20.1 Introduction

20.2 Phylogeny and Classification

20.2.1 Sister Group Relationships

20.2.2 Chromosome Numbers

20.3 Morphology

20.3.1 Shell and Operculum

20.3.1.1 Shell Structure

20.3.1.2 Larval or Embryonic Shell

20.3.1.3 Shell Reduction and Loss

20.3.1.4 Shell Variation

20.3.1.5 Operculum

20.3.2 External Body Morphology

20.3.2.1 Head Tentacles

20.3.2.2 Cephalic (Head) Shield

20.3.2.3 Epithelial Modifications

20.3.2.4 Oral Veil

20.3.2.5 The Foot, Pedal Glands, and Parapodia

20.3.2.6 The Mantle

20.3.3 Mantle Cavity and Associated Structures

20.3.3.1 Heterobranch Gills

20.3.3.2 Opposed Ciliary Ridges (Raphes) and Mantle Tentacles

20.3.3.3 The Hypobranchial Gland

20.3.3.4 The Pigmented Mantle Organ (PMO)

20.3.3.5 Other Glands Associated with the Mantle Cavity

20.3.4 Digestive System

20.3.5 The Renopericardial System

20.3.6 Nervous System and Sense Organs

20.3.6.1 Sense Organs

20.3.7 Reproductive System

20.3.7.1 Terminology

20.3.7.2 The Ovotestis and Gametes

20.3.7.3 The Reproductive System of ‘Lower Heterobranchs’

20.3.7.4 Some Small Euthyneurans Previously Thought to Be Lower Heterobranchs

20.3.7.5 Euthyneuran Reproductive Systems

20.4 Biology, Ecology, and Behaviour

20.4.1 Habits and Habitats

20.4.2 Locomotion

20.4.3 Food and Feeding

20.4.3.1 Herbivory

20.4.3.2 Fungivory

20.4.3.3 Active Carnivory

20.4.3.4 Parasitism

20.4.3.5 Suspension Feeding

20.4.3.6 Coprophagy

20.4.4 Reproduction, Development, and Growth

20.4.4.1 Self-Fertilisation

20.4.4.2 Hypodermic Insemination

20.4.4.3 Aphally

20.4.4.4 Asexual Reproduction

20.4.4.5 Life Cycles

20.4.4.6 Reproductive Behaviour

20.4.4.7 Development

20.4.4.8 Spawn

20.4.4.9 Larvae and Larval Shells

20.4.5 Behaviour

20.4.6 Predation and Defence

20.4.6.1 The Shell as a Means of Defence

20.4.6.2 Mucus

20.4.6.3 Defence without a Shell

20.4.6.4 Use of Nematocysts

20.4.6.5 Aposematic (Warning) Colouration and Batesian and Müllerian Mimicry

20.4.6.6 Autotomy

20.4.6.7 Predators of Heterobranchs

20.4.7 Associations

20.4.8 Economic and Ecological Values

20.5 Diversity and Fossil History

20.5.1 Major Groups within the Heterobranchia

20.5.1.1 ‘Lower Heterobranchs’

20.5.1.2 Euthyneura