The Lives of Bees
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Page 101: To make a detailed dissection of each nest and conduct an accurate census of the bees within it, I put to death each colony on the day on which its tree was felled. In doing so, I worked to minimize the bees’ suffering. My procedure was to arrive at the bee tree in early morning, when
it was still cool and just starting to get light, so the bees were not yet flying out. I then climbed up the tree, plugged all but one of the nest’s entrance openings with rags, spooned several table-spoons of Cyanogas powder (calcium cyanide) in the one entrance still open, and then plugged it
too. Each time, I heard a rapid rise of buzzing sounds coming from inside the sealed- up nest, but within two minutes there was silence. It was grim work, and I doubt that I could do it today, but I believe that the benefits of the information gained from this study justify the deaths of the 21
colonies studied.
Page 101: Analysis of the distribution of compass directions of the entrances of the bee trees using circular statistics yielded the following results: mean vector bearing, 192° (ca. SSW); mean
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orientation vector, 0.39. This shows that their distribution is biased toward the south and is unques-tionably nonrandom; p < 0.01, Rayleigh test for nonrandomness. See Batschelet (1981).
Page 103: The three surveys of wild colonies in the Arnot Forest are reported in Visscher and Seeley (1982), Seeley (2007), and Seeley, Tarpy et al. (2015). The survey of the wild colonies in the
Shindagin Hollow State Forest is reported in Radcliffe and Seeley (2018).
Page 105: One of the bee- tree nests was odd in having both an immense (204 cm2/31.6 sq. in.) nest entrance and a huge (448 liter/118.3 gal.) nest cavity. The entrance was a gaping opening at the base of a large beech tree ( Fagus grandifolia), and the cavity was a ca. 5 m (16.4 ft.) tall space inside it. The colony had built its nest near the top of this cavity, so even though both entrance and cavity were unusually large, the colony occupied a good, snug nesting site.
Page 108: I regret that we did not take more systematic measurements of the thickness of the wooden walls of these tree cavities, for this turns out to be of great importance for understanding how colony thermoregulation works in these natural cavities. See Mitchell (2016).
Page 108: The high percentage of drone comb that we found in these nests (10%–24%, average 17%)
has been confirmed in another study that described the nests built by colonies whose comb building was not manipulated. The range reported in this second study is 11%–23%, with an average of 20%.
See Smith, Ostwald et al. (2016).
Page 108: My study of small- cell combs as a means of controlling infestations of Varroa mites, which includes data on the cells built on small- cell foundation, is reported in Seeley and Griffin (2011).
Page 109: Edgell (1949) reports an estimated average of 8.5 kg (ca. 19 lb.) of honey collected from 56
bee- tree colonies in central New Hampshire. Their nests were taken up at various times across the summer.
Page 111: The seasonal pattern of swarm emergence for Ithaca was measured over six years, when 126
swarms were collected; see Fell et al. (1977).
Page 111: Rangel, Griffin et al. (2010) document how nest- site scouts can start searching for potential homesites even before their colony casts a swarm. This paper reports scouts inspecting potential nest sites starting two to three days before their colony swarms.
Page 111: For a detailed description of the behavior of a nest- site scout as she inspects a potential nesting cavity to assess its size and other properties, see Seeley (1977). For detailed information about how the scout bees in a swarm work together in a process of collective decision- making to choose a new homesite, see Lindauer (1955) and Seeley (2010).
Page 112: The article written by a French beekeeper on how to build bait hives that are attractive to honey bee swarms is Marchand (1967). The long history of the quest by beekeepers for a perfect
hive is described beautifully by Kritsky (2010).
Page 112: For details of the methods and results of my study of the nest- site preferences of the wild honey bees around Ithaca, see Seeley and Morse (1978a).
Page 116: The paper by Tibor I. Szabo on the benefits of a south- facing nest entrance for bees living where winters are cold and snowy is Szabo (1983a).
Page 116: The paper by Derek M. Mitchell on the benefits to the bees of not having a top entrance is Mitchell (2017).
Pages 116–119: My studies of how scout bees measure the volume of a prospective nest cavity and
what they prefer regarding cavity volume are reported in Seeley (1977). The related studies
done by Elbert R. Jaycox and Stephen G. Parise are reported in Jaycox and Parise (1980, 1981).
The paper that reports the large- scale investigation by Thomas E. Rinderer and colleagues is
Rinderer, Tucker et al. (1982). The report of the related work by Justin O. Schmidt is Schmidt and Hurley (1995).
Page 119: For the full report of the study by Tibor I. Szabo on the benefits of installing swarms in hives whose frames are filled with comb, see Szabo (1983b). That tree beekeepers in Russia placed a high Seeley.indb 300
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value on tree cavities that have already been occupied by colonies is described in Galton (1971).
Three good references on bait hives for honey bees are Marchand (1967), Guy (1971), and Seeley
(2017a).
Page 119–120: The dimensions of the entrance openings used in the test of vertical slit vs. circle were 1 × 7 cm (0.4 × 2.75 in.) for the slit and 3 cm (1.2 in.) in diameter for the circle. In the test of cavity dryness, each nest box received its 2 liters (0.5 gal.) of sawdust (wet or dry) just before it was mounted on its tree or power- line pole. The wet sawdust was created by mixing 2 liters of
sawdust with 1 liter (0.26 gal.) of water. Another liter of water was poured into each box in the wet- sawdust treatment group each time the nest boxes were inspected, which was about every 10
days. For more details on the methods used in the study of the bees’ real- estate preferences, see Seeley and Morse (1978a).
Pages 121–122: For a review of how the wax- gland epithelium (and thus the wax production) changes with age in worker honey bees, see chapter 4 in Hepburn (1986). This chapter also reviews the
studies that have shown that the elderly bees in a colony, which are normally functioning as foragers and have only a thin wax- gland epithelium, can rejuvenate their wax glands when they are members of a swarm.
Page 122: The formula for calculating the area of a hexagon is a2·6 tan (30°), or a2·0.8655, where a is the wall- to- wall dimension (the apothem) of the hexagon. Using this formula, one can calculate that there are approximately 82,051 worker cells in 1.92 m2 (20.7 sq. ft.) of worker comb (apothem
= 5.20 mm/0.205 in., hence the area of a worker cell is 23.40 mm2/0.0363 sq. in.) and
approximately 13,125 drone cells in 0.48 m2 (5.2 sq. ft.) of drone comb (apothem = 6.50
mm/0.256 in., hence the area of a drone cell is 36.57 mm2/0.0567 sq. in.). These areas of worker and drone comb were calculated as 80% and 20%, respectively, of 2.4 m2 (25.8 sq. ft.) of total
comb surface (i.e., counting both surfaces on a comb). This is the average amount of total comb
surface that I have found in nests of wild colonies.
Page 122: Wojciech Skowronek (cited by Hepburn 1986, p. 39) found that a worker honey bee can
produce approximately 20 mg (0.0007 oz.) of beeswax; 60,000 bees × 20 mg of beeswax/bee =
1,200,000 mg, or 1.2 kg (2.6 lb.) of beeswax. The estimate that bees need about 1 g (0.04 oz.) of beeswax to build 20 cm2 (3.1 sq. in.) of comb surface (i.e., 10 cm2/1.6 sq. in. of double- sided comb) comes from Wojciech Skowronek (cited by Hepburn 1986, p. 64).
Page 122: The figures regarding amount of s
ugar carried per swarm bee come from Combs (1972). The mean number of worker bees in a swarm comes from Fell et al. (1977). The estimate of efficiency
of beeswax synthesis from sugar comes from Horstmann (1965) and Weiss (1965).
Page 123: For detailed information on the likelihood of winter survival by colonies that were started by swarms the previous summer (founder colonies), see Seeley (1978) and Seeley (2017b).
Page 125: For information on the circular cross- section shape of the cells in the nests of solitary bees and social bees other than honey bees (e.g., bumble bees and stingless bees), see Michener (1974) or Michener (2000). In the nests of all these non- Apis species, the cells serve only as brood cells.
The social bees that are not honey bees build special honey pots for holding their honey stores.
Page 126: For more information on how worker bees use their antennae to judge the thickness of a cell’s walls during cell construction, see Martin and Lindauer (1966).
Pages 127–128: For a full report on the patterns of nest construction and colony population dynamics after a swarm moves into an empty nest cavity, see Smith, Ostwald et al. (2016). This paper also reports how the colony’s comb area increases, its populations of workers and drones fluctuate, and its honey and pollen stores rise and fall over the 14 months following the colony’s occupation of a new nesting cavity. The dimensions of the large observation hives used in this study are 4.3 cm deep, 88 cm wide, and 100 cm tall (1.7 in. × 34.6 in. × 39.7 in.).
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of a colony’s first- year comb building that occurs within the first four to six weeks of living in a new nest cavity. Smith, Ostwald et al. (2016) report 57% within the first four weeks. Lee and Winston (1985) report 90% within the first six weeks.
Page 127: The tanking up on honey by the worker bees in swarms is described in Combs (1972).
Pages 129–131: The theoretical and experimental studies by Stephen C. Pratt on optimal timing of comb construction are found in Pratt (1999) and are reviewed in Pratt (2004).
Pages 131: Rösch (1927) and Seeley (1982) have reported the match in the age ranges of the bees responsible for comb building and the bees responsible for unloading nectar from incoming foragers and storing it in comb cells.
Pages 131: See Pratt (1998a) for the study with paint- marked bees that showed that the worker bees engaged in nectar receiving do not form a large percentage of the bees engaged in comb building.
Pages 132: That swarms newly settled in their homesites build only worker cells for the first several weeks of construction has been reported by Free (1967), Taber and Owens (1970), and Lee and
Winston (1985), as well as Smith, Ostwald et al. (2016).
Pages 132–135: See Pratt (1998b) for the study of the information pathways used by bees to decide the kind of comb (worker or drone) to build. See also Pratt (2004) for a review of work on this subject and on the matter of when to build comb.
Pages 135–136: See table 12.5 in Crane (1990) for an excellent summary of the plants reported to be sources of propolis collected by the honey bee Apis mellifera. See also Simone- Finstrom and Spivak (2010) for references to more recent papers on this subject.
Page 136: See Meyer (1954) and Meyer (1956) for detailed descriptions of the process whereby a resin collector packs her corbiculae with resin.
Pages 136–138: See Nakamura and Seeley (2006) for detailed information on the close observations of resin collectors and resin users made by Jun Nakamura.
Pages 139: See Simone- Finstrom, Gardner et al. (2010) for the study that discovered better associative learning of tactile stimuli in resin collectors than pollen foragers.
CHAPTER 6. ANNUAL CYCLE
Page 140: The Robert Frost quotation is from his poem “A Prayer in Spring”; see Latham (1969).
Page 142: See Avitabile (1978) for information about the weights of honey bee colonies living in central Connecticut from late autumn to early spring. The climate in central Connecticut is a close match to that of central New York State.
Page 142: See Southwick (1982) for information about the metabolic rate of a typical overwintering colony as a function of ambient temperatures.
Page 143: See McLellan (1977) for a critical evaluation of using change in a honey bee colony’s weight as an indicator of its acquisition (or loss) of energy.
Page 143: See Milum (1956), Mitchener (1955), and Koch (1967) for reviews of records of weight
changes of honey bee colonies across a summer or throughout a year for the United States, Canada, and Germany, respectively.
Page 143: See Seeley and Visscher (1985) for more information about the study in which the weekly weight changes of two unmanaged colonies (simulating wild colonies) were recorded from November 1980 to June 1983.
Page 144: See Farrar (1936) for the details of his study of the influence of pollen reserves on brood rearing and weight losses of honey bee colonies in winter.
Page 148: See Nolan (1925), Allen and Jeffree (1956), Jeffree (1956), and Winston (1981) for examples of describing colony growth patterns by measuring the number of cells of brood in the nest. See
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Jeffree (1955) and Loftus et al. (2016) for examples of doing this by repeatedly censusing a colony’s population of adult bees.
Page 146: See Simpson (1957a) and Gary and Morse (1962) for information on the occurrence of false starts in queen rearing in preparation for swarming.
Page 148: In Fig. 6.4, the data on low levels of brood rearing from mid- November to the end of
February come from Avitabile (1978). The swarm dates data are an extension of the results reported in Fell et al. (1977). Air temperature data come from Brumbach (1965).
Page 148: The evidence that colonies adjust the start of brood rearing in the winter in response to increasing day length comes from a study by Kefuss (1978).
Page 148: For more information on the annual patterns of brood rearing in temperate climates see Nolan (1925) and Jeffree (1955, 1956).
Page 148: See Louveaux (1973) and Strange et al. (2007) for more information about the colony
transplant experiments conducted in France that have shown that the annual cycles of brood rearing are partly under genetic control.
Page 150: For more information about the timing of drone production in colonies at the Rothamsted Experimental Station, see Free and Williams (1975).
Page 151: For more reports on the seasonal timing of swarming see Mitchener (1948) for Manitoba; Murray and Jeffree (1955) for Scotland; Simpson (1957b) for southern England; Fell et al. (1977) and Caron (1980) for northeastern United States; and Page (1982) for central California.
Page 151: The probabilities presented here of colony survival over summer and over winter, and for new and established colonies, come from two long- term studies of wild colony survival that I
conducted in the 1970s and in the 2010s. See Seeley (1978) and Seeley (2017b).
Page 151: To postpone the onset of brood rearing from midwinter to mid- spring in 6 of the 12 colonies used in this study, we caged each colony’s queen between two sheets of plastic queen excluder
material, spaced 8 mm (0.3 in.) apart by strips of wood and inserted between the upper and lower stories of each colony’s hive. This arrangement enabled each queen to move laterally during the
winter and thereby maintain contact with the winter cluster but also prevented her from gaining
access to combs in which to lay eggs. In each of the other six colonies in the study, we did not restrict the queen’s movements, thus we did not limit the colony’s winter brood rearing.
Pages 152: The studies reviewed here, on the critical timing of colony growth and reproduction, are reported in Seeley and Visscher (1985).
; Page 153: For more information on the life cycle of bumble bee colonies, see Heinrich (1979) and Goulson (2010).
Page 153: For more information on how insects overwinter in temperate, subarctic, and even arctic regions, where they are exposed to subfreezing temperatures for some part of the year, see
Chapman (1998), pp. 518–520.
Page 153: For more information on the tundra- dwelling bumble bee, Bombus polaris, see Richards (1973).
Page 153: Michener (1974) provides excellent summaries of the natural history and biogeography of both stingless bees and honey bees.
Page 154: The studies of latitudinal differences in rates of ant predation on undefended wasp larvae are reported in Jeanne (1979).
CHAPTER 7. COLONY REPRODUCTION
Page 155: The George C. Williams quotation is from his book Adaptation and Natural Selection. See the opening sentence of chapter 6 (“Reproductive Physiology and Behavior”) in Williams (1966).
Page 155: A sexual organism (plant or animal) is a simultaneous hermaphrodite if it produces both female and male gametes (eggs and sperm) in each breeding season. See chapter 2 in Charnov
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(1982) for a full discussion of this matter. The gametes of a honey bee colony are its virgin queens and unmated drones.
Page 157: Detailed information on development times for honey bee queens and drones (16 and 24
days), and on the sexual maturation times for queens and drones following emergence from their
cells (minimum values: 6 and 10 days), is found in chapters 4 and 5 of Koeniger, Koeniger, Ellis et al. (2014) and Koeniger, Koeniger, and Tiesler (2014).
Page 159: To convert an area of capped drone comb (in Page 1981), or an area of brood- filled drone comb (in Smith, Ostwald et al. 2016), into the number of cells of drone brood, I multiplied the