Biology and Evolution of the Mollusca, Volume 2

Biology and Evolution of the Mollusca, Volume 2

 
Edition number and title: Volume 2
Edition number: 1
Publisher: CRC Press
Date of Publication:
 
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Product details:

ISBN13:9780815361848
ISBN10:081536184X
Binding:Hardback
No. of pages:890 pages
Size:280x216 mm
Weight:2599 g
Language:English
Illustrations: 993 Illustrations, black & white; 250 Illustrations, color; 110 Tables, black & white
201
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Short description:

This text will provide the first general account of molluscs in decades and will include hundreds of colour figures.

Long description:

This volume provides individual treatments of the major molluscan taxa. Each chapter provides an overview of the evolution, phylogeny and classification of a group of molluscs, as well as more specific and detailed coverage of their biology (reproduction, feeding and digestion, excretion, respiration etc.), their long fossil record and aspects of their natural history. The book is illustrated with hundreds of colour figures. In both volumes, concepts are summarised in colour-coded illustrations. 


Key selling features:



  • Comprehensively reviews molluscan biology and evolutionary history

  • Includes a description the anatomy and physiology of anatomical systems

  • Up to date treatment with a comprehensive bibliography

  • Reviews the phylogenetic history of the major molluscan lineages


"? an immensely useful 'go-to resource' ? on almost any mollusk-related topic. ? This is a brilliant work that should be on the desk of any serious 'student' of mollusks, at any career level." - American Malacological Bulletin, 38 (1) 2020.



"The two-volume set is a treasure trove for anyone lucky enough to be teaching malacology, a reference work for investigators in a multitude of fields, and a true companion for students and advanced researchers.? It will stand for decades, joining immediately the list of classic volumes covering the Mollusca.? A tour d?force that speaks to the knowledge, dedication, and talent of the authors, illustrator, and also the publisher that had the vision to make such an epic possible in today?s publishing world." - Sandra E. Shumway in National Shellfisheries Association Quarterly Newsletter 2020 (2).


"The scholarship and deep thought that the authors bring to this monumental work are beyond staggering... It is unlikely that the scientific achievement manifested in the publication of this work will be surpassed in the foreseeable future. Whichever field of molluscan study a student or seasoned researcher is contemplating, this publication is the resource to consult first for an authoritative synthesis. Anyone with an interest in living and extinct molluscs will want to own and consult this landmark set of volumes." - Geerat J. Vermeij in The Quarterly Review of Biology

"In case the rest of my review is not emphatic enough, Biology and Evolution of the Mollusca (hereafter BEM) is the most wonderful molluscan work I have ever seen. It is an intellectual tour de force, truly astounding in its breadth and depth of subject coverage; the sampling of the literature is comprehensive and up to date; and the figures are works of art while simultaneously being ? like the text ? exceptionally clear and informative. The authors ? writers Winston F. Ponder and David R. Lindberg, together with illustrator Juliet M. Ponder ? reviewers and publisher are all to be congratulated on such an outstanding result. ? [BEM] will be the standard reference work about the Mollusca for years to come. It undoubtedly will, but I changed my mind; it can do better! Instead, I would like to think that BEM will stimulate such a flowering of molluscan research that it will soon be seen as a more foundational document." - Hamish G. Spencer in Molluscan Research


Table of Contents:

Contents


About the Authors......................................................................................................................................................................xvii


Introduction to Volume Two.......................................................................................................................................................xix


Chapter 12 Molluscan Relationships.......................................................................................................................................... 1


12.1 The Hypothetical Ancestral Mollusc............................................................................................................. 1


12.2 Hypotheses of Phylogenetic Relationships of the Major Molluscan Groups................................................ 1


12.2.1 Aculifera ? Conchifera, Testaria, and Serialia Hypotheses............................................................. 4


12.2.2 The Search for the Molluscan Sister Group..................................................................................... 6


12.3 Repetition of Organs..................................................................................................................................... 8


12.4 Ano-Pedal Flexure and a Change in Body Orientation................................................................................. 9


12.5 Reduction and Paedomorphosis.................................................................................................................... 9


Chapter 13 Early History and Extinct Groups.......................................................................................................................... 11


13.1 Introduction................................................................................................................................................. 11


13.1.1 The First Molluscs?........................................................................................................................ 12


13.1.2 Shell Morphology........................................................................................................................... 13


13.1.2.1 Beyond Shell Morphology............................................................................................ 14


13.1.2.2 Muscle Scars................................................................................................................. 16


13.1.3 Phylogenetics and Fossils............................................................................................................... 18


13.2 Originations, Extinctions, and Recoveries.................................................................................................. 18


13.3 The Molluscan Fossil Record...................................................................................................................... 20


13.3.1 Ediacaran........................................................................................................................................ 20


13.3.2 Cambrian........................................................................................................................................ 22


13.3.2.1 Biomineralisation.......................................................................................................... 24


13.3.2.2 Taxa............................................................................................................................... 25


13.3.3 Ordovician...................................................................................................................................... 51


13.3.3.1 Taxa............................................................................................................................... 52


13.3.4 Silurian........................................................................................................................................... 55


13.3.4.1 Taxa............................................................................................................................... 55


13.3.5 Devonian........................................................................................................................................ 59


13.3.5.1 Taxa............................................................................................................................... 59


13.3.6 Carboniferous................................................................................................................................. 61


13.3.6.1 Taxa............................................................................................................................... 62


13.3.7 Permian.......................................................................................................................................... 64


13.3.7.1 Extinction...................................................................................................................... 64


13.3.7.2 Taxa............................................................................................................................... 65


Chapter 14 Polyplacophora, Monoplacophora, and Aplacophorans......................................................................................... 67


14.1 Introduction................................................................................................................................................. 67


14.2 Polyplacophora (Chitons, Placophora, Loricata, Amphineura in Part)....................................................... 67


14.2.1 Introduction.................................................................................................................................... 67


14.2.2 Phylogeny and Classification.......................................................................................................... 67


14.2.2.1 Sister Group Relationships............................................................................................ 69


14.2.3 Morphology.................................................................................................................................... 69


14.2.3.1 Shell and Girdle............................................................................................................. 70


14.2.3.2 Head-Foot, Muscles, and Mantle.................................................................................. 70


14.2.3.3 Mantle Groove (Mantle Cavity).................................................................................... 72


14.2.3.4 Digestive System........................................................................................................... 72


14.2.3.5 Renopericardial System................................................................................................. 75


14.2.3.6 Reproductive System..................................................................................................... 76


14.2.3.7 Nervous System and Sense Organs............................................................................... 78


14.2.3.8 Chromosomes................................................................................................................ 82


14.2.4 Biology, Ecology, and Behaviour................................................................................................... 82


14.2.4.1 Habits and Habitats....................................................................................................... 82


14.2.4.2 Locomotion................................................................................................................... 82


14.2.4.3 Feeding and Diet........................................................................................................... 82


14.2.4.4 Reproduction, Development, and Growth..................................................................... 83


14.2.4.5 Associations................................................................................................................... 84


14.2.4.6 Predators, Defence, and Behaviour............................................................................... 84


14.2.5 Diversity and Fossil History........................................................................................................... 84


14.3 The Monoplacophora (= Tryblidia, Neopilinida, Tergomya in part)........................................................... 84


14.3.1 Introduction.................................................................................................................................... 84


14.3.2 Phylogeny and Classification.......................................................................................................... 84


14.3.2.1 Sister Group Relationships............................................................................................ 85


14.3.3 Morphology.................................................................................................................................... 86


14.3.3.1 Shell and Protoconch..................................................................................................... 86


14.3.3.2 Head-Foot and Mantle................................................................................................... 87


14.3.3.3 Mantle Groove (Mantle Cavity).................................................................................... 88


14.3.3.4 Digestive System........................................................................................................... 89


14.3.3.5 Renopericardial and Reproductive Systems.................................................................. 90


14.3.3.6 Nervous System and Sense Organs............................................................................... 90


14.3.4 Biology, Ecology, and Behaviour................................................................................................... 91


14.3.4.1 Habits and Habitats....................................................................................................... 91


14.3.4.2 Feeding and Diet........................................................................................................... 92


14.3.4.3 Reproduction, Development, and Growth..................................................................... 92


14.3.4.4 Associations................................................................................................................... 92


14.3.4.5 Predation, Defence, and Other Behavioural Attributes................................................. 93


14.3.5 Diversity and Fossil History........................................................................................................... 93


14.4 Caudofoveata and Solenogastres: The Aplacophoran Groups.................................................................... 93


14.4.1 Introduction.................................................................................................................................... 93


14.4.2 Phylogeny and Classification.......................................................................................................... 94


14.4.3 Morphology.................................................................................................................................... 96


14.4.3.1 Head-Foot and Mantle................................................................................................... 96


14.4.3.2 Digestive System........................................................................................................... 97


14.4.3.3 Renopericardial and Reproductive Systems.................................................................. 97


14.4.3.4 Nervous System............................................................................................................. 97


14.4.3.5 Development.................................................................................................................. 98


14.4.3.6 Habits and Habitats....................................................................................................... 98


14.5 Caudofoveata (or Chaetodermomorpha, Scutopoda).................................................................................. 99


14.5.1 Introduction.................................................................................................................................... 99


14.5.2 Phylogeny and Classification.......................................................................................................... 99


14.5.2.1 Sister Group Relationships............................................................................................ 99


14.5.3 Morphology.................................................................................................................................... 99


14.5.3.1 Spicules......................................................................................................................... 99


14.5.3.2 Head-Foot and Mantle................................................................................................... 99


14.5.3.3 Mantle Cavity................................................................................................................ 99


14.5.3.4 Digestive System......................................................................................................... 100


14.5.3.5 Renopericardial and Reproductive Systems.................................................................101


14.5.3.6 Nervous System and Sense Organs..............................................................................101


14.5.4 Biology, Ecology, and Behaviour..................................................................................................101


14.5.4.1 Habits and Habitats......................................................................................................101


14.5.4.2 Feeding and Diet..........................................................................................................101


14.5.4.3 Reproduction, Development, and Growth................................................................... 102


14.5.4.4 Associations................................................................................................................. 102


14.5.5 Diversity and Fossil History......................................................................................................... 102


14.6 Solenogastres (or Neomeniomorpha)........................................................................................................ 102


14.6.1 Introduction.................................................................................................................................. 102


14.6.2 Phylogeny and Classification........................................................................................................ 102


14.6.2.1 Sister Group Relationships.......................................................................................... 102


14.6.3 Morphology.................................................................................................................................. 102


14.6.3.1 Spicules....................................................................................................................... 102


14.6.3.2 Body............................................................................................................................ 102


14.6.3.3 Mantle Cavity.............................................................................................................. 103


14.6.3.4 Digestive System......................................................................................................... 103


14.6.3.5 Renopericardial and Reproductive Systems................................................................ 104


14.6.3.6 Nervous System and Sense Organs............................................................................. 105


14.6.4 Biology and Ecology.................................................................................................................... 105


14.6.4.1 Habits and Habitats..................................................................................................... 105


14.6.4.2 Feeding and Diet......................................................................................................... 105


14.6.4.3 Reproduction, Development, and Growth................................................................... 105


14.6.4.4 Associations................................................................................................................. 107


14.6.5 Diversity and Fossil History......................................................................................................... 107


Chapter 15 The Bivalvia......................................................................................................................................................... 109


15.1 Introduction............................................................................................................................................... 109


15.2 Classification............................................................................................................................................. 109


15.3 Morphology................................................................................................................................................112


15.3.1 The Shell.......................................................................................................................................113


15.3.1.1 Orientation....................................................................................................................113


15.3.1.2 Sculpture.......................................................................................................................113


15.3.1.3 Shell Structure..............................................................................................................113


15.3.1.4 Periostracum.................................................................................................................115


15.3.1.5 Ligament.......................................................................................................................117


15.3.1.6 Hinge............................................................................................................................119


15.3.1.7 Pallial Line and Sinus.................................................................................................. 120


15.3.1.8 Shell Reduction, Accessory Shell Plates, and Secondary Shells................................. 120


15.3.2 Muscles and Their Scars.............................................................................................................. 124


15.3.3 The Foot....................................................................................................................................... 124


15.3.4 The Byssus ? Structure and Function.......................................................................................... 125


15.3.5 Mantle and Siphons...................................................................................................................... 127


15.3.6 Mantle Cavity............................................................................................................................... 132


15.3.6.1 The Ctenidia ? Their Structure and Ciliation.............................................................. 132


15.3.6.2 The Labial Palps.......................................................................................................... 139


15.3.6.3 Other Features of the Mantle Cavity............................................................................141


15.3.7 Digestive System...........................................................................................................................141


15.3.8 Renopericardial System................................................................................................................143


15.3.9 Reproductive System.....................................................................................................................143


15.3.10 Nervous System and Sense Organs.............................................................................................. 145


15.4 Biology and Ecology................................................................................................................................. 145


15.4.1 Habits and Habitats...................................................................................................................... 145


15.4.1.1 Locomotion................................................................................................................. 145


15.4.1.2 Attachment.................................................................................................................. 146


15.4.1.3 Secondary Soft-Bottom Dwellers.................................................................................147


15.4.1.4 Burrowers and Borers.................................................................................................. 149


15.4.2 Feeding......................................................................................................................................... 151


15.4.3 Life History and Growth.............................................................................................................. 152


15.4.4 Associations................................................................................................................................. 156


15.4.4.1 Bacterial and Algal Symbionts.................................................................................... 156


15.4.5 Relationships with Other Animals............................................................................................... 158


15.4.5.1 Predators of Bivalves................................................................................................... 159


15.5 Economic Values....................................................................................................................................... 159


15.6 Diversity and Fossil History...................................................................................................................... 159


15.6.1 The First Bivalves......................................................................................................................... 159


15.6.2 The Evolutionary Radiation of Bivalves...................................................................................... 159


15.6.2.1 Freshwater Bivalves.....................................................................................................161


15.6.3 Patterns of Bivalve Diversity.........................................................................................................161


15.7 The Protobranch Bivalves......................................................................................................................... 162


15.7.1 Phylogeny and Classification........................................................................................................ 162


15.7.2 Biology and Ecology.................................................................................................................... 163


15.7.3 Diversity and Fossil History......................................................................................................... 164


15.8 The Autobranch Bivalves.......................................................................................................................... 164


15.8.1 Pteriomorphia............................................................................................................................... 164


15.8.1.1 Phylogeny and Classification...................................................................................... 164


15.8.1.2 Biology and Ecology................................................................................................... 165


15.8.1.3 Diversity and Fossil History........................................................................................ 167


15.8.2 Heteroconchia............................................................................................................................... 167


15.8.3 Palaeoheterodonta........................................................................................................................ 168


15.8.3.1 Trigoniida.................................................................................................................... 168


15.8.3.2 Unionida...................................................................................................................... 168


15.8.4 Heterodonta...................................................................................................................................170


15.8.4.1 Archiheterodonta......................................................................................................... 172


15.8.5 Euheterodonta.............................................................................................................................. 172


15.8.5.1 Phylogeny and Classification.......................................................................................173


15.8.5.2 Biology and Ecology....................................................................................................174


15.8.5.3 Diversity and Fossil History.........................................................................................175


15.8.5.4 The Rudists...................................................................................................................175


15.8.6 Anomalodesmata (= Poromyata)...................................................................................................178


Chapter 16 The Scaphopoda................................................................................................................................................... 183


16.1 Introduction............................................................................................................................................... 183


16.2 Phylogeny and Classification.................................................................................................................... 183


16.2.1 Sister Group Relationships.......................................................................................................... 183


16.3 Morphology............................................................................................................................................... 184


16.3.1 Shell............................................................................................................................................. 184


16.3.2 Head-Foot, Captacula, and Mantle.............................................................................................. 185


16.3.3 Mantle Cavity.............................................................................................................................. 186


16.3.4 Digestive System......................................................................................................................... 188


16.3.5 Renopericardial System............................................................................................................... 189


16.3.6 Reproductive System................................................................................................................... 189


16.3.7 Nervous System and Sense Organs............................................................................................. 189


16.4 Biology and Ecology................................................................................................................................. 190


16.4.1 Habits and Habitats...................................................................................................................... 190


16.4.2 Locomotion.................................................................................................................................. 190


16.4.3 Feeding and Diet.......................................................................................................................... 190


16.4.4 Reproduction, Development, and Growth................................................................................... 192


16.4.5 Associations................................................................................................................................. 192


16.4.6 Predation, Defence, and Other Behavioural Attributes............................................................... 193


16.5 Diversity and Fossil History...................................................................................................................... 193


16.5.1 The Major Groups of Scaphopods............................................................................................... 193


16.6 Human Uses.............................................................................................................................................. 193


Chapter 17 The Cephalopoda................................................................................................................................................. 195


17.1 Introduction............................................................................................................................................... 195


17.2 Phylogeny and Classification....................................................................................................................200


17.3 Morphology............................................................................................................................................... 202


17.3.1 Shell.............................................................................................................................................. 202


17.3.2 Head, Arms, and Funnel.............................................................................................................. 205


17.3.2.1 Embryological Evidence for the Derivation of Head Structures................................. 208


17.3.3 Epidermis (Skin)........................................................................................................................... 208


17.3.4 Head and Funnel Retractor Muscles............................................................................................ 209


17.3.4.1 Muscular Attachments, Shell Growth and the Addition of Septa.................................210


17.3.5 The Fossil Record of Soft Tissues................................................................................................ 212


17.3.6 Mantle Wall.................................................................................................................................. 213


17.3.7 Mantle Cavity................................................................................................................................214


17.4 Biology and Ecology..................................................................................................................................218


17.4.1 Habits and Habitats.......................................................................................................................218


17.4.1.1 The Habits and Habitats of Extinct Ectocochleate Cephalopods.................................218


17.4.2 Locomotion and Buoyancy.......................................................................................................... 220


17.4.2.1 Locomotion................................................................................................................. 220


17.4.2.2 Buoyancy..................................................................................................................... 222


17.4.2.3 The Shell as a Buoyancy Device................................................................................. 222


17.4.2.4 Other Buoyancy Strategies.......................................................................................... 227


17.4.3 The Ink Sac and Inking................................................................................................................ 228


17.4.4 Colour Changes ? Chromatophores and Other Special Skin Structures..................................... 228


17.4.4.1 Chromatophores.......................................................................................................... 229


17.4.4.2 Structural Reflectors (Iridophores, Silvery Reflectors, and Leucophores).................. 229


17.4.5 Luminescent Organs.................................................................................................................... 231


17.5 Digestive System....................................................................................................................................... 233


17.5.1 The Jaws....................................................................................................................................... 233


17.5.2 The Radula................................................................................................................................... 233


17.5.3 Feeding......................................................................................................................................... 233


17.6 Renopericardial System............................................................................................................................. 236


17.7 Courtship, Reproduction, Development, and Growth............................................................................... 239


17.7.1 Reproductive System.................................................................................................................... 239


17.7.1.1 Sexual Characters and Sexual Dimorphism................................................................ 242


17.7.2 Courtship and Mating.................................................................................................................. 243


17.7.3 Life History..................................................................................................................................244


17.7.3.1 Spawn and Parental Care............................................................................................. 245


17.7.3.2 Development................................................................................................................ 245


17.7.4 The Nervous System, Sense Organs, and Behaviour................................................................... 247


17.7.5 The Endocrine System................................................................................................................. 249


17.7.6 Learning and Intelligence............................................................................................................ 249


17.7.7 Behaviour..................................................................................................................................... 249


17.7.8 Symbiotic Relationships............................................................................................................... 250


17.7.9 Predators....................................................................................................................................... 251


17.8 Economic Values....................................................................................................................................... 251


17.9 The Origin of Cephalopods....................................................................................................................... 251


17.10 Diversity and Fossil History...................................................................................................................... 256


17.10.1 The Ordovician Radiation and Beyond........................................................................................ 256


17.11 The Major Groups of Cephalopoda........................................................................................................... 258


17.11.1 The Nautiliforms.......................................................................................................................... 258


17.11.1.1 Ellesmeroceratia and Plectronoceratia...................................................................... 259


17.11.1.2 Orthoceratia............................................................................................................... 260


17.11.1.3 Discosorida................................................................................................................ 261


17.11.1.4 Endoceratia................................................................................................................ 261


17.11.1.5 Actinoceratia.............................................................................................................. 261


17.11.1.6 Nautilia...................................................................................................................... 262


17.12 Bactrita and Ammonitia (= Ammonoidea)................................................................................................ 262


17.12.1 Bactrita (= Bactritoidea)............................................................................................................... 262


17.12.2 Ammonitia (Ammonites)............................................................................................................. 264


17.12.2.1 The Embryonic Shell (Ammonitella)........................................................................ 267


17.12.2.2 Ammonite Sutural Patterns........................................................................................ 267


17.12.2.3 Life Habits................................................................................................................. 268


17.12.2.4 Aptychus and Jaws.................................................................................................... 269


17.12.2.5 Phylogeny, Classification, Diversity, and Fossil History of Ammonites................... 271


17.12.2.6 The Major Groups of Ammonites............................................................................. 272


17.13 Coleoida.................................................................................................................................................. 273


17.13.1 Phylogeny, Classification, Diversity, and Fossil History............................................................ 275


17.13.1.1 Belemnitidia (The Belemnites)................................................................................ 279


17.13.1.2 Decabrachia............................................................................................................. 279


17.13.1.3 Squid ? A Grade Not a Clade ? The Myopsida, Oegopsida, Bathyteuthoidea


(?Teuthida?) -......................................................................................................... 284


17.13.1.4 Octobrachia (= Octopodiformes)............................................................................. 285


17.13.1.5 Giant Squid and Octopuses...................................................................................... 287


Chapter 18 Gastropoda I ? Introduction and the Stem Groups.............................................................................................. 289


18.1 Introduction to the Gastropoda.................................................................................................................. 289


18.1.1 Phylogeny and Classification........................................................................................................ 289


18.1.2 Morphology.................................................................................................................................. 290


18.1.2.1 Shell, Operculum and External Body.......................................................................... 293


18.1.2.2 Shell Muscles.............................................................................................................. 294


18.1.2.3 Operculum................................................................................................................... 294


18.1.2.4 Head-Foot.................................................................................................................... 294


18.1.2.5 Mantle Cavity and Its Organs...................................................................................... 296


18.1.2.6 Digestive System......................................................................................................... 296


18.1.2.7 Renopericardial System............................................................................................... 297


18.1.2.8 Reproductive System...................................................................................................300


18.1.2.9 Nervous System and Sense Organs.............................................................................300


18.1.3 Biology, Ecology, and Behaviour.................................................................................................300


18.1.3.1 Habits and Habitats.....................................................................................................300


18.1.3.2 Locomotion and Behaviour......................................................................................... 301


18.1.3.3 Life History................................................................................................................. 301


18.1.3.4 Feeding........................................................................................................................ 301


18.1.3.5 Associations................................................................................................................. 302


18.1.4 Economic Values.......................................................................................................................... 303


18.1.5 Diversity and Fossil History......................................................................................................... 303


18.2 The Initial Radiation ? Eogastropods and Orthogastropods ............................................................  303


18.2.1 Hypothetical Reconstructions...................................................................................................... 303


18.2.1.1 A Hypothetical Ancestral Gastropod........................................................................... 307


18.2.1.2 Eogastropoda............................................................................................................... 308


18.2.1.3 Orthogastropoda...........................................................................................................310


18.3 The Lower Groups of Gastropods..............................................................................................................312


18.4 Patellogastropoda (= Docoglossa, Cyclobranchia, Eogastropoda in Part).................................................312


18.4.1 Introduction...................................................................................................................................312


18.4.2 Phylogeny and Classification.........................................................................................................312


18.4.2.1 Sister Group Relationships...........................................................................................313


18.4.2.2 Outline of Classification...............................................................................................313


18.4.3 Morphology...................................................................................................................................313


18.4.3.1 Shell and Protoconch....................................................................................................313


18.4.3.2 Head-Foot, Mantle, and Operculum.............................................................................315


18.4.3.3 Mantle Cavity...............................................................................................................315


18.4.3.4 Digestive System..........................................................................................................317


18.4.3.5 Reproductive System................................................................................................... 320


18.4.3.6 Renopericardial System............................................................................................... 320


18.4.3.7 Nervous System and Sense Organs............................................................................. 321


18.4.4 Biology, Ecology, and Behaviour................................................................................................. 322


18.4.4.1 Habits and Habitats..................................................................................................... 322


18.4.4.2 Locomotion and Behaviour......................................................................................... 322


18.4.4.3 Life History................................................................................................................. 322


18.4.4.4 Feeding and Diet......................................................................................................... 322


18.4.4.5 Associations................................................................................................................. 323


18.4.5 Diversity and Fossil History......................................................................................................... 323


18.5 Vetigastropoda........................................................................................................................................... 324


18.5.1 Introduction.................................................................................................................................. 324


18.5.2 Phylogeny and Classification........................................................................................................ 324


18.5.2.1 Outline of Classification.............................................................................................. 326


18.5.2.2 Deep-Sea Enigmas; The Neomphaloidean and Cocculinoidean Taxa........................ 332


18.5.3 Morphology.................................................................................................................................. 333


18.5.3.1 Shell and Protoconch................................................................................................... 333


18.5.3.2 Head-Foot, Mantle, Operculum, and Locomotion...................................................... 337


18.5.3.3 Mantle Cavity.............................................................................................................. 338


18.5.3.4 Digestive System......................................................................................................... 338


18.5.3.5 Reproductive System................................................................................................... 343


18.5.3.6 Renopericardial System............................................................................................... 344


18.5.3.7 Nervous System and Sense Organs............................................................................. 344


18.5.4 Biology, Ecology, and Behaviour................................................................................................. 345


18.5.4.1 Habits and Habitats..................................................................................................... 345


18.5.4.2 Feeding and Diet......................................................................................................... 345


18.5.4.3 Life History................................................................................................................. 346


18.5.4.4 Associations................................................................................................................. 346


18.5.4.5 Behaviour.................................................................................................................... 346


18.5.5 Diversity and Fossil History......................................................................................................... 346


18.6 Neritimorpha (= Neritopsina).................................................................................................................... 347


18.6.1 Introduction.................................................................................................................................. 347


18.6.2 Phylogeny and Classification........................................................................................................ 347


18.6.2.1 Sister Group Relationships.......................................................................................... 349


18.6.2.2 Outline of Classification.............................................................................................. 349


18.6.3 Morphology.................................................................................................................................. 350


18.6.3.1 Shell and Protoconch................................................................................................... 350


18.6.3.2 Head-Foot, Mantle, Operculum, and Locomotion.......................................................351


18.6.3.3 Mantle Cavity.............................................................................................................. 353


18.6.3.4 Digestive System......................................................................................................... 354


18.6.3.5 Renopericardial System............................................................................................... 355


18.6.3.6 Reproductive System................................................................................................... 355


18.6.3.7 Nervous System and Sense Organs............................................................................. 360


18.6.4 Biology and Ecology.................................................................................................................... 360


18.6.4.1 Habits and Habitats..................................................................................................... 360


18.6.4.2 Feeding and Diet......................................................................................................... 361


18.6.4.3 Life History................................................................................................................. 361


18.6.4.4 Associations................................................................................................................. 362


18.6.4.5 Behaviour.................................................................................................................... 362


18.6.5 Diversity and Fossil History......................................................................................................... 363


Chapter 19 Gastropoda II ? The Caenogastropoda................................................................................................................ 365


19.1 Introduction............................................................................................................................................... 365


19.2 Phylogeny and Classification.................................................................................................................... 365


19.2.1 Sister Group Relationships........................................................................................................... 365


19.2.2 Outline of Classification............................................................................................................... 365


19.3 Morphology............................................................................................................................................... 366


19.3.1 Shell and Protoconch.................................................................................................................... 377


19.3.2 Head-Foot, Mantle, and Operculum............................................................................................ 379


19.3.2.1 Head Structures........................................................................................................... 379


19.3.2.2 The Foot...................................................................................................................... 379


19.3.2.3 The Operculum............................................................................................................ 380


19.3.2.4 Mantle Edge................................................................................................................ 380


19.3.3 Mantle Cavity............................................................................................................................... 380


19.3.4 Digestive System.......................................................................................................................... 381


19.3.5 Renopericardial System............................................................................................................... 394


19.3.6 Reproductive System.................................................................................................................... 394


19.3.6.1 The Female System..................................................................................................... 395


19.3.6.2 The Male System......................................................................................................... 396


19.3.7 Nervous System and Sense Organs..............................................................................................400


19.4 Biology, Ecology, and Behaviour.............................................................................................................. 403


19.4.1 Habits and Habitats...................................................................................................................... 403


19.4.2 Locomotion.................................................................................................................................. 403


19.4.3 Feeding and Diet..........................................................................................................................404


19.4.4 Life History and Growth.............................................................................................................. 405


19.4.4.1 Spawn, Development, and Larvae............................................................................... 405


19.4.5 Associations..................................................................................................................................410


19.4.5.1 The Evolution of Parasitism in Eulimidae....................................................................410


19.4.6 Predators and Defence..................................................................................................................412


19.4.7 Other Behavioural Attributes........................................................................................................412


19.5 Economic and Ecological Values...............................................................................................................412


19.6 Diversity and Fossil History.......................................................................................................................413


19.7 Major Groups within Caenogastropoda.....................................................................................................414


19.7.1 The Architaenioglossans...............................................................................................................414


19.7.1.1 Campanilida.................................................................................................................415


19.7.2 Sorbeoconcha................................................................................................................................415


19.7.2.1 Cerithiimorpha.............................................................................................................415


19.7.2.2 Hypsogastropoda..........................................................................................................416


Chapter 20 Gastropoda III ? The Heterobranchia...................................................................................................................419


20.1 Introduction................................................................................................................................................419


20.2 Phylogeny and Classification.....................................................................................................................419


20.2.1 Sister Group Relationships........................................................................................................... 421


20.2.2 Chromosome Numbers................................................................................................................. 422


20.3 Morphology............................................................................................................................................... 429


20.3.1 Shell and Operculum.................................................................................................................... 435


20.3.1.1 Shell Structure............................................................................................................. 439


20.3.1.2 Larval or Embryonic Shell.......................................................................................... 439


20.3.1.3 Shell Reduction and Loss............................................................................................ 439


20.3.1.4 Shell Variation.............................................................................................................440


20.3.1.5 Operculum...................................................................................................................440


20.3.2 External Body Morphology.......................................................................................................... 441


20.3.2.1 Head Tentacles............................................................................................................ 441


20.3.2.2 Cephalic (Head) Shield............................................................................................... 442


20.3.2.3 Epithelial Modifications.............................................................................................. 442


20.3.2.4 Oral Veil.......................................................................................................................444


20.3.2.5 The Foot, Pedal Glands, and Parapodia.......................................................................444


20.3.2.6 The Mantle.................................................................................................................. 445


20.3.3 Mantle Cavity and Associated Structures.................................................................................... 447


20.3.3.1 Heterobranch Gills...................................................................................................... 450


20.3.3.2 Opposed Ciliary Ridges (Raphes) and Mantle Tentacles............................................ 451


20.3.3.3 The Hypobranchial Gland........................................................................................... 451


20.3.3.4 The Pigmented Mantle Organ (PMO)......................................................................... 452


20.3.3.5 Other Glands Associated with the Mantle Cavity....................................................... 452


20.3.4 Digestive System.......................................................................................................................... 452


20.3.5 The Renopericardial System........................................................................................................ 472


20.3.6 Nervous System and Sense Organs.............................................................................................. 474


20.3.6.1 Sense Organs............................................................................................................... 477


20.3.7 Reproductive System.................................................................................................................... 479


20.3.7.1 Terminology................................................................................................................ 479


20.3.7.2 The Ovotestis and Gametes......................................................................................... 481


20.3.7.3 The Reproductive System of ?Lower Heterobranchs?.