Bemisia: Bionomics and Management of a Global Pest

Contents
Contributors
Introduction
	Literature Cited
Section I Taxonomy, Molecular Systematics, and Gene Flow in the Bemisia tabaci Complex and Bemisia Relatives
	Introduction
1 Systematics of Bemisia and Bemisia Relatives: Can Molecular Techniques Solve the Bemisia tabaci Complex Conundrum  A Taxonomists Viewpoint
	Introduction
		History of the Problem
	The Problem
	Problems in Whitefly Systematics
	Morphological Studies: A Versus B Biotype
	What All This Means
	Conclusions
	Literature Cited
2 Phylogenetic Biology of the Bemisia tabaci Sibling Species Group
	Introduction
	TheBemisia tabaci Sibling Species Group or Assemblage
		Historical Underpinnings
			The Invasion of the B Biotype
			Taxonomic Conundrums
			From Concept to Working Definition
		Biological Criteria for B. tabaci Biotypes
			Biotype Classification and Nomenclature
	Molecular Markers and Phylogeography
		Mitochondrial Molecular Markers
			Cytochrome Oxidase I
			Phlogenetic Relationships Are Generally Congruent with Extant Geography
			Global Genetic Divergence
			Selection and Differentiation
		Nuclear Markers
			18S rRNA Gene
			Non-coding Nuclear Sequences -- Ribosomal ITS1
			Sodium Voltage Gated Channel Gene Intron
	Some Biological Characteristics Used to Differentiate Biotypes
		Host Range and Preference
			Monophagous Biotypes
			Polyphagous Biotypes
			Habitat
			Host Range
			Dispersal Behavior
			Silvering Phenotype in Cucurbita spp. (and Other Hosts that Exhibit Phytotoxic-Like Symptoms)
		Gene Flow and Reproductive Isolation
			Evidence from Reciprocal Crosses Under Laboratory Conditions
			Specific Results and Trends
			Sympatry and Sister Clades
			Allopatry
		Endosymbionts of the B. tabaci Complex
			Secondary Symbionts
			Wolbachia and Cardinium Infection
	Conclusions
	Literature Cited
3 Tools and Recent Progress in Studying Gene Flow and Population Genetics of the Bemisia tabaci Sibling Species Group
	Introduction
	Protein Markers
		Enzyme Electrophoresis
			Uses, Assumptions, and Limitations
			Applications of Allozyme Analysis to Bemisia tabaci
			Protein Polymorphisms
			Esterases as Markers
	DNA-Level Markers
		RFLP Analysis
			Uses, Assumptions, and Limitations
			Applications of RFLP Analysis to Bemisia tabaci
	DNA-Level, PCR-Based Markers
		RAPD Analysis
			Uses, Assumptions, and Limitations
			Applications of RAPD Analysis to Bemisia tabaci
		AFLP Analysis
			Uses, Assumptions, and Limitations
			Applications of AFLP Analysis to in Bemisia tabaci
		Microsatellite Analysis
			Uses, Assumptions, and Limitations
			Applications of Microsatellite Analysis in Bemisia tabaci
	Additional Considerations and Future Directions
	Conclusions
	Literature Cited
Section II Biology and Ecology of Bemisia tabaci
	Introduction
	Literature Cited
4 Life History, Functional Anatomy, Feeding and Mating Behavior
	Introduction
	Morphology of Life Stages
		Egg Stage
		Nymphal Stage
		\'\'Pupal\'\' Stage
		Adult
			Head
			Thorax
			Abdomen
	Molting
	Feeding Apparatus and Feeding
		Mouthpart Morphology
			Adults
			Nymphs
		Mechanics of Stylet Penetration
			Adults
			Nymphs
		Precibarium and Cibarial Pump
		Saliva
			Salivary Glands
			Sheath Saliva
			Watery Saliva
			Salivary Components
		Feeding Behavior
	Alimentary Canal
	Courtship and Mating
	Specific Mating Behaviors of Three Bemisia tabaci Biotypes
		Male Searching and Initial Contact Between Sexes
		Parallel Positioning
		Antennal Drumming
		Male Abdominal Undulation
		Body Pushing
		Male Positioning
		Copulation
		Post-copulation
		Male Interference
	The Mating Behavior Cascade and Mate Discrimination
	Mating Behavior and Competitive Advantage of Biotype B
	Conclusions
	Literature Cited
5 Mutualistic and Dependent Relationships with Other Organisms
	Introduction
	Symbiotic Relationships
		Bacteriocyte Associated Endosymbionts
		Types of Endosymbiotic Relationships
	Endoymbionts Role in Nutrient Provisioning
		Provisioning in Aphids
		Provisioning in Whiteflies
	The Role of Secondary Symbionts in Whitefly Biology
	Potential for Symbiont Manipulation and Pest Management
	Role of Whitefly Endosymbionts in Virus Transmission
		Begomovirus GroEL-Relationships
		Use of B. tabaci Endosymbiotic GroEL in Diagnostic Tests
		Transgenic Plants Expressing the GroEL Confer Broad-Range Virus Resistance
	Intra- and Interspecific Interactions with Herbivores
		Importance of Intra- and Interspecific Interactions in B. tabaci Biology
		Positive Interactions of Bemisia feeding
		Competition Among Whiteflies
		Competition Between Whiteflies and Other Herbivores
	Conclusions
	Literature Cited
6 Population Dynamics, Demography, Dispersal and Spread of Bemisia tabaci
	Introduction
	Sampling Populations
	Demography
		Life History Studies
		Sex Ratio
		Life Tables
	Dispersal, Migration and Seasonality
		Measuring Movement and Flight Behavior
		Seasonality and Metapopulations
	Invasion and Spread
		The Invasion Process
		Mating Interactions
		Host Plant Effects
		Insecticide Resistance
		Whitefly--Begomovirus Interactions
		Future Invasion Threats
	Population Outbreaks
		Historical Outbreaks
			India -- 1920s
			Israel -- 1930s and 1940s
			Brazil -- 1970s
		Factors Contributing to Outbreaks
			Climate
			Agriculture
			Biotic Potential
			Management
	Population Models
	Conclusions
	Literature Cited
Section III Biology and Epidemiology of Bemisia -Vectored Viruses
	Introduction
	Viruses Transmitted by Bemisia spp
		Begomoviruses
		Criniviruses
		Ipomoviruses
		Other Viruses
	Whitefly Diversity and Virus Transmission
	Conclusions
	Literature Cited
7 Epidemiology of a Whitefly-Transmitted Cassava Mosaic Geminivirus Pandemic in Africa
	Introduction
	An Overview of Cassava Mosaic Geminiviruses, Whiteflies and the Epidemiology of the CMD Pandemic
		Cassava Mosaic Geminivirus Biology
		CMG Transmission
		Vector Host Interactions
		The CMD Pandemic
	The CMD Pandemic: New Insights and Research Gaps
		What Caused the Changes in the Impact of CMD-causing Viruses on Cassava Plants?
			A Recombinant Virus Is Associated with the Pandemic
			Virus--Virus Synergism Enhances Symptom Severity
			Molecular Complementation Facilitates Synergism
			Virus-Host Dynamics Change Following the Passage of the Pandemic \'Front\'
		Why Are B.tabaci Whiteflies Super-Abundant in the Pandemic-Affected Zone?
			B. tabaci Has Adapted Rapidly to Cassava
			Both Genetic and Environmental Factors Have Been Proposed as Causes of Super-Abundance
		What Are the Factors Behind the Rapid Local Spread of the Pandemic?
			Two Key Changes in the CMD Pathosystem Have Driven the Pandemic
			Evidence for the Origins of Changes in Both Virus and Vector Remains Inconclusive
		What Are the Factors Behind the Rapid Regional Spread of the Pandemic?
			The Flight Capabilities of B. tabaci Permit Mid-range Migration
			Patterns of Whitefly Migration and Resulting Regional Pandemic Spread Are Influenced by Environmental Factors
			The CMD Pandemic Is not Spread by Growers Carrying Infected Cassava Cuttings from Pandemic Affected to Unaffected Regions
		Can the Spread of the CMD Pandemic Be Halted?
			Host Plant Resistance Provides Effective CMD Control
			Pre-emptive Deployment of Resistant Varieties Has Been Used to Enhance the Response Times of CMD Mitigation Programmes
			Biological and Socio-political Factors Preclude the Halting of the Pandemic
			The Pandemic Can Be Confined to, and Managed in Africa
	Conclusions
		Questions to Direct Future Research Efforts
	Literature Cited
8 Tomato Yellow Leaf Curl Disease Epidemics
	Introduction
	Genetic Diversity of TYLCD-Associated Viruses
	Virus-Vector Interactions in the TYLCD Complex
	Cultivated Hosts and Wild Reservoirs of TYLCD-Associated Viruses
	Recombination as a Key Force Driving Evolution of TYLCD-Associated Virus Populations
	Diagnosis of TYLCD-Associated Viruses
		Polymerase Chain-Reaction
		RFLP Analysis of PCR-Amplified Products
		Nucleic Acid Hybridization
		Rolling Circle Amplification (RCA) sing 29 RNA Polymerase
	Management of TYLCD Epidemics
	Conclusions
	Literature Cited
9 Distribution and Dissemination of Begomoviruses in Latin America and the Caribbean
	Introduction
	Historical Background
		The Original Begomoviruses
		Original Begomovirus Reservoirs
		The Vector:Bemisia tabaci
		The Origin of Begomovirus Epidemics in Latin America
	Main Crops Affected by Begomoviruses in Latin America
		Common Bean
			South America
			Central America
			Northern Mexico
		Tomato
			South America
			Mexico
			Central America
			Caribbean Region
		Sweet and Hot Peppers
		Cucurbits
		Potato
		Tobacco
		Soybean
		Fruit Crops
		Integrated Whitefly and Begomovirus Management
	Literature Cited
10 Transmission Efficiency and Epidemiology of Criniviruses
	Introduction
	Crinivirus Epidemiology Is Influenced by Vector Whitefly Population Shifts
	Unique Vector Transmission Characteristics of Tomato Criniviruses
	Factors Influencing Crinivirus Transmission Efficiency
		Does the Host Plant Influence Crinivirus Transmission by Whitefly Vectors?
	Literature Cited
11 A Review of Ipomoviruses and Watermelon Vine Declinein Florida
	Ipomoviruses
	Watermelon Vine Decline in Florida
	Squash Vein Yellowing Virus
	Literature Cited
12 Transovarial Transmission of Begomoviruses in Bemisia tabaci
	Introduction
	Are Begomoviruses Transmitted Transovarially?
	Conclusions
	Literature Cited
Section IV Management of Bemisia in Diverse Cropping Systems
	Introduction
13 Optical Manipulation for Control of Bemisia tabaci and Its Vectored Viruses in the Greenhouse and Open Field
	Introduction
	Light Manipulation in Protected Crops
		UV-Absorbing Films Protect Greenhouses from Insect Invasion and Spread of Virus Diseases
		The Effect of the Chemical Attributes of UV-Blocking Films on Their Protection Capacity
		Effect of UV Absorbing Films on Natural Enemies
		Effect of UV-Absorbing Films on Pollinators
		Putative Mechanism of UV-Blocking Films
	Light Manipulation in the Open Field
		Soil Mulches Protect Crops in the Open Field
		The Putative Mechanism of Action of Colored Soil Mulches
	Conclusions
	Literature Cited
14 Host Plant Resistance for the Management of Bemisia tabaci: A Multi-crop Survey with Emphasis on Tomato
	Introduction
	Innate Resistance in Tomato: The MI-1 Gene
	Induced Resistance in Tomato
	Plant Resistance to B. tabaci in Other Crops
	Final Remarks
	Literature Cited
15 Natural Enemies of Bemisia tabaci: Predators and Parasitoids
	Introduction
	Predator Biology and Ecology
		Coleoptera
		Heteroptera
		Neuroptera
		Diptera
		Acarina
	Parasitoid Biology and Ecology
		Encarsia
			Encarsia formosa
			Encarsia bimaculata
			Encarsia porteri (Mercet)
			Encarsia sophia (= En. transvena Timberlake)
			Other Encarsia species
		Eretmocerus
			Eretmocerus mundus
			Eretmocerus eremicus (= Er. nr. californicus   )
			Eretmocerus queenslandensis Naumann and Schmidt
			Eretmocerus sp. nr. furuhashii
			Eretmocerus emiratus Zolnerowich and Rose (Ethiopia)
			Other Eretmocerus sp.
		Behavior
			Dispersal
			Functional Responses and Handling Times
			Influence of Host Volatiles and Chemical Cues on Behaviour
			Foraging on the Leaf
			Ovipositional Marking
			Host Feeding and Egg Production
	Natural Enemy Interaction
		Intraguild Predation
		Parasitoid--Parasitoid Interactions
		Entomopathogen-Parasitoid and Predator Interactions
		Natural Enemy-Plant Interactions
		Relevance of Interactions Between Natural Mortality Factors to Biological Control
	Utilization, Monitoring, and Assessing the Impact of Natural Enemies
		Utilization
			Predators
			Parasitoids
		Monitoring and Impact Assessment
			Predators
			Parasitoids
		Life Table Studies
	Conclusions
	Literature Cited
16 Ecological Determinants of Bemisia tabaci Resistanceto Insecticides
	Introduction
	Ecological Characteristics of B. tabaci as a Resistance Recidivist
		Polyphagy
		r-Selection
		Dispersal
		Adaptability
		Integrated Control
	Agro-Ecology of Resistance
		Agro-Environment
			Closed Versus Open Systems
			Intermittent Monoculture Versus Continuous Polyculture
	Patterns of Resistance
		Stable Versus Unstable Resistance
		Rates of Resistance Evolution
		Cross Resistance
	Biotype
	Resistance Management
		Minimize Insecticide Use
		Diversify Insecticide Use
		Refine Insecticide Use
	Conclusions
	Literature Cited
17 Integrated Systems for Managing Bemisia tabaci in Protected and Open Field Agriculture
	Introduction
	Biologically Based Management of B. tabaciin Protected Vegetable Crops
		Key Pests of Greenhouse Vegetables
		Damage to Vegetable Crops Caused by Bemisia tabaci
		Greenhouse Exclusion Technology
		Host Plant Resistance
		Biological Control of Bemisia tabaci
			Entomopathogenic Fungi
			Hymenoptera: Aphelinidae
			Heteroptera: Miridae
			Acari: Phytoseiidae
		Compatibility of Various Pest Control Practices
		Compatibility of Pesticides with Biological Control
	Area-Wide Management of Whitefly in Open Field Crops
		Management System for Cotton in the Desert Southwest USA
			Cultural Control
			Host Plant Resistance
			Biological Control
			Monitoring and Treatment Decisions
		Management in Alfalfa in the Desert Southwest USA
		Vegetables in the USA Desert Southwest and Elsewhere
			Action Thresholds for Whiteflies in Open-Field Vegetables
			Role of Biological Control and Adaptation of Augmentative Control Practices
	Conclusions
	Literature Cited
Section V Prospects for the Application of Genomics
	Introduction
	Consortium Partner Countries and Participants (2009)
	History of Synergistic Activities
	Literature Cited
18 The Whitefly Genome  White Paper: A Proposal to Sequence Multiple Genomes of Bemisia tabaci
	Introduction
	Homopteran Model
		Cryptic Species
		Systematics Model -- Sibling Species Group
		Subtropical Homopteran Model -- Comparative Biology
		Reproduction -- Haplodiploidy, Reproductive Isolation, and Speciation
			B. tabaci Biotypes as a Model for Ongoing Speciation
	Whitefly-Begomovirus Interactions: A Cross Kingdom Model
	The Whitefly Transcriptome
		Genome Size of the Whitefly,    B. tabaci
		Activation and Repression of B. tabaci Stress-Response Gene from Insecticide Application, Parasitism by Natural Enemies and High Temperature
			Insecticide Application
			Parasitism by Natural Enemies
			High Temperatures
		Knowledge Base and Available Tools
			Colonies of B. tabaci
			Nuclear Genome Size of B. tabaci
			Expressed Sequence Tags
			Spotted cDNA Microarrays -- Functional Genomics
			Gene Silencing
		Top Candidate Biotypes for Genome-Transcriptome Sequencing
			B Biotype
			A Biotype
			Q Biotype
			Others
		Available Annotation Tools
			Model Insects
		Sequencing and De Novo Assembly of Whitefly Endosymbiont Genomes -- The Microbiome
	Synergy  Other Genome Projects and User Communities
		The User Community is Global
		Insect Genome Projects
		Some Relevant Websites
	Conclusions
	Literature Cited
Index