The genealogy of honey bees in the United States
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by Scott McArt
The genealogy of honey bees in the United States
T
he Western honey bee, Apis that the pedigree of your bees matters this diversity and differentiation that
mellifera, is not native to the since those genes are passed down provides the raw material necessary
United States. Most beekeep- from the previous generation. In other for breeding.
ers know this. words, knowing the genetics and ge- So, when and where were honey
But do you know the genealogy nealogy of honey bees can help us un- bees first introduced to the United
of A. mellifera (hereafter, the “honey derstand their history, but it can also States? 1622 in Jamestown, Virginia
bee”) in the United States? In other allow us to understand their future in (see Figure 1). If you’re sitting in Vir-
words, do you know the full story of terms of the diversity of traits that can ginia right now and reading this ar-
when honey bees came to the USA, be improved through breeding. ticle, you’re sitting in the region that’s
how many times they were intro- For their study, Carpenter and enjoyed the longest history of honey
duced, where each of those intro- Harpur tapped into two major re- bees in the USA, starting with the first
ductions occurred, how quickly they sources. First, they did some good introduction by the Virginia Compa-
spread from one part of our country to old-fashioned work in the library. ny from England.
another, and which lineages are most Because honey bees have been in- Three other locations were relative-
abundant in your neighborhood as a tensively managed for nearly 11,000 ly quick to follow. Honey bees were
result of this history? These are the years, we humans have been pretty introduced by English colonists to
topics for our fortieth Notes from the good about noticing them and writ- Plymouth and Boston, Massachusetts
Lab, where we summarize “Genetic ing down our observations. So, if in 1637 and 1640, respectively, and by
past, present, and future of the hon- you want to know when and where English colonists to Long Island, New
ey bee (Apis mellifera) in the United honey bees were present, you can York in 1670 (Figure 1). Between 1670
States of America,” written by Made- read old documents to see when and and 1845, only two additional impor-
line Carpenter and Brock Harpur and where someone mentions them in tation records exist; Pensacola, Florida
published in Apidologie [2021]. historical records. The authors pored in 1763, and Mobile, Alabama in 1773.
More than 25 million people have over thousands of documents, ul- During this time, honey bees spread
used 23andMe, Ancestry.com, or oth- timately finding nearly 200 records west to the Mississippi by two mecha-
er genetic tests to trace their genealo- that indicated the first time honey nisms: trade among humans and nat-
gies. Many of us are clearly curious bees were observed in a particular ural swarming. For example, in the
about where our great-great-great- geographic location in the USA. 1700s, it was common for “bee trees”
grandparents came from! Similarly, Second, nearly two dozen honey to be found throughout the Carolinas
I can assure you from talking with bee population genetic studies have and Pennsylvania. Then, in the mid-
hundreds of beekeepers over the been conducted in the United States 1800s, importations started to pick up
years that every single one of us is in- to date, and more studies are in the in the western United States. In 1845,
terested in the genealogy of our bees. works. Carpenter and Harpur sum- honey bees were imported to Texas by
Some beekeepers are simply curious marized all existing scientific litera- German settlers, and numerous intro-
to know the lineage of their bees. Do ture on this topic. Ultimately, these ductions occurred on the west coast
I have Italian bees? Are my bees Carnio- genetic studies allow us to under- (California, Oregon, Washington) and
lans? But many beekeepers also know stand how introductions of various Hawaii in the 1850s (Figure 1).
that specific traits, such as varroa resis- lineages have shaped the genetic di- Which lineages were introduced?
tance, have a genetic basis (e.g., Pur- versity and differentiation of honey At least nine subspecies from four of
due “mite biting” bees). This means bee populations across the USA. It is the five A. mellifera lineages have been
April 2021 417
Fig. 1 Estimated A. mellifera population expansions in the mainland United States. Locations have been recorded or estimated from digitized importation records of European colonialists or their accounts of swarm sightings. Sightings are given a specific location if possible; otherwise, the date text is larger and imposed over the state or area the sighting occurred. Color gradients are proposed ranges estimated by combining known sightings and introductions with an estimation of average yearly spread as (50 years – es- tablishment date) x 16.5 km. Alaska has been excluded because honey bees were first introduced to the state in 1924 and cannot be attributed to a source population. Dates before 1859 are exclusively A.m. mellifera sightings, but dates after 1859 can be attributed to A.m. ligustica, A.m. mellifera, A.m. ligustica x A.m. mellifera hybrids, and/or various A. mellifera hybrids. introduced to the USA. But for over northern Europe and central Asia were the next to arrive, imported in 200 years (between 1622 and 1859), and include the English “dark bees,” 1859 to New York City and Philadel- the only honey bee subspecies pres- which were the first honey bees to ar- phia from Germany and Switzerland ent in the United States was A.m. mel- rive in Jamestown. (Figure 2, red square). In the 1860s, lifera (Apis mellifera mellifera). Bees from Italian bees (A.m. ligustica; lineage Italian queens were imported to sev- this lineage (lineage M) come from C, from central and southern Europe) eral locations in the USA from the Fig. 2 Seven subspecies were introduced to the mainland United States between 1859 and 1922: A.m. ligustica (1859), A.m. la- marckii (1866), A.m. carnica (1877), A.m. cypria (1880), A.m. syriaca (1880), A.m. caucasica (1882), and A.m. intermissa (1891). Here, we have highlighted the first 5 years of a recorded importation of a given subspecies into the mainland United States. The excep- tions are A.m. caucasica and A.m. cypria, both of which were imported sporadically or for a short time before importations ceased and were continued under Benton. One series of A.m. cypria and A.m. syriaca importations to Beeton (now New Tecumseth), Canada is recorded because these were the source of importations to the USA. The boxes represent large-scale operations in Ohio (Rev. L.L. Langstroth); Washington, D.C. (Frank Benton); and Canada (D. A. Jones). 418 American Bee Journal
Fig. 3 Summary of locations, methods, and sample sizes of honey bee genetic studies in the U.S., as well as lineages or subspe- cies (when identifiable) detected. Both managed and feral populations are included in each state, but there is considerable — if not complete — overlap between lineages and/or mitotypes found within a state’s feral and commercial colonies. The studies represented in this figure specified the ancestry of individual samples. Overall, the figure highlights two major shortcomings in U.S. honey bee genomics: small sample sizes (11 states have fewer than 11 samples) and biases toward either managed or feral colonies within states. Furthermore, despite the documented presence of AHB in Louisiana, Alabama, Mississippi, and Georgia, sampling methods have missed them there. Lake Maggiore region of northern ing, and/or defensive behavior. This potentially arrive. Finally, since 2009 Italy, until it became more common in is why most readers in the USA have there have been at least 11 USDA-ap- the 1890s to import from Milan and probably rarely heard of Cyprian, proved importations of germplasm to Bologna. Syrian, Egyptian, or Tellian bees; they Washington State for use in breeding Carniolan bees (A.m. carnica; lin- are not common lineages of bees in efforts. eage C) were first imported in 1877 the United States due to their impor- Which subspecies of honey bees to Hamilton, IL (Figure 2, red circle) tation history. are most common in the USA right by none other than Charles Dadant, Well, there were certainly a lot now? Italians and Carniolans. Or, the founder of Dadant & Sons, which of introductions, especially in the more precisely, current genetic evi- publishes the fine magazine you’re 1800s! What’s happened since then? dence shows the two most common currently reading. Carniolan bees Two events in the early 1900s reduced mitotypes of honey bees in the United were imported throughout the late honey bee importations in the USA. States are C1 (A.m. ligustica) and C2 1800s from areas in Dalmatia and Isle of Wight disease (1906-1916) and (A.m. carnica), as shown in Figure 3. Carniola (modern-day Croatia and World War I (1914-1918) dramatically The fact that lineage C (A.m. li- Slovenia) and Hungary. reduced the number of colonies im- gustica and A.m. carnica) is currently Caucasian bees (A.m. caucasica; ported from Europe; the former ul- more common than lineage M (A.m. lineage O, from the Middle East and timately led to the Bee Act of 1922, mellifera) likely says something about western Asia) were first imported in which forbade the importation of live how popular Italian and Carnio- 1883 to upstate New York (Figure 2, honey bees and hive products to the lan bees were in the late 1800s and blue circle). These bees were initially USA. This act was expanded in the how popular some of their traits are imported from beekeepers in Ger- 1960s to include germplasm, due to with beekeepers today. Even though many, but in the early 1900s, queens concern about Africanized honey A.m. mellifera was the only subspe- were imported directly from opera- bees (AHB) from South and Central cies present in the USA for over 200 tions on the shores of the Black and America. In 1987, importations from years (between 1622 and 1859), the Caspian seas. Canada were also restricted over con- late 1800s saw a dramatic increase Four additional subspecies of A. cerns about AHB. in importation and distribution of mellifera have been imported to the Limited importations commenced A.m. ligustica and A.m. carnica. That USA: The Cyprian honey bee (A.m. again in 1989 after the identification period in the late 1800s appears to cypria; lineage O), Syrian honey bee of varroa-resistant strains in Europe. have had a large and lasting impact (A.m. syriaca; lineage O), Egyptian Importation of varroa-tolerant strains on present-day honey bee genetics in honey bee (A.m. lamarkii; lineage A, were also allowed from Russia be- the United States. from Africa), and Tellian honey bee tween 1997 and 2002. From 2004- Is there enough genetic diver- (A.m. intermissa; lineage A). Records 2010, importations from New Zea- sity and differentiation to select for indicate they weren’t distributed land and Australia occurred, but were traits that will keep my bees strong widely because each of the lineages halted in 2010 out of concern that the and healthy? This, of course, is the was prone to swarming, abscond- Eastern honey bee, Apis cerana, could million-dollar question. Can it be April 2021 419
answered? Well, on the one hand, Reference:
the study by Carpenter and Harpur Carpenter, M. H. and B. A. Harpur. Genet-
shows we definitely need more genet- ic past, present, and future of the honey
bee (Apis mellifera) in the United States
ic and genomic data for honey bees in
of America. Apidologie (2021). https://doi.
the United States (for example, see org/10.1007/s13592-020-00836-4
the big blank areas in Figure 3).
But on the other hand, genetic di- Scott McArt, an
versity of honey bees in the USA is Assistant Professor
high. Several studies have shown that of Pollinator Health,
genetic diversity of bees in the USA is helps run the Dyce
just as high as (or higher than) genetic Lab for Honey Bee
Studies at Cornell
diversity in Europe. In addition, new University in Ithaca,
genomic studies show there are re- New York. He is par-
gional differences in honey bees, and ticularly interested
genetically unique feral populations in scientific research that can inform man-
exist. Thus, the evidence to date indi- agement decisions by beekeepers, growers
cates honey bees in the USA have high and the public.
genetic diversity and differentiation. Email: shm33@cornell.edu
In other words, breeders have quite Lab website: blogs.cornell.edu/mcartlab
a bit of raw material to work with Pollinator Network: pollinator.cals.cornell.edu
when selecting for traits in their bees. Facebook: facebook.com/dycelab
While it’s impossible to know wheth-
er it will be enough raw material giv-
en all the current and future stresses
that bees may face, it certainly seems
promising. Especially if we recognize
the importance of good breeders (and
genetic/genomic researchers!) and
give them as much support as we can.
Until next time, bee well and do
good work.
Scott McArt
420 American Bee Journal
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