The Lives of Bees
Page 39
reported values of drone comb (in cm2) by 2.73 cells/cm2. This is the density of hexagonal cells that have an apothem (wall- to- wall dimension) of 6.5 mm (0.24 in.), which is the average dimension of drone cells for European honey bees. To calculate the total number of drone brood cell- days for a colony, I calculated the areas under the curves reported in Page (1981) and Smith, Ostwald et al.
(2016) of the number of capped (or brood- filled) drone cells in a colony’s nest over the entire summer. To calculate the total number of drones reared by a colony across the summer, I divided
the total number of drone cell- days for a colony by the number of drone cell- days per drone (either 14 days for capped cells of drone brood or 21 days for all cells with drone brood). This assumes every cell that contains a developing drone yields a viable drone.
Page 159: For more information about the study that tested whether honey bee colonies seasonally switch which type of comb (drone or worker) they use preferentially for honey storage, so their
drone comb is empty and ready for brood rearing in the spring, see Smith, Ostwald et al. (2015).
Page 161: For more information about the behavior of worker bees shaking the queen and how it
functions to (among other things) prepare queens to fly out of the hive, either with a swarm or on a mating flight, see Allen (1956, 1958, 1959a), Hammann (1957), and Schneider (1989, 1991).
Page 161: The figure of a 25% reduction in the queen’s weight in preparation for swarming comes from Fell et al. (1977).
Page 161: The evidence that only about a quarter of a colony’s workers are left behind when the swarm containing the mother queen departs is discussed in detail later in the chapter.
Page 161: One wonders how a worker bee in a swarming colony decides whether to stay at home and
support a young (sister) queen or leave in the prime swarm and support the old (mother) queen.
Does she prefer the former option if the new queen could be a full sister but the latter option if all the young queens are only half sisters? A study (Rangel, Mattila et al. 2009) has checked whether the workers are more inclined to stay at home if some of the immature queens are their full sisters, but no evidence of this was found. Thus, it seems clear that there is no intracolonial nepotism during swarming in Apis mellifera.
Page 161: The wondrous process whereby the scout bees in a swarm select their new homesite is
described in detail in Seeley (2010).
Page 162: The information about the large dispersal distances of honey bee swarms is found in Seeley and Morse (1978b) and Kohl and Rutschmann (2018).
Page 162: The value of 0.87 for the probability of swarming by an unmanaged colony during a summer comes from my study with simulated wild colonies living in the woods around Ithaca, reported in
Seeley (2017b).
Page 163: For an impressively detailed description of the mechanisms by which all but one of the unmated queens produced during the swarming process are eliminated from the parental nest, see
Gilley and Tarpy (2005). See also Winston (1980) for detailed information on the occurrence of
swarming and afterswarming in unmanaged honey bee colonies living in Lawrence, Kansas.
Page 164: The probabilities of a colony producing a first afterswarm (0.70) and a second afterswarm (0.60) were calculated based on what Winston (1980) and Gilley and Tarpy (2005) have reported.
Each study describes the pattern of swarming and afterswarming for five colonies.
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Notes to Chapter 7 305
Page 164: The work by M. Delia Allen mentioned here is found in Allen (1956).
Page 164: The probability of survival to the following summer of the mother queen that departs in a prime swarm ( p = 0.23) and of the daughter queen that inherits the original nest ( p = 0.81) comes from Seeley (2017b, see fig. 5).
Page 165: The methods and results of the two long- term studies of wild- colony demography that are mentioned here are described in detail in Seeley (1978) and Seeley (2017b).
Page 165: I describe how I use bait hives to capture swarms in Seeley (2012, 2017a).
Page 166: The bimodal seasonal pattern of swarming in the Ithaca area is described in Fell et al. (1977).
Page 167: Afterswarms are delayed, relative to prime swarms, in building their nests because they leave the parental nest well after the prime swarm has departed. Detailed information about the delayed departures of afterswarms is found in table I in Gilley and Tarpy (2005). They report first
afterswarms departing 5–7 days after the prime swarm and second afterswarms departing 12–18
days after the prime swarm.
Page 169: The formula for calculating how long, on average, a site will be continuously occupied by a honey bee colony is as follows, where A represents site age in years:
20
0.5 + ∑ A[(0.23)(0.81)A–1][0.19] = age in years
A=0
Page 171: For more information about the evolution of parental allocation of resources to male and female offspring, see Charnov (1982).
Page 172: The values used here for the average dry weights of a drone and a worker come from the study by Henderson (1992).
Page 174: How large a food reserve is carried off by the worker bees in a swarm when they leave home?
Combs (1972) reports that, on average, a worker bee in a swarm holds 37 mg (0.001 oz.) of a 67%
sugar solution in her crop (honey stomach). (For comparison, on average, a worker bee in a non-
swarming colony holds only 10 mg/0.0003 oz. of a 39% sugar solution.) Therefore, an average
swarm with 12,000 workers will be stocked with some 300 g (10.6 oz.) of sugar. (12,000 bees ×
37 mg sugar solution per bee × 67% sugar = 297.5 grams of sugar.)
Page 174: I have studied the cost to a colony of producing and maintaining a population of drones (Seeley 2002). To do so, I compared the honey yields of colonies that were managed for honey
production, and were either with or without drone comb, and found that colonies with full- scale drone production produced, on average, some 20 kg (44 lb.) less honey than colonies with severely limited drone production.
Page 174: How many mating flights does a drone make across his life? A drone lives for about 20 days after reaching sexual maturity, and he makes two to four mating flights on a day of good weather (Winston 1987, pp. 56 and 202). If we assume that half the days in summer have weather good
enough for drones and queens to conduct mating flights, then we can estimate that a typical drone makes 20–40 mating flights in his brief (and probably sexually unfulfilled) life.
Page 175: The two- part investigation that is mentioned here is Rangel, Reeve et al. (2013).
Page 176: For more information on how we measured the winter survival probabilities for mother-
queen and sister- queen colonies as a function of swarm fraction, see Rangel and Seeley (2012).
Page 176: The three studies that report measurements of the swarm fraction are Martin (1963), Getz et al. (1982), and Rangel and Seeley (2012).
Pages 178–182: The fascinating biology of mating by honey bees is now thoroughly reviewed in two books: Koeniger, Koeniger, and Tiesler (2014), in German, and Koeniger, Koeniger, Ellis et al.
(2014), in English.
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306 Notes to Chapter 8
Page 178: The discovery that 9- oxo- 2- decenoic acid is the sex attractant pheromone of honey bees is reported in Gary (1962).
Page 180: The figures for the average distance to a queen’s mating site (2–3 km/1.2–1.9 mi.) and for the maximum distances to which drones fly in search of a mating opportunity (5–7 km/3.0–
4.2 mi.) come from Ruttner and Ruttner (1972).
Pages 180: The original report of the impressive mark- and- recapture study of the distance of drone mating flights is Ruttner and Ruttner (1966).
Page 181: The elegant experimental study of the maximum mating range of honey bees, conducted in Ontario, Canada, is reported in Peer (1957).
Page 182: The evolution of polyandry (multiple mating by queens) in the genus Apis is reviewed by Palmer and Oldroyd (2000). For a report of the average paternity numbers of all eight species of Apis, see Tarpy et al. (2004). For Apis mellifera, they report 12.0 ± 6.3 (mean ± one standard deviation) for the observed insemination number, and 12.1 ± 8.6 or 11.6 ± 7.9 for the effective
paternity frequency. There are two estimates of the effective paternity frequency because there are two ways to estimate this number.
Page 182: The multistage process whereby drones inject their sperm into the queen’s oviducts and then a small portion of it is transferred to the queen’s spermatheca for long- term storage is described well in chapter 10 in Koeniger, Koeniger, Ellis et al. (2014).
Page 182: There are many studies that show how a honey bee colony benefits from the queen acquiring sperm from numerous drones and then producing a genetically diverse workforce. For information
about improved disease resistance, see Tarpy (2003), Seeley and Tarpy (2007), and Simone- Finstrom, Walz et al. (2016); about greater nest microclimate stability, see Jones et al. (2004); and about increased productivity through enhanced acquisition of food resources, see Mattila and Seeley
(2007) and Mattila et al. (2008).
Page 183: Heather Mattila’s discovery that queen promiscuity helps ensure that a colony possesses a critical minority of workers who are social facilitators of foraging- related activities is reported in Mattila and Seeley (2010).
Pages 183–186: For more information on the study that compared the mating frequencies of queens
living in the wild, where colonies are widely dispersed, and queens living in apiaries, where colonies are tightly bunched, see Tarpy, Delaney et al. (2015).
CHAPTER 8. FOOD COLLECTION
Page 187: The Thomas Smibert quotation is from his poem “The Wild Earth- Bee”; see Smibert (1851).
Page 187: The evidence that worker bees will travel 14 km (8.7 mi.) to collect food comes from a study conducted in a semidesert region of Wyoming. Colonies were placed at various distances (up to 14
km/8.7 mi.) from irrigated fields of alfalfa and yellow sweet clover, and it was found that even the most distant colonies collected nectar and pollen from these fields. See Eckert (1933).
Page 187: The statement that a colony will field about a third of its members as foragers is based on the results reported in Thom et al. (2000).
Page 189: That pollen foragers preferentially unload pollen near cells containing brood (esp. cells containing eggs and larvae) was shown by Dreller and Tarpy (2000). For broader analyses of the
behavioral rules of honey bees that create the consistent pattern of brood, pollen, and honey in the combs of their nests—brood at the bottom, surrounded by pollen, and honey above—see Camazine
(1991), Johnson (2009), and Montovan et al. (2013).
Page 190: Even during a strong honey flow—a time of intense nectar collection—water collectors
constitute a small percentage of the bees coming into a hive bearing a load of liquid cargo. See, for example, fig. 2 in Seeley (1986).
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Page 190: For more information about the age distributions of the nectar receivers and water receivers in a colony, see Seeley (1989) and Kühnholz and Seeley (1997).
Page 192: The details of the study of these undisturbed colonies living in Connecticut are described in Seeley and Visscher (1985).
Page 192: There are approximately 7,700 bees per kg (3,500 bees/lb.); see Otis (1982).
Page 193: The figure of 130 mg (0.004 oz.) of pollen to produce a worker bee comes from Haydak
(1935).
Page 193: The statement that, on average, a worker bee lives about one month in summer is based on Sekiguchi and Sakagami (1966) and Sakagami and Fukuda (1968).
Page 193: The estimates of total brood rearing and food consumption by colonies managed for honey production come from Brünnich (1923) and Nolan (1925), regarding brood production; Eckert
(1942), Hirschfelder (1951), and Louveaux (1958), regarding pollen consumption; and Weipple
(1928) and Rosov (1944), regarding honey consumption.
Page 193–194: The average weight of a pollen load comes from Parker (1926) and Fukuda et al. (1969).
The average flight distance (= twice the average flower- patch distance) comes from Visscher and Seeley (1982). The estimate of flight cost comes from Scholze et al. (1964) and Heinrich (1980).
The energy value for pollen comes from Southwick and Pimentel (1981). The average sugar
concentration of nectar comes from Park (1949), Southwick et al. (1981), and Seeley (1986). The
average sugar concentration of honey comes from White (1975). The average weight of a nectar
load comes from Park (1949) and Wells and Giacchino (1968).
Page 195: The figure of 6 km (3.7 mi.) for the foraging range of a colony is based on the work reported in Visscher and Seeley (1982), which described the spatial patterns of foraging by a full- size colony living in the Arnot Forest. It found that a circle large enough to enclose 95% of the sites indicated by the waggle dances of this colony’s foragers had a radius of 6 kilometers.
Page 195: The indicated flight speed of a forager—30 km/hr (18.6 mph)—is the approximate average of the cruising flight speed of nectar foragers for their empty, outbound flights (34.2 km/hr, 21.3
mph) and their laden, homebound flights (24.2 km/hr, 15.0 mph). For details on how these flight
speeds were measured, see Seeley (1986) or Biology Box 5 in Seeley (2016).
Pages 195–196: For examples of the foraging- range studies using the standard mark- and- recapture method, see Berlepsch (1860, p. 176), Levin et al. (1960), Levin (1961), and Robinson (1966). The study conducted in the semidesert region of Wyoming is Eckert (1933). The method of magnetic
retrieval of steel labels in a capture- recapture system for honey bees is described in Gary (1971).
A good example of a study in which the method of magnetic retrieval of steel labels was used to
determine the distribution of a colony’s foragers is Gary et al. (1978).
Page 197: The pioneering work by Herta Knaffl on the spatial range of a honey bee colony’s foraging operation based on reading the bees’ waggle dances is described in detail in Knaffl (1953).
Pages 197–200: For a detailed description of the study conducted with Kirk Visscher of the spatial and temporal patterns in the foraging work of a full- size colony living in the Arnot Forest, see Visscher and Seeley (1982).
Page 201: The remarkable study of long- range foraging by bees flying to the heather in the high moors outside of Sheffield, England, is that of Beekman and Ratnieks (2000). For more remarkable studies conducted in England that have used the technique of spying on waggle dances to explore the effects of colony size and season on foraging range and dynamics, see Beekman et al. (2004) and Couvillon et al. (2014, 2015).
Pages 201–204: The treasure hunt study of the abilities of colonies to conduct reconnaissance for profitable food sources far from their nests is described in Seeley (1987).
Page 207: For more examples of maps that depict the day- by- day dynamics of the recruitment targets of the colony living in the Arnot Forest, see fig. 3 in Visscher and Seeley (1982).
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308 Notes to Chapter 9
Pages 206–209: The experimental study of the ability of a colony to choose among a changing array of potential food sources by skillfully adjusting the number of foragers engaged at these sources is described in detail in Seeley, Camazine et al. (1991). For a review of all the studies on this topic, see chapters 3 and 5 in Seeley (1995).
Page 210: For an example of the distribut
ion of sugar concentrations in the loads of liquid collected by worker bees in Connecticut, see fig. 2.12 in Seeley (1995). The range of sugar concentrations shown in this figure extends from about 15% to 65% and averages around 40%. Park (1949) and Southwick
et al. (1981) report the same average concentration of sugar in nectar collected by worker bees in Iowa and New York, respectively.
Page 210: For examples of studies that have looked at what stimulates robbing among colonies in
apiaries, see Butler and Free (1952) and Ribbands (1954).
Pages 211–212: The study of the speed and occurrence of robbing among widely separated wild
colonies is reported in Peck and Seeley (forthcoming).
Page 212: The bait hive on the shed attached to my barn is a small, five- frame Langstroth hive. To make my bait hives conspicuous and attractive to scout bees, I insert five frames filled with dark, aromatic comb. See Seeley (2012) and Seeley (2017a).
Page 213: The paper that describes the spectacular skill of Varroa mites at climbing onto worker bees is Peck et al. (2016). The paper that shows that when Varroa mites are in colonies that are weakened by mite- transmitted diseases, they no longer discriminate against climbing onto foragers (including robbers) is Cervo et al. (2014).
Page 214: I believe that all the colonies in the Arnot Forest are now infested with Varroa mites because every swarm that I capture in this forest is infested with mites, as discussed in chapter 2.
CHAPTER 9. TEMPERATURE CONTROL
Page 215: The Thomas Hood quotation is from his poem “November”; see Hood (1873), p. 332.
Page 215: There are several good sources of information about the temperatures in broodless winter clusters. See Hess (1926), Owens (1971), Fahrenholz et al. (1989), and Stabentheiner et al. (2003).
For detailed information about the brood- nest temperatures of honey bee colonies, see Himmer
(1927), Owens (1971), Levin and Collison (1990), and Kraus et al. (1998).
Pages 216: Two studies that have shown that the proper behavioral performance of worker bees depends on their being kept at 34.5°–35.5°C (94°–96°F) throughout their pupal development are Tautz et