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Gastropods

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378 FOSSIL INVERTEBRATES/Gastropods
exploiting the more specialist boring and cementing
habit has continued to grow and there has been a
further taxonomic explosion of the shallow burrowers
at lower taxonomic levels.
See Also
Biological Radiations and Speciation. Fossil Inverte-
brates: Molluscs Overview; Gastropods; Cephalopods
(Other Than Ammonites); Ammonites. Mesozoic: End
Cretaceous Extinctions. Palaeoecology.
Further Reading
Beesley PL, Ross GJB, and Wells A (1998) Mollusca: the
Southern Synthesis, vol. A. Collingwood, Australia:
CSIRO Publishing.
Harper EM (1998) The fossil record of bivalve molluscs. In:
Donovan SK and Paul CRC (eds.) The Adequacy of the Fossil
Record, pp. 243–267. Chichester: John Wiley and Sons.
Harper EM and Skelton PW (1993) The Mesozoic Marine
Revolution and epifaunal bivalves. Scripta Geologica,
Special Issue 2: 127–153.
Gastropods
J Frýda, Czech Geological Survey, Prague,
Czech Republic
ß 2005, Elsevier Ltd. All Rights Reserved.
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 gas-
tropods occur mostly in shallow-water benthic com-
munities; however, some gastropod species have
also lived in the deep sea (e.g., faunas associated
with hydrothermal vents), and others, such as holo-
planktic animals, have spent their whole lives as free-
swimming gastropods. The terrestrial gastropods
colonized most land environments, ranging from low-
lands 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 morpho-
logical and functional diversity of their bodies and
Harper EM, Taylor JD, and Crame JA (2000) The Evolu-
tionary Biology of the Bivalvia. Geological Society of
London Special Publication 177. London: Geological
Society of London.
Johnston PA and Haggart JW (1998) Bivalves: an Eon of
Evolution. Paleobiological Studies Honoring Norman
D. Newell. Calgary: Calgary University Press.
Skelton PW, Crame JA, Morris NJ, and Harper EM (1990)
Adaptive divergence and taxonomic radiation in post-
Palaeozoic bivalves. In: Taylor PD and Larwood GP
(eds.) Major Evolutionary Radiations. The Systematics
Association Special Volume 42, pp. 91–117. Oxford:
Clarendon Press.
Stanley SM (1970) Relation of shell form to life habits of
the Bivalvia. Geological Society of America Memoir 125:
1–296.
Taylor JD (1996) Origin and Evolutionary Radiation of the
Mollusca. Oxford: Oxford University Press.
Vermeij GJ (1987) Evolution and Escalation. An Ecological
History of Life. Princeton, NJ: Princeton University
Press.
Vermeij GJ (1993) A Natural History of Shells. Princeton,
NJ: Princeton University Press.
shells. The gastropods comprise one of the most di-
verse 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 biogeograph-
ical 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 diver-
sity of gastropod bodies is also reflected in their anat-
omy. Generally, the body consists of a large foot, a
 FOSSIL INVERTEBRATES/Gastropods 379

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 dif-
ferent organs. The circulatory system comprises a
contractile heart with one (Caenogastropoda and
Heterobranchia) or two (Patellogastropoda, Archae-
ogastropoda, 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 gas-
tropods, 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
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 Archae-
ogastropoda) 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 develop-
ment is characterized by a spiral cleavage, which
differs slightly in the main gastropod groups. The
subsequent larval stage is called the trochophore

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: Chicor-
eus 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) Heterobran-
chia; (l) Sorbeoconcha (Caenogastropoda); (M) Vetigastropoda (Archaeogastropoda).
380 FOSSIL INVERTEBRATES/Gastropods
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 gastro-
pods. The last larval stage is termed veliger, which
typically bears two ciliate paddles (velum), sometimes
subdivided into several lobes. If free-swimming gas-
tropod larvae use planktic organisms for their nutri-
tion, their development is termed planktotrophic.
Marine gastropods with such development have
small eggs, but numbering over half a million. Plank-
totrophic larvae may stay planktic for several months
and thus can be carried for long distances by oceanic
currents. The gastropods, however, developed an-
other 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) typic-
ally 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 simpli-
fied 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 gas-
tropod 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 embry-
onic shell (protoconch I), which is followed by a
teleoconch (Figures 3B and 3F–3H). On the other
hand, the protoconchs of more advanced gastropods
(Neritimorpha, Caenogastropoda, and Heterobran-
chia) 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 3K and 3L),
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 op-
posite direction to the teleoconchs (Figures 3J and 4).
Such shells are termed heterostrophic. The Neritimor-
pha form typical, strongly convolute protoconchs
during planktotrophic development, which are
homeostrophic (Figures 3A, 3N, and 4). Higher gas-
tropods 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 opercu-
lum is mostly horny (corneous) and may be tightly
(multispiral) or loosely (paucispiral) coiled or concen-
tric. Some gastropod groups have calcareous oper-
cula, and the oldest operculum known is from the
Ordovician (Macluritoidea).
Shell Structure
Most gastropod shells are composed of an outer or-
ganic layer (periostracum) and an inner, mostly much
thicker, calcified layer. The colour pattern typical of
many gastropod shells (Figure 1) is formed by differ-
ent organic pigments which are limited to the perios-
tracum 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 com-
plex shell structure. On the other hand, the majority
of the higher gastropods have developed simple
 FOSSIL INVERTEBRATES/Gastropods 381

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 symmet-
rical 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: Vosko-
piella barborae (Onychochilidae). (H) Naticiform shell: Eifelcyrtus blodgetti (Vltavielidae). (I) Fusiform shell: Havlicekiela parvula (Perune-
loidea). High-spired shells: (J) Pragozyga costata; (K) Palaeozyga bohemica (Loxonematoidea). (L, M) Bilaterally symmetrical shell:
Bellerophon vasulites (Bellerophontoidea). (N) Limpet: Pragoscutula wareni (Pragoscutulidae). (P) 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) Mimos-
pirina; (H) Cyrtoneritimorpha; (L, M, Q) Bellerophontida; (I) Perunelomorpha; (E, J, K, N) Order uncertain; (P) Euomphalomorpha.

aragonitic shells with a crossed lamellar structure
(Figures 6A and 6B). Some structural types are re-
stricted to certain groups (e.g., nacre) and this may be
used for their identification in fossils (Figure 6C).
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 (Figures 1M, 2C, 2L, and 2M). The limpet-
shaped shells were independently developed within
all main gastropod groups from the asymmetrically
coiled shells of their ancestors. In contrast, planispi-
rally coiled shells are known only in several groups,
such as the Palaeozoic Porcellioidea and Bellerophon-
toidea (Figures 2L and 2M) or the Holocene Planor-
bioidea. Some gastropods may change the coiling
382 FOSSIL INVERTEBRATES/Gastropods

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 Euom-
phalus 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 (Caenogas-
tropoda and Neritimorpha) is termed homeostrophic. If the handedness of the shells is opposite (Heterobranchia), the coiling is termed
heterostrophic.

of their shells (Figure 4) from sinistral to dextral
(dextral heterostrophy), or vice versa (sinistral het-
erostrophy), 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 3J), or later (e.g.,
Porcellioidea; Figure 3M).
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 de-
veloped a spiral operculum. The spiral operculum of
anatomically dextral gastropods is coiled counter-
clockwise (viewed externally), and vice versa in sinis-
tral gastropods. Thus, there are four possible
relationships between shell coiling and body asym-
metry 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 major-
ity 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).
Figure 5 Apical (A) and lateral (B) views of the Middle Devon-
ian (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).
FOSSIL INVERTEBRATES/Gastropods 383
Muscle Scars
Gastropod shells are attached to the soft body by
muscles, which may leave distinct scars on the inner
shell surface. The geometry of muscle scars has fre-
quently been used as a diagnostic feature for distinc-
tion 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 inde-
pendently. Thus, the muscle scar pattern sometimes
observable in the fossil molluscan shells may be a
good ecological indicator, but has no systematic
significance.
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 (see Famous
Geologists: Cuvier). Since then, scientists have tried
to classify them by using different features of their
bodies. However, the classification of such a numer-
ous 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 Gastro-
poda 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
384 FOSSIL INVERTEBRATES/Gastropods
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).
Thiele, integrated earlier classifications and divided
the gastropods into three subclasses: Prosobranchia,
Opisthobranchia, and Pulmonata. In addition, the
Prosobranchia were divided into three orders:
Archaeogastropoda, Mesogastropoda, and Neogas-
tropoda. 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 fea-
tures. The evaluation of this newly gathered data in
the light of the existing classification revealed a need
for its revision. Recent analyses of numerous morpho-
logical and developmental characters of living gas-
tropods have resulted in a new classification scheme
(Figure 8), which has 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) represents the sister
group to all other living gastropods. Living patellogas-
tropods with limpet-shaped shells are exclusively
marine and occur mostly on rocky shores in all con-
tinents. The Patellogastropoda and their coiled ances-
tors have been united into the subclass Eogastropoda.
All other living gastropods and their ancestors have
been placed in the subclass Orthogastropoda, com-
prising four main groups of living gastropods:
Neritimorpha, Archaeogastropoda, Caenogastro-
poda, and Heterobranchia (Figure 8).
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 Cyrtoneritimorpha,
with openly coiled early shells (Figures 2H and
3D), may represent a closely related group. The
living Archaeogastropoda unites the Vetigastropoda
(Figures 3B and 3C) and several smaller groups, such
as the Neomphaloidea, which occur in faunas asso-
ciated with deep-sea hydrothermal vents. The Archae-
ogastropoda have colonized almost all marine and
estuarine environments. There are also a number of
extinct, mainly Palaeozoic groups (Figure 2) with un-
certain relationships to living archaeogastropods. The
Palaeozoic Euomphaloidea (¼ Euomphalomorpha;
Figures 2P and 3E), 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 Apogas-
tropoda. Both groups are highly diverse and have
colonized almost all marine, freshwater, and terres-
trial environments. The Palaeozoic Subulitoidea and
Peruneloidea (Perunelomorpha) (Figures 2I and 3L)
may be ancestral or basal groups of the Caenogastro-
poda or of all Apogastropoda. The extant Caenogas-
tropoda unites the two orders Architaenioglossa
and Sorbeoconcha. Terrestrial Cyclophoroidea and
freshwater Ampullarioidea form the Architaenio-
glossa. On the other hand, the mostly marine Sor-
beoconcha represents a highly diverse group uniting

Figure 8 Recent classification scheme of living gastropods, illustrating their phylogenetic relationships and the distribution of
freshwater and terrestrial groups. Based on 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.
more than 25 superfamilies of living gastropods
(Figure 1).
The Heterobranchia encompasses the gastropod
groups placed by Thiele’s classification into the
‘Opisthobranchia’ and ‘Pulmonata’, as well as some
‘prosobranch’ groups, such as the Valvatoidea and
Architectonicoidea. The Valvatoidea is an ancient
group of freshwater gastropods, but the highly di-
verse Architectonicoidea represents a marine group
(Figure 1H). The majority of the lower Hetero-
branchia (Opisthobranchia or sea slugs) are also
marine gastropods, typically with their shells re-
duced or absent. They are extraordinarily variable
and are divided into about 30 superfamilies of nine
orders. 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 three orders: Systellommatophora,
Basommatophora, and Eupulmonata. The ancient
marine Basommatophora have been separated into
the Archaeopulmonata and the freshwater Basomma-
tophora into the Brachiopulmonata. The Stylommato-
phora 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 2L and 2M), Macluritoidea, and Mimospir-
ina are amongst the most discussed extinct groups,
and the gastropod nature of the Bellerophontida and
Pelagiellida is still a frequently discussed problem.
FOSSIL INVERTEBRATES/Gastropods 385
The phylogenetic relationships of the Macluritoidea
and Mimospirina (Figure 2G), 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 difficul-
ties are that the phylogenetic positions and relation-
ships 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).
However, the number of extinct gastropod species and
genera is much higher than those living. 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 (homoplastic similarity) which has been
documented in many living gastropods.
Origin and Early History of the Gastropoda
Since 1970, many new mollusc-like fossils from the
Cambrian have been discovered (e.g., Halkieria, Mer-
ismoconcha, 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 Cephalo-
poda (see Fossil Invertebrates: Cephalopods (Other
Than Ammonites)) or Tryblidiida (‘Monoplaco-
phora’). The latter have been combined with the
386 FOSSIL INVERTEBRATES/Gastropods

Scaphopoda and Bivalvia (see Fossil Invertebrates:
Bivalves) 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 Gastro-
poda. For this reason, the majority of the models of
gastropod origin have been based on different inter-
pretations of this anatomical feature in the extinct
gastropod-like molluscs. The Early Palaeozoic Hel-
cionelloida, Bellerophontida (Figures 2L and 2M),
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 en-
abled 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 (Figure 9). The Macluritoidea with large
shells, together with different groups of Archaeogas-
tropoda, Euomphaloidea, Bellerophontida, and
Mimospirina, were typical elements of gastropod
faunas of the tropical regions. In contrast, higher
latitude faunas were composed mainly of the Beller-
ophontida and Archaeogastropoda. This arrange-
ment 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, Subu-
litoidea, 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
(Figure 9). The end of the Ordovician saw a dra-
matic 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 in-
creased, together with an increase in the morpho-
logical variability of their shells. This suggests an
increase in their ecological adaptation to specific

Figure 9 Diagram illustrating the evolution of the Gastropoda and the types of early shell ontogeny. The bars show the stratigraph-
ical 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, and Frýda 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.

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
2G), new groups appeared (Heterobranchia), and
many groups underwent rapid radiation and special-
ization (Caenogastropoda and Neritimorpha). Thus,
the Devonian faunas contained representatives of all
extant gastropod orders (Archaeogastropoda, Neriti-
morpha, Caenogastropoda, and Heterobranchia), as
well as many Palaeozoic groups (Figure 9). The Dev-
onian was also the time when the protoconch morph-
ology of several gastropod groups underwent
considerable change. Gastropods with openly coiled
protoconchs (Perunelomorpha, Cyrtoneritimorpha,
and Euomphalomorpha; Figures 2H, 2P, 3D, and
3E) formed a considerable, sometimes even domin-
ant, part of the Ordovician and Silurian gastropod
communities. During the Early Devonian, their
numbers rapidly decreased and none survived the
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 Heterobran-
chia (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 Euomphalomor-
pha and Cyrtoneritimorpha (Figures 2 and 3), as well
as many groups of Archaeogastropoda, Neritimor-
pha, 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 Neri-
toidea), caenogastropods (Ctenoglossa, Cerithimor-
pha, Architaenioglossa, and Littorinimorpha), and
heterobrachs (Allogastropoda and Archaeopulmo-
nata). 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 bearing coiled shells) have not yet been
recognized amongst Palaeozoic gastropods (Figure 9).
The composition of the Jurassic and Early Cret-
aceous marine gastropod faunas was roughly the
FOSSIL INVERTEBRATES/Gastropods 387
same as in the Late Triassic. The characteristic feature
of Mesozoic and Cenozoic gastropods was the devel-
opment 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 ac-
tivities by other animals. During the Cretaceous, more
advanced caenogastropod groups (higher Mesogas-
tropoda and Neogastropoda) appeared (Figure 9),
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 holo-
planktic 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 repro-
duction. External fertilization, which occurs in the
ancient Patellogastropoda and Archaeogastropoda,
limited them to marine environments. The freshwater
and terrestrial environments were colonized by gas-
tropods with egg capsules and internal fertilization
(Neritimorpha, Caenogastropoda, and Heterobran-
chia). Even though members of these groups are
known from the Early (Neritimorpha and Caenogas-
tropoda) or Middle (Heterobranchia) Palaeozoic
(Figure 9), the first freshwater and terrestrial gastro-
pods are recorded from Late Palaeozoic strata
(Archaeopulmonata). The first freshwater Basomma-
tophora appeared during Jurassic time and, in the
Cretaceous, the Stylommatophora started their inva-
sion of the land and soon became the most diversified
group of terrestrial gastropods.
Glossary
Archaeogastropoda Group of extant gastropods.
Bellerophontida Extinct group of Palaeozoic mol-
luscs with bilaterally symmetrical shells.
Caenogastropoda Group of extant gastropods.
embryonic shell (protoconch I) Gastropod shell
formed during embryonic development.
 388 FOSSIL INVERTEBRATES/Gastropods
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 dex-
tral 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 Hetero-
branchia.
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.
operculum Lid-like structure used for closing of the
aperture in gastropod shells.
Opisthobranchia Gastropod subclass of Thiele’s
classification.
orthostrophy Condition in which anatomically dex-
tral (or sinistral) animals occupy dextrally (or sin-
istrally) 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 classi-
fication.
protoconch Gastropod shell formed during larval
and/or embryonic development.
Pulmonata Gastropod subclass of Thiele’s classifica-
tion.
teleoconch Post-larval gastropod shell.
trochophore larva Gastropod larva formed during
early larval development which may be free swim-
ming or occurs in egg capsules.
veliger Gastropod larva formed during later larval
development before metamorphosis to post-larval
stages.
See Also
Biological Radiations and Speciation. Evolution.
Famous Geologists: Cuvier. Fossil Invertebrates:
View publication stats
Molluscs Overview; Bivalves; Cephalopods (Other Than
Ammonites). Palaeoecology. Palaeozoic: End Permian
Extinctions.
Further Reading
Bandel K (1997) Higher classification and pattern of evolu-
tion of the Gastropoda. Courier Forschungsinstitut 201:
57–81.
Beesley PL, Ross GJB, and Wells A (eds.) (1998) 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.
Fretter V and Graham A (1994) British Prosobranch Mol-
luscs. Their Functional Anatomy and Ecology. London:
Ray Society.
Frýda J and Rohr DM (2004) Gastropoda, 184–195. In:
Webby BD, Droser ML, Paris F, and Percival IG (eds.)
The Great Ordovician Biodiversification Event, p. 408.
New York: Columbia University Press.
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. Law-
rence, KS: Geological Society of America and University
of Kansas Press.
Lindberg DR and Ponder WF (2001) The influence of clas-
sification 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 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 classi-
fication of the phylum Mollusca. Iberus 9: 1–33.
von Salvini-Plawen L and Haszprunar G (1987) The Veti-
gastropoda and the systematics of streptonerous Gastro-
poda (Mollusca). Journal of Zoology 11: 747–770.
Taylor JD (ed.) (1996) Origin and Evolutionary Radiation
of the Mollusca. Oxford, New York, Tokyo: Oxford
University Press.
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 Paläozoologie, p. 1639. Berlin: Borntraeger.
Zilch A (1959–1960) Gastropoda; Teil 2, Euthyneura. In:
Schindewolf OH (ed.) Handbuch der Paläozoologie,
p. 834. Berlin: Zehlendorf.