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1 Chapter 1Figure 1.1 Gathering honey, a beekeeping scene from the Tomb of Rekhmire. Eg...Figure 1.2 Grooming, or mite‐chewing, is a heritable trait in which honey be...Figure 1.3 Hygienic behavior or Varroa Sensitive Hygiene (VSH), is a form of...Figure 1.4 An illustration comparing the structure and organization of a hon...Figure 1.5 A research station beside the Shindagin Hollow State Forest in up...Figure 1.6 A month‐long comparison of temperatures (°C) inside a thin‐walled...Figure 1.7 Polyandry, or the multiple matings of a queen with drones from di...Figure 1.8 Bait hives are small nest boxes that are filled with empty comb a...

2 Chapter 2Figure 2.1 Charles Darwin marveled at the superorganism. Recognizing the rem...Figure 2.2 Trophyllaxis, or the transfer of food from bee to bee, augments d...Figure 2.3 Allogrooming, or the grooming of one bee by another nestmate, is ...Figure 2.4 Tree saps or resins are collected from leaf buds and packed onto ...

3 Chapter 3Figure 3.1 A photograph showing the three members of the honey bee hive – dr...Figure 3.2 External anatomy of the honey bee.Figure 3.3 Magnified view of a honey bee compound eye. The hexagonal shape o...Figure 3.4 The three ocelli are indicated by the small arrows.Figure 3.5 Closeup of a honey bee antenna, showing the segments.Figure 3.6 Wing hooklets, greatly magnified.Figure 3.7 (a, b) Magnified view of the corbicula (pollen basket) on the lat...Figure 3.8 Honey bee foraging on prairie spiderwort Tradescantia occidentali...Figure 3.9 Antenna cleaner.Figure 3.10 Stinger of a worker honey bee. The queen's is barbless.Figure 3.11 Circulatory and respiratory system.Figure 3.12 Ventral nerve cord of the honey bee, shown in purple.Figure 3.13 Digestive system.Figure 3.14 Overview of the glands of the honey bee. The presence or absence...Figure 3.15 Wax scales emerging from the wax glands.Figure 3.16 Ovary of a laying queen. Individual ovarioles can be observed wi...

4 Chapter 4Figure 4.1 Comparative anatomy of the vertebrate equine (a) with the inverte...Figure 4.2 Physical communication in the honey bee using the Waggle Dance. T...Figure 4.3 Life inside a honey bee hive is complex with many work activities...Figure 4.4 A colony of honey bees survives in large part because of the pres...

5 Chapter 5Figure 5.1 The queen functions not only as the ovary of the honey bee supero...Figure 5.2 Whenever a “good” queen pauses on the comb, an ad hoc group of ad...Figure 5.3 A well‐fed queen is an egg‐laying machine, capable of producing a...Figure 5.4 To the untrained eye, virgin queens are difficult to spot. Look f...Figure 5.5 The leading edge of a “drone comet” chasing a “virgin” queen. At ...Figure 5.6 A freshly‐returned no‐longer‐a‐virgin exhibiting “mating sign” – ...Figure 5.7 Typically 10–14 days after emergence (dependent upon weather), a ...Figure 5.8 A histogram of colony performance (weight gain) of 35 colonies, s...Figure 5.9 Preconstructed natural “queen cups.” It is not unusual to see the...Figure 5.10 A typical supersedure cell. Supersedure cells are generally foun...Figure 5.11 A dissected supersedure cell after emergence, showing that the e...Figure 5.12 Two laying queens on the same comb face (the lower one is a bit ...Figure 5.13 Shaking bees through a sieve box to recover any queens. This is ...Figure 5.14 The thorax of a queen (center) or drone (lower right) is too bro...Figure 5.15 This aged queen (note the well‐polished thorax) became a “drone ...Figure 5.16 Typical queen cells, some sealed, some in development, along the...Figure 5.17 Typical swarm cells along the bottom bar of the upper brood cham...Figure 5.18 Bottom view of swarm cells prior to sealing. Note the huge amoun...Figure 5.19 An emerged queen cell, indicating that the colony has recently s...Figure 5.20 A sharp‐eyed vet can really impress a beekeeper when they walk u...Figure 5.21 An aged or damaged queen may not be able to fly far, and may lan...Figure 5.22 A beekeeper about to shake a low‐hanging swarm into an empty box...Figure 5.23 The author has too many times observed a novice beekeeper inadve...Figure 5.24 Typical emergency queen cells at about a day after initiation, s...Figure 5.25 Typical eggs from laying workers. Unlike those laid by a queen (...Figure 5.26 A nice egg‐laying pattern by a queen – each egg centered in its ...Figure 5.27 Raised drone cells scattered among worker cells, as opposed to g...Figure 5.28 Drone cells are normally built in discrete patches, rather than ...Figure 5.29 A brood frame with scattered drone cells. In this case, there we...Figure 5.30 A case of a young queen commencing egg laying without having mat...Figure 5.31 A solid brood pattern, indicating an excellent queen, and a well...Figure 5.32 An example of “spotty brood.” Although beekeepers often blame th...Figure 5.33 The queen is most often to be found on a brood frame, and not su...Figure 5.34 Queens are typically held by both wings. If you only get one win...Figure 5.35 Even when gently handled, some queens may play possum. Do not le...Figure 5.36 Placing a queen headfirst into a queen cage for her own protecti...Figure 5.37 Bee behavior indicating acceptance of a caged queen. The bees wi...Figure 5.38 Bees that are not accepting a queen. These workers are attemptin...Figure 5.39 The round opening at the bottom of the inserted queen cell indic...

6 Chapter 6Figure 6.1 Lineages of bees: Apis mellifera scutellata Lepeletier, the Afric...Figure 6.2 Examples of the Carniolan Bee.Figure 6.3 An example of a Caucasian Queen.Figure 6.4 An Italian bee.Figure 6.5 An Africanized honey bee apiary in Bolivia. Note the stone walled...Figure 6.6 A frame of brood from an Africanized hive. Note the extensive bro...Figure 6.7 Working defensive bees in Bolivia.

7 Chapter 7Figure 7.1 Cladogram depicting the phylogenetic relationships between “sphec...Figure 7.2 Structures for pollen transport in different bee species. (a) The...Figure 7.3 Diversity of natural history traits among solitary non‐Apis bee s...

8 Chapter 8Figure 8.1 Relationship between pollen availability and colony population dy...Figure 8.2 Age structure of a colony over a year in Manitoba, Canada. Number...Figure 8.3 Growth rates of both package and overwintered colonies, generally...Figure 8.4 Comparison between the weight gain of package bees vs. that of a ...Figure 8.5 The availability of trace elements in the soil varies greatly acr...Figure 8.6 Pollen foragers pack their loads into empty cells near young larv...Figure 8.7 The degree of distension of the bees' abdomens is a quick field s...Figure 8.8 This colony is enjoying a surplus of nectar and pollen, as eviden...Figure 8.9 Wax flakes produced by the eight wax glands on a worker's abdomen...Figure 8.10 The presence of “white wax” indicates to the beekeeper that a st...Figure 8.11 The above is an example of a colony suffering from dysentery, re...Figure 8.12 It requires microscopic examination of a bee's diluted gut conte...Figure 8.13 Honey bees are not native to California, and thus did not evolve...Figure 8.14 Bees collecting sawdust on a sunny morning in January, when ther...Figure 8.15 Almond pollen and nectar both contain bee‐toxic amygdalin, yet n...Figure 8.16 Typical layers of pollen from different returning foragers. The ...Figure 8.17 Beebread exhibiting a range of colors generally guarantees a col...Figure 8.18 Plants of the sunflower family, such as Dandelion, although very...Figure 8.19 The owner of this prune orchard seeded mustard, knowing that bee...Figure 8.20 Although is colony is packed with what looks like pollen, the or...Figure 8.21 The above beebread at 400x magnification. The roundish grains ar...Figure 8.22 Fat bodies in the abdomen of a forager (left) and nurse or winte...Figure 8.23 Nurse bees sharing jelly via trophallaxis. Only colonies with we...Figure 8.24 A queen cell opened to show a second‐instar larva in a queen cel...Figure 8.25 The abdomen of the returning forager at the lower left is disten...Figure 8.26 This colony is said to be “shaking nectar,” meaning it is bringi...Figure 8.27 A typical central brood comb from the center of a brood chamber ...Figure 8.28 The interface above tells you most all you need to know about th...Figure 8.29 Pollen and nectar are initially placed throughout the brood area...Figure 8.30 There are two dynamic interfaces with which one can assess the s...Figure 8.31 Note the unripened shiny nectar surrounding the brood, the abund...Figure 8.32 A colony living hand‐to‐mouth, as indicated by the small amount ...Figure 8.33 A frame of healthy brood with a broad ring of pollen stored on t...Figure 8.34 Decent larval survivorship is indicated by an even‐age grouping ...Figure 8.35 The above would generally be considered to be a “solid brood pat...Figure 8.36 Colonies often build drone cells in the space between the two br...Figure 8.37 This rapidly growing colony was hit by a snowstorm during its sp...Figure 8.38 In the final stages of starvation due to lack of honey, the work...Figure 8.39 Typical “spotty brood” during a pollen dearth after the main hon...Figure 8.40 First‐ and second‐instar larvae “swimmin' in jelly.” Nurse bees ...Figure 8.41 Adequately‐fed larvae. Notice the shimmering reflection of jelly...Figure 8.42 The larvae above are being fed far less than optimally. Note the...Figure 8.43 Another example of adequately‐fed larvae. Although the cells are...Figure 8.44 Nutritional stress. The upper cells contain eggs, the lower two ...Figure 8.45 Emergency feeding of colonies during the spring buildup can make...Figure 8.46 The author's sons feeding chunks of pollen sub in an almond orch...Figure 8.47 Testing a hive for weight in order to assess the adequacy of win...Figure 8.48 Colonies being fed sugar syrup during a nectar dearth.Figure 8.49 Close‐up of supplemental sugar feeding jars on colonies. Note th...Figure 8.50 Commercial pollen substitutes (“pollen subs”) in patty form. Hig...Figure 8.51 Comparison of colony buildup during an extended natural pollen d...Figure 8.52 A strong, hungry colony can easily consume a 3‐lb pollen sub pat...Figure 8.53 Commercial pollen sub fed in paper‐sandwiched patty form. Note t...Figure 8.54 As opposed to feeding paper‐covered preformed patties, many comm...Figure 8.55 Feeding chunks of pollen sub in the rain. The author's sons are ...Figure 8.56 Pollen sub fed outside the hive in dry powder form is eagerly ga...Figure 8.57 The “core” gut bacteria exhibit rather strict niche fidelity wit...

9 Chapter 9Figure 9.1 Structure and organization of the honey bee worker gut microbiome...Figure 9.2 The dominant bacteria found in the gut of workers, drones, and qu...Figure 9.3 Factors that can alter the composition of the honey bee gut micro...Figure 9.4 (a) Average bacterial abundance in the gut of honey bees that wer...

10 Chapter 10Figure 10.1 Honey Bee Methods of Detoxifying Drugs, Insecticides, and Other ...Figure 10.2 Routes of Exposure by Honey Bees to Drugs, Insecticides, and Oth...Figure 10.3 Honey bees foraging for water on a leaf.

11 Chapter 11Figure 11.1 A modern Langstroth bee hive equipped with medium hive bodies an...Figure 11.2 Bee space incorporated into a hive design allows for bees to mov...Figure 11.3 The bottom board landing provides a space for bees to alight and...Figure 11.4 Solid (left) and screened (right) bottom boards commonly used by...Figure 11.5 Four‐way pallets used by commercial beekeepers incorporate the b...Figure 11.6 Entrance reducer installed on a Langstroth hive.Figure 11.7 Hive bodies in 8 and 10 frame designs are widely used by beekeep...Figure 11.8 Bee hives using two deep hive bodies for the brood chamber (bott...Figure 11.9 Some apiaries may incorporate several approaches to hive body us...Figure 11.10 Removable frames allow for easy management of the hive by divid...Figure 11.11 This hive had a top deep hive body (removed here) that had medi...Figure 11.12 A frame with plastic foundation installed (left) and the format...Figure 11.13 Drone frame broodFigure 11.14 Queen excluder.Figure 11.15 The inner cover allows for proper bee space and ventilation at ...Figure 11.16 Top covers protect hives from the elements and potential robber...Figure 11.17 Hive stands keep hives off the ground to reduce moisture accumu...Figure 11.18 The Flow Hive uses a honey super and proprietary plastic frames...Figure 11.19 Veils that incorporate coveralls provide maximum protection to ...Figure 11.20 Gloves provide protection from stings. The most common styles u...Figure 11.21 The J Hook hive tool incorporates a ledge on the prying end of ...Figure 11.22 A smoker is a tool that uses smoke directed at bees through a b...Figure 11.23 A top feeder is a common way to feed bees sugar syrup during th...Figure 11.24 Supplemental feeding of dry sugar, candy boards, and fondant du...Figure 11.25 Pollen traps allow a beekeeper to manage the amount of pollen e...Figure 11.26 An apiary site should be on flat ground with all tripping hazar...Figure 11.27 An apiary site that is accessible by car or off‐road vehicle wi...

12 Chapter 12Figure 12.1 Hobby beekeeper apiary.Figure 12.2 Commercial beekeeper apiary with 32 colonies.Figure 12.3 Blooming monoculture fields of planted sunflowers (a) and canola...Figure 12.4 Commercial beekeeper loading honey bee colonies onto a truck.Figure 12.5 Commercial beekeeper colonies covered with a net to prevent exce...Figure 12.6 Commercial beekeeper holding yard used to store colonies awaitin...Figure 12.7 In locations where bears are known to be present, apiaries may h...Figure 12.8 Entrance to an apiary that requires undoing a knot to open the g...Figure 12.9 If it has recently rained, take caution when entering a location...Figure 12.10 Transporting strapped colonies into a yard with difficult acces...Figure 12.11 Colonies in the floodplain of a nearby river. Access to the loc...Figure 12.12 Colonies placed in almond orchards just before bloom (a), durin...Figure 12.13 Smaller scale beekeepers may use hand‐powered tools to extract ...

13 Chapter 13Figure 13.1 Hives placed on stands can keep the colonies out of potentially ...Figure 13.2 When space allows hives can be placed to maximize distance betwe...Figure 13.3 Sometimes the beekeeper is forced to lay out hives in a linear f...Figure 13.4 Painting hives different colors, large geometric shapes on the h...Figure 13.5 Hives enclosed within electric fencing.

14 Chapter 14Figure 14.1 The bee yard on a nice day.Figure 14.2 Lighting a smoker. A variety of fuels can be used as long as the...Figure 14.3 Full bee suit with gloves.Figure 14.4 (a) Standing in front of the hive entrance (not to do). Returnin...Figure 14.5 Observe activity at the entrance, including the pollen loads bei...Figure 14.6 A few puffs of smoke at the entrance will help calm the defensiv...Figure 14.7 Lift the lid and apply a few puffs of smoke.Figure 14.8 A hive tool.Figure 14.9 The top of the hive placed upside down (inside is up) on an empt...Figure 14.10 Applying a few puffs of smoke over the top of the exposed frame...Figure 14.11 Tipping the box to apply smoke.Figure 14.12 If the box is a honey super, remove and place on the upturned l...Figure 14.13 If there is one, remove the queen excluder.Figure 14.14 (a–c) Remove the outer frame first and set it down on end. This...Figure 14.15 Sequentially remove and examine each frame.Figure 14.16 Examining a frame with the sun at your back to best visualize e...Figure 14.17 Pollen and nectar.

15 Chapter 15Figure 15.1 All three antibiotics approved for honey bees can be fed by mixi...Figure 15.2 Example VFD form designed by the author and available on the int...Figure 15.3 Identification of animals treated with antibiotics to avoid resi...Figure 15.4 Example of a Canadian provincial (Ontario) VCPR form.

16 Chapter 16Figure 16.1 A beekeeper performing a routine hive inspection using an electr...

17 Chapter 17Figure 17.1 Warning sign.

18 Chapter 18Figure 18.1 Bees robbing a hive.Figure 18.2 Monitoring a hive for varroa mites.Figure 18.3 Honey bee foraging alongside a sweat bee on Echinacea flower. Pa...Figure 18.4 Parasite prevalence in bee (a) and on flower (b) genera across t...

19 Chapter 19Figure 19.1 Steady honey bee colony numbers in the United States since 1996....Figure 19.2 Self‐reported causes of bee colony losses, by year (www.beeinfor...Figure 19.3 Colonies marked in a commercial apiary for having CCD symptoms....Figure 19.4 Field collection of honey bee samples for laboratory analysis....

20 Chapter 20Figure 20.1 An adult female Varroa destructor perched on an experimentally r...Figure 20.2 An adult mite perched on a toothpick.Figure 20.3 A mite on the author's finger, extending her chemosensory forele...Figure 20.4 The ventral surface of a mite suspended in oil. The large dorsal...Figure 20.5 Most feeding phoretic mites position themselves between the thir...Figure 20.6 A normal sized worker bee for scale next to two stunted pupae fo...Figure 20.7 This bee developed in a terminally mite‐infested colony exhibiti...Figure 20.8 Vertical and horizontal mechanisms of mite transmission between ...Figure 20.9 An experimentally heavily infested colony in a glass‐walled obse...Figure 20.10 “Parasitic Mite Brood Syndrome” in honey bees is characterized ...Figure 20.11 The author's sugar shake sampling kit arranged on the tailgate ...Figure 20.12 A yellow ball of pollen and three red mites floating in dissolv...

21 Chapter 21Figure 21.1 Symptomatic deformed wing virus infection of worker honey bees. ...Figure 21.2 Symptomatic larvae and adult bees infected by acute bee paralysi...Figure 21.3 Type 2 symptoms of chronic bee paralysis virus. (a) Typical symp...Figure 21.4 Modes of virus transmission.

22 Chapter 22Figure 22.1 Michigan State University researchers study colonies affected wi...Figure 22.2 A brood frame from a colony infected with EFB exhibiting a typic...Figure 22.3 A frame from a colony with severe EFB disease. Note the multiple...Figure 22.4 Perforated cappings in a colony infected with AFB disease. As th...Figure 22.5 a (left) and Figure 22‐5b (right) Comparison of matchstick test ...

23 Chapter 23Figure 23.1 (A) Schematic representation of the microsporidia spore. (B) Lig...Figure 23.2 Life cycle of a Nosema spore. Infection begins when a bee ingest...Figure 23.3 Brownish‐yellow fecal spotting on the exterior of a beehive.Figure 23.4 Honey bee Chalkbrood disease. (A) Chalkbrood mummy.(B) Brood...

24 Chapter 24Figure 24.1 A dissected honey bee showing a blotchy, darkened discolouration...Figure 24.2 Tracheal mites can be visualized within the trachea of a honey b...Figure 24.3 Adult wax moth and cocoons on a frame.Figure 24.4 Wax moth larvae.Figure 24.5 Wax moth cocoons on the top frames of a hive. Note the gouges in...Figure 24.6 A heavy wax moth infestation leads to extensive damage to the fr...Figure 24.7 Small Hive Beetle larvae.Figure 24.8 Adult small hive beetle. Note the club shaped antennae and the s...Figure 24.9 Tropilaelaps mites on a developing bee.Figure 24.10 Vespa mandarinia.Figure 24.11 Ants feeding on a honey bee colony.Figure 24.12 Entrance reducers will decrease the area that the colony needs ...Figure 24.13 Note the increased activity around the hive entrance with bees ...Figure 24.14 A heavy rock is placed on top of the lid to prevent raccoons fr...Figure 24.15 Mice have chewed through the wax frames to make room for a nest...

25 Chapter 25Figure 25.1 Mock pesticide label with bee guidance in the Environmental Haza...Figure 25.2 Topical application of pesticides to individual honey bees to de...Figure 25.3 Honey bee larvae reared in vitro for pesticide testing.Figure 25.4 Dead bees at the entrance of colonies following a moderate bee‐k...Figure 25.5 Dead bee traps used for quantifying dead bees resulting from a b...Figure 25.6 Dead and dying larvae observed in a colony fed the organophospha...

26 Chapter 26Figure 26.1 The contents of a cell roping out at least 2 cm indicates a posi...Figure 26.2 Holst Milk Test.Figure 26.3 Remove the contents of a suspect cell with the spatula provided ...Figure 26.4 Place the test material into the solution container from the kit...Figure 26.5 After shaking sample bottle for 20 seconds, use the provided pip...Figure 26.6 The appearance of only the control line (bottom) indicates the t...Figure 26.7 A simple kit for performing alcohol was to assess varroa mite in...Figure 26.8 Quart jar with isopropyl alcohol and approximately 300 bees read...Figure 26.9 After shaking for one minute, pour bees and alcohol through coar...Figure 26.10 Dislodged Varroa mites are visible against a light‐colored wash...Figure 26.11 After counting mites in the wash tub, the alcohol should be pou...Figure 26.12 The strained alcohol is poured over the previously washed bees ...Figure 26.13 After completing a field alcohol wash, bees can be collected in...Figure 26.14 Bees covered in powdered sugar in the sugar roll jar and the su...Figure 26.15 Bee bread samples should be collected from cells using a small ...Figure 26.16 When bees need to be collected kept alive in transit to the lab...Figure 26.17 A popsicle stick can be used to scrape wax from the foundation ...

27 Chapter 27Figure 27.1 Loss of colonies.Figure 27.2 Four annual bee colony phases.Figure 27.3 Personal protective equipment – bee suit with veil and gloves....Figure 27.4 Dead colony (note remains of cluster top center‐right) showing f...Figure 27.5 An example of a spotty brood pattern.Figure 27.6 Photo of drone cells occupying worker cells.Figure 27.7 Photo of tiny dead spring cluster (smaller than softball in size...Figure 27.8 Bee nest organization in Fall; capped brood central, bee bread s...Figure 27.9 An example of starvation. Dead bees in a tightly packed cluster ...Figure 27.10 American foul brood.Figure 27.11 European foul brood.Figure 27.12 American Foulbrood “ropiness test.”Figure 27.13 Emergency queen cell.Figure 27.14 Large capped supersedure queen cell.Figure 27.15 (a) Bees are robbing the hive on the left. (b) Close‐up of Figu...Figure 27.16 Advanced case of PMBS – spotty capped brood, disorganized brood...

28 Chapter 28Figure 28.1 Curious cows.Figure 28.2 This colony was particularly defensive, perhaps because of racco...Figure 28.3 Some beekeepers, upon spotting the queen, carefully contain her ...Figure 28.4 This single deep frame will yield about as many bees as arrived ...Figure 28.5 Two beekeepers worked two days relocating this colony, construct...Figure 28.6 Bearding in the late afternoon of a very steamy summer day.Figure 28.7 Bees working sugar blocks on an unusually warm Michigan day. The...Figure 28.8 Various wraps and windbreaks for a midwest winter.Figure 28.9 This infrared camera photo reveals differences in cluster size a...

29 Chapter 29Figure 29.1 A variety of grafting tools.Figure 29.2 Grafting larvae into queen cups.Figure 29.3 Queen cells on the frame.Figure 29.4 Queens emerge from the end of the cell. Queen cells with a hole ...Figure 29.5 Photograph of an artificially inseminated queen laying an egg su...