BUZZING COMMUNITY: Colony Life Of The Honeybee CLASSIFICATION
Apis mellifera
Honeybee
BUZZING COMMUNITY : Colony Life Of The Honeybee
Copyright 2010 by Princeton University Press Published by Princeton University Press 41 William Street Princeton, New Jersey 08540 In the United Kingdom:Princeton University Press 6 Oxford Street Woodstock, Oxfordshire OX20 1TW press.princeton.edu All Rights Reserved Library of Congress Cataloging-in-Publication Data ISBN 978-0-691-14721-5 Printed in the United States of America
For the preservation of bees.
TABLE OF CONTENTS
01 CHAPTER PAGE
The Colony
13
02 CHAPTER PAGE
Collecting Food
21
03 CHAPTER PAGE
Waggle Dance
29
04 CHAPTER PAGE
Nesting Architecture
37
INTRODUCTION The honeybee exhibits a combination of individual traits and social cooperation which is unparalleled in the animal kingdom. The multiple levels at which the honey bee expresses adaptations to its world provide one of the richest sources for study and knowledge among all organisms, made even more enriching by the economic benefits the honey bee provides. A glimpse into the nest will show the infrastructure of the honeybee nest. This exquisitely uniform and functional comb, is composed of beeswax produced by the workers and constructed into a repeating series of hexagonal cells. The comb provides the substrate for interactions between colony members and is used for almost everything, from larval nursery to pantry to message center. As the stage for colony activites, it provides us with a rich arena for observing the individual and social behaviors that are at the heart of honey bee society. At the individual level, honeybees have not one but three types of colony members: queens, drones, and workers, each with their own specializations and place in honey bee society. The queen reigns over the nest, surrouneded by attendants and fed the rich food she requires to perform her few but crucial tasks in the colony. Her slim lines hide the huge ovaries which make her an egg-laying machine, capable of laying thousands of eggs a day, and her calm behavior masks her powerful pheromones, chemical signals to recipient workers which control many of their behaviors and provide part of the social glue which holds honeybee life together.
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The other sexual members of this society, the drones are tended and fed by the workers. They perform only one function, which is mating with the queen, after which they die. With their large eyes, flight muscles, and powerful mating urge, the drones are magnificently constructed for this task. The workers perform endless and diverse tasks in the nest, sometimes dying from stinging their colony's opponents, and rarely reproducing. At any time workers may be found walking on the comb surface tending the brood, cleaning debris from the nest, capping cells, rippening or storing honey, organizing pollen for storage near the brood, feeding or grooming the queen, or any one of a myriad other activities. Not only have their individual behavior manifested by the three castes prompted our investigations into honey bee life, but their sociality and adaptability has fueled the study of this insect to an extent not found in other organisms on earth. The social nature of this organism, its ability to regulate it functions as a colony of individuals according to events within and outside the nest, provides the key to its success and makes the contribution of individuals much greater than the sum of their indivudal behaviors. We are futher motivated to study this insect by an economic stake, since the products of the hive are crucial to our agricultural systems. Not only do honeybees provide us with honey, wax, propolis, royal jelly, and pollen, but they also pollinate our crops, including such diverse agricultural plants as fruit trees, oilseeds, small berries, and forage crops.
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01 CHAPTER
THE COLONY CLASSIFICATION
Apis Mellifevra
Honeybee
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
01 THRIVING COMMUNITY The honeybee, Apis mollifier, is one of 20,000 species of bees that exist in the world. Although a widely diverse species, all bees are descended from one ancestral species of vegetarian wasp that lived 100 million years ago in the Early Cretaceous period. Unlike their predecessors who developed a carnivorous behavior because of the lack of vegetation, bees instead collect protein-rich pollen from flowers. This pollenivorous habit explains the fuzzy, hairy look many bees have because of the plumose hairs that cover their bodies, which are used to catch pollen grains from flowers. Bees depend on flowers for adequate nourishment and feed on their nectar for energy, and in turn many flowering plants depend on bees for sexual reproduction. In a honeybee hive there are thousands and thousands of bees living together. They are all female worker bees, all of whom are daughters of the one queen bee that lives in their midst. These worker bees are fully equipped to care for offspring, but have poorly developed ovaries and they rarely lay any eggs. The queen slightly resembles the worker bees, but is has a bigger and longer abdomen with longer legs. Her greater size is impressive sets her apart as she moves slowly, indeed majestically, across the combs The queen lays all the eggs, depositing them into cells at a rate of one or more per minute, or more than 1,500 per day (with a combined weight nearly equal to her own) in late spring and early summer. Over an entire summer she will produce 150,000 eggs and about half a million during the two to three years of her lifespan.
14
The C olony
Most of the eggs laid by the queen will be fertilized, but some will be unfertilized. In the first week of her life, the queen flew from the hive and mated with 10 to 20 males from other hives in the area. This procured a lifetime supply of approximately five million sperm. The queen stores the sperm inside a spherical organ called a spermatheca, which lies in the rear of her abdomen, behind her ovaries. With each egg she lays, the queen decides whether to dispense a few fertilizing sperm or to hold back, which in this process determines the sex of her offspring. The fertilized eggs become female and the unfertilized eggs become male. Whether a fertilized egg develops into a non-breeding worker or an egg-laying queen depends on how the egg is treated. If the egg is deposited into a standard-size cell in the combs, and is fed by workers with standard-quality larval food after it hatches, then it will develop into a non-breeding worker. If the fertilized egg is deposited into a large, specially built queen cell, which hangs from the bottom of the comb, then the larva will be fed with royal jelly. This larva will be reared into a developmental path to become a queen. The food feed to the budding larvas define their place in the hive.
Queen
Lays eggs
Worker
Clean cells, feed larvae, build combs, gather food, ripen honey, ventilate hive
Drone
Mates with Queen
15
Just as a human body f integrated unit even th of cells, the super organ colony operates as a sin though it is a multitude
functions as a single hough it is a multitude nism of a honeybee ngle coherent whole even e of bees.
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
The majority of the hive's population consists of worker bees. Worker bees complete a various amount of tasks to keep the hive functioning as a whole.
LIKE THE QUEEN, WORKER BEES ARE ALL FEMALE.
WORKER BEES:
FEMALE
01
Clean the cell she emerged from and other empty cells to receive new eggs and to store nectar and pollen.
02
Worker bees feed and care for larvae laid by the queen. A nurse bee can check a single larvae 1,300 times a day.
03
Not only do worker bees tend to the larvae, but to the queen as well. They groom and feed her.
04
Nectar is collected from forager bees and the pollen is then packed into cells by the worker bees.
Produce beeswax for the hive. Used to build new wax comb, capping of ripened honey, and cells containing pupae.
05
WORKER BEES PROVE TO BE AN EXTREMELY IMPORTANT ASSET TO THE SURVIVAL OF THE HIVE.
A B C D E F G H I J K
A Forewing B Hindwing C Antenna D Compound Eye E Head F Legs G Thorax H Femur I Tibia J Claw K Abdomen
18
The C olony
UNIFIED, INTEGRATED UNIT A queen withholds sperm from less than five percent of her eggs. These unfertilized eggs are important for they give rise to her sons, the colony’s drones. These are the colony’s brawniest bees, endowed with huge eyes for spying young queens out on nuptial flights and massive flight muscles for chasing after the queen at speeds up to 35 kilometers an hour (about 22 miles per hour). They are also a colony’s laziest bees. Unlike the workers who perform all the household tasks inside their hive—clean cells, feed larvae, build combs, ripen honey, ventilate hive, guard entrance, etc— the drones spend their time at home simply hanging out in restful leisure, from time to time helping themselves to meals from the colony’s honey reserves or begging feedings from their worker sisters. Nevertheless, they make a fundamentally important contribution to their colony’s success, for in mating with the young queens of the neighboring colonies they help their colony win in the ceaseless evolutionary competition to pass genes on to future generations. To understand the distinctive biology of this species of bee, it is helpful to think of a colony, not just as thousands of separate bees, but also as a single living entity that functions as a unified whole. In other words, it can help to think of a honeybee colony as a superorganism. Just as a human body functions as a single integrated unit even though it's a multitude of cells, the super organism of a honeybee colony operates as a single coherent whole even though it's a multitude of bees. The fact that both perspectives— colony as a superorganism and as a society—are valid reflects the way which evolution has repeatedly built higher-level units. For example, during the origins of multicellular organisms, natural selection favored some societies of cells whose members cooperated rather than competed. Bit by bit, this selection for close cooperation produced the thoroughly integrated societies of cells that we know today, including human beings. The same sort of selection for extreme cooperation also happened with some societies that we can call super organisms.
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02 CHAPTER
COLLECTING FOOD NUTRITION COLLECTED FROM: Nec t ar
Pol l e n
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
02 NUTRITIONAL REQUIREMENTS The three honey bee castes have somewhat different nutritional needs and feeding mechanisms to satisfy their food requirements, as do the larval and adults stages withing each caste. But the starting materials for brood and adult bees, whether workers, queens, or drones, are the same: nectar and pollen. These two floral products provide all of the food necessary for larval growth and metamorphosis and for adult development and functions. Nectar provides carbohydrates in the form of sugars, and pollen provides protein, lipids, vitamins, and minerals. Honey bees have evolved various mechanisms for processing nectar and pollen so that the food fed to each stage and case are ideally suited to their needs. Honey bees obtain most of their energy from carbohydrates in the form of plant-produced sugars, primarily from nectar produced in flowers. Floral nectar is an aqueous plant secretion which contains anywhere from 5 to 80% sugars, and small quantities of nitrogenous compounds and minerals, organic acids, vitamins, lipids, pigments, and aromatic substances. The nectar collected by foraging workers can be fed to brood and adults directly, but it is more commonly processed into honey first. The nectar is carried back to the nest in the honey stomach and transferred to nest workers for processing. Enzymes from the hypopharyngeal glands are added to the nectar in the crop. They break down the sugars into simple inverted forms, which are the most easily digested by bees and also protect the stored honey from bacterial attack. The nectar is then evaporated on the worker's tongue and placed into cells for further evaporation by fanning. When the enzymatic activity and water evaporation is complete, the nectar is considered to be "ripened" and can be called honey.
22
Collecti ng Food
Pollen is the male germ plasm of plants, but in addition to being necessary for reproduction, many pollens have evolved so as to be attractive and edible for bees. The most important constituent of pollen is protein. Pollen contain anywhere from 6 to 28% protein and are virtually the only source of protein naturally available to bees. Pollen contains lipids that are important for bee nutrition. Sterols are also present, which are required for bee metabolism since bees cannot synthesize cholesterol without the precursors obtained by pollen. Pollens also contain sugars, starches, vitamins, and minerals, which are important to bee nutrition. Once the pollen has been brought back to the colony by foragers, the workers treat it to prevent germination, begin the digestive process, and prepare it for the long-tern storage. A phytocidal acid is added when polled is packed into the comb, which prevents germination and deleterious bacterial activity.
Larvae
Fed primarily brood food produced by the hypopharyngeal and mandibular glands of nurse bees. Larvae fed on the fourth or fifth day of larval development.
Worker
Nutrients for workers are honey and pollen although some brood food is transmitted during food exchange between workers and used for nutrients. The nectar or honey eaten by workers provides sugars for energy; pollen cannot substitute as an energy source, and workers will die without an adequate supply of honey.
Drone
Drones receive more food than workers because of their larger size. Their brood food has more diverse proteins than does worker brood food. Food of older drone larvae contains more carbohydrates, riboflavin, and folic acid than that of younger larvae, and less thiamin, biotin, pantothenic acid, choline, pyridoxine, protein, fat, ash, and niacin. Adult drones are fed by workers for the first few days and gradually begin to feed themselves.
Queen
Quantity and quality fed to female larvae determines whether larvae will develop into workers or queens. Adult queen are fed by workers for brood food producing age and receive mostly brood, possibly with additional honey. Isolated queens feed themselves on sugar candy and survive for weeks, but queens in colonies seldom feed themselves.
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Workers tend to visit the same location at the same time of the day for many consecutive trips or days, as long as the floral rewards continue.
A worker may return to locations only a few meters in diameter in homogeneous environments such as apple orchards or alfalfa fields. Many flowers secrete nectar or pollen at certain times of the day, and workers visiting these flowers will reappear at the same time on consecutive days expecting to find a "reward".
Collecting Nectar and Pollen Plants Crops Aster Apples Asylum Alfalfa Buttercup Blueberry Clover Broccoli Dandelion Cashew Germaniums Kiwi Milkweed Lemon Poppy Mango Sage Papaya Sunflower Peach Thyme Strawberry Tangerine Watermelon
One of the most important choices for a forager is what to collect. As with most foraging characteristics, this is partly determined by the availability of flowers. Workers tend to "specialize" on one type of foraging task at a time and often exhibit constancy for nectar or pollen. Workers will visit only one species of flower on a trip, and they continue to visit that flower for prolonged periods until it ceases to produce nectar or pollen or a superior source becomes available.
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
26
Collecti ng Food
A worker leaving a foraging trip can face an overwhelming array of flowers to choose from. The quantity and quality of nectar and/or pollen produced by useful flowers can vary tremendously, both between plant species and within patches of the same species. In some regions plants can produce nectar so copiously that colonies collecet 5 kg or more of nectar daily and then produce a harvestable honey surplus of more than 200 kg annually. Numerous plants worldwide are known for their nectar production. Plants that produce over 500 kg/hectare of honey, includes the common maple, milkweed, phacelia, sage, thyme, acacia, and figwort. All of the good honeyproducing plants are characterized by well-developed floral and extrafloral nectaries which concentrate and secrete sugars, and their flowers are designed so that bees are easily attracted to it and the nectar is accessible. Many plants invest energy in nectar production to attract bees, which serve to transfer pollen from one plant to another. Bees use the pollen for food, but in the process of collecting nectar and pollen sufficient pollen grains are transferred between flowers to effect pollination. Pollen is produced in the stamen of the plant and the quality and amount produced varies within and between plant species. Workers choose which pollens to collect not by their nutritive value, age, moisture, content, or color, but on the odor and physical configuration of the pollen grains.
Foraging Bees Nectar vs. Pollen Flowers Visited Per Trip
1 to 1,000
1 to 500
Trips Per Day
10-150
10-15
Time Per Trip
30-80 mins
10 mins
Weight of Load
25-45 mg
10-30 mg
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03 CHAPTER
WAGGLE DANCE COMMUNICATES D i st anc e A n d D i r e ct i on
Food S our ce N e w N e st S i t e
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
03 COMMUNICATION THROUGH MOVEMENT The worker honeybee, informs her hive mates of the direction and distance to a rich food source by means of dance behavior. When a lone forager finds a rich food source of nectar, she returns excitedly to her hive and performs a conspicuous “waggle dance.” In performing this, the dancer walks straight ahead on the vertical surface of a comb, waggling her body for side to side, then she stops the “waggle run” and turns left or right to make a semicircular “return run” back to her starting point, whereupon she produces another waggle run followed by another return run, and so on. Each waggle dance consists of a series of dance circuits, and each dance circuit contains a waggle run and return run. A bee may continue dancing for seconds or even minutes, all the while trailed by unemployed foragers that “take part in each of her maneuverings so that the dancer herself, in her madly wheeling movements, appears to carry behind her a perpetual comet’s tail of bees.” After a dance-follower has tripped along behind the dancer throughout several circuits of her dance, she rushes out of the hive to search
Duration of waggle dance (sec)
for the bonanza announced by the dancing bee.
4.0
3.0
2.0
1.0
1000
3000
5000
Distance (m)
One second of the combined body-waggling/ wing-buzzing represents some 1,000 meters.
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Waggle Dance
Round Dance
Informing a resource in close proximity Less than 15 m away Closely followed by attending bees Repeated small circles, reversing Dances last for seconds up to minutes
In 1945, zoology professor Karl von Frisch discovered that when a bee performs a waggle run inside a dark hive, she produces a reenactment of her recent flight, and in this way indicates the location of the rich food source she has just visited. The duration of the waggle run—made conspicuous despite the darkness by the dancer audibly buzzing her wings while waggling her body—is directly proportional to the length of the outward journey. On average, one second of the combined body-waggling/wing buzzing represents 1,000 meters (six-tenths of a mile) of flight. And the angle of the waggle run, relative to straight up on the vertical comb, represents the angle of the outward journey relative to the direction of the sun. For example, if a successful forager walks directly upward while producing a waggle run, she indicates that “the feeding place is in the same direction as the sun.” Or, if the waggling bee heads 40 degrees to the right of vertical, her message is, “The feeing place is 40 degrees to the right of the sun.” Perhaps most remarkably, the bees that follow a dancer, monitoring her waggle runs, are able to decode her dance and put her flight instructions into action.
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Nest-site scouts announce an inferior nesting place with a "faint-hearted dance" while those from a superior nesting place "solicited with a lively and long-lasting dance."
32
S P N
S [Sun] N [Nest] P
[Potential Site]
Waggle Dance Position / 40째
33
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
SCOUT DANCING Good decision making by a honeybee swarm depends critically on the scouts adjusting dance strength in relation to site quality, so that scouts advocating higher quality properties are better at attracting additional supporters. A honeybee colony deploys its foragers among nectar sources by a group decision-making process that depends on a colony's foragers making graded advertisements of the various nectar sources—the richer the nectar source that a bee exploits, the greater number of dance circuits she produces when she returns to the hive. In short, the richer the nectar source, the stronger the waggle dance. A bee adjusts the number of dance circuits that she produces in relation to nectar-source richness by adjusting two aspects of her dancing: the rate and duration of the dance circuit production. The total number of dance circuits produced in a dancing bee's advertisement is the product of the rate and duration of her dancing. Richer nectar sources elicit longer-lasting and livelier dances than poorer nectar sources. Inferior nesting places are announced with a "faint-hearted dance" while those from a superior nesting place are "solicited with a lively and long-lasting dance."
Waggle Dance
Foraging Communicates distance and direction 50-150 meters Vigorous abdomen movement Emits buzzing sound
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The potential for decision-making errors arising from individual-level noise in reporting on sites is great when each site is discovered. If the scout that discovered a site fails to report on it with a waggle dance, the site won't be entered into the scout bee's debate. It will be lost from the swarm's attention unless another scout happens to find and report the same site, which is highly unlikely. A solution to this problem would be to have each scout bee report on the site they discover and thereby enter it into the debate.
Waggle Dance
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04 CHAPTER
NEST ARCHITECTURE REQUIRES Ch o o si ng A N e w S i t e
C om b C on st r uct i on
B uzzi ng C ommuni ty: C olony Li fe Of The Hon ey b e e
04 CONSTRUCTING A SOCIETY A typical worker bee spends the first 15-20 days of her life entirely inside the colony's nest, and even when she begins to forage, she spends only a few hours a day outdoors. The nest, almost city-like in the diversity of its functions, is constructed inside a long cavity with a small easily defended entrance. Inside, the bees construct wax combs with hexagonal cells in which they rear brood and store nectar, honey, and pollen. Bees patrol the nest on the comb surface, performing or following the dances of foraging bees, passing food to each other, fanning to cool the nest and evaporate water from nectar, and clustering for warmth, among other activities. Honey bees select a new colony site at the last stage of swarming, or colony reproduction. Colonies generally swarm in late spring, when the old colony has an excess of workers and has become overcrowded. At that time a majority of the workers leave the nest with a queen and forms a cluster, usually under an overhanging limb or in a snarl of branches. The swarm then faces a critical problem; it must quickly find a new nest site before the workers run out of honey carried in their honey stomachs or the swarm population will begin to dwindle as workers die. The swarm also has to choose a site in which the new colony can survive and grow for many years.
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Nest Ar chi tectur e
CHOOSING A NEW NEST SITE
A. A swarm must be faced with numerous potential sites to scout and choose from, and the workers must reach a consensus to the preferred site before the swarm leaves the nest.
ONCE A SWARM HAS SETTLED INTO ITS INTERIM CLUSTERING SITE, SCOUTS LEAVE THE SWARM CLUSTER ALMOST IMMEDIATELY AND BEGIN LOOKING FOR NESTS AS MUCH AS 3 DAYS BEFORE THE SWARM LEAVES THE OLD NEST.
B. When a scout bee has found a potential cavity, she spends a considerable amount of time examining it. During the first phase of examination, the scout alternates brief inspections of the nest interior with short examination periods outside the nest cavity.
OUTSIDE BEHAVIORS CONSIST OF EXTENSIVE CRAWLING AROUND THE OUTER SURFACE OF THE POTENTIAL NEST. INTERIOR INSPECTIONS INVOLVE RAPID WALKING ABOUT THE CAVITY'S INNER SURFACES.
C. When a scout bee finds a reasonable site, she returns to the swarm and does dances similar to those used by returning foragers to communicate the location of nectar and pollen sources.
NEST SCOUTS RARELY BRING BACK NECTAR OR POLLEN, AND THEY ARE ONLY COMMUNICATING POTENTIAL NEST LOCATIONS.
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In the first half of the decisio the scouts reported numero located at various directions the swarm cluster. Then, in the debate, the dances of the and smoothly became focus Ultimately, there was virtua the dancing bees, and the sw agreed-upon site.
DAY 1
Foraging bees report potential new nest sites A-M
DAY 2
Sites with the strongest vote are sites G and I
DAY 3
Site I wins with a majority vote and the bees take off
on-making process, ous candidate sites s and distances from the second half of e scout bees quickly sed on just one site. al unanimity among warm moved to the
DAY 1 DAY 2 DAY 3 A
B
C
D
E
F
G
H
I
J
K
L
M
B uzzi ng Democr ac y: The Democr ati c Syst e m O f Hon ey b e e s
COMB CONSTRUCTION
Once the swarm has arrived at its final nest site, comb construction begins immediately. Many of the workers in the swarm have already begun producing beeswax for comb building, which can be seen as wax flakes protruding ventrally from between the abdominal segments. Rapid comb construction is essential to the swarm, since none of a colony's brood rearing or honey and pollen storage can be done without the comb. Over 90% of a feral colony's comb building is completed withing 45 days of nest colonization, indicating the priority the workers give to comb construction. The comb itself is one of the marvels of animal architecture. It consists of a regular back-to-back array of hexagonal cells, arranged in parallel series, each comb precise distance from its neighbor. There are two types of hexagonal cells which make up the comb; the smaller cells are used for rearing worker brood, and drones are reared in the larger ones.
New Nest Criteria Determined By Scout Bees
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Size of entrance
12.5 > 75 cm²
Colony defense and thermoregulation
Direction of entrance
South > North facing
Colony defense and thermoregulation
Height of entrance
5>1m
Colony defense
Position of entrance
Bottom > top of cavity
Colony thernoregulation
Volume of cavity
10 < 40 > 100 Liters
For honey storage and colony thermoregulation
Nest Ar chi tectur e
The hexagonal shape of cells is common among cell-building social insects and there is a sound architectural reason for this style. Round, octagonal, or pentagonal cell arrangements leave empty spaces between cells, and triangles or squares have a greater circumference than hexagons. Thus, the greatest number of cells per area can be arranged in a comb using a hexagonal shape. By staggering each side of the comb slightly from the opposing side, the bees can construct cells back-to-back, further maximizing the number of cells per area. Unlike most social insects, honey bees build their cells horizontally rather than hanging vertically, although they are angled up at about 13째 from base to opening to prevent honey from running out. When construction begins, the workers hang together in tight chains forming a dense cluster in which they maintain a temperature of 35째C, the best temperature for wax secretion and manipulation. Flakes of wax are removed by the enlarged first tarsal joint of the hind leg from four paired glands on the underside of the abdomen and passed forward for construction manipulations by the front legs and mandibles. The wax is mixed with saliva and kneaded to the proper consistency and degree of plasticity at which it can be best molded. The process of removing and manipulating each scale takes about 4 minutes to produce the 77,000 cells which can be constructed with 1 kg of wax. Construction progresses randomly as several cells are under construction simultaneously by several bees contributing to the building of any one cell. Workers begin construction on the roof or side of the nest cavity, with two or three construction sites initially for each comb. Thick layers of wax are placed at the base of each comb, and these are gradually drawn out into cells elongating and thinning the wax into the cell wall. A single worker may add wax or smooth it, moving from cell to cell building site to site in no evident order. Each section of comb is linked with the other so that no traces of their separate beginning are visible.
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Seeley, T.D. "Social Foraging in Honeybees: How Nectar Forag ers Assess Their Colony's Nutritional Status." Behavioral Ecology and Sociobiology 24 (1989): 181-99. Web. Seeley, Thomas D. The Wisdom of the Hive: The Social Physiology of Honey Bee Colonies. Cambridg e, MA: Harvard UP, 1995. Print. Seeley, T.D. "Social Foraging in Honeybees: How Nectar Forag ers Assess Their Colony's Nutritional Status." Behavioral Ecology and Sociobiology 24 (1989): 181-99. Web. Seeley, Thomas D. The Wisdom of the Hive: The Social Physiology of Honey Bee Colonies. Cambridg e, MA: Harvard UP, 1995. Print. Seeley, T.D., and C.A. Tovey. "Why Search Time to Find a Food-storer Bee A ccurately Indicates the Relatie Rates of Nectar Collecting and Nectar Processing in Honey Bee Colonies." Animal Behaviour 47 (1994): 311-16. Web. Waddington, K.D. "Honey Bee Foraging Profitability and round Dance Correlates." Journal of Comparative Physiology 148 (1982): 297-301. Web. Winston, Mark L. The Biology of the Honey Bee. Cambridg e, MA: Harvard UP, 1987. Print.
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Ty p efaces C hron ic le design ed by Jon ath an Hoef ler and Tobias Frere-Jon es f or Hoef ler & Co. Akz idenz G rotes k by Herman Berth old f or Berth old Type Fou nd ry. K n oc kout by Jon ath an Hoef ler an d Tobias Frere-Jo nes for Hoef ler & Co. S oftwar e A dobe Creative Clou d InDesign , Illu strator, Ph otosh op E qu i ptment Mac Book Pro 15-in c h , 2.6 G Hz Epson Stylus Ph oto R2880 Pa p er R ed River Premium Matte 50 lb B ind i ng C alif orn ia Of fic e, San Fran c isc o, Calif orn ia May 12, 2015 Publi s her P rin c eton Un iversity Press ( Fic tic iou s book n ot pu blish ed by Prin c eton U n iversity Press) D es i gner Mallory A dams Photgr ap hy and Illu str ati ons P h otography: Flic kr Il lu stration s: Mallory A dams Ab out The Pr oject Tis is a stu den t projec t on ly. Not part of thi s b ook or any oth er part of th e projec t was pr o d u c ed for c ommerc ial u se.
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LIFE IN THE COLONY These tiny insects do more than just forage for honey; if you take a closer look you will see a colony buzzing with life. Although honeybees are spilt up into three different castes, they all participate in everyday tasks and decision that help keep the colony alive. Every job and every action they make ultimately affects the life of the entire colony. These incredible insects not only spend their lifespan collecting food for the entire colony, but also raise the larvae born from their queen, construct a unique living arrangement, and work together for the overall survival of the colony. Learn how these tiny creatures function in a colony as a single, unified unit.