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Bees as Superorganisms
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Table of Contents

1 What Is a Superorganism?.- 1.1 From Cells to Metazoan Organisms.- 1.2 What Makes a Social Group a Superorganism?.- 1.3 Diversity of Superorganisms.- 1.3.1 Termites.- 1.3.2 Superorganismic Ants.- 1.3.3 Bees.- 1.3.4 The Naked Mole Rat.- 1.4 What Not to Expect from Superorganisms.- 2 Evolution of Superorganisms.- 2.1 From Solitary Individuals to Superorganisms: The Theories.- 2.1.1 Darwin's Dilemma.- 2.1.2 Individual Selection Models.- 2.1.2.1 Kin Selection.- 2.1.2.2 Assured Fitness Returns.- 2.1.2.3 Parental Manipulation.- 2.1.2.4 Mutualism and Reciprocal Altruism.- 2.1.2.5 Testing Theories.- 2.1.3 Group Selection.- 2.1.3.1 Classical Group Selection.- 2.1.3.2 Competitive Group Selection.- 2.1.3.3 Colony-Level Selection.- 2.1.3.4 Testing Theories of Group Selection.- 2.2 From Solitary Individuals to Superorganisms: The Evidence.- 2.2.1 Sociality in Bees.- 2.2.2 Solitary and Subsocial Bees.- 2.2.2.1 Megachile rotundata: The Individualist.- 2.2.2.2 Carpenter Bees: The Hierarchists.- 2.2.3 Primitively Social Bees.- 2.2.3.1 Exoneura bicolor Smith: The Mutualist.- 2.2.3.2 Halictine Bees: Getting Social.- 2.2.3.3 Bombus: Controlling Environment.- 2.2.4 Superorganismic Bees.- 2.2.4.1 Stingless Bees.- 2.2.4.2 Evolution Within the Genus Apis.- 3 Physiology.- 3.1 Development.- 3.1.1 From Egg to Adult.- 3.1.2 Caste.- 3.1.3 Birth, Aging, and Death of a Superorganism.- 3.1.3.1 Birth.- 3.1.3.2 Aging and Age Variation.- 3.1.3.3 Death and Immortality.- 3.2 Glands and Secretions.- 3.2.1 Endocrine System.- 3.2.2 Exocrine System.- 3.3 Feeding and Digestion.- 3.3.1 Nutrition.- 3.3.2 Food Exchange and the Meniscus Effect.- 3.3.3 Pollen Stores.- 3.3.4 Making Honey.- 3.4 Respiration.- 3.4.1 Respiration in Individuals.- 3.4.2 Respiration in Colonies.- 3.5 Circulation.- 3.5.1 Haemolymph Circulation.- 3.5.2 Circulation in the Colony.- 3.6 Colonial Excretion and Water Balance.- 3.7 Metabolic Physiology.- 3.7.1 Temperature Control.- 3.7.1.1 Cooling.- 3.7.1.2 Heating.- 3.7.1.3 Maintenance of Nest Temperature.- 3.7.1.4 Optimal Environmental Temperature and Nest Choice.- 3.7.2 Mass-Metabolism Relationships and Colony Fitness.- 3.7.3 Seasonal Relationships.- 3.8 Neurophysiology.- 3.8.1 Vision.- 3.8.2 Chemical Sense.- 3.8.3 Acoustic and Mechanical Reception.- 3.8.4 Special Sense, Learning and Integration.- 3.9 Muscle Function.- 3.9.1 Locomotion and Flight.- 3.9.2 Stinging and Biting.- 3.10 Circadian Rhythms and Sleep.- 3.10.1 Cyclic Metabolism.- 3.10.2 Cyclic Locomotion.- 3.10.3 Cyclic Ventilation.- 3.10.4 Sleep.- 4 Communication Network of the Superorganism.- 4.1 The Analysis of Communication Networks.- 4.2 Division of Labour.- 4.3 Food Procurement and Temperature Regulation.- 4.3.1 The Search for Food.- 4.3.1.1 Individual Search Pattern of a Scout.- 4.3.1.2 Foraging a Patch of Flowers.- 4.3.2 Communication Mechanisms.- 4.3.2.1 Round Dance.- 4.3.2.2 Waggle Dance.- 4.3.2.3 Acoustic Signals.- 4.3.2.4 Odour Signals.- 4.3.2.5 Dorsoventral Abdominal Vibration (DVAV) Dance.- 4.3.3 Central Food Handling and Social Integration.- 4.3.3.1 Amoeboid Foraging Pattern.- 4.3.3.2 Nectar Storage.- 4.3.3.3 Queueing.- 4.3.3.4 Comb Utilization.- 4.3.3.5 Water Handling.- 4.4 Search for Housing.- 4.4.1 Nest Site Selection.- 4.4.2 Swarm Orientation.- 4.4.3 Migration.- 4.5 Pheromones.- 4.5.1 Superorganismal Control.- 4.5.2 Queen Pheromones Inside the Nest.- 4.5.2.1 Queen Retinue Behaviour.- 4.5.2.2 Colony Stabilization and Ovary Suppression in Workers.- 4.5.3 Queen Pheromones Outside the Nest.- 4.5.3.1 Swarm Attraction.- 4.5.3.2 Sex Attractant.- 4.5.4 Worker Pheromones.- 4.5.4.1 Orientation.- 4.5.4.2 Colony Defence.- 4.5.5 Other Pheromones and Odours.- 4.6 Learning.- 4.7 Social Intelligence and Network Analysis.- 4.7.1 Social Intelligence.- 4.7.2 Network Analysis.- 5 Ecology.- 5.1 Demography of Natural Populations.- 5.1.1 Life History Strategies.- 5.1.1.1 Sessile Phase.- 5.1.1.2 Mobile Phase.- 5.1.1.3 Life Cycle.- 5.1.2 Spatial and Temporal Distribution.- 5.1.2.1 Colony Defence.- 5.1.2.2 Removal of Debris and the Dead.- 5.1.2.3 Home Range.- 5.1.2.4 Mating Distance.- 5.1.2.5 Overdispersion of Colonies.- 5.1.3 Distribution of Honeybees.- 5.1.4 Intracolonial Structure.- 5.2 Community Structure.- 5.2.1 Plant/Bee Symbioses.- 5.2.1.1 Foraging and Pollination.- 5.2.1.2 Energy Flow and Material Turnover.- 5.2.1.3 Honeydew.- 5.2.2 Selective Forces and Diversity.- 5.2.2.1 Intraspecific Competition.- 5.2.2.2 Interspecific Competition Among Superorganisms.- 5.2.2.3 Competition with Individual Organisms.- 5.2.3 Diseases, Parasites, and Predators.- 5.2.3.1 Superorganism Pathology.- 5.2.3.2 Specific Examples.- 5.3 Superorganism Size.- 5.3.1 Temporal Variation in Group Size.- 5.3.1.1 Environmental Determinants of Group Size.- 5.3.1.2 Intrinsic Determinants of Group Size.- 5.3.2 Fitness and Group Size.- 5.3.2.1 r and K Selection.- 5.3.2.2 Queen Fecundity.- 5.3.2.3 Periods of Dearth.- 5.3.2.4 Defence and Group Size.- 5.3.3 Optimization Model of Group Size.- 5.4 Ecological Impact of Superorganismic Honeybees.- 5.4.1 Natural Areas.- 5.4.2 Agricultural Land.- 6 Reproduction.- 6.1 Individual Reproduction.- 6.1.1 Sexual Reproduction.- 6.1.1.1 Sex Determination and Sex Ratios.- 6.1.1.2 The Queen.- 6.1.1.3 The Drones.- 6.1.1.4 Mating Behaviour.- 6.1.2 Parthenogenesis.- 6.1.2.1 Production of Drones by the Queen.- 6.1.2.2 Production of Drones by Workers.- 6.1.2.3 Production of Females by Workers.- 6.1.3 Negative Reproduction.- 6.1.3.1 Brood Cannibalism.- 6.1.3.2 Worker Policing.- 6.2 Colonial Reproduction.- 6.2.1 The Seasonal Development of a Colony.- 6.2.1.1 The Yearly Life Cycle in a Temperate Climate.- 6.2.1.2 Somatic Growth and Worker Force Dynamics.- 6.2.2 Sexual Reproduction of Colonies.- 6.2.2.1 Production of Gametes and the Zygote.- 6.2.2.2 The Sex of a Colony.- 6.2.3 Asexual Reproduction of Colonies.- 6.2.3.1 Swarming Behaviour.- 6.2.3.2 How Many Swarms?.- 6.2.3.3 The Primary Swarm: Join the Old or the New Queen?.- 6.2.3.4 The Afterswarms: Join the Super or Half Sister?.- 6.2.3.5 Is Swarming = Colony Budding?.- 7 Genetics.- 7.1 Gene Expression vs Genotype Expression.- 7.1.1 The Limbs and the Stomach Model.- 7.1.2 Intraorganismic Genetic Variation - the Principal Difference.- 7.1.3 Gametes - the Functional Difference.- 7.2 Genetic Basis of Honeybees.- 7.2.1 Male Haploidy.- 7.2.2 Sex Determination.- 7.2.2.1 The Sex Locus.- 7.2.2.2 Diploid Males.- 7.2.3 The Terminology Problem.- 7.2.3.1 What Is a Generation?.- 7.2.3.2 Who Is Who in a Honeybee Family?.- 7.3 Genetic Variation in Honeybee Colonies.- 7.3.1 Origins for Intracolonial Genetic Variance.- 7.3.2 Measures of Genetic Relationship.- 7.3.2.1 Coefficients of Relationship Based on Pedigree Information.- 7.3.2.2 Relationships in Natural Populations.- 7.3.2.3 Weighted Coefficients of Relatedness.- 7.3.3 Genetic Relationship Between the Members of a Colony.- 7.3.3.1 Relationship Between Queen and Her Offspring.- 7.3.3.2 Relationship Between Workers and Drones.- 7.3.4 Relationship Between Groups.- 7.3.4.1 Average Individual Relationship.- 7.3.4.2 Group Relationship.- 7.4 Quantitative Genetics.- 7.4.1 The Problems in Analyzing Social Characters.- 7.4.2 The Classical Parameters.- 7.4.2.1 Repeatability.- 7.4.2.2 Selectability.- 7.4.2.3 Non-Linear Interactions.- 7.4.3 Similarities Between Colony Members.- 7.4.3.1 Genetic Covariance Among Workers.- 7.4.3.2 Genetic Covariance Among Drones.- 7.4.3.3 Covariance Between Queen and Offspring.- 7.4.3.4 Covariance Between Drone and Offspring.- 7.4.3.5 Covariance Between Mid-Parent and Offspring.- 7.4.4 Similarities Between Related Colonies.- 7.4.4.1 Worker Effects.- 7.4.4.2 Combined Effects.- 7.4.5 Selection Index.- 7.4.6 Empirical Examples.- 7.4.6.1 Analysis of Individual Characters.- 7.4.6.2 Analysis of Social Traits of Small Worker Groups.- 7.5 Behavioural Genetics.- 7.5.1 Analysis of Individual Behaviour.- 7.5.1.1 Learning Behaviour.- 7.5.1.2 Trophallactic Behaviour.- 7.5.1.3 Reproductive Dominance.- 7.5.2 Analysis of Social Behaviour.- 7.5.2.1 Hoarding Behaviour.- 7.5.2.2 Hygienic Behaviour.- 7.5.2.3 Defensive Behaviour.- 7.5.2.4 Division of Labour.- 7.6 Population Genetics.- 7.6.1 The Effective Population Size of Superorganisms.- 7.6.2 Polymorphisms in Nuclear Genome.- 7.6.2.1 Isozyme Polymorphism.- 7.6.2.2 Molecular DNA Markers.- 7.6.3 Inbreeding.- 7.6.3.1 Inbreeding and Practical Breeding Schemes.- 7.6.3.2 Inbred Superorganisms.- 7.6.4 Variation of Mitochondrial Genes.- 7.6.4.1 Mitochondrial DNA of Honeybees.- 7.6.4.2 Cyto-Nuclear Disequilibrium.- 8 Natural Selection.- 8.1 Selection at the Individual Level.- 8.1.1 Selection of Sexual Reproductives.- 8.1.2 Intracolonial Selection Among Laying Workers.- 8.2 Kin Selection.- 8.2.1 Discrimination Among Queens.- 8.2.2 Kin Discrimination Among Workers.- 8.3 Selection on Phenotypes of Superorganisms.- 8.3.1 Genetic Analysis of Division of Labour.- 8.3.2 Genetic Variability and Task Specialization.- 8.3.3 Non-Linear Worker Interactions.- 8.4 Africanized Honeybees: A Large-Scale Evolutionary Experiment.- 8.4.1 The History of Africanization.- 8.4.2 Reproductive Advantage of Africanized Honeybees.- 8.4.3 Genetic Evidence from Field Studies.- 8.5 Selection in Superorganisms: Too Complex to Study?.- References.

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