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
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’................................................. 482 20.3.7.4 Some Small Euthyneurans Previously Thought to Be Lower Heterobranchs............. 485 20.3.7.5 Euthyneuran Reproductive Systems............................................................................ 486 20.4 Biology, Ecology, and Behaviour.............................................................................................................. 503 20.4.1 Habits and Habitats...................................................................................................................... 503 20.4.2 Locomotion.................................................................................................................................. 503 20.4.3 Food and Feeding......................................................................................................................... 504 20.4.3.1 Herbivory..................................................................................................................... 505 20.4.3.2 Fungivory.................................................................................................................... 505 20.4.3.3 Active Carnivory.......................................................................................................... 505 20.4.3.4 Parasitism.................................................................................................................... 507 20.4.3.5 Suspension Feeding..................................................................................................... 507 20.4.3.6 Coprophagy................................................................................................................. 507 20.4.4 Reproduction, Development, and Growth.................................................................................... 507 20.4.4.1 Self-Fertilisation.......................................................................................................... 508 20.4.4.2 Hypodermic Insemination........................................................................................... 508 20.4.4.3 Aphally........................................................................................................................ 508 20.4.4.4 Asexual Reproduction................................................................................................. 509 20.4.4.5 Life Cycles.................................................................................................................. 509 20.4.4.6 Reproductive Behaviour.............................................................................................. 509 20.4.4.7 Development................................................................................................................ 509 20.4.4.8 Spawn...........................................................................................................................510 20.4.4.9 Larvae and Larval Shells..............................................................................................514 20.4.5 Behaviour..................................................................................................................................... 515 20.4.6 Predation and Defence................................................................................................................. 515 20.4.6.1 The Shell as a Means of Defence................................................................................ 515 20.4.6.2 Mucus...........................................................................................................................516 20.4.6.3 Defence without a Shell...............................................................................................516 20.4.6.4 Use of Nematocysts......................................................................................................517 20.4.6.5 Aposematic (Warning) Colouration and Batesian and Müllerian Mimicry.................518 20.4.6.6 Autotomy......................................................................................................................519 20.4.6.7 Predators of Heterobranchs..........................................................................................519 20.4.7 Associations................................................................................................................................. 520 20.4.8 Economic and Ecological Values................................................................................................. 520 20.5 Diversity and Fossil History...................................................................................................................... 520 20.5.1 Major Groups within the Heterobranchia.................................................................................... 521 20.5.1.1 ‘Lower Heterobranchs’................................................................................................ 521 20.5.1.2 Euthyneura.................................................................................................................. 521 Chapter 21 Molluscan Research – Present and Future Directions......................................................................................... 527 21.1 Molluscan Research Publications.............................................................................................................. 527 21.1.1 Provision of Basic Information.................................................................................................... 527 21.1.1.1 Use of Museum Collections........................................................................................ 529 21.1.1.2 Research Workshops................................................................................................... 529 21.1.1.3 DNA Barcoding........................................................................................................... 529 21.1.1.4 Genetics....................................................................................................................... 529 21.2 Palaeobiology............................................................................................................................................ 530 21.3 Taphonomy.................................................................................................................................................531 21.4 Systematics................................................................................................................................................ 532 21.5 Systematic, Phylogenetic, and Evolutionary Studies................................................................................ 532 21.5.1 Different Datasets......................................................................................................................... 532 21.5.1.1 Molecular Datasets...................................................................................................... 533 21.6 Biogeography............................................................................................................................................ 535 21.7 Discovery................................................................................................................................................... 535 21.8 The Future................................................................................................................................................. 535 21.8.1 Text Mining.................................................................................................................................. 536 21.8.2 New Technology........................................................................................................................... 536 21.8.3 Phylogenetic and Evolutionary Research..................................................................................... 536 21.8.4 Biodiversity.................................................................................................................................. 538 21.8.5 Citizen Science............................................................................................................................. 540 Appendix Higher Classification of the Extant Classes of Mollusca..................................................................................... 543 References................................................................................................................................................................................. 725 Index...........................................................................................................................................................................................817
Winston Frank Ponder, David R. Lindberg, Juliet Mary Ponder
Reviews for Biology and Evolution of the Mollusca, Volume 2
... 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
