Biology and Evolution of the Mollusca, Volume 2

著者名：Ponder, Winston Frank/Lindberg, David R./Ponder, Juliet Mary

CRC Press（2020/02/14発売）

• 言語：ENG
• ISBN：9781032173542
• eISBN：9781351115230

Description

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

Key selling features:

• Comprehensively reviews molluscan biology and evolutionary history
• Includes a description the anatomy and physiology of anatomical systems
• Up to date treatment with a comprehensive bibliography
• Reviews the phylogenetic history of the major molluscan lineages

Table of Contents

<p><strong>Contents</strong></p><p>About the Authors......................................................................................................................................................................xvii</p><p>Introduction to Volume Two.......................................................................................................................................................xix</p><p>Chapter 12 Molluscan Relationships.......................................................................................................................................... 1</p><p>12.1 The Hypothetical Ancestral Mollusc............................................................................................................. 1</p><p>12.2 Hypotheses of Phylogenetic Relationships of the Major Molluscan Groups................................................ 1</p><p>12.2.1 Aculifera – Conchifera, Testaria, and Serialia Hypotheses............................................................. 4</p><p>12.2.2 The Search for the Molluscan Sister Group..................................................................................... 6</p><p>12.3 Repetition of Organs..................................................................................................................................... 8</p><p>12.4 Ano-Pedal Flexure and a Change in Body Orientation................................................................................. 9</p><p>12.5 Reduction and Paedomorphosis.................................................................................................................... 9</p><p>Chapter 13 Early History and Extinct Groups.......................................................................................................................... 11</p><p>13.1 Introduction................................................................................................................................................. 11</p><p>13.1.1 The First Molluscs?........................................................................................................................ 12</p><p>13.1.2 Shell Morphology........................................................................................................................... 13</p><p>13.1.2.1 Beyond Shell Morphology............................................................................................ 14</p><p>13.1.2.2 Muscle Scars................................................................................................................. 16</p><p>13.1.3 Phylogenetics and Fossils............................................................................................................... 18</p><p>13.2 Originations, Extinctions, and Recoveries.................................................................................................. 18</p><p>13.3 The Molluscan Fossil Record...................................................................................................................... 20</p><p>13.3.1 Ediacaran........................................................................................................................................ 20</p><p>13.3.2 Cambrian........................................................................................................................................ 22</p><p>13.3.2.1 Biomineralisation.......................................................................................................... 24</p><p>13.3.2.2 Taxa............................................................................................................................... 25</p><p>13.3.3 Ordovician...................................................................................................................................... 51</p><p>13.3.3.1 Taxa............................................................................................................................... 52</p><p>13.3.4 Silurian........................................................................................................................................... 55</p><p>13.3.4.1 Taxa............................................................................................................................... 55</p><p>13.3.5 Devonian........................................................................................................................................ 59</p><p>13.3.5.1 Taxa............................................................................................................................... 59</p><p>13.3.6 Carboniferous................................................................................................................................. 61</p><p>13.3.6.1 Taxa............................................................................................................................... 62</p><p>13.3.7 Permian.......................................................................................................................................... 64</p><p>13.3.7.1 Extinction...................................................................................................................... 64</p><p>13.3.7.2 Taxa............................................................................................................................... 65</p><p>Chapter 14 Polyplacophora, Monoplacophora, and Aplacophorans......................................................................................... 67</p><p>14.1 Introduction................................................................................................................................................. 67</p><p>14.2 Polyplacophora (Chitons, Placophora, Loricata, Amphineura in Part)....................................................... 67</p><p>14.2.1 Introduction.................................................................................................................................... 67</p><p>14.2.2 Phylogeny and Classification.......................................................................................................... 67</p><p>14.2.2.1 Sister Group Relationships............................................................................................ 69</p><p>14.2.3 Morphology.................................................................................................................................... 69</p><p>14.2.3.1 Shell and Girdle............................................................................................................. 70</p><p>14.2.3.2 Head-Foot, Muscles, and Mantle.................................................................................. 70</p><p>14.2.3.3 Mantle Groove (Mantle Cavity).................................................................................... 72</p><p>14.2.3.4 Digestive System........................................................................................................... 72</p><p>14.2.3.5 Renopericardial System................................................................................................. 75</p><p>14.2.3.6 Reproductive System..................................................................................................... 76</p><p>14.2.3.7 Nervous System and Sense Organs............................................................................... 78</p><p>14.2.3.8 Chromosomes................................................................................................................ 82</p><p>14.2.4 Biology, Ecology, and Behaviour................................................................................................... 82</p><p>14.2.4.1 Habits and Habitats....................................................................................................... 82</p><p>14.2.4.2 Locomotion................................................................................................................... 82</p><p>14.2.4.3 Feeding and Diet........................................................................................................... 82</p><p>14.2.4.4 Reproduction, Development, and Growth..................................................................... 83</p><p>14.2.4.5 Associations................................................................................................................... 84</p><p>14.2.4.6 Predators, Defence, and Behaviour............................................................................... 84</p><p>14.2.5 Diversity and Fossil History........................................................................................................... 84</p><p>14.3 The Monoplacophora (= Tryblidia, Neopilinida, Tergomya in part)........................................................... 84</p><p>14.3.1 Introduction.................................................................................................................................... 84</p><p>14.3.2 Phylogeny and Classification.......................................................................................................... 84</p><p>14.3.2.1 Sister Group Relationships............................................................................................ 85</p><p>14.3.3 Morphology.................................................................................................................................... 86</p><p>14.3.3.1 Shell and Protoconch..................................................................................................... 86</p><p>14.3.3.2 Head-Foot and Mantle................................................................................................... 87</p><p>14.3.3.3 Mantle Groove (Mantle Cavity).................................................................................... 88</p><p>14.3.3.4 Digestive System........................................................................................................... 89</p><p>14.3.3.5 Renopericardial and Reproductive Systems.................................................................. 90</p><p>14.3.3.6 Nervous System and Sense Organs............................................................................... 90</p><p>14.3.4 Biology, Ecology, and Behaviour................................................................................................... 91</p><p>14.3.4.1 Habits and Habitats....................................................................................................... 91</p><p>14.3.4.2 Feeding and Diet........................................................................................................... 92</p><p>14.3.4.3 Reproduction, Development, and Growth..................................................................... 92</p><p>14.3.4.4 Associations................................................................................................................... 92</p><p>14.3.4.5 Predation, Defence, and Other Behavioural Attributes................................................. 93</p><p>14.3.5 Diversity and Fossil History........................................................................................................... 93</p><p>14.4 Caudofoveata and Solenogastres: The Aplacophoran Groups.................................................................... 93</p><p>14.4.1 Introduction.................................................................................................................................... 93</p><p>14.4.2 Phylogeny and Classification.......................................................................................................... 94</p><p>14.4.3 Morphology.................................................................................................................................... 96</p><p>14.4.3.1 Head-Foot and Mantle................................................................................................... 96</p><p>14.4.3.2 Digestive System........................................................................................................... 97</p><p>14.4.3.3 Renopericardial and Reproductive Systems.................................................................. 97</p><p>14.4.3.4 Nervous System............................................................................................................. 97</p><p>14.4.3.5 Development.................................................................................................................. 98</p><p>14.4.3.6 Habits and Habitats....................................................................................................... 98</p><p>14.5 Caudofoveata (or Chaetodermomorpha, Scutopoda).................................................................................. 99</p><p>14.5.1 Introduction.................................................................................................................................... 99</p><p>14.5.2 Phylogeny and Classification.......................................................................................................... 99</p><p>14.5.2.1 Sister Group Relationships............................................................................................ 99</p><p>14.5.3 Morphology.................................................................................................................................... 99</p><p>14.5.3.1 Spicules......................................................................................................................... 99</p><p>14.5.3.2 Head-Foot and Mantle................................................................................................... 99</p><p>14.5.3.3 Mantle Cavity................................................................................................................ 99</p><p>14.5.3.4 Digestive System......................................................................................................... 100</p><p>14.5.3.5 Renopericardial and Reproductive Systems.................................................................101</p><p>14.5.3.6 Nervous System and Sense Organs..............................................................................101</p><p>14.5.4 Biology, Ecology, and Behaviour..................................................................................................101</p><p>14.5.4.1 Habits and Habitats......................................................................................................101</p><p>14.5.4.2 Feeding and Diet..........................................................................................................101</p><p>14.5.4.3 Reproduction, Development, and Growth................................................................... 102</p><p>14.5.4.4 Associations................................................................................................................. 102</p><p>14.5.5 Diversity and Fossil History......................................................................................................... 102</p><p>14.6 Solenogastres (or Neomeniomorpha)........................................................................................................ 102</p><p>14.6.1 Introduction.................................................................................................................................. 102</p><p>14.6.2 Phylogeny and Classification........................................................................................................ 102</p><p>14.6.2.1 Sister Group Relationships.......................................................................................... 102</p><p>14.6.3 Morphology.................................................................................................................................. 102</p><p>14.6.3.1 Spicules....................................................................................................................... 102</p><p>14.6.3.2 Body............................................................................................................................ 102</p><p>14.6.3.3 Mantle Cavity.............................................................................................................. 103</p><p>14.6.3.4 Digestive System......................................................................................................... 103</p><p>14.6.3.5 Renopericardial and Reproductive Systems................................................................ 104</p><p>14.6.3.6 Nervous System and Sense Organs............................................................................. 105</p><p>14.6.4 Biology and Ecology.................................................................................................................... 105</p><p>14.6.4.1 Habits and Habitats..................................................................................................... 105</p><p>14.6.4.2 Feeding and Diet......................................................................................................... 105</p><p>14.6.4.3 Reproduction, Development, and Growth................................................................... 105</p><p>14.6.4.4 Associations................................................................................................................. 107</p><p>14.6.5 Diversity and Fossil History......................................................................................................... 107</p><p>Chapter 15 The Bivalvia......................................................................................................................................................... 109</p><p>15.1 Introduction............................................................................................................................................... 109</p><p>15.2 Classification............................................................................................................................................. 109</p><p>15.3 Morphology................................................................................................................................................112</p><p>15.3.1 The Shell.......................................................................................................................................113</p><p>15.3.1.1 Orientation....................................................................................................................113</p><p>15.3.1.2 Sculpture.......................................................................................................................113</p><p>15.3.1.3 Shell Structure..............................................................................................................113</p><p>15.3.1.4 Periostracum.................................................................................................................115</p><p>15.3.1.5 Ligament.......................................................................................................................117</p><p>15.3.1.6 Hinge............................................................................................................................119</p><p>15.3.1.7 Pallial Line and Sinus.................................................................................................. 120</p><p>15.3.1.8 Shell Reduction, Accessory Shell Plates, and Secondary Shells................................. 120</p><p>15.3.2 Muscles and Their Scars.............................................................................................................. 124</p><p>15.3.3 The Foot....................................................................................................................................... 124</p><p>15.3.4 The Byssus – Structure and Function.......................................................................................... 125</p><p>15.3.5 Mantle and Siphons...................................................................................................................... 127</p><p>15.3.6 Mantle Cavity............................................................................................................................... 132</p><p>15.3.6.1 The Ctenidia – Their Structure and Ciliation.............................................................. 132</p><p>15.3.6.2 The Labial Palps.......................................................................................................... 139</p><p>15.3.6.3 Other Features of the Mantle Cavity............................................................................141</p><p>15.3.7 Digestive System...........................................................................................................................141</p><p>15.3.8 Renopericardial System................................................................................................................143</p><p>15.3.9 Reproductive System.....................................................................................................................143</p><p>15.3.10 Nervous System and Sense Organs.............................................................................................. 145</p><p>15.4 Biology and Ecology................................................................................................................................. 145</p><p>15.4.1 Habits and Habitats...................................................................................................................... 145</p><p>15.4.1.1 Locomotion................................................................................................................. 145</p><p>15.4.1.2 Attachment.................................................................................................................. 146</p><p>15.4.1.3 Secondary Soft-Bottom Dwellers.................................................................................147</p><p>15.4.1.4 Burrowers and Borers.................................................................................................. 149</p><p>15.4.2 Feeding......................................................................................................................................... 151</p><p>15.4.3 Life History and Growth.............................................................................................................. 152</p><p>15.4.4 Associations................................................................................................................................. 156</p><p>15.4.4.1 Bacterial and Algal Symbionts.................................................................................... 156</p><p>15.4.5 Relationships with Other Animals............................................................................................... 158</p><p>15.4.5.1 Predators of Bivalves................................................................................................... 159</p><p>15.5 Economic Values....................................................................................................................................... 159</p><p>15.6 Diversity and Fossil History...................................................................................................................... 159</p><p>15.6.1 The First Bivalves......................................................................................................................... 159</p><p>15.6.2 The Evolutionary Radiation of Bivalves...................................................................................... 159</p><p>15.6.2.1 Freshwater Bivalves.....................................................................................................161</p><p>15.6.3 Patterns of Bivalve Diversity.........................................................................................................161</p><p>15.7 The Protobranch Bivalves......................................................................................................................... 162</p><p>15.7.1 Phylogeny and Classification........................................................................................................ 162</p><p>15.7.2 Biology and Ecology.................................................................................................................... 163</p><p>15.7.3 Diversity and Fossil History......................................................................................................... 164</p><p>15.8 The Autobranch Bivalves.......................................................................................................................... 164</p><p>15.8.1 Pteriomorphia............................................................................................................................... 164</p><p>15.8.1.1 Phylogeny and Classification...................................................................................... 164</p><p>15.8.1.2 Biology and Ecology................................................................................................... 165</p><p>15.8.1.3 Diversity and Fossil History........................................................................................ 167</p><p>15.8.2 Heteroconchia............................................................................................................................... 167</p><p>15.8.3 Palaeoheterodonta........................................................................................................................ 168</p><p>15.8.3.1 Trigoniida.................................................................................................................... 168</p><p>15.8.3.2 Unionida...................................................................................................................... 168</p><p>15.8.4 Heterodonta...................................................................................................................................170</p><p>15.8.4.1 Archiheterodonta......................................................................................................... 172</p><p>15.8.5 Euheterodonta.............................................................................................................................. 172</p><p>15.8.5.1 Phylogeny and Classification.......................................................................................173</p><p>15.8.5.2 Biology and Ecology....................................................................................................174</p><p>15.8.5.3 Diversity and Fossil History.........................................................................................175</p><p>15.8.5.4 The Rudists...................................................................................................................175</p><p>15.8.6 Anomalodesmata (= Poromyata)...................................................................................................178</p><p>Chapter 16 The Scaphopoda................................................................................................................................................... 183</p><p>16.1 Introduction............................................................................................................................................... 183</p><p>16.2 Phylogeny and Classification.................................................................................................................... 183</p><p>16.2.1 Sister Group Relationships.......................................................................................................... 183</p><p>16.3 Morphology............................................................................................................................................... 184</p><p>16.3.1 Shell............................................................................................................................................. 184</p><p>16.3.2 Head-Foot, Captacula, and Mantle.............................................................................................. 185</p><p>16.3.3 Mantle Cavity.............................................................................................................................. 186</p><p>16.3.4 Digestive System......................................................................................................................... 188</p><p>16.3.5 Renopericardial System............................................................................................................... 189</p><p>16.3.6 Reproductive System................................................................................................................... 189</p><p>16.3.7 Nervous System and Sense Organs............................................................................................. 189</p><p>16.4 Biology and Ecology................................................................................................................................. 190</p><p>16.4.1 Habits and Habitats...................................................................................................................... 190</p><p>16.4.2 Locomotion.................................................................................................................................. 190</p><p>16.4.3 Feeding and Diet.......................................................................................................................... 190</p><p>16.4.4 Reproduction, Development, and Growth................................................................................... 192</p><p>16.4.5 Associations................................................................................................................................. 192</p><p>16.4.6 Predation, Defence, and Other Behavioural Attributes............................................................... 193</p><p>16.5 Diversity and Fossil History...................................................................................................................... 193</p><p>16.5.1 The Major Groups of Scaphopods............................................................................................... 193</p><p>16.6 Human Uses.............................................................................................................................................. 193</p><p>Chapter 17 The Cephalopoda................................................................................................................................................. 195</p><p>17.1 Introduction............................................................................................................................................... 195</p><p>17.2 Phylogeny and Classification....................................................................................................................200</p><p>17.3 Morphology............................................................................................................................................... 202</p><p>17.3.1 Shell.............................................................................................................................................. 202</p><p>17.3.2 Head, Arms, and Funnel.............................................................................................................. 205</p><p>17.3.2.1 Embryological Evidence for the Derivation of Head Structures................................. 208</p><p>17.3.3 Epidermis (Skin)........................................................................................................................... 208</p><p>17.3.4 Head and Funnel Retractor Muscles............................................................................................ 209</p><p>17.3.4.1 Muscular Attachments, Shell Growth and the Addition of Septa.................................210</p><p>17.3.5 The Fossil Record of Soft Tissues................................................................................................ 212</p><p>17.3.6 Mantle Wall.................................................................................................................................. 213</p><p>17.3.7 Mantle Cavity................................................................................................................................214</p><p>17.4 Biology and Ecology..................................................................................................................................218</p><p>17.4.1 Habits and Habitats.......................................................................................................................218</p><p>17.4.1.1 The Habits and Habitats of Extinct Ectocochleate Cephalopods.................................218</p><p>17.4.2 Locomotion and Buoyancy.......................................................................................................... 220</p><p>17.4.2.1 Locomotion................................................................................................................. 220</p><p>17.4.2.2 Buoyancy..................................................................................................................... 222</p><p>17.4.2.3 The Shell as a Buoyancy Device................................................................................. 222</p><p>17.4.2.4 Other Buoyancy Strategies.......................................................................................... 227</p><p>17.4.3 The Ink Sac and Inking................................................................................................................ 228</p><p>17.4.4 Colour Changes – Chromatophores and Other Special Skin Structures..................................... 228</p><p>17.4.4.1 Chromatophores.......................................................................................................... 229</p><p>17.4.4.2 Structural Reflectors (Iridophores, Silvery Reflectors, and Leucophores).................. 229</p><p>17.4.5 Luminescent Organs.................................................................................................................... 231</p><p>17.5 Digestive System....................................................................................................................................... 233</p><p>17.5.1 The Jaws....................................................................................................................................... 233</p><p>17.5.2 The Radula................................................................................................................................... 233</p><p>17.5.3 Feeding......................................................................................................................................... 233</p><p>17.6 Renopericardial System............................................................................................................................. 236</p><p>17.7 Courtship, Reproduction, Development, and Growth............................................................................... 239</p><p>17.7.1 Reproductive System.................................................................................................................... 239</p><p>17.7.1.1 Sexual Characters and Sexual Dimorphism................................................................ 242</p><p>17.7.2 Courtship and Mating.................................................................................................................. 243</p><p>17.7.3 Life History..................................................................................................................................244</p><p>17.7.3.1 Spawn and Parental Care............................................................................................. 245</p><p>17.7.3.2 Development................................................................................................................ 245</p><p>17.7.4 The Nervous System, Sense Organs, and Behaviour................................................................... 247</p><p>17.7.5 The Endocrine System................................................................................................................. 249</p><p>17.7.6 Learning and Intelligence............................................................................................................ 249</p><p>17.7.7 Behaviour..................................................................................................................................... 249</p><p>17.7.8 Symbiotic Relationships............................................................................................................... 250</p><p>17.7.9 Predators....................................................................................................................................... 251</p><p>17.8 Economic Values....................................................................................................................................... 251</p><p>17.9 The Origin of Cephalopods....................................................................................................................... 251</p><p>17.10 Diversity and Fossil History...................................................................................................................... 256</p><p>17.10.1 The Ordovician Radiation and Beyond........................................................................................ 256</p><p>17.11 The Major Groups of Cephalopoda........................................................................................................... 258</p><p>17.11.1 The Nautiliforms.......................................................................................................................... 258</p><p>17.11.1.1 Ellesmeroceratia and Plectronoceratia...................................................................... 259</p><p>17.11.1.2 Orthoceratia............................................................................................................... 260</p><p>17.11.1.3 Discosorida................................................................................................................ 261</p><p>17.11.1.4 Endoceratia................................................................................................................ 261</p><p>17.11.1.5 Actinoceratia.............................................................................................................. 261</p><p>17.11.1.6 Nautilia...................................................................................................................... 262</p><p>17.12 Bactrita and Ammonitia (= Ammonoidea)................................................................................................ 262</p><p>17.12.1 Bactrita (= Bactritoidea)............................................................................................................... 262</p><p>17.12.2 Ammonitia (Ammonites)............................................................................................................. 264</p><p>17.12.2.1 The Embryonic Shell (Ammonitella)........................................................................ 267</p><p>17.12.2.2 Ammonite Sutural Patterns........................................................................................ 267</p><p>17.12.2.3 Life Habits................................................................................................................. 268</p><p>17.12.2.4 Aptychus and Jaws.................................................................................................... 269</p><p>17.12.2.5 Phylogeny, Classification, Diversity, and Fossil History of Ammonites................... 271</p><p>17.12.2.6 The Major Groups of Ammonites............................................................................. 272</p><p>17.13 Coleoida.................................................................................................................................................. 273</p><p>17.13.1 Phylogeny, Classification, Diversity, and Fossil History............................................................ 275</p><p>17.13.1.1 Belemnitidia (The Belemnites)................................................................................ 279</p><p>17.13.1.2 Decabrachia............................................................................................................. 279</p><p>17.13.1.3 Squid – A Grade Not a Clade – The Myopsida, Oegopsida, Bathyteuthoidea</p><p>(‘Teuthida’) �......................................................................................................... 284</p><p>17.13.1.4 Octobrachia (= Octopodiformes)............................................................................. 285</p><p>17.13.1.5 Giant Squid and Octopuses...................................................................................... 287</p><p>Chapter 18 Gastropoda I – Introduction and the Stem Groups.............................................................................................. 289</p><p>18.1 Introduction to the Gastropoda.................................................................................................................. 289</p><p>18.1.1 Phylogeny and Classification........................................................................................................ 289</p><p>18.1.2 Morphology.................................................................................................................................. 290</p><p>18.1.2.1 Shell, Operculum and External Body.......................................................................... 293</p><p>18.1.2.2 Shell Muscles.............................................................................................................. 294</p><p>18.1.2.3 Operculum................................................................................................................... 294</p><p>18.1.2.4 Head-Foot.................................................................................................................... 294</p><p>18.1.2.5 Mantle Cavity and Its Organs...................................................................................... 296</p><p>18.1.2.6 Digestive System......................................................................................................... 296</p><p>18.1.2.7 Renopericardial System............................................................................................... 297</p><p>18.1.2.8 Reproductive System...................................................................................................300</p><p>18.1.2.9 Nervous System and Sense Organs.............................................................................300</p><p>18.1.3 Biology, Ecology, and Behaviour.................................................................................................300</p><p>18.1.3.1 Habits and Habitats.....................................................................................................300</p><p>18.1.3.2 Locomotion and Behaviour......................................................................................... 301</p><p>18.1.3.3 Life History................................................................................................................. 301</p><p>18.1.3.4 Feeding........................................................................................................................ 301</p><p>18.1.3.5 Associations................................................................................................................. 302</p><p>18.1.4 Economic Values.......................................................................................................................... 303</p><p>18.1.5 Diversity and Fossil History......................................................................................................... 303</p><p>18.2 The Initial Radiation – Eogastropods and Orthogastropods ............................................................ 303</p><p>18.2.1 Hypothetical Reconstructions...................................................................................................... 303</p><p>18.2.1.1 A Hypothetical Ancestral Gastropod........................................................................... 307</p><p>18.2.1.2 Eogastropoda............................................................................................................... 308</p><p>18.2.1.3 Orthogastropoda...........................................................................................................310</p><p>18.3 The Lower Groups of Gastropods..............................................................................................................312</p><p>18.4 Patellogastropoda (= Docoglossa, Cyclobranchia, Eogastropoda in Part).................................................312</p><p>18.4.1 Introduction...................................................................................................................................312</p><p>18.4.2 Phylogeny and Classification.........................................................................................................312</p><p>18.4.2.1 Sister Group Relationships...........................................................................................313</p><p>18.4.2.2 Outline of Classification...............................................................................................313</p><p>18.4.3 Morphology...................................................................................................................................313</p><p>18.4.3.1 Shell and Protoconch....................................................................................................313</p><p>18.4.3.2 Head-Foot, Mantle, and Operculum.............................................................................315</p><p>18.4.3.3 Mantle Cavity...............................................................................................................315</p><p>18.4.3.4 Digestive System..........................................................................................................317</p><p>18.4.3.5 Reproductive System................................................................................................... 320</p><p>18.4.3.6 Renopericardial System............................................................................................... 320</p><p>18.4.3.7 Nervous System and Sense Organs............................................................................. 321</p><p>18.4.4 Biology, Ecology, and Behaviour................................................................................................. 322</p><p>18.4.4.1 Habits and Habitats..................................................................................................... 322</p><p>18.4.4.2 Locomotion and Behaviour......................................................................................... 322</p><p>18.4.4.3 Life History................................................................................................................. 322</p><p>18.4.4.4 Feeding and Diet......................................................................................................... 322</p><p>18.4.4.5 Associations................................................................................................................. 323</p><p>18.4.5 Diversity and Fossil History......................................................................................................... 323</p><p>18.5 Vetigastropoda........................................................................................................................................... 324</p><p>18.5.1 Introduction.................................................................................................................................. 324</p><p>18.5.2 Phylogeny and Classification........................................................................................................ 324</p><p>18.5.2.1 Outline of Classification.............................................................................................. 326</p><p>18.5.2.2 Deep-Sea Enigmas; The Neomphaloidean and Cocculinoidean Taxa........................ 332</p><p>18.5.3 Morphology.................................................................................................................................. 333</p><p>18.5.3.1 Shell and Protoconch................................................................................................... 333</p><p>18.5.3.2 Head-Foot, Mantle, Operculum, and Locomotion...................................................... 337</p><p>18.5.3.3 Mantle Cavity.............................................................................................................. 338</p><p>18.5.3.4 Digestive System......................................................................................................... 338</p><p>18.5.3.5 Reproductive System................................................................................................... 343</p><p>18.5.3.6 Renopericardial System............................................................................................... 344</p><p>18.5.3.7 Nervous System and Sense Organs............................................................................. 344</p><p>18.5.4 Biology, Ecology, and Behaviour................................................................................................. 345</p><p>18.5.4.1 Habits and Habitats..................................................................................................... 345</p><p>18.5.4.2 Feeding and Diet......................................................................................................... 345</p><p>18.5.4.3 Life History................................................................................................................. 346</p><p>18.5.4.4 Associations................................................................................................................. 346</p><p>18.5.4.5 Behaviour.................................................................................................................... 346</p><p>18.5.5 Diversity and Fossil History......................................................................................................... 346</p><p>18.6 Neritimorpha (= Neritopsina).................................................................................................................... 347</p><p>18.6.1 Introduction.................................................................................................................................. 347</p><p>18.6.2 Phylogeny and Classification........................................................................................................ 347</p><p>18.6.2.1 Sister Group Relationships.......................................................................................... 349</p><p>18.6.2.2 Outline of Classification.............................................................................................. 349</p><p>18.6.3 Morphology.................................................................................................................................. 350</p><p>18.6.3.1 Shell and Protoconch................................................................................................... 350</p><p>18.6.3.2 Head-Foot, Mantle, Operculum, and Locomotion.......................................................351</p><p>18.6.3.3 Mantle Cavity.............................................................................................................. 353</p><p>18.6.3.4 Digestive System......................................................................................................... 354</p><p>18.6.3.5 Renopericardial System............................................................................................... 355</p><p>18.6.3.6 Reproductive System................................................................................................... 355</p><p>18.6.3.7 Nervous System and Sense Organs............................................................................. 360</p><p>18.6.4 Biology and Ecology.................................................................................................................... 360</p><p>18.6.4.1 Habits and Habitats..................................................................................................... 360</p><p>18.6.4.2 Feeding and Diet......................................................................................................... 361</p><p>18.6.4.3 Life History................................................................................................................. 361</p><p>18.6.4.4 Associations................................................................................................................. 362</p><p>18.6.4.5 Behaviour.................................................................................................................... 362</p><p>18.6.5 Diversity and Fossil History......................................................................................................... 363</p><p>Chapter 19 Gastropoda II – The Caenogastropoda................................................................................................................ 365</p><p>19.1 Introduction............................................................................................................................................... 365</p><p>19.2 Phylogeny and Classification.................................................................................................................... 365</p><p>19.2.1 Sister Group Relationships........................................................................................................... 365</p><p>19.2.2 Outline of Classification............................................................................................................... 365</p><p>19.3 Morphology............................................................................................................................................... 366</p><p>19.3.1 Shell and Protoconch.................................................................................................................... 377</p><p>19.3.2 Head-Foot, Mantle, and Operculum............................................................................................ 379</p><p>19.3.2.1 Head Structures........................................................................................................... 379</p><p>19.3.2.2 The Foot...................................................................................................................... 379</p><p>19.3.2.3 The Operculum............................................................................................................ 380</p><p>19.3.2.4 Mantle Edge................................................................................................................ 380</p><p>19.3.3 Mantle Cavity............................................................................................................................... 380</p><p>19.3.4 Digestive System.......................................................................................................................... 381</p><p>19.3.5 Renopericardial System............................................................................................................... 394</p><p>19.3.6 Reproductive System.................................................................................................................... 394</p><p>19.3.6.1 The Female System..................................................................................................... 395</p><p>19.3.6.2 The Male System......................................................................................................... 396</p><p>19.3.7 Nervous System and Sense Organs..............................................................................................400</p><p>19.4 Biology, Ecology, and Behaviour.............................................................................................................. 403</p><p>19.4.1 Habits and Habitats...................................................................................................................... 403</p><p>19.4.2 Locomotion.................................................................................................................................. 403</p><p>19.4.3 Feeding and Diet..........................................................................................................................404</p><p>19.4.4 Life History and Growth.............................................................................................................. 405</p><p>19.4.4.1 Spawn, Development, and Larvae............................................................................... 405</p><p>19.4.5 Associations..................................................................................................................................410</p><p>19.4.5.1 The Evolution of Parasitism in Eulimidae....................................................................410</p><p>19.4.6 Predators and Defence..................................................................................................................412</p><p>19.4.7 Other Behavioural Attributes........................................................................................................412</p><p>19.5 Economic and Ecological Values...............................................................................................................412</p><p>19.6 Diversity and Fossil History.......................................................................................................................413</p><p>19.7 Major Groups within Caenogastropoda.....................................................................................................414</p><p>19.7.1 The Architaenioglossans...............................................................................................................414</p><p>19.7.1.1 Campanilida.................................................................................................................415</p><p>19.7.2 Sorbeoconcha................................................................................................................................415</p><p>19.7.2.1 Cerithiimorpha.............................................................................................................415</p><p>19.7.2.2 Hypsogastropoda..........................................................................................................416</p><p>Chapter 20 Gastropoda III – The Heterobranchia...................................................................................................................419</p><p>20.1 Introduction................................................................................................................................................419</p><p>20.2 Phylogeny and Classification.....................................................................................................................419</p><p>20.2.1 Sister Group Relationships........................................................................................................... 421</p><p>20.2.2 Chromosome Numbers................................................................................................................. 422</p><p>20.3 Morphology............................................................................................................................................... 429</p><p>20.3.1 Shell and Operculum.................................................................................................................... 435</p><p>20.3.1.1 Shell Structure............................................................................................................. 439</p><p>20.3.1.2 Larval or Embryonic Shell.......................................................................................... 439</p><p>20.3.1.3 Shell Reduction and Loss............................................................................................ 439</p><p>20.3.1.4 Shell Variation.............................................................................................................440</p><p>20.3.1.5 Operculum...................................................................................................................440</p><p>20.3.2 External Body Morphology.......................................................................................................... 441</p><p>20.3.2.1 Head Tentacles............................................................................................................ 441</p><p>20.3.2.2 Cephalic (Head) Shield............................................................................................... 442</p><p>20.3.2.3 Epithelial Modifications.............................................................................................. 442</p><p>20.3.2.4 Oral Veil.......................................................................................................................444</p><p>20.3.2.5 The Foot, Pedal Glands, and Parapodia.......................................................................444</p><p>20.3.2.6 The Mantle.................................................................................................................. 445</p><p>20.3.3 Mantle Cavity and Associated Structures.................................................................................... 447</p><p>20.3.3.1 Heterobranch Gills...................................................................................................... 450</p><p>20.3.3.2 Opposed Ciliary Ridges (Raphes) and Mantle Tentacles............................................ 451</p><p>20.3.3.3 The Hypobranchial Gland........................................................................................... 451</p><p>20.3.3.4 The Pigmented Mantle Organ (PMO)......................................................................... 452</p><p>20.3.3.5 Other Glands Associated with the Mantle Cavity....................................................... 452</p><p>20.3.4 Digestive System.......................................................................................................................... 452</p><p>20.3.5 The Renopericardial System........................................................................................................ 472</p><p>20.3.6 Nervous System and Sense Organs.............................................................................................. 474</p><p>20.3.6.1 Sense Organs............................................................................................................... 477</p><p>20.3.7 Reproductive System.................................................................................................................... 479</p><p>20.3.7.1 Terminology................................................................................................................ 479</p><p>20.3.7.2 The Ovotestis and Gametes......................................................................................... 481</p><p>20.3.7.3 The Reproductive System of ‘Lower Heterobranchs’................................................. 482</p><p>20.3.7.4 Some Small Euthyneurans Previously Thought to Be Lower Heterobranchs............. 485</p><p>20.3.7.5 Euthyneuran Reproductive Systems............................................................................ 486</p><p>20.4 Biology, Ecology, and Behaviour.............................................................................................................. 503</p><p>20.4.1 Habits and Habitats...................................................................................................................... 503</p><p>20.4.2 Locomotion.................................................................................................................................. 503</p><p>20.4.3 Food and Feeding......................................................................................................................... 504</p><p>20.4.3.1 Herbivory..................................................................................................................... 505</p><p>20.4.3.2 Fungivory.................................................................................................................... 505</p><p>20.4.3.3 Active Carnivory.......................................................................................................... 505</p><p>20.4.3.4 Parasitism.................................................................................................................... 507</p><p>20.4.3.5 Suspension Feeding..................................................................................................... 507</p><p>20.4.3.6 Coprophagy................................................................................................................. 507</p><p>20.4.4 Reproduction, Devel