The Lives of Bees
Page 41
CHAPTER 11. DARWINIAN BEEKEEPING
Page 277: The Leslie Bailey quotation is from his book Honey Bee Pathology; see Bailey (1981), p. 7.
Page 277: The concept of Darwinian beekeeping is an application of the ideas of Darwinian medicine, as discussed by Williams and Nesse (1991) and Nesse and Williams (1994), to the subject of beekeeping. The fundamental insight of both Darwinian medicine and Darwinian beekeeping is that
living systems experience differences between their modern environment (current environment)
and the environment that they evolved to live in (environment of evolutionary adaptation) and that these differences cause many problems because living systems are often poorly equipped to deal
with the novelties of their modern environments.
Page 278: The experiments that have shown that the unusual annual brood cycle of colonies in the Landes region of southwestern Frances is an adaptive, genetically based trait are reviewed in
Louveaux (1973) and Strange et al. (2007). Hatjina et al. (2014) describe a large- scale study that investigated locally adaptive differences in the timing of colony development. They describe an
experimental analysis that used five European subspecies of Apis mellifera: carnica, ligustica, macedonica, mellifera, and siciliana.
Page 280: A study that has looked specifically at the effects of crowding colonies in apiaries on the problems of colony reproduction and disease transmission is Seeley and Smith (2015). Brosi et al.
(2017) present a model of infectious disease epidemiology that shows the key role of hive/nest
density on the spread of infectious diseases.
Page 280: A study that has looked explicitly at the effects of hive size on both the production of honey and the problems of brood diseases is Loftus et al. (2016).
Page 280: An experimental investigation of the effects on the immune systems of worker bees of having a propolis coating on the walls of their colony’s nest cavity (or hive) is Borba et al. (2015).
Page 281: The best source of information about the differences in insulation value between walls of tree cavities and standard wooden hives and about the effects of these differences on the energetics of colony thermoregulation is Mitchell (2016).
Page 281: To the best of my knowledge, there are no published studies of the effect of nest entrance height on the riskiness of taking cleansing flights in winter when there is snow on the ground. I have, however, performed a pilot study in which I placed two colonies on the gently sloping roof of the storage shed at my laboratory and placed two more colonies nearby on hive stands at ground level. When ca. 20 cm (8 in.) of snow covered the ground, the bees exiting the higher hives started out approximately 200 cm (ca. 6.5 ft.) above the snow, whereas those exiting the lower hives started out only a few cm (1–2 in.) above the snow. On three sunny days in winter, when the air warmed
enough for the bees to make cleansing flights, I counted the number of bees that had flown from
and then crashed in the snow outside each of these four hives. The average for each of the two high hives was 8 bees per warm day, while for the low hives it was 113 bees.
Page 281: The evidence that inhibiting drone production boosts a colony’s honey production is reported and reviewed in Seeley (2002) and that it slows reproduction by Varroa destructor is described in Martin (1998).
Page 281: Several investigators have explored the behavioral rules of worker bees that produce the Seeley.indb 313
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314 Notes to Chapter 11
cell- allocation pattern in honey bee nests. The pioneering studies are reported in Camazine (1991) and Camazine et al. (1990), which show that bees can produce these patterns by following simple
rules that do not require the bees to have global knowledge (a “blueprint”) of their nest’s final layout.
Johnson (2009) added a gravity- based rule that biases the movement of the nectar storers toward the top of the nest, to produce the pattern of nectar storage primarily in the top of the nest. The most recent work, by Montovan et al. (2013), adds two more behavioral rules: 1) the consumption
of nectar and pollen by the workers is brood- density dependent (strongest near the brood) and 2) the movements by the queen are biased toward the center of the comb by responding to temperature gradients. The richness of mechanisms for building and maintaining the cell allocation pattern in honey bee nests is a strong indicator of the adaptive value of this pattern to the bees.
Pages 282: The evidence that moving a colony overnight to a new location can reduce a colony’s weight gain for the following week is reported in Moeller (1975).
Page 282: The study that measured the effects of colony disturbance on colony weight gain (honey production) for the day is Taber (1963).
Page 283: The statement that Varroa destructor (and the viruses it spreads) has killed millions of honey bee colonies comes from Martin et al. (2012).
Page 283: The study of the effects of pollen diversity in the diets of nurse bees is Di Pasquale et al.
(2013).
Pages 283: A study that compares the effectiveness of various pollen substitutes to that of real pollen is Oliver (2014). See also Randy Oliver, A comparative test of the pollen subs, ScientificBeekeeping.
com (n.d.), http://scientificbeekeeping.com/a-comparative-test-of-the-pollen-sub/. Oliver found that “natural pollen still reigns supreme.” The study that has demonstrated that workers reared in pollen- stressed colonies become poor foragers is Scofield and Mattila (2015).
Page 283: One study that has documented the high levels of agrochemicals in honey bee colonies in North America is Mullin et al. (2010). Traynor et al. (2016) have reported that colonies incurred higher risks of brood being poisoned by insecticides and fungicides during development when the
colonies were performing pollination services (for apples, blueberries, cranberries, citrus fruits, and cucumbers) than when they were producing honey or were sitting in holding yards. Other
studies have shown that commercial pollination environments almost invariably expose managed
honey bee colonies to higher levels of pesticide residues because drift of pesticides onto
noncultivated plants nearby creates a summer- long route of pesticide exposure (Botías et al. 2015).
Page 283: For a review of the known populations of wild honey bee colonies that are surviving without treatments for Varroa mites, see Locke (2016). For a paper that examines how treating honey bee colonies with miticides and antibiotics can alter their microbiomes, see Engel et al. (2016).
Page 284: A study by Loftus et al. (2016) compared colonies living in large hives to ones living in small hives in terms of their vulnerability to population explosions of Varroa mites and agents of disease that reproduce in cells containing brood, such as chalkbrood (causative agent the fungus Ascosphaera apis) and American foulbrood (causative agent the bacterium Paenibacillus larvae).
Page 284: The figure of 5 unit weights of honey consumed to produce 1 unit weight of beeswax comes from data of Weiss (1965) as analyzed by Hepburn (1986).
Page 285: The study that found that workers can favor larvae of certain patrilines when they rear queens is that of Moritz, Lattorff et al. (2005). Sometimes these are even subfamilies that are poorly
represented among the workers.
Page 286: The construction and use of bait hives is described in papers by Seeley (2012) and Seeley (2017a) and in the book by Magnini (2015).
Page 287: The evidence of the effectiveness of spacing colonies for reducing the spread of parasites and pathogens between colonies is presented in Seeley and Smith (2015) and the references cited in it.
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Notes to Chapter 11 315
Page 287: The effectiveness of reducing hive size in lowering a colony’s disease load is described in Loftus et al. (2016).
Page 287: The evidence that having a thick coating of propolis on the inte
rior wall surfaces of hives is reviewed in Simone- Finstrom and Spivak (2010).
Page 287: The influence of thick insulation of a hive’s ceiling and walls on the cost of colony
thermoregulation is discussed in Mitchell (2016, 2017).
Page 289: The behaviors of the bees that create the “mite bomb” phenomenon have recently been
analyzed (Peck and Seeley, forthcoming). The main mechanism by which colonies around a
collapsing colony suddenly acquire higher infestations of Varroa mites is robbing of honey from the dying colony by foragers from the healthy colonies nearby. The mites are quite skilled at climbing onto worker bees when they are standing still, filling themselves with a dying colony’s honey (Peck and Seeley, forthcoming).
Page 291: The preeminent importance of Apis mellifera as a crop pollinator is reported in the detailed study by Kleijn et al. (2015) of crop pollination services worldwide. In this study, the crop
production values of different species of bees were calculated based on data from 90 studies
conducted on 1,394 crop fields distributed across five continents. In each study, the investigators measured the abundance and density of the bees visiting the flowers of a crop that depends on bee pollination for maximum yield. Twenty different crops were examined. On average, honey bees
contributed $2,913 per ha ($1,179 per ac.) to the production of the crops, while the community
of “wild bees” (= non- Apis bees) contributed $3,251 per ha ($1,316 per ac.). This indicates that honey bees contributed nearly as much value to the production of these crops as all the other kinds of bees combined.
Page 292: For a discussion of the various names that have been used to describe a type of beekeeping that differs from conventional forms of honey bee management, see Phipps (2016). He points out
that despite their different names, they all refer to the methods used by beekeepers who want to let their bees live in homes made of natural materials, build their combs freely, swarm as they see fit, and handle diseases on their own.
Page 292: A book by Heaf (2010) provided the first detailed discussion of the health consequences of conventional beekeeping and of the range of attitudes of beekeepers to their bees. Neumann and
Blacquière (2016) and Seeley (2017c) were the first to review systematically the various ways in which the practices of conventional beekeeping—such as treatments against disease, artificial
selection against propolis usage, and crowding colonies in apiaries—interfere with natural selection for healthy honey bee colonies and to suggest that lasting solutions to the problems of beekeeping are most likely to come by making full use of the power of natural selection. Blacquière and Panziera (2018) make an explicit plea to let natural selection, rather than artificial selection, be the main way forward to acquire bees that have natural resistance to the mite Varroa destructor and other environmental threats.
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