that this area could be a secondary contact zone between
the two branches... [The] higher haplotype A variability observed
in Spanish and Portuguese samples compared to that found in Africa
is explained by a higher mutation rate and multiple and recent
introductions. Selection appears as the best explanation to the
morphological and allozymic clines and to the diffusion and maintenance
of African haplotypes in Spain and Portugal." ["The
origin of West European subspecies of honeybees: new insights
from mitochondrial DNA and microsatellite data"; Evolution
52(4):1119-1134. (abstract):
http://www.ensam.inra.fr/URLB/apis/evolution/evolution.html]
Whether the remnant populations
in Florida will continue to survive in the face of numerous stresses
remains unresolved in light of similar scenarios, including the
fate of feral honey bees in Arizona and Texas as observed by
Baum, K. A. et al. (199?): "Few studies have documented
population trends for feral colonies, but Loper (1997) reported
an 82 percent decline in spring populations from 1992 to 1997
[in Arizona]...The observed population trends [in Texas] correspond
to the arrival of Varroa mites and Africanized honey bees. Varroa
mites were first recorded on the Welder Refuge in 1995. By 1996,
the feral populations had declined dramatically. Africanization
began to increase in 1995, with all the colonies Africanized
by 1998...These data suggest populations of European feral honey
bees were decimated by Varroa mites in 1995, and then replaced
by Africanized honey bees in the following years." ["Effects
of Landscape Pattern on the Distribution of Feral Honey Bee Colonies
in South Texas"; http://kelab.tamu.edu/standard/honeybees/].
However, recent findings (pers. communication, W. L. Rubink)
of ~25% European mtDNA persisting in the Texas feral population
are encouraging.
Likewise, reports of mite tolerant
populations from Arizona give one reason for optimism, as anecdotal
evidence shows our study population in northeast Florida approaches
infestation levels over 5 years without treatments (pers. observations,
T. Mozer), comparable to those of Erickson, E. et al. (1999):
"A long term study was undertaken to determine whether a
varroa-tolerant honey bee population could be developed and maintained
via selective breeding and conventional beekeeping practices,
and without the use of other mite control strategies. The results
of this study conducted at an isolated site
demonstrate that it is relatively easy to find varroa-tolerant
colonies, and to produce and maintain varroa-tolerant strains
of honey bees. This Varroa tolerant population has survived for
nearly five years with mean annual infestation level between
6 and 7 percent." ["Varroa-Tolerant Honey Bees Are
A Reality", (abstract):
http://www.nps.ars.usda.gov/publications/
publications.htm?lognum=0000106275]
Research is currently underway
at the University of Florida beelab
[http://www.ifas.ufl.edu/~hgh/deptpage/hgh_biog.htm]
on quantitative trait loci (QTL) involved in suppression of Varroa
mite reproduction (SMR): "SMR is latent in bee populations
and can be greatly enhanced through selective breeding. The goal
of this project is to find DNA markers associated with SMR, thereby
locating the genes responsible (QTL) along genetic maps. Markers
associated with the trait would facilitate selecting from different
sources of bees and combining SMR with other desirable traits.
This project will use bees from Louisiana already selected for
SMR and will select for SMR from Florida bees that have survived
the mite without treatment." [Abstract 2001-02839;
http://www.reeusda.gov/nri/pubs/abstracts/2001/entnem.pdf]
Disclaimer: the views expressed
do not represent the State and/or University of Florida's official
positions. Copyright © 2002 by Tomas Mozer, all rights reserved.
Copyleft: Verbatim copying of this document is permitted, in
any medium, under Design Science License terms and conditions
(see
http://dsl.org/copyleft/dsl.txt).
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