Description
Book SynopsisThis 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
Trade Review"… 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 ContentsContents
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
