Fossil Invertebrates: Gastropods☆

J Frýda, Czech University of Life Sciences, Prague, Czech Republic

ã 2013 Elsevier Inc. All rights reserved.

Introduction 1
Definition and General Description 2
Anatomical Features 2
Ontogeny 3
The Gastropod Shell 3
Protoconch and Teleoconch 3
Operculum 3
Shell Structure 4
Shell Coiling 4
Muscle Scars 6
Classification of the Gastropoda 8
Evolution of the Gastropoda 9
Origin and Early History of the Gastropoda 9
Palaeozoic Era 9
Mesozoic and Cenozoic Eras 10
Evolution of Freshwater and Terrestrial Gastropods 11

Glossary Operculum Lid-like structure used for closing of the

Archaeogastropoda Group of extant gastropods.
Bellerophontida Extinct group of Palaeozoic molluscs with
bilaterally symmetrical shells.
Caenogastropoda Group of extant gastropods.
Embryonic shell (protoconch I) Gastropod shell formed
during embryonic development.
Heterobranchia Group of extant gastropods.
Heterostrophic Condition of the protoconch when its
whorls coil in the opposite direction to those of
the teleoconch.
Homeostrophic Protoconch and teleoconch whorls coil in
the same direction.
Hyperstrophy Condition in which anatomically dextral
animals occupy sinistrally coiled shells, and vice versa.
larval shell (protoconch II) Gastropod shell formed during
larval development in members of the Neritimorpha,
Caenogastropoda, and Heterobranchia.
Lecithotrophic Form of development in which larvae use
yolk in egg for their nutrition.
Mimospirina Extinct group of Early and Middle Palaeozoic
gastropods with sinistrally coiled, homeostrophic shells.
Neritimorpha Group of extant gastropods.
aperture in gastropod shells.
Opisthobranchia Gastropod subclass of
Thiele’s classification.
Orthostrophy Condition in which anatomically dextral
(or sinistral) animals occupy dextrally (or sinistrally)
coiled shells.
Patellogastropoda Group of extant gastropods with limpet-
shaped shells.
Periostracum Outer organic layer of gastropod shells.
Planktotrophic Form of development in which free-
swimming larvae use planktic organisms for their nutrition.
Prosobranchia Gastropod subclass of
Thiele’s classification.
Protoconch Gastropod shell formed during larval and/or
embryonic development.
Pulmonata Gastropod subclass of Thiele’s classification.
Teleoconch Post-larval gastropod shell.
Trochophore larva Gastropod larva formed during early
larval development which may be free swimming or occurs
in egg capsules.
Veliger Gastropod larva formed during later larval
development before metamorphosis to post-larval stages.

Introduction

Gastropods are well-known animals which have been associated with humans since the dawn of civilization. Their bodies were
gathered for food and their shells were used as tools, ornaments, and later as money. Their widespread occurrence is clear evidence
of their successful adaptation to different environments. During a long evolution, they are the only molluscan class to have
colonized the majority of marine, freshwater, and terrestrial environments. Marine gastropods occur mostly in shallow-water

☆
Change History: July 2013. J Frýda updated Abstract, Classification of the Gastropoda, Evolution of the Gastropoda, the reference section, Figures 4, 7, 11, and
added Figures 8, 9 and 10.

Reference Module in Earth Systems and Environmental Sciences http://dx.doi.org/10.1016/B978-0-12-409548-9.02806-2 1

2 Fossil Invertebrates: Gastropods

benthic communities; however, some gastropod species have also lived in the deep sea (e.g., faunas associated with hydrothermal
vents), and others, such as holoplanktic animals, have spent their whole lives as free-swimming gastropods. The terrestrial
gastropods colonized most land environments, ranging from lowlands to high mountains, and including humid to arid biotopes
of tropical to subarctic areas. Such adaptive radiation is quite exceptional amongst all animal phyla and is linked to the
extraordinary morphological and functional diversity of their bodies and shells. The gastropods comprise one of the most diverse
groups of living animals (the second after Insecta). All these facts, together with their long and rich fossil record, make gastropods a
unique animal group for evolutionary, ecological, and biogeographical investigations. There follows a brief review of gastropod
anatomy, shell morphology, classification, and more than 500 million years of evolution.

Definition and General Description

The Gastropoda forms one of eight molluscan classes, and is defined by several unique anatomical features which support its
interpretation as a molluscan group derived from the same ancestor (i.e., monophyly). The most characteristic feature of
gastropods is torsion of their soft bodies during early larval stages, producing a crossing of their nerve connectives, bending
of the intestine, and twisting of the mantle cavity (together with associated structures, including the ctenidia, anus, kidney
openings, etc.) anteriorly over the gastropod head.

Anatomical Features
The exceptional morphological and functional diversity of gastropod bodies is also reflected in their anatomy. Generally, the body
consists of a large foot, a visceral mass, and a head with a mouth, tentacles, and eyes. The visceral mass is mostly enclosed, together
with the mantle cavity, in a calcareous shell (Figure 1). Gastropods as soft-bodied animals use the pressure of their blood and
muscles for movements of different organs. The circulatory system comprises a contractile heart with one (Caenogastropoda and
Heterobranchia) or two (Patellogastropoda, Archaeogastropoda, and Neritimorpha) auricles, and a ventricle, as well as a system of
arteries and veins. Gastropod blood transports oxygen using the copper-bearing pigment haemocyanin. In most marine gastro-
pods, one or two gills (ctenidia) situated in the mantle cavity are used for respiration. Some marine and freshwater gastropods
developed secondary gills after the loss of their ctenidia in previous evolution. In the terrestrial gastropods (e.g., Pulmonata), a

Figure 1 Some variations in shell form of living gastropods. (a) High-spired: Mitra mitra (Muricoidea). (b) Strombiform: Lambis chiragra
(Stromboidea). (c) Turbiniform: Liguus vittatus (Orthalicoidea). (d) Convolute: Cyprea tigris (Cypraeoidea). (e) Spinose fusiform: Chicoreus ramosus
(Muricoidea). (f) Fusiform: Pleuroploca trapezium (Muricoidea). (g) Conoidal: Conus litteratus (Conoidea). (h) Discoidal: Architectonica perspectiva
(Architectonicoidea). (i) Turriculate: Terebra sp. (Conoidea). (j) Ovoid: Olivancillaria gibbosa (Olivoidea). (k) Involute: Cypraecassis rufa (Tonnoidea). (l)
Irregularly coiled: Siliquaria ponderosa (Cerithioidea). (m) limpet: Megathura crenulata (Fissurelloidea). (a, e–g, i, j) Neogastropoda (Caenogastropoda);
(b, d, k) Littorinimorpha (Caenogastropoda); (c, h) Heterobranchia; (l) Sorbeoconcha (Caenogastropoda); (m) Vetigastropoda (Archaeogastropoda).
 Fossil Invertebrates: Gastropods 3

highly vascularized internal wall of the mantle cavity (lung) is used for respiration. One or, rarely, two (Patellogastropoda and
Vetigastropoda) kidneys serve for excretion through the mantle cavity. The digestive system starts with a mouth containing a tooth-
bearing ribbon (radula). The organization of gastropod radulae and stomach, as well as additional parts of the digestive system,
reflects their different feeding habits (herbivory, detritivory, carnivory, or parasitism). The anus opens into the mantle cavity. The
gastropod nerve system includes paired ganglia which are linked with different sensory receptors by connectives and commissures.
The morphology of the reproductive organs and the reproductive strategies are highly diverse. Generally, more ancient gastropod
groups are gonochoristic with a simple reproductive system and external (Patellogastropoda and Archaeogastropoda) or internal
(Neritimorpha) fertilization. Caenogastropoda also use internal fertilization with complex reproductive morphology, and some
may be simultaneous hermaphrodites. The Heterobranchia have the most complex and variable reproductive system and
are hermaphroditic.

Ontogeny
Gastropods, like all Mollusca, have a biphasic life cycle (i.e., larval and post-metamorphosis stages), and this feature is shared with
closely related animal phyla (Kamptozoa, Sipunculida, Polychaeta, etc.). Like other molluscan groups, the embryonic development
is characterized by a spiral cleavage, which differs slightly in the main gastropod groups. The subsequent larval stage is called the
trochophore larva, and a similar larval type is developed in all molluscan groups. The trochophore larvae may be free swimming, as
in the ancient gastropod groups (Patellogastropoda and Archaeogastropoda), or may occur in egg capsules, as in more advanced
gastropods. The last larval stage is termed veliger, which typically bears two ciliate paddles (velum), sometimes subdivided into
several lobes. If free-swimming gastropod larvae use planktic organisms for their nutrition, their development is termed plankto-
trophic. Marine gastropods with such development have small eggs, but sometimes numbering over half a million. Planktotrophic
larvae may stay planktic for several months and thus can be carried for long distances by oceanic currents. The gastropods, however,
developed another ontogenetic strategy in which their larvae were not dependent on an external food source, but on the yolk of
their eggs. Gastropods with such a non-planktotrophic development (lecithotrophic) typically produce fewer eggs, which are
relatively large. The larval stages end with a metamorphosis that involves anatomical and physiological reorganization of the larval
body into the juvenile, post-larval body. Terrestrial and freshwater gastropods have simplified their development, and their
embryonic and larval stages are fixed on egg capsules or the female body (direct development). Such ontogenetic changes
considerably decreased their dispersal potential.

The Gastropod Shell

Gastropods are not only one of the most diverse animal groups, but the morphology of their shells is extremely varied (Figures 1
and 2). During more than 500 million years of evolution, they developed shells with various shapes and ornament, ranging in size
from about 1 mm up to more than 1 m (Eocene Campaniloidea, Caenogastropoda). The shell and its ornament may be broadly
linked to the mode of gastropod life (e.g., origin of limpet-shaped shells in unrelated gastropod groups). Generally, the most ornate
shells occur in tropical marine environments, but freshwater and terrestrial gastropods are often less ornate.

Protoconch and Teleoconch

In shell-bearing gastropods, the shell grows during almost the whole of their ontogeny. The part of the shell formed during the
embryonic and larval stages is called a protoconch (Figure 3), and that growing after metamorphosis is termed a teleoconch. The
main gastropod groups differ in their early development, which is reflected in their protoconch morphology. The more ancient
gastropod groups (Patellogastropoda and Archaeogastropoda) have the simplest shell ontogeny and their protoconchs have only
an embryonic shell (protoconch I), which is followed by a teleoconch (Figures 3(b) and 3(f)–3(h)). On the other hand, the
protoconchs of more advanced gastropods (Neritimorpha, Caenogastropoda, and Heterobranchia) consist of an embryonic shell
(protoconch I) and a subsequent larval shell (protoconch II). In most caenogastropods, the larval shells have different ornament
from the teleoconchs (Figures 3(k) and 3(l)), and both shells are coiled in the same direction (such a condition is termed
homeostrophic; Figure 4). In contrast, in the Heterobranchia with planktotrophic development, the protoconchs are coiled in the
opposite direction to the teleoconchs (Figures 3(j) and 4). Such shells are termed heterostrophic. The Neritimorpha form typical,
strongly convolute protoconchs during planktotrophic development, which are homeostrophic (Figures 3(a), 3(n), and 4). Higher
gastropods with non-planktotrophic development (some marine, freshwater, and terrestrial gastropods) have simplified their early
ontogeny and thus also the morphology of their protoconchs. The latter strategy is documented from the Devonian (400 Ma).

Operculum
The majority of gastropods have a lid-like structure (operculum) to close their aperture. This operculum is present in all living
gastropods during their larval stages, but is lost in some adults (e.g., limpets and the majority of terrestrial gastropods). The
operculum is mostly horny (corneous) and may be tightly (multispiral) or loosely (paucispiral) coiled or concentric. Some
gastropod groups have calcareous opercula, and the oldest operculum known is from the Ordovician (Macluritoidea).
4 Fossil Invertebrates: Gastropods

Figure 2 Some variations in shell form in the main groups of middle Palaeozoic gastropods. (a) High-spired shell: Murchisonia coronata
(Murchisonioidea). (b) Trochiform, slit-bearing shell: Devonorhineoderma orbignyana (Eotomarioidea). (c) Bilaterally symmetrical shell with a
prominent selenizone: Kolihadiscus tureki (Cyrtolitoidea). (d) Turbiniform shell: Gyronema armata (Gyronematidae). (e) Openly coiled shell:
Pragoserpulina tomasi (Pragoserpulinidae). (f) Discoidal shell: Stusakia pulchra. (g) Sinistrally coiled shell: Voskopiella barborae (Onychochilidae).
(h) Naticiform shell: Eifelcyrtus blodgetti (Vltavielidae). (i) Fusiform shell: Havlicekiela parvula (Peruneloidea). High-spired shells: (j) Pragozyga
costata; (k) Palaeozygopleura bohemica (Loxonematoidea). (l, m) Bilaterally symmetrical shell: Bellerophon vasulites (Bellerophontoidea).
(n) Limpet: Pragoscutula wareni (Pragoscutulidae). (o) Sinistrally coiled shell: Alaskiella medfraensis (Porcellioidea). (p) Discoidal shell:
Nodeuomphalus labadyei (Euomphaloidea); (q) Bilaterally symmetrical shell covered by secondary shell deposits: Branzovodiscus bajae
(Bellerophontoidea). (a, b, d, f, o) Archaeogastropoda; (c) Cyrtonellida; (g) Mimospirina; (h) Cyrtoneritimorpha; (l, m, q) Bellerophontida;
(i) Perunelomorpha; (e, j, k, n) Order uncertain; (p) Euomphalomorpha.

Shell Structure
Most gastropod shells are composed of an outer organic layer (periostracum) and an inner, mostly much thicker, calcified layer. The
colour pattern typical of many gastropod shells (Figure 1) is formed by different organic pigments which are limited to the
periostracum and the uppermost calcified layer. This shell feature, sometimes reflecting the mode of life, has been known since the
Palaeozoic (Figure 5). The inner layers of gastropod shells consist of minute calcium carbonate crystals (aragonite or calcite) in an
organic matrix. There are over 20 structural types of gastropod shell and, in general, more ancient groups exhibit more diverse shell
structures. The Patellogastropoda (Eogastropoda) had the most complex shell structure. On the other hand, the majority of the higher
gastropods have developed simple aragonitic shells with a crossed lamellar structure (Figures 6(a) and 6(b)). Some structural types are
restricted to certain groups (e.g., nacre) and this may be used for their identification in fossils (Figure 6(c)). Nacreous and crossed
lamellar structures have been known since the Palaeozoic.

Shell Coiling
The majority of the shell-bearing gastropods have right-handed (dextral) shells, but some have left-handed (sinistral) shells
(Figures 1 and 2). Only a few gastropods have bilaterally symmetrical shells which may be uncoiled (limpets) or planispirally coiled
 Fossil Invertebrates: Gastropods 5

Figure 3 Variety of protoconch shape. Strongly convolute larval shell (protoconch II), Neritimorpha: (a) Holocene Smaragdia sp. (Neritoidea); (n)
Triassic Pseudorthonychia alata (Pseudorthonychiidae). Embryonic shell (protoconch I) followed by teleoconch, Archaeogastropoda: (b) Triassic
Wortheniella coralliophila (Vetigastropoda); (c) Holocene Anatoma proxima (Vetigastropoda); (f) Devonian Zlichomphalina sp. (Eotomarioidea); (g, h)
Devonian Diplozone innocens (Murchisonioidea). (d) Openly coiled larval shell (protoconch II), Cyrtoneritimorpha, Carboniferous Orthonychia parva
(Orthonychiidae). (e) Openly coiled early shell, Permian Euomphalus sp. (Euomphaloidea). (i) Openly coiled larval shell (protoconch II) of the Silurian
Peruneloidea. (j) Heterostrophic larval shell (protoconch II), Jurassic Mathilda sp. (Architectonicoidea, Heterobranchia). Larval shell (protoconch II),
Caenogastropoda: (k) Holocene Hipponix sp. (Vanikoroidea); (l) Devonian Balbiniconcha cerinka (Subulitoidea). (m) Heterostrophic early shell,
Devonian Alaskiella medfraensis (Porcellioidea, Archaeogastropoda).

Figure 4 Schematic diagram showing the relationship between the coiling of larval (protoconch II) and post-larval (teleoconch) shells in planktotrophic
Caenogastropoda, Neritimorpha, and Heterobranchia. Coiling of both shells in the same direction (Caenogastropoda and Neritimorpha) is termed
homeostrophic. If the handedness of the shells is opposite (Heterobranchia), the coiling is termed heterostrophic.

(Figures 1(m), 2(c), 2(l), and 2(m)). The limpet-shaped shells were independently developed within all main gastropod groups from
the asymmetrically coiled shells of their ancestors. In contrast, planispirally coiled shells are known only in several groups, such as the
Palaeozoic Porcellioidea and Bellerophontoidea (Figures 2(l) and 2(m)) or the Holocene Planorbioidea. Some gastropods may
change the coiling of their shells (Figure 4) from sinistral to dextral (dextral heterostrophy), or vice versa (sinistral heterostrophy),
during ontogeny. Such a change may occur at a developmental stage, when gastropods undergo a metamorphosis from larval to post-
larval stages (e.g., Heterobranchia; Figure 3(j)), or later (e.g., Porcellioidea; Figure 3(m)). Dextrality or sinistrality of the shell is
independent of the coiling of the soft body, and the asymmetrical soft body of gastropods may be dextral or sinistral. Anatomical
dextrality or sinistrality may be easily recognized, even in fossil gastropods, if they developed a spiral operculum. The spiral operculum
of anatomically dextral gastropods is coiled counterclockwise (viewed externally), and vice versa in sinistral gastropods. Thus, there are
four possible relationships between shell coiling and body asymmetry in the shell-bearing gastropods (Figure 7). If anatomically
dextral (or sinistral) animals occupy dextrally (or sinistrally) coiled shells, such a condition is called dextral (or sinistral) orthostrophy.
If the handedness of the shell and soft body is different, the term hyperstrophy is used. All four kinds of coiling (Figure 7) have occurred
in gastropods, but their frequencies are very different. The great majority of living gastropods are dextrally orthostrophic, and sinistral
orthostrophy is uncommon. Dextral or sinistral hyperstrophy is very rare (e.g., Ordovician Macluritoidea or some
Holocene Planorbioidea).
6 Fossil Invertebrates: Gastropods

Figure 5 Apical (a) and lateral (b) views of the Middle Devonian (about 400 Ma) neritimorph, Paffrathopsis subcostata, showing the colour pattern.

Figure 6 Examples of shell structure in fossil gastropods. Aragonitic crossed lamellar structure in the Carboniferous (about 300 Ma) Amphiscapha
catilloides (Euomphaloidea): views perpendicular to (a) and parallel to (b) the shell surface. (c) Nacreous structure (columnar nacre) in the Late
Cretaceous (about 80 Ma) Sensuitrochus ferreri (Porcellioidea, Archaeogastropoda).

Muscle Scars
Gastropod shells are attached to the soft body by muscles, which may leave distinct scars on the inner shell surface (Figure 8). The
geometry of muscle scars has frequently been used as a diagnostic feature for distinction between torted (i.e., gastropods) and
untorted states in the Palaeozoic molluscs (Monoplacophora, Helcionelloida, Cyrtonellida, etc.). However, new anatomical studies
of living gastropods have shown that the larval muscles taking part in torsion and the post-larval muscles are developed quite
independently. Thus, the muscle scar pattern sometimes observable in the fossil molluscan shells may be a good ecological
indicator, but has no systematic significance.
 Fossil Invertebrates: Gastropods 7

Figure 7 Schematic diagram showing the four possible relationships between shell coiling and body asymmetry. Orthostrophy means that
anatomically dextral (or sinistral) animals occupy dextrally (or sinistrally) coiled shells. The term hyperstrophy is used when the handedness of the shell
and soft body is different (see text for explanation).

Figure 8 Muscle scars in the Early Devonian gastropod Orthonychia protei. Gastropod shells are attached to the soft body by muscles, which may
leave distinct scars on the inner shell surface. (a) – inner cast of a shell showing the position of the muscle scar; (b) specimen with partly preserved
thick shell with characteristic growth lines; (c,d) detailed views of two specimens of the same species showing intraspecific variability in the
muscle scars.
8 Fossil Invertebrates: Gastropods

Classification of the Gastropoda

Gastropods as an independent group of molluscs were recognized and named by the French naturalist, Georges Cuvier, more than
200 years ago. Since then, scientists have tried to classify them by using different features of their bodies. However, the classification
of such a numerous group with extraordinary morphological and anatomical variability of their bodies and shells has encountered
many problems. During the nineteenth century, several different classifications of the Gastropoda were published, based on the
shape of the shells, position of the mantle cavity, or on the arrangement of various organs (e.g., gills or head). Generally, these
classification schemes used only a limited number of distinguishing characters. At the beginning of the twentieth century, the
German zoologist, Johannes Thiele, integrated earlier classifications and divided the gastropods into three subclasses: Prosobran-
chia, Opisthobranchia, and Pulmonata. In addition, the Prosobranchia were divided into three orders: Archaeogastropoda,
Mesogastropoda, and Neogastropoda. Thiele’s system was used by zoologists and palaeontologists for most of the twentieth
century. However, during recent decades, numerous new data on the anatomy of various gastropod groups have been accumulated,
mainly by the application of new methods (e.g., transmission electron microscopy). At the same time, studies of the deep-sea
faunas associated with hydrothermal vents have brought the discovery of new gastropod groups with unusual anatomical features.
The evaluation of this newly gathered data in the light of the existing classification revealed a need for its revision. Recent analyses
of numerous morphological and developmental characters of living gastropods have resulted in new classification schemes
(Figure 9), which have been independently supported by results from molecular studies. The placement of fossil gastropods
into this classification of living gastropods has been difficult because of the lack of necessary anatomical characters.
Recent studies have revealed that Patellogastropoda (¼ Docoglossa, Cyclobranchia) may represent the sister group to all other
living gastropods. Living patellogastropods with limpet-shaped shells are exclusively marine and occur mostly on rocky shores in
all continents. The Patellogastropoda and their coiled ancestors have been united into the subclass Eogastropoda (Figure 9(a)). All
other living gastropods and their ancestors have been placed in the subclass Orthogastropoda, comprising four main groups of
living gastropods: Archaeogastropoda, Neritimorpha, Caenogastropoda, and Heterobranchia (Figure 9).
The Neritimorpha (¼ Neritopsina) is an ancient gastropod group with a long fossil record (Figures 2, 5, and 9), which
colonized many different marine (shallow- and deep-water), freshwater, and terrestrial environments. The Palaeozoic Cyrtoner-
itimorpha, with openly coiled early shells (Figures 2(h) and 3(d)), may represent a closely related group. The living Archaeogas-
tropoda unites the Vetigastropoda (Figures 3(b) and 3(c)) and several smaller groups, such as the Neomphaloidea, which occur in
faunas associated with deep-sea hydrothermal vents. The Archaeogastropoda have colonized almost all marine and estuarine
environments. There are also a number of extinct, mainly Palaeozoic groups (Figure 2) with uncertain relationships to living
archaeogastropods. The Palaeozoic Euomphaloidea (¼ Euomphalomorpha; Figures 2(p) and 3(e)), known mainly from shallow-
water, marine environments, may be a sister or basal group of the Archaeogastropoda.
The Caenogastropoda and Heterobranchia are sister groups which are united in the taxon Apogastropoda. Both groups are highly
diverse and have colonized almost all marine, freshwater, and terrestrial environments. The Palaeozoic Subulitoidea and Peruneloi-
dea (Perunelomorpha) (Figures 2(i) and 3(l)) may be ancestral or basal groups of the Caenogastropoda or of all Apogastropoda. The
extant Caenogastropoda unites the three clades Architaenioglossa, Sorbeoconcha, and Hypsogastropoda. Terrestrial Cyclophoroidea
and freshwater Ampullarioidea form the Architaenioglossa. On the other hand, the mostly marine Sorbeoconcha and Hypsogas-
tropoda represent a highly diverse groups uniting several tens of superfamilies of living gastropods (Figure 1).
The Heterobranchia encompasses the gastropod groups placed by Thiele’s classification into the ‘Opisthobranchia’ and ‘Pulmo-
nata’, as well as some ‘prosobranch’ groups, such as the Valvatoidea and Architectonicoidea. The Valvatoidea is an ancient group of
freshwater gastropods, but the highly diverse Architectonicoidea represents a marine group (Figure 1(h)). The majority of the

Figure 9 Two topologies of higher-level relationships among Gastropoda as derived from analyses of combined morphology and molecular data.
Based on Aktipis SW, Giribet G, Lindberg D and Ponder WF (2008) Gastropoda an overview and analysis, 201–237 In: Ponder W and Lindberg DL (eds),
Phylogeny and Evolution of the Mollusca, University of California Press, 488 pp., Berkeley and Los Angeles, California.
 Fossil Invertebrates: Gastropods 9

Heterobranchia are also marine gastropods, some with their shells reduced or absent. The lower Heterobranchia (Opisthobranchia)
uniting seven large clades (Cephalaspidea, Thecosomata, Gymnosomata, Aplysiomorpha, Sacoglossa, Umbraculida, and Nudipleura)
are extraordinarily variable. The higher Heterobranchia (Pulmonata) form a dominant group of terrestrial gastropods, but also occur in
freshwater environments. There are several classifications of the Pulmonata, which may be divided into two clades: Basommatophora,
and Eupulmonata. The ancient marine Basommatophora have been separated into the Archaeopulmonata and the freshwater
Basommatophora into the Brachiopulmonata. The Stylommatophora is a dominant group of terrestrial gastropods and is the most
numerous group in the Eupulmonata. The higher classification of extinct gastropods is less stable than that for living groups. The
Palaeozoic, exclusively marine Pelagiellida, Bellerophontida (Figures 2(l) and 2(m), 10 and 11), Macluritoidea, and Mimospirina are
amongst the most discussed extinct groups, and the gastropod nature of the Bellerophontida and Pelagiellida is still a frequently
discussed problem. The phylogenetic relationships of the Macluritoidea and Mimospirina (Figure 2(g)), with sinistrally coiled shells,
are uncertain and both groups may be sister groups to more advanced gastropods.

Evolution of the Gastropoda

The more than 500 million years of evolution of the Gastropoda is still poorly known. The main difficulties are that the
phylogenetic positions and relationships of extinct gastropods can be inferred only from the limited number of characters
observable in their fossilized hard body parts (i.e., shell and operculum). In addition, some belong to extinct higher taxa of family
or order levels with unknown anatomy. Another complication is the development of similar shells in unrelated groups (homo-
plastic similarity) which has been documented in many living gastropods. However, because a third of all gastropod families are
extinct, understanding of gastropod evolution and phylogeny necessarily involves study of both fossil and living species.

Origin and Early History of the Gastropoda

During last 50 years, many new mollusc-like fossils from the Cambrian strata have been discovered (e.g., Halkieria, Merismoconcha,
etc.). Their interpretation has given rise to different models of evolutionary relationships within the Mollusca. Even though these
models are controversial, the Gastropoda has been generally accepted to be the sister group of the classes Cephalopoda or Tryblidiida
(‘Monoplacophora’). The latter have been combined with the Scaphopoda and Bivalvia within the group Conchifera, which unites
the higher Mollusca. Whether or not the Conchifera is monophyletic is uncertain. Torsion of the soft body has been considered to be
one of the main diagnostic characters of the Gastropoda. For this reason, the majority of the models of gastropod origin have been
based on different interpretations of this anatomical feature in the extinct gastropod-like molluscs. The Early Palaeozoic Helcionel-
loida, Bellerophontida (Figures 2(l) and 2(m)), and Tryblidiida, with bilaterally symmetrical shells, as well as the Pelagiellida and
Macluritida, with asymmetrically coiled shells, are the most frequently discussed groups, and have been variously interpreted as
untorted or torted molluscs. However, there is no reliable method of recognizing torsion in extinct fossil molluscs. Thus, the unknown
nature of the bodies in the Early Palaeozoic gastropod-like fossils has enabled controversial speculations to be made about the origin
of the Gastropoda. Generally, it is accepted that the first undoubted gastropods appeared in the Late Cambrian.

Palaeozoic Era
During the Early Ordovician radiation, the diversity of gastropod groups which had appeared in the Late Cambrian
(Archaeogastropoda, Euomphaloidea, Macluritoidea, Mimospirina, Peruneloidea) rapidly increased (Figures 10 and 11). The Maclur-
itoidea with large shells, together with different groups of Archaeogastropoda, Euomphaloidea, Bellerophontida, and Mimospirina,
were typical elements of gastropod faunas of the tropical regions. In contrast, higher latitude faunas were composed mainly of the
Bellerophontida and Archaeogastropoda. This arrangement survived until the early Middle Ordovician, when the diversity of some
groups (Macluritoidea and Euomphaloidea) decreased and some new groups appeared (the slit-lacking Archaeogastropoda, Sub-
ulitoidea, Platyceratoidea, Loxonematoidea, etc.). During the Middle Ordovician, gastropod diversity rapidly increased and, in the Late
Ordovician, reached its maximum. Middle and Late Ordovician faunas consisted of members of all the main groups of Palaeozoic
gastropods, except the Heterobranchia (Figures 10 and 11). The end of the Ordovician saw a dramatic decrease in gastropod diversity,
as well as the extinction of the Macluritoidea. The Silurian was a period of increasing diversity of many gastropod groups (e.g.,
Archaeogastropoda, Bellerophontida, and Platyceratoidea), when some gastropods in all marine communities continually increased,
together with an increase in the morphological variability of their shells. This suggests an increase in their ecological adaptation to
specific environments. In comparison with the Ordovician, during the Silurian some gastropods with high-spired shells (mainly
Loxonematoidea, Murchisonoidea and Subulitoidea; Figure 2) increased considerably. The Devonian was a time of distinct changes in
marine gastropod communities. Some Ordovician– Silurian groups became extinct (Mimospirina; Figure 2(g)), new groups appeared
(Heterobranchia), and many groups underwent rapid radiation and specialization (Caenogastropoda and Neritimorpha). Thus, the
Devonian faunas contained representatives of all extant gastropod orders (Archaeogastropoda, Neritimorpha, Caenogastropoda, and
Heterobranchia), as well as many Palaeozoic groups (Figure 11). The Devonian was also the time when the protoconch morphology of
several gastropod groups underwent considerable change. Gastropods with openly coiled protoconchs (Perunelomorpha, Cyrtoner-
itimorpha, and Euomphalomorpha; Figures 2(h), 2(p), 3(d), and 3(e)) formed a considerable, sometimes even dominant, part of the
Ordovician and Silurian gastropod communities. During the Early Devonian, their numbers rapidly decreased and none survived the
10 Fossil Invertebrates: Gastropods

Figure 10 Diagram showing hypothetical phylogenies of the Palaeozoic gastropods derived mainly from the patterns of their early shell ontogenies (see
text for details). Modified from Frýda J (2012) Phylogeny of Paleozoic gastropods inferred from their ontogeny, 395–435. In: Earth and Life – Global
Biodiversity, Extinction Intervals and Biogeographic Perturbations Through Time, John A. Talent (Ed.), Series: International Year of Planet Earth, Springer,
1100pp.

Permian / Triassic extinction. Carboniferous and Permian faunas had a similar composition of marine gastropod communities. A
characteristic feature of Late Palaeozoic gastropod faunas was the fast radiation of different groups of Apogastropoda. The dominance
of diverse groups of Caenogastropoda with high-spired shells (mainly Ctenoglossa and Cerithiomorpha) and Heterobranchia
(Allogastropoda) was typical of shallow-water, muddy bottom communities.

Mesozoic and Cenozoic Eras

The Permian/Triassic crisis affected gastropods as well as all other marine animals. The Euomphalomorpha and Cyrtoneritimorpha
(Figures 2 and 3), as well as many groups of Archaeogastropoda, Neritimorpha, and Caenogastropoda, became extinct. During the
Triassic, the last members of the Bellerophontida disappeared. The Late Triassic was a time of fast radiation of neritimorphs
(Neritopsoidea and Neritoidea), caenogastropods (Ctenoglossa, Cerithimorpha, Architaenioglossa, and Littorinimorpha), and
heterobranch (Allogastropoda and Archaeopulmonata). From Triassic strata, the oldest limpets of the subclass Patellogastropoda
are documented. The Patellogastropoda is considered to represent the most ancient gastropod group, but their ancestors (probably
 Fossil Invertebrates: Gastropods 11

Figure 11 Diagram illustrating the evolution of the Gastropoda and the types of early shell ontogeny. The bars show the stratigraphical ranges of
each gastropod group. Based on Bandel K (1997) Higher classification and pattern of evolution of the Gastropoda. Courier Forschungsinstitut
Senckenberg 201: 57–81, Fryda J and Rohr DM (2004) Gastropoda, 184–195 In: Webby BD, Droser ML, Paris F, and Percival IG (eds.) The Great
Ordovician Biodiversification Event, pp. 408. New York: Columbia University Press, and Ponder W and Lindberg DL (eds), Phylogeny and Evolution
of the Mollusca, University of California Press, 488 pp., Berkeley and Los Angeles, California.

bearing coiled shells) have not yet been recognized amongst Palaeozoic gastropods (Figures 10 and 11). The composition of the
Jurassic and Early Cretaceous marine gastropod faunas was roughly the same as in the Late Triassic. The characteristic feature of
Mesozoic and Cenozoic gastropods was the development of more ornamented shells in most groups, as well as the lesser
occurrence of openly coiled shells, by comparison with Palaeozoic gastropods (Figures 1 and 2). Both macro-evolutionary trends
have been interpreted as adaptation to increasing predation activities by other animals. During the Cretaceous, more advanced
caenogastropod groups (higher Mesogastropoda and Neogastropoda) appeared (Figure 11), which underwent fast radiation and
diversification after the Cretaceous / Tertiary faunal crises. Both groups developed the possibility of extending their planktotrophic
larval stages and, from the beginning of the Tertiary, they formed one of the dominant groups of marine gastropods. During the
Cretaceous, some gastropods (lower Heterobranchia) started to reduce their shells, enabling their adaptation to holoplanktic life
(e.g., pteropods). The Early Cenozoic marine gastropod faunas are very similar to extant gastropods in higher taxonomic composition.

Evolution of Freshwater and Terrestrial Gastropods

In contrast with marine gastropods, the fossil record for freshwater and terrestrial forms is less complete, limiting our knowledge of
their evolution. Successful invasion to freshwater and land habitats has been closely linked with the mode of gastropod
reproduction. External fertilization, which occurs in the ancient Patellogastropoda and Archaeogastropoda, limited them to marine
environments. The freshwater and terrestrial environments were colonized by gastropods with egg capsules and internal fertiliza-
tion (Neritimorpha, Caenogastropoda, and Heterobranchia). Even though members of these groups are known from the Palaeo-
zoic (Figure 11), the first freshwater and terrestrial gastropods are recorded from Late Palaeozoic strata (Archaeopulmonata). The
first freshwater Basommatophora appeared during Jurassic time and, in the Cretaceous, the Stylommatophora started their
invasion of the land and soon became the most diversified group of terrestrial gastropods.

Further Reading
Aktipis SW, Giribet G, Lindberg DR, and Ponder WF (2008) Gastropoda – an overview and analysis. In: Ponder W and Lindberg DL (eds.) Phylogeny and evolution of the Mollusca,
pp. 201–237. California: University of California Press, 488 pp., Berkeley and Los Angeles.
Bandel K (1997) Higher classification and pattern of evolution of the gastropoda. Courier Forschungsinstitut Senckenberg 201: 57–81.
Beesley PL, Ross GJB, and Wells A (eds.) (1998) In: Mollusca: The Southern Synthesis, Part B, Fauna of Australia, Vol.5, pp. 565–1234. Melbourne: CSIRO Publishing.
Bieler R (1992) Gastropod phylogeny and systematics. Annual Review of Ecology and Systematics 23: 311–338.
12 Fossil Invertebrates: Gastropods

Fretter V and Graham A (1994) British prosobranch molluscs. Their functional anatomy and ecology. London: Ray Society.
Bouchet P, Rocroi JP, Frýda J, Hausdorf B, Ponder W, Valdes A, and Warén A (2005) Classification and nomenclator of gastropod families. Malacologia 47(1–2): 1–368.
Frýda J (2012) Phylogeny of Paleozoic gastropods inferred from their ontogeny. In: Talent John A (ed.) Earth and life – global biodiversity, extinction intervals and biogeographic
perturbations through time. International Year of Planet Earthpp. 395–435. Springer, 1100pp.
Frýda J, Nützel A, and Wagner PJ (2008) Paleozoic gastropods. In: Ponder W and Lindberg DL (eds.) Phylogeny and evolution of the Mollusca, pp. 239–270. California: University of
California Press, 488 pp., Berkeley and Los Angeles.
Frýda J and Rohr DM (2004) Gastropoda. In: Webby BD, Droser ML, Paris F, and Percival IG (eds.) The great ordovician biodiversification event, pp. 184–195. New York: Columbia
University Press, 408.
Geiger DL, Nützel A, and Sasaki T (2008) Vetigastropoda. In: Ponder W and Lindberg DL (eds.) Phylogeny and evolution of the Mollusca, pp. 297–330. California: University of
California Press, 488 pp., Berkeley and Los Angeles.
Knight JB, Cox LR, Keen AM, et al. (1960) Systematic descriptions. In: Moore RC (ed.) Treatise on invertebrate paleontology, part I, Mollusca 1, pp. I169–I324. Lawrence, KS:
Geological Society of America and University of Kansas Press.
Lindberg DR (2008) Patellogastropoda, neritimorpha, and cocculinoidea. In: Ponder W and Lindberg DL (eds.) Phylogeny and evolution of the Mollusca, pp. 271–296. California:
University of California Press, 488 pp., Berkeley and Los Angeles.
Lindberg DR and Ponder WF (2001) The influence of classification on the evolutionary interpretation of structure – a re-evaluation of the evolution of the pallial cavity of gastropod
molluscs. Organisms, Diversity and Evolution 1: 273–299.
Peel JS (1991) The classes Tergomya and Helcionelloida, and early molluscan evolution. Groenlands Geologiske Undersoegelse, Bulletin 161: 11–65.
Ponder WF, Colgan DJ, Healy JM, Nützel A, Simone LRL, and Strong EE (2008) Caenogastropoda. In: Ponder W and Lindberg DL (eds.) Phylogeny and Evolution of the Mollusca,
pp. 331–383. California: University of California Press, 488 pp., Berkeley and Los Angeles.
Ponder WF and Lindberg DR (1997) Towards a phylogeny of gastropod molluscs: An analysis using morphological characters. Zoological Journal of the Linnean Society
119: 83–256.
von Salvini-Plawen L (1990) Origin, phylogeny and classification of the phylum mollusca. Iberus 9: 1–33.
von Salvini-Plawen L and Haszprunar G (1987) The vetigastropoda and the systematics of streptonerous gastropoda (mollusca). Journal of Zoology 11: 747–770.
Taylor JD (ed.) (1996) Origin and evolutionary radiation of the Mollusca. Oxford: Oxford University Press, New York, Tokyo.
Wägele H, Klussmann-Kolb A, Vonnemann V, and Medina M (2008) Heterobranchia I – the opisthobranchia. In: Ponder W and Lindberg DL (eds.) Phylogeny and evolution of the
Mollusca, pp. 385–405. California: University of California Press, 488 pp., Berkeley and Los Angeles.
Waren A and Bouchet P (1993) New records, species, genera, and a new family of gastropods from hydrothermal vents and hydrocarbon seeps. Zoologica Scripta 22(1): 1–90.
Wenz W (1938–1944) Gastropoda. In: Schindewolf OH (ed.) Handbuch der Palä ozoologie, p 1639. Berlin: Borntraeger.
Zilch A (1960) Gastropoda: Teil 2, Euthyneura. In: Schindewolf OH (ed.) Handbuch der Paläozoologie, Berlin: Zehlendorf, p. 834.