The Behavior, Ecology and Evolution of Cichlid Fishes

Dedication
In Memoriam
	References
Preface
	Reference
Contents
Chance, Choice, and Cichlids
	References
Frontiers in Cichlid Research: A History of Scientific Advancement
	1 Cichlids in Natural History
		1.1 Traditions in Taxonomy: 1750-1800
		1.2 The Golden Age of Natural History: 1800-1900
	2 Pioneers of Cichlid Research
		2.1 The Shift from Natural History and the Legacy of The Crown: 1900-1930
		2.2 Tropical Fisheries and the Growth of Cichlid Research: 1930-1960
		2.3 Decolonization and the Struggle for Sustainable Fisheries: 1960-1980
	3 A Model Species in Evolutionary Biology
		3.1 The Evolutionary Synthesis and Specialization in Cichlid Research: 1930-1970
		3.2 Sexual Selection and the Curious Adaptive Radiation of Cichlids: 1970-1980
	4 The Future of Cichlid Research
		4.1 The Gene and Cichlid Research: 1980-1990
		4.2 The Tragedy and Promise of Fisheries and Aquaculture: 1980-2000
		4.3 The Continuing Role of Genetics in Cichlid Phylogeny: 1990-2000
		4.4 Looking Back and Moving Forward
	References
Ecological Opportunity, Genetic Variation, and the Origins of African Cichlid Radiations
	1 Introduction
	2 What Predicts Cichlid Adaptive Radiation and Diversity?
		2.1 Making Links to Speciation Mechanisms
		2.2 Predictors of Species Richness of African Lake Cichlid Assemblages
	3 The Origins and Maintenance of Genetic Variation in African Cichlids
		3.1 Genomic Data Facilitate Major Advances in Understanding Cichlid Evolutionary History
		3.2 The Maintenance of Genetic Variation During Adaptive Radiation
			3.2.1 Hybridization and Adaptive Radiation
			3.2.2 Incomplete Lineage Sorting
		3.3 Spatial Genetic Structure and the Origin of Cichlid Species
	4 Conclusion: Ecological Opportunity and Adaptive Radiation
	References
Evolution in the Fast Lane: Diversity, Ecology, and Speciation of Cichlids in the Lower Congo River
	1 Introduction
	2 Characterization of the LCR: A Geo-hydrological Context for Cichlid Diversification
	3 Geological Context of the LCR
	4 Living on the Edge: Lamprologus lethops
	5 Molecular Phylogenetics: Patterns of Colonization and Radiation in the LCR
	6 Biogeographic Divisions Within the LCR
	7 Hybridization and Reticulate Evolution
	8 Conclusions and Future Studies
	References
Neotropical Riverine Cichlids: Adaptive Radiation and Macroevolution at Continental Scales
	1 Neotropical Cichlids Then and Now
	2 The Riverine Neotropical Cichlid Fauna
		2.1 Taxonomic Diversity and Geographic Distribution
		2.2 Phylogeny, Times of Divergence, and Macroevolutionary Analyses
		2.3 Ecological and Morphological Diversity: Cichlinae Functional Morphospace
			2.3.1 Elongation or Ram-Suction Feeding Axis
			2.3.2 Suction-Biting and Maneuverability Axis
			2.3.3 Adaptive Landscape of Neotropical Cichlid Evolution
	3 Neotropical Cichlids as Ancient, Continental Adaptive Radiations
		3.1 Decoupled Divergence Patterns: Temporal and Phylogenetic Scale of Divergence in Neotropical Cichlids
		3.2 South America: Early Burst of Divergence in Lineages and Phenotypes
		3.3 Central America: Renewed Ecological Opportunity and Convergence
	4 Divergence in Rivers and Lakes: Are Processes the Same?
	References
Sympatric and Allopatric Diversification in the Adaptive Radiations of Midas Cichlids in Nicaraguan Lakes
	1 Introduction
		1.1 Discovery of the Midas Cichlid Species Complex
	2 The Ecological Setting
		2.1 Other Fish Species in Nicaraguan Lakes
	3 Parallel Phenotypic Divergence and Allopatric Speciation: Interlacustrine Radiation of Midas Cichlids
		3.1 Parallel Evolution in Body Shape After Crater Lake Colonization
		3.2 Parallel Evolution of the Visual System
	4 Sympatric Speciation: Midas Cichlids´ Intralacustrine Radiations
		4.1 Rapid and Parallel Intralacustrine Speciation Along the Benthic-Limnetic Axis
		4.2 Midas Cichlids with Hypertrophied Lips
		4.3 The Pharyngeal Jaws and Their Role in Midas Cichlid Sympatric Speciation
	5 The Intriguing Case of Color Polymorphism: Stable Polymorphism or In Vivo Speciation?
	6 Why Midas Cichlids?
	References
The Consequences of Anthropogenic Stressors on Cichlid Fish Communities: Revisiting Lakes Victoria, Kyoga, and Nabugabo
	1 Introduction: Cichlid Diversity and Adaptive Radiation
	2 The Lake Victoria Region: Past Environment and Fish Fauna
		2.1 Fish Fauna
		2.2 Exposure and Sensitivity to Multiple Anthropogenic and Environmental Stressors
	3 Relationship Between Multiple Stressors and Cichlid Diversity and Abundance in the Lake Victoria Region
		3.1 Overfishing
		3.2 Non-native Fish Species
			3.2.1 Impact on Native Oreochromine Cichlids
			3.2.2 Impact on Haplochromine Cichlids
		3.3 Habitat Degradation
		3.4 Invasive Plants
		3.5 Climate Variability and Change
	4 Persistence of Cichlid Fishes Under Multiple Stressors
	5 Conclusions
	References
Rapid Evolutionary Responses in Cichlids: Genetics of Adaptation, Morphology and Taxonomic Implications
	1 Introduction
	2 Molecular Tools
	3 Case Study: Haplochromis (Yssichromis) pyrrhocephalus
		3.1 Ecology: Horizontal Distribution and Diet
			3.1.1 Horizontal Distribution
			3.1.2 Diet
		3.2 Morphology
			3.2.1 External Measurements
			3.2.2 Gill Rakers and Gill Surface
			3.2.3 Premaxilla and Pharyngeal Jaws
			3.2.4 Eyes
		3.3 Mechanism Behind Morphological Changes
	4 Implications for Distinguishing Lake Victoria Species and Assessing Species Richness
	5 Future Directions
	References
Identifying and Conserving Tilapiine Cichlid Species in the Twenty-First Century
	1 Identifying Species
		1.1 Morphological Characterization
		1.2 Delimiting Sympatric Populations with Diet Analyses
		1.3 Advances in Molecular Approaches
		1.4 Integrated Analysis
	2 Citizen Science and Mobile Technology
	3 Defining Management Units for Tilapia
		3.1 Species Conservation Priorities, Threats, and Opportunities
		3.2 Advances in Identifying Tilapia in the Wild
		3.3 New Methods in Biomonitoring: Environmental DNA
	4 Conclusions
	References
Introduced Cichlids in the Americas: Distribution Patterns, Invasion Ecology, and Impacts
	1 Introduction
	2 Distribution of Non-native Cichlids in the Americas
		2.1 Distribution of Non-native Cichlids in the American River Basins
	3 Cichlid Introduction Vectors in the Americas
	4 Reasons for Success
	5 Impacts of Non-native Cichlids
	6 Final Remarks
	References
The Evolutionary Ecology of Cichlid Vision
	1 Visual Ecology
	2 Cichlid Vision
		2.1 The Cichlid Eye
		2.2 Groundbreaking Studies of Cichlid Vision
	3 Visual Ecology
		3.1 Visual Environment: Heterogeneity and Water Clarity
		3.2 Visual Signals
		3.3 Visual Sensitivity
	4 Evolution of Cichlid Visual Systems
		4.1 Sensory Drive and Speciation
		4.2 Spectral Tuning and Adaptive Radiation
			4.2.1 Adaptive Radiation
	5 Cichlid Vision and Human-Induced Environmental Change
		5.1 Lake Victoria Eutrophication
		5.2 Compensatory Responses
	6 Future Research
	7 Summary
	References
The Mechanosensory Lateral Line System of Cichlid Fishes: From Anatomy to Behavior
	1 Introduction
	2 Overview of the Anatomy of the Lateral Line System
		2.1 The Anterior (Cranial) Lateral Line System of Cichlids
		2.2 The Posterior (Trunk) Lateral Line System of Cichlids
	3 Neuroanatomy, Physiology, and Central Processing
		3.1 Peripheral Processing of Mechanosensory Information
		3.2 Central Processing of Mechanosensory Information
			3.2.1 Hindbrain Input and Processing
			3.2.2 Midbrain and Forebrain Input and Processing
	4 Lateral Line-Mediated Behaviors
		4.1 Feeding Behavior
		4.2 Mechanosensory Signaling During Social Interactions
			4.2.1 Role of Mechanoreception During Agonistic Behaviors
			4.2.2 Role of Mechanoreception in Reproduction and Parental Care
		4.3 Neural Processing of Lateral Line-Mediated Social Behaviors
	5 Conclusions and Prospects for Future Research
	References
Sonic Cichlids
	1 Introduction
	2 Checklist of Sound Producing Cichlid Species
		2.1 Calling Activity and Caller Sex
	3 Sound Producing Mechanism
		3.1 Functional Morphology of the Cichlid Sound Mechanisms
		3.2 Pharyngeal Teeth
		3.3 Pharyngeal Jaw Complex
		3.4 Fin-Girdle Muscles (Buckling Mechanism)
		3.5 Evolutionary Implications
	4 Sound Characteristics
		4.1 Acoustic Patterns
		4.2 Typical Sound Characteristics
		4.3 Aggression and Courtship
	5 Playback Experiments
	6 Species-Specific Sounds
		6.1 Sonic Diversity and Color Patterns
		6.2 Silent Behavior and/or Non-acoustic Cichlids
	7 Sound Detection and Hearing
		7.1 Background Noise Interference
	8 Future Directions
	References
The Evolution of Enhanced Cichlid Hearing: Functional Morphology and the Role of Ecoacoustical Factors
	1 Introduction
		1.1 General Structure and Function of the Inner Ear in Fishes
		1.2 The Role of the Swimbladder in Hearing
	2 Hearing Abilities in Cichlids: Auditory Measurements
		2.1 Baseline Auditory Sensitivities
		2.2 Extrinsic (Noise) and Intrinsic (Hormones) Effects on Hearing
	3 Diversity of Swimbladder and Inner Ears in Cichlids
		3.1 Morphology of the Swimbladder and Ancillary Auditory Structures
		3.2 Inner Ear Morphology
	4 Hearing in Cichlids: Functional Morphology and Evolution
		4.1 Relationship between Swimbladder Morphology and Hearing Abilities
		4.2 Relationship between Inner Ear Morphology and Hearing Abilities
		4.3 Evolution of Hearing Enhancement in Cichlids
	5 Suggestions for Future Research
	References
Parental Care in Cichlid Fishes
	1 Introduction
	2 Parental Care Strategies
		2.1 Gamete Provisioning
		2.2 Nest Preparation
		2.3 Defense
		2.4 Embryo Directed Behavior (Cleaning and Fanning)
		2.5 Provisioning of Young
	3 Where Does Care for Young Occur?
		3.1 Substrate Guarders
		3.2 Mouthbrooders
		3.3 Delayed (Primitive) Mouthbrooding
	4 Who Looks After Young? The Sex of the Caregiver
		4.1 Biparental Care
		4.2 Male Care
		4.3 Female Care
	5 Evolution of Care in Cichlids
	6 Alloparental Care
		6.1 Brood Mixing
			6.1.1 Why Should Foster Parents Adopt Free-Swimming Non-Descendants?
		6.2 Evolution of Cooperative Breeding
	7 Future Directions
	8 Summary and Conclusions
	References
Cichlids as a Model System for Studying Social Behaviour and Evolution
	1 The Beautiful Complexity of Cichlids
	2 Cichlid Social Groups
		2.1 Ecology: Defendable, Stable Resources
		2.2 From Fission-Fusion to Long-Term `Closed´ Groups
		2.3 Reproduction
		2.4 Social Group Demography and Dominance Hierarchies
		2.5 Cooperation
	3 Behavioural Complexity
		3.1 Social Interactions Generate Selection for Behavioural Complexity
		3.2 Communication Modalities
		3.3 Aggressive/Agonistic/Territorial Interactions in Cichlids
		3.4 Courtship and Mating
		3.5 Signalling Danger (Predation)
		3.6 Helping and Cooperation
	4 Cichlids as a Model for Studying Social Behaviour
		4.1 Phylogeny and the Evolution of Sociality
		4.2 Plasticity, Behavioural Development and Individual Differences
		4.3 The (Co-)Evolution of Social Structure and Behaviour
	5 The Past, Present, and Future of Cichlid Behavioural Research
	References
Integrative Neurobiology of Social Behavior in Cichlid Fish
	1 Introduction
	2 Cichlids as Models for the Study of Social Behavior
	3 The Cichlid Social Brain: Social Complexity and Brain Evolution
		3.1 Comparing Teleost Fishes and Mammals: SDMN Brain Homologies
		3.2 The Cichlid Social Decision-Making Network
	4 Cognitive Adaptations for Social Living
		4.1 Social Skills in Cichlids
	5 Neuroendocrine Regulation of Social Behavior
		5.1 Hormonal Actions on Behavior
		5.2 Social Feedback on Neuroendocrine Mechanisms
	6 Neurogenomics of Social Behavior
		6.1 Microarray Studies
		6.2 New Approaches to Neurogenomics
	7 Conclusions
	References
Respiratory Ecology of Cichlids
	1 Introduction
	2 Measuring Hypoxic Environments
	3 Cichlid Distribution in Hypoxic Habitats
	4 Strategies for Persisting Under Hypoxia
		4.1 Behavioral Responses to Hypoxia
		4.2 Mechanisms to Increase Oxygen Uptake and Delivery
			4.2.1 Gill Plasticity
			4.2.2 Increasing Blood Oxygen Transport
		4.3 Mechanisms to Reduce Aerobic Demands
	5 Hypoxia and Reproductive Traits in Cichlids
		5.1 Effects on Growth, Fecundity, and Secondary Sex Characteristics
	6 Hypoxic Refugia from Piscine Predators
	7 Summary
	References
Feeding Ecology of Lake Tanganyika Cichlids
	1 Introduction
	2 Food Web Structure of the Cichlid Community
	3 Community Structure and Trophic Guilds of Cichlids in Lake Shore Areas
		3.1 Trophic Niches
		3.2 Aufwuchs (Algae) Feeders
		3.3 Shrimp Eaters and Zoobenthos Feeders
		3.4 Piscivores
		3.5 Scale Eaters
	4 Future Directions
		4.1 Food Web Structure and Topology of Food Webs
		4.2 Symbiosis with Microorganisms
		4.3 Evo-Devo in Morphology in Relation to Foraging
		4.4 Reconstruction of Ancient Food Web and Morphological Evolution from Preserved Collections
		4.5 Personality/Behavioral Syndromes in Foraging Behaviors
	5 Conclusion
	References
Patterns of Trophic Evolution: Integration and Modularity of the Cichlid Skull
	1 Key Innovations in the Evolutionary Success of Cichlid Diversification
	2 Modularity as a Driving and Constraining Force in Morphological Evolution
	3 Modularity at Multiple Levels
		3.1 Phenotypic Modularity
		3.2 Genetic Modularity
		3.3 Functional Modularity
		3.4 Evolutionary Modularity
	4 Integrating Modularity at Multiple Levels: Lessons from the Hedgehog Pathway
	5 Feedback in the System
	6 Conclusions
	References
An Evo-devo View of Post-genomic African Cichlid Biology: Enhanced Models for Evolution and Biomedicine
	1 Introduction
	2 From Past to Present: Ground Breaking Phylogenetic and Genomic Cichlid Studies
	3 Cichlid Genomics in the Context of Current Evolutionary Thinking
	4 Combined Approaches Towards Understanding the Genetics of Cichlid Adaptations
	5 The Growing Relevance of Non-traditional Model Organisms for Medicine
		5.1 Craniofacial Genetics in Cichlids: From Embryonic Patterning to Adult Morphology
		5.2 Sex Determination: A Genomic Puzzle in Cichlids
		5.3 Genetic and Developmental Basis of Behaviour, Brain, and Neurobiology
	6 Conclusions: The Genome as the Basis for a Way Forward
	References
Index