Pages 36–39: The study of the density of colonies in two national parks, and a third rural area, in Germany, done with an indirect, genetic method, is that of Moritz, Kraus et al. (2007). The more recent study of colony density in two areas of natural beech forests in Germany, done by means of bee hunting (bee lining) and inspecting the nest cavities of black woodpeckers, is that of Kohl and Rutschmann (2018).
Page 39: The study for three national parks in Australia is that of Hinson et al. (2015).
Page 41: The survey of the wild colonies in part of the Shindagin Hollow State Forest, in New York State, by means of bee hunting, is reported in Radcliffe and Seeley (2018). For a study of the density of honey bee colonies in various locations across the natural range of Apis mellifera (Europe, Africa, and central Asia), see Jaffé et al. (2009). It should be noted, however, that the estimates of colony density reported in Jaffé et al. (2009) include both managed and wild colonies.
Pages 42–44: The biology of Varroa destructor is reviewed by De Jong (1997). See also, Varroa mite, Featured Creatures, University of Florida, http://entnemdept.ufl.edu/creatures/misc/bees
/varroa_mite.htm (accessed 10 July 2017). For detailed information on this mite’s native distribution on Apis cerana in eastern Asia, the history of its host shift to Apis mellifera in eastern Russia, and its subsequent dispersal with the aid of beekeepers to Europe, Africa, and South
America, see De Jong et al. (1982). Anderson and Trueman (2000) found that the Varroa mites infesting Apis cerana consist of two species— V. jacobsoni and V. destructor—and that the mites infesting Apis mellifera nearly worldwide are all members of the species V. destructor. This original host species of V. destructor was A. cerana on mainland Asia; the host species for V. jacobsoni remains A. cerana on the islands of the Malaysia–Indonesia–New Guinea region.
Page 44: The introduction of Varroa destructor to North America and its rapid spread across the continent are reviewed by Wenner and Bushing (1996) and Sanford (2001). The information on Africanized
bee swarms found in Florida on eight ships from Central and South America between 1983 and
1989 comes from the records “Florida Africanized Bee Interceptions,” provided to me by Dave
Westervelt, of the Bureau of Plant and Apiary Inspection for the state of Florida.
Page 46: The paper by Bernhard Kraus and Robert E. Page Jr. that reported the extremely grim news about California’s population of wild colonies is Kraus and Page (1995).
Pages 46–47: Gerald Loper describes his studies of a population of wild colonies living north of Tucson in Loper (1995, 1997, and 2002). He also provides a follow- up report in Loper et al. (2006).
Pages 48–56: Full reports on the mapping of the wild honey bee colonies in the Arnot Forest in the autumn of 2002, and on the testing of these colonies for infestations with Varroa destructor in 2003
and 2004, are found in Seeley (2007). The earliest report of this work is found in Seeley (2003).
CHAPTER 3. LEAVING THE WILD
Page 57: The Euell Gibbons quotation is from his book Stalking the Wild Asparagus; see Gibbons (1962), p. 235.
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Page 57: The view that our prehuman ancestors (other members of the genus Homo and their direct ancestors, including Australopithecines) consumed and enjoyed honey is supported by the fact that several species of nonhuman primates—including baboons, macaques, chimpanzees, gorillas, and
orangutans—are also successful honey hunters. This topic is reviewed by Crittenden (2011).
Chimpanzees in Gabon are so fond of honey that they will prepare a three- piece tool kit for raiding the nest of a honey bee colony. It includes a pounder (a heavy stick for breaking open the nest
entrance), an enlarger (to enlarge the opening into the nest), and a collector (a stick with a frayed end to poke into the honey combs and then into the mouth, slurping the honey off). See Boesch et al. (2009)
Page 57: The first correct assignment of a fossil bee to the honey bee genus Apis is that of Cockerell (1907). For more information about fossil honey bees, see the monograph on the fossil bees of the world by Zeuner and Manning (1976), the recent review of fossil honey bees by Engel (1998), and
the definitive work on the evolution of insects by Grimaldi and Engel (2005).
Page 58: The oldest well- dated evidence of Homo sapiens is a fossil skull found during mining operations in the Jebel Irhoud massif near the west coast of Morocco. Its age is 315,000 years, plus or minus 34,000 years, as determined by thermoluminescence dating. For a recent analysis of this skull, see Hublin et al. (2017).
Page 58: The complex story of the origins of Homo sapiens in Africa and subsequent migrations into Asia and Europe, and eventually the entire world, is reviewed by Wenke (1999). Gibbons (2017) and
Hublin et al. (2017) provide updates.
Page 58: For analyses of the caloric content of honey, see White et al. (1962) or Murray et al. (2001).
The latter reference provides results for honey collected by the Hadza of Tanzania, thus for honey as it would have been eaten by early humans: fortified with proteins and fats from crushed bee larvae and pupae.
Pages 58–59: For detailed reports on honey hunting by the Hadza of Tanzania, see Marlowe et al. (2014) and Wood et al. (2014). A recent paper (Smits et al. 2017) describes how the gut microbiome of
the Hadza hunter- gatherers changes between the wet and dry seasons so they maintain a healthy
gut as they shift from heavy consumption of meat in the dry season to consumption of honey, ber-
ries, and other fruits in the wet season. For detailed reports on honey hunting by the Efe (and their close relatives, the Mbuti) of the Ituri Forest in the Democratic Republic of the Congo, see Turnbull (1976), Ichikawa (1981), and Terashima (1998). To enjoy a short novel on honey hunting and rock
art painting by early people of southern Africa, see Dixon (2015).
Page 60–61: The evidence from rock art of humans gathering honey from wild colonies of honey bees is thoroughly reviewed in Crane (1999). The monograph by Hernández- Pacheco (1924) is the
definitive reference for the prehistoric paintings found in 1917 in the Araña cave complex in Spain.
Dams and Dams (1977) report their discovery in 1976 of additional rock art depicting honey
gathering during the Mesolithic in eastern Spain.
Page 62: The stone bas- relief in pharaoh Nyuserre’s temple to the sun god Re is discussed in further detail in chapter 20 of Crane (1999) and chapter 2 of Kritsky (2015). Another rich source of well-preserved scenes of beekeeping and other activities of daily life in ancient Egypt is the lavishly decorated tomb (at Thebes) of Rekhmire, the vizier for two pharaohs from about 1470 to 1445 bce; see Garis Davies (1944).
Page 62–63: For more information on the Iron Age apiary found in Tel Rehov in northern Israel, see Mazar and Panitz- Cohen (2007) and Bloch et al. (2010).
Page 64: The final chapter of Kritsky (2015), cleverly titled “The Afterlife of Ancient Egyptian Beekeeping,” contains excellent photos and detailed descriptions of the tools and techniques of
traditional beekeeping in modern- day Egypt. These probably resemble those used by beekeepers
during Pharaonic times, some 2,000 years ago.
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Page 64–66: For a translation of Columella’s writings on beekeeping in book 9 of his work De re rustica (On agriculture), see Columella (1968).
Pages 66–69: For general information about tree beekeeping in northern Europe, see chapter 16 in Crane (1999). The best source of detailed information about tree beekeeping in Russia is Galton
(1971). The estimate by Prokopovich of the honeycomb per bee tree harvested by tree beekeepers
is reported in Galton (1971), p. 27.
Page 69: Additional information about
tree beekeeping in the South Ural area of Bashkortostan is found in Ilyasov et al. (2015).
Pages 69–72: For excellent reviews of traditional beekeeping in northwestern Europe, where the most widespread traditional hive was an inverted basket, or skep, see chapter 27 in Crane (1999) and
chapters 3 and 4 in Kritsky (2010). The woodcut of two skeps in a shelter shown in Fig. 3.6 is from Münster (1628), p. 1415.
Pages 72–74. The story of Lorenzo L. Langstroth’s discovery of the bee space and his use of it when designing the first fully functional movable- frame hive is told with admirable precision in the biog-raphy of Langstroth by Naile (1976). For more information about the context of Langstroth’s dis-
covery, especially the many parallel attempts in Europe to develop a movable- frame hive, see
Kritsky (2010).
Pages 74–76: For the 30 October 1851 entry from Langstroth’s journal, see Naile (1976), p. 75. The Langstroth quotation, “will give the apiarian perfect control over his bees,” is from the 26 November 1851 entry of his journal; see Naile (1976), p. 79.
Pages 76–78: For reviews of the impact of Langstroth’s movable- frame hive on beekeeping worldwide, and of the follow- on inventions of tools and methods that made beekeeping with movable- frame
hives so productive, see chapters 41 and 43 in Crane (1999).
CHAPTER 4. ARE HONEY BEES DOMESTICATED?
Page 79: The Lorenzo L. Langstroth quotation is from the title of chapter 2 of his book Langstroth on the Hive and the Honey- Bee; see Langstroth (1853).
Page 79: To learn more about domestication in general, see Roberts (2017) and DeMello (2012).
Page 79: The fascinating story of the domestication of the industrial yeasts ( Saccharomyces cerevisiae) used to brew beers, wine, spirits, sake, and bioethanol is reported by Gallone et al. (2016).
Page 80: For more information about the evidence of the exploitation of honey bees by early farmers of the Middle East, see Roffet- Salque et al. (2015).
Page 81: The biblical quotation “a land flowing with milk and honey” is from the Book of Exodus, chapter 3, verses 8 and 17.
Page 81: The evidence that wild colonies of honey bees of European ancestry preferentially occupy rather modest- size nesting cavities, in the range of 20–40 liters (5.3–10.6 gallons), is found in papers by Seeley and Morse (1976, 1978a), Jaycox and Parise (1980, 1981), and Rinderer, Tucker
et al. (1982).
Page 81: To the best of my knowledge, Eva Crane is the first person to hypothesize that hive beekeeping began with swarms of honey bees occupying empty pots and baskets of Neolithic farmers; see Crane (1999), p. 161.
Page 82: The engorgement of worker honey bees before swarming is described in detail by Combs
(1972). Free (1968) describes engorgement by worker honey bees whose colony has been smoked.
Page 82: That colonies started by larger swarms have a higher probability of surviving the first winter is shown in Rangel and Seeley (2012).
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the central nervous system of bees, but the effects of smoke on the bee’s sensory (olfactory) system also play a role. Worker honey bees have reduced sensitivity to the smell of their alarm pheromones when smoked. Smoke probably interferes with the bees’ olfaction generally, not just their detection of alarm pheromones. See Visscher, Vetter et al. (1995).
Page 83–84: For more information on how wild colonies of honey bees living in the Cape Point Nature Reserve in South Africa survived a wildfire there, see Tribe et al. (2017).
Page 85: The archaeological evidence regarding when humans learned to control fire and make use of it at will is described by Gowlett (2016).
Page 85: For more information on industrial dairy farming in New York State, see Kurlansky (2014).
Page 86: Biologists use the concept of heritability to describe the degree to which a particular trait of an organism (or, for honey bees, a colony) is genetically influenced. It varies between 0 and 1 and is used to assess the reliability of using a certain trait as a guide to judging the breeding value of an individual. Table 1 in Collins (1986) lists heritability estimates for traits of both individual honey bees (e.g., worker longevity) and whole colonies of honey bees (e.g., honey production). See also Bienefeld and Pirchner (1990) and Oxley and Oldroyd (2010) for estimates of heritability for
several colony traits (growth rate in spring, wax production, calmness, etc.); the estimates of
heritability of honey production, for example, range from 0.15 to 0.54.
Page 87: For the original description of instrumental insemination, see Watson (1928). The subsequent improvements needed to make instrumental insemination a reliable technique are described in
Laidlaw (1944). An up- to- date description is that of Harbo (1986). A video showing how
instrumental insemination is done is available; see, Instrumental insemination of honey bee queens
– Susan Cobey, YouTube video, 5:23, posted by “tlawrence53,” on 6 January 2009, https://www
.youtube.com/watch?v=Csjy020fpyI (accessed 22 December 2017).
Page 87: American foulbrood (AFB) is the only pathogen- based disease of honey bees that is virulent—
i.e., that can rapidly overcome a colony’s defensive mechanisms and kill it. Fries and Camazine
(2001) explain how this virulence has evolved because the spores of AFB are easily transmitted from one colony to an unrelated colony (horizontal transmission) when a weakened, infected colony is
robbed by bees from other colonies or when a swarm occupies a nest site where the previous colony succumbed to AFB. Ewald (1994) provides a clear evolutionary explanation of why some
pathogens—including those that cause malaria, smallpox, tuberculosis, AIDS in humans, and AFB
in bees—are extremely deadly, while others are not.
Page 87–89: Rothenbuhler (1958) provides a detailed and well- referenced review of the successful breeding program for resistance to American foulbrood conducted by O. Wallace Park and colleagues in the 1930s and 1940s, as well as briefer descriptions of other breeding programs, such as that by Brother Adam in England for resistance to acarine disease (presumed causative agent,
Acarapis woodi). For another excellent, and more recent, review of the breeding for resistance to AFB, see Spivak and Gilliam (1998a).
Page 89–90: Spivak and Gilliam (1998b) provide a detailed and well- referenced review of the excellent studies on hygienic behavior, mainly as a mechanism of defense against chalkbrood, European
foulbrood, and Varroa, since the eras of O. Wallace Park and Walter C. Rothenbuhler.
Page 90–91: To learn more about the remarkable story of the alfalfa bee, see Mackensen and Nye (1966) and Nye and Mackensen (1968, 1970). The breeding of these bees by commercial producers of
alfalfa seed is reported by Cale (1971).
Page 91: Oxley and Oldroyd (2010) and Oldroyd (2012) discuss in greater detail the absence of distinct breeds of honey bees and the fact that honey bees have never been truly domesticated.
Page 91–92: Roberts (2017) reviews the deep history of the domestication of dogs and cattle, along with eight other animals (chickens, horses, and humans) and plants (wheat, apples, potatoes, rice, and maize) that used to live wild.
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Page 92: Some will say that humans have made a fundamental change in the genetics of Apis mellifera, because the honey bee populations that have been established in North America, New Zealand, and
Australia are mixtures of most of the honey bee subspecies of Europe (as shown for Canada by
Harpur et al. 2012) and because the migratory activities of beekeepers and the shipping of queens between countries are starting to homogenize the honey bee subspecies within Eur
ope (De la Rúa
et al. 2009). However, I am not treating these genetic changes as fundamental, because they have not produced distinct subpopulations (breeds) of honey bees that are inbred.
Page 93: Box 5.2 in DeMello (2012) contains a list of all 18 domesticated animals, besides dogs, along with an estimate of the time of domestication for each. The list comprises sheep, cats, goats, pigs, cattle, chickens, guinea pigs, donkeys, ducks, horses, llamas, Bactrian camels, dromedary camels, water buffalo, yaks, alpacas, and turkeys—and, also, honey bees.
Page 94: For more information about the movement of honey bee colonies to the heather moorlands
in Scotland, see Manley (1985) and Badger (2016).
Page 94–95: For a general overview of the migratory beekeeping that is required to meet the demand for pollination during California’s almond bloom, see Ferris Jabr, The mind- boggling math of migra tory beekeeping, Scientific American, 1 September 2013, https://www.scientificamerican.com
/article/migratory-beekeeping-mind-boggling-math/ (accessed 23 December 2017). Jacobsen (2008) and Nordhaus (2011) provide more detailed portraits of the trucking of millions of colonies each year to farmers across the United States who need pollinators.
Page 95–98: For more information about the tools and methods of modern beekeeping, see Flottum
(2014) and Sammataro and Avitabile (2011).
CHAPTER 5. THE NEST
Page 99: The Charles Darwin quotation is from his book On the Origin of Species; see Darwin (1964), p. 224.
Page 99: The concept of looking at the nests of animals as part of their survival equipment is best described by Richard Dawkins when he discusses how the structures built by organisms are part of their “extended phenotypes”; see the final chapter in either Dawkins (1982) or Dawkins (1989).
Page 100: The full report of the study of the natural nests of honey bees living in the woods around Ithaca is found in Seeley and Morse (1976). It describes the methods of nest dissection, including how we measured the nest cavities by filling them with sand after removing the combs. Avitabile et al. (1978) report a related study that was conducted in Connecticut and collected information about nest entrance height, size, and orientation for 108 colonies living in trees. These authors also report finding mostly nest entrances that were low (< 5 m/16.4 ft.), small (< 60 cm2/9.3 sq. in.), and with a southerly orientation.
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