The Evolution and Fossil Record of Parasitism: Identification and Macroevolution of Parasites

Foreword
Preface
Contents
Chapter 1: Parasites of Fossil Vertebrates: What We Know and What Can We Expect from the Fossil Record?
	1.1 Introduction
		1.1.1 Cestodes (Tapeworms)
		1.1.2 Trematodes (Flukes)
		1.1.3 Nematodes (Roundworms)
		1.1.4 Acanthocephalans (Thorny-Headed Worms)
		1.1.5 Monogeneans
		1.1.6 Parasitic Copepods
		1.1.7 Parasitic Isopods
		1.1.8 Pentastomids (Tongue Worms)
		1.1.9 Ticks
		1.1.10 Ectoparasitic Insects (Fleas and Lice)
	1.2 A Note of Caution Regarding Fossil Parasites
	1.3 Framework for Inferring Parasite Communities of Extinct Vertebrate Groups
	1.4 Future Research Directions and Further Questions
	References
Chapter 2: Fossil Record of Viruses, Parasitic Bacteria and Parasitic Protozoa
	2.1 Introduction
	2.2 Virus Fossils
		2.2.1 Direct Evidence of Fossil Viral Infections
		2.2.2 Indirect Evidence of Virus Infections
			2.2.2.1 Parasitic Wasps with Polydnaviruses
			2.2.2.2 Tumors in Lepidoptera
			2.2.2.3 Iridoviridae
	2.3 Fossil Pathogenic Bacteria
		2.3.1 Direct Evidence of Fossil Pathogenic Bacteria
		2.3.2 Indirect Evidence of Fossil Symbiotic-Pathogenic Bacteria
	2.4 Protozoan Fossils
		2.4.1 Direct Evidence of Fossil Protozoan Parasites
			2.4.1.1 Fossil Trypanosome Parasites
			2.4.1.2 Fossil Malaria Parasites
			2.4.1.3 Other Fossil Records of Protozoan Parasites
				Fossil Piroplasmid Parasites
				Fossil Gregarine Parasites
		2.4.2 Indirect Evidence of Fossil Protozoan Parasites
			2.4.2.1 Fossil Parasites in Coprolites
	2.5 Conclusions
	References
Chapter 3: Fungi as Parasites: A Conspectus of the Fossil Record
	3.1 Introduction
	3.2 Identifying Fungal Parasitism in the Fossil Record
		3.2.1 Finding Fossil Fungi
		3.2.2 Tracing Fungal Parasitism in the Fossil Record
	3.3 Fossils of Fungi as Parasites
		3.3.1 Fungal Parasites of Land Plants
			3.3.1.1 Early Land Plants
			3.3.1.2 Plant Structural Alterations in Response to Fungal Intrusion
			3.3.1.3 Host Responses in Woody Plants
			3.3.1.4 Host Plant Preservation and Fungal Distribution
			3.3.1.5 Epiphyllous Fungi
			3.3.1.6 Dispersed Remains and Plant Pathogens
		3.3.2 Fungal Parasites of Algae
		3.3.3 Fungal Parasites of Other Fungi
			3.3.3.1 Rhynie Chert Interfungal Interactions
			3.3.3.2 Fossil Fungal “Sporocarps”
			3.3.3.3 Hyperparasitism
		3.3.4 Fungal Parasites of Animals
			3.3.4.1 Rhynie Chert
			3.3.4.2 Amber Inclusions
			3.3.4.3 Cordycipitaceae Interactions with Arthropods
			3.3.4.4 Dinosaurs
	3.4 Concluding Remarks
	References
Chapter 4: Evolution, Origins and Diversification of Parasitic Cnidarians
	4.1 Introduction
	4.2 Parasitic Cnidarians Other than Endocnidozoans
	4.3 The Endocnidozoa
		4.3.1 General Biology
		4.3.2 Comparative Development and Body Plans
	4.4 Evolution and Life Cycles of Endocnidozoans
		4.4.1 Preadaptations to Parasitism
		4.4.2 Life Cycle Speculations
	4.5 Origins and Fossil Records of Endocnidozoa and Their Recognised Major Host Groups
		4.5.1 Cnidarian Origins and Fossil Record
		4.5.2 Vertebrate Origins and Fossil Record
		4.5.3 Lophotrochozoan Origins and Fossil Record
		4.5.4 Annelid Origins and Fossil Record
		4.5.5 Bryozoan Origins and Fossil Record
		4.5.6 Other Potential Ancient Invertebrate Hosts
		4.5.7 Summary of Origins and Ancient Hosts
	4.6 Inferring Endocnidozoan Origins and Acquisition of Early Hosts
		4.6.1 The Process of Host Acquisition
		4.6.2 Molecular Clock and Cophylogenetic Investigations
			4.6.2.1 Some General Pitfalls
			4.6.2.2 Endocnidozoan Origins and Host Use Over Time
			4.6.2.3 The Endocnidozoa
		4.6.3 Scenarios of Endocnidozoan Evolution and Recommendations for Future Study
	4.7 Adaptation and Diversification of Endocnidozoans
		4.7.1 Adaptations to a Parasitic Life Style
		4.7.2 Patterns of Diversification
	4.8 Conclusions
	References
Chapter 5: Evolutionary History of Bivalves as Parasites
	5.1 Introduction
		5.1.1 Bivalves in Relations with Other Organisms
		5.1.2 Galeommatoidea
		5.1.3 Unionida
	5.2 Functional Anatomy of Adult Mussels and the Larvae
		5.2.1 Adults
		5.2.2 Larvae
	5.3 Life History, with the Review of the Adaptations to Attracting the Host
		5.3.1 Life History
		5.3.2 Adaptations to Host Infection
	5.4 Phylogeny of the Unionida
		5.4.1 Origin of Unionida
		5.4.2 Classification and Diversity
		5.4.3 Phylogeny
	5.5 Evolution of Parasitism
		5.5.1 Parental Care
		5.5.2 Parasitic Larvae
		5.5.3 Origin and Evolution of Unionoid Parasitism
			5.5.3.1 Phoresis
			5.5.3.2 Encapsulation
			5.5.3.3 Suitable Host
			5.5.3.4 Duration of Encapsulation and Metamorphosis
			5.5.3.5 In Search of the Host of Early Unionids
	5.6 Conclusions
	References
Chapter 6: Gastropods as Parasites and Carnivorous Grazers: A Major Guild in Marine Ecosystems
	6.1 Introduction
	6.2 How to Infer Parasitism in Fossil Gastropods
		6.2.1 Direct Observations
		6.2.2 Taxonomic Uniformitarianism
			6.2.2.1 Eulimidae
			6.2.2.2 Epitoniidae
			6.2.2.3 Coralliophilinae (Muricidae)
			6.2.2.4 Pyramidellidae
			6.2.2.5 Architectonicidae and Mathildidae
			6.2.2.6 Triphoridae and Cerithiopsidae: Carnivorous Grazers (Sponge Feeders)
				Triphoridae
				Cerithiopsidae
			6.2.2.7 Ovulidae
		6.2.3 Functional Shell Morphology
		6.2.4 Analogy Based on Associated Phenomena
	6.3 Conclusions
	References
Chapter 7: Fossil Constraints on the Timescale of Parasitic Helminth Evolution
	7.1 Introduction
	7.2 Phylogenetic Affinities and Distribution of Parasitic “Helminths”
	7.3 Preservation Potential
	7.4 Time Constraints on the Origin of Eumetazoan Helminths
		7.4.1 Fossil Record of Parasitic Annelids
		7.4.2 Fossil Record of Parasitic Pentastomids
		7.4.3 Fossil Record of Parasitic Platyhelminths
		7.4.4 Fossil Record of Parasitic Nematodes
		7.4.5 Fossil Record of Nematomorphs
		7.4.6 Fossil Record of Syndermata
	7.5 Discussion
	7.6 Conclusions
	References
Chapter 8: Thorny-Headed Worms (Acanthocephala): Jaw-Less Members of Jaw-Bearing Worms That Parasitize Jawed Arthropods and Jawed Vertebrates
	8.1 Introduction
	8.2 Acanthocephalans in Hominoids and Potential Reservoirs for Human Infections
	8.3 Solid-Parts and Their Preservation Potential
		8.3.1 Acanthocephalan Propagules: Eggs in Space and Time
		8.3.2 Hooks
		8.3.3 Copulatory Cap
	8.4 Soft Tissue, Functional Morphology and the Ideal Fossil
		8.4.1 Outer Contour and Tegument
		8.4.2 Presomal Musculature and Anchoring
		8.4.3 Presomal Sensory Organs
		8.4.4 Lack of an Intestinal Tract
	8.5 Pathological Manifestations of Infections with Acanthocephalans
	8.6 Phylogenetic Relationships of Acanthocephala and Taxonomic Implications
	8.7 Evolution of Acanthocephalan Endoparasitism: A Conditional Hypothesis
	8.8 Acanthocephala and Gnathifera: Fossil Report and Time Line
	8.9 Cambroclavida: Microfossils of Questionable Acanthocephalan Affiliation
	8.10 Conclusions
	References
Chapter 9: Chelicerates as Parasites
	9.1 Introduction
		9.1.1 The Chelicerate Fossil Record
	9.2 Sea Spiders
		9.2.1 Cambropycnogon
	9.3 Horseshoe Crabs and Eurypterids
	9.4 Arachnids
		9.4.1 Acariform Mites
			9.4.1.1 Trombidiform Mites
			9.4.1.2 Sarcoptiform Mites
		9.4.2 Parasitiform Mites
			9.4.2.1 Opilioacarids
			9.4.2.2 Holothyrids
			9.4.2.3 Ticks (Ixodida)
			9.4.2.4 Mesostigmatids
	9.5 The Origins of Parasitic Behaviour
		9.5.1 Nest Associates
		9.5.2 From Phoresy to Parasitism
			9.5.2.1 Phoresy in the Fossil Record
	9.6 Chelicerates as Victims
		9.6.1 Arachnid Parasites and Parasitoids
	References
Chapter 10: Evolutionary History of Crustaceans as Parasites
	10.1 Introduction
	10.2 Amphipoda
		10.2.1 General Aspects
		10.2.2 Phylogenetic Inference of Appearance and Molecular Estimations of Early Evolution
		10.2.3 Fossil Representatives
	10.3 Isopoda
		10.3.1 General Aspects
		10.3.2 Phylogenetic Inference of Appearance and Molecular Estimations of Early Evolution
		10.3.3 Fossil Representatives
	10.4 Copepoda
		10.4.1 General Aspects
		10.4.2 Phylogenetic Inference of Appearance and Molecular Estimations of Early Evolution
		10.4.3 Fossil Representatives
	10.5 Thecostraca
		10.5.1 General Aspects
		10.5.2 Phylogenetic Inference of Appearance and Molecular Estimations of Early Evolution
		10.5.3 Fossil Representatives
	10.6 Branchiura
		10.6.1 General Aspects
		10.6.2 Phylogenetic Inference of Appearance and Molecular Estimations of Early Evolution
		10.6.3 Fossil Representatives
	10.7 Pentastomida
		10.7.1 General Aspects
		10.7.2 Phylogenetic Inference of Appearance and Molecular Estimations of Early Evolution
		10.7.3 Fossil Representatives
	10.8 Conclusion and Outlook
	References
Chapter 11: The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
	11.1 Introduction
	11.2 Defining the Insect Consumption of Animals
		11.2.1 Predation
		11.2.2 Parasitism
		11.2.3 Parasitoidism
		11.2.4 Parasitoidism: A History of the Term
		11.2.5 Types of Parasitoidism
			11.2.5.1 Ectoparasitoidism Versus Endoparasitoidism
			11.2.5.2 Koinobiont Parasitoidism Versus Idiobiont Parasitoidism
			11.2.5.3 Solitary Versus Gregarious Parasitoidism
			11.2.5.4 Superparasitoidism Versus Multiparasitoidism
			11.2.5.5 Hyperparasitoidism
			11.2.5.6 Egg Parasitoidism, Larval Parasitoidism, Pupal Parasitoidism and Adult Parasitoidism
			11.2.5.7 Cleptoparasitoidism
	11.3 The Evidence: Distinguishing Predation, Parasitism and Parasitoidism
		11.3.1 Biomolecular Data
		11.3.2 Taxonomic Affiliation
		11.3.3 Structural and Functional Attributes
		11.3.4 Host Tissue Damage
		11.3.5 Plant–Insect Interactions
		11.3.6 Gut Contents
		11.3.7 Coprolites
		11.3.8 Sedimentary Ichnological Evidence
	11.4 Evolutionary and Ecological Biology
	11.5 Parasite and Parasitoid Taxa
		11.5.1 Parasite Taxa
			11.5.1.1 Blattodea (Cockroaches)
			11.5.1.2 Dermaptera (Earwigs)
			11.5.1.3 Psocoptera (Booklice, Psocids)
			11.5.1.4 Phthiraptera (Parasitic Lice)
			11.5.1.5 Hemiptera (Bugs)
			11.5.1.6 Coleoptera (Beetles)
			11.5.1.7 Siphonaptera (Fleas)
			11.5.1.8 Diptera (Flies)
			11.5.1.9 Lepidoptera (Moths)
			11.5.1.10 Hymenoptera (Wasps)
		11.5.2 Parasitoid Taxa
			11.5.2.1 Neuroptera (Mantidflies)
			11.5.2.2 Coleoptera (Beetles)
			11.5.2.3 Strepsiptera (Twisted-Wing Parasites)
			11.5.2.4 Diptera (Flies)
			11.5.2.5 Trichoptera (Caddisflies)
			11.5.2.6 Lepidoptera (Moths)
			11.5.2.7 Hymenoptera (Wasps)
	11.6 Modern Food Webs and the Mid-Mesozoic Parasitoid Revolution (MMPR)
		11.6.1 Ovipositors and Host-Seeking First Instar Larvae: Vetting the Parasitoid Taxa
		11.6.2 The Trophic Cascade and Resource Concentration Hypotheses of Food Webs
		11.6.3 The Importance of Parasites and Parasitoids in Food Webs
		11.6.4 Top–Down Control of Food Webs by Parasitoids in Modern Ecosystems
		11.6.5 Insect Faunas Before the Mid-Mesozoic Parasitoid Revolution
		11.6.6 Insect Faunas During the Mid-Mesozoic Parasitoid Revolution
		11.6.7 Insect Faunas After the Mid-Mesozoic Parasitoid Revolution
	11.7 Parasitoid Clade Diversification in the Early Mid-Mesozoic Parasitoid Revolution
		11.7.1 Stephanoidea (Stephanid Wasps)
		11.7.2 Evanioidea (Ensign Wasps)
	11.8 Discussion: Trophic Specialization and the Mid Mesozoic Parasitoid Diversification
	11.9 Summary and Conclusions
	11.10 An Outlook Toward the Future
	References
Index