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Gastropoda
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Gastropods (/ˈɡæstrəpɒdz/), commonly known as slugs

Gastropoda
and snails, belong to a large taxonomic class of
Temporal range:
invertebrates within the phylum Mollusca called
Gastropoda (/ɡæsˈtrɒpədə/).[5]

This class comprises snails and slugs from saltwater,

freshwater, and from the land. There are many thousands of
species of sea snails and slugs, as well as freshwater snails,
freshwater limpets, land snails and slugs.

The class Gastropoda is a diverse and highly successful class

of mollusks within the phylum Mollusca. It contains a vast
total of named species, second only to the insects in overall Various gastropods from different types:
number. The fossil history of this class goes back to the Late Black slug (a slug), Haliotis asinina (an
Cambrian. As of 2017, 721 families of gastropods are known, abalone), Cornu aspersum (a land snail),
of which 245 are extinct and appear only in the fossil record, Notarchus indicus (a seahare), Patella
while 476 are currently extant with or without a fossil
vulgata (a limpet), and Polycera
record.[6]
aurantiomarginata (a nudibranch).
Gastropoda (previously known as univalves and sometimes Scientific classification
spelled "Gasteropoda") are a major part of the phylum
Domain: Eukaryota
Mollusca, and are the most highly diversified class in the
phylum, with 65,000 to 80,000[3][4] living snail and slug Kingdom: Animalia
species. The anatomy, behavior, feeding, and reproductive Phylum: Mollusca
adaptations of gastropods vary significantly from one clade
Class: Gastropoda
or group to another, so stating many generalities for all
Cuvier, 1795[2]
gastropods is difficult.
Subclasses
The class Gastropoda has an extraordinary diversification of
habitats. Representatives live in gardens, woodland, deserts, Caenogastropoda
and on mountains; in small ditches, great rivers, and lakes;
Heterobranchia
in estuaries, mudflats, the rocky intertidal, the sandy
subtidal, the abyssal depths of the oceans, including the Neomphaliones
hydrothermal vents, and numerous other ecological niches, Neritimorpha
including parasitic ones.
Patellogastropoda
Although the name "snail" can be, and often is, applied to all Vetigastropoda
the members of this class, commonly this word means only
Diversity[3][4]
those species with an external shell big enough that the soft
65,000 to 80,000 species

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Gastropods are found in a wide range of aquatic and terrestrial

habitats, from deep ocean trenches to deserts.

Some of the more familiar and better-known gastropods are

terrestrial gastropods (the land snails and slugs). Some live in
fresh water, but most named species of gastropods live in a marine
environment.

Gastropods have a worldwide distribution, from the near Arctic

and Antarctic zones to the tropics. They have become adapted to
Cepaea nemoralis: a European
almost every kind of existence on earth, having colonized nearly pulmonate land snail, which has
every available medium. been introduced to many other
countries
In habitats where not enough calcium carbonate is available to
build a really solid shell, such as on some acidic soils on land,
various species of slugs occur, and also some snails with thin, translucent shells, mostly or entirely
composed of the protein conchiolin.

Snails such as Sphincterochila boissieri and Xerocrassa seetzeni have adapted to desert conditions.
Other snails have adapted to an existence in ditches, near deepwater hydrothermal vents, in oceanic
trenches 10,000 meters below the surface,[17] the pounding surf of rocky shores, caves, and many
other diverse areas.

Gastropods can be accidentally transferred from one habitat to another by other animals, e.g. by
birds.[18]

Anatomy
Snails are distinguished by an anatomical process known as
torsion, where the visceral mass of the animal rotates 180° to one
side during development, such that the anus is situated more or
less above the head. This process is unrelated to the coiling of the
shell, which is a separate phenomenon. Torsion is present in all
gastropods, but the opisthobranch gastropods are secondarily
The anatomy of a common air-
untorted to various degrees.[19][20]
breathing land snail: much of this
anatomy does not apply to
Torsion occurs in two stages. The first, mechanistic stage is
gastropods in other clades or
muscular, and the second is mutagenetic. The effects of torsion are
groups.
primarily physiological. The organism develops by asymmetrical
growth, with the majority of growth occurring on the left side. This
leads to the loss of right-side anatomy that in most bilaterians is a duplicate of the left side anatomy.
The essential feature of this asymmetry is that the anus generally lies to one side of the median plane.
The gill-combs, the olfactory organs, the foot slime-gland, nephridia, and the auricle of the heart are
single or at least are more developed on one side of the body than the other. Furthermore, there is
only one genital orifice, which lies on the same side of the body as the anus.[21] Furthermore, the anus
becomes redirected to the same space as the head. This is speculated to have some evolutionary

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The nervous system of gastropods includes the peripheral nervous

system and the central nervous system. The central nervous
system consists of ganglia connected by nerve cells. It includes
paired ganglia: the cerebral ganglia, pedal ganglia, osphradial
ganglia, pleural ganglia, parietal ganglia and the visceral ganglia.
There are sometimes also buccal ganglia.[23]

Digestive system
The radula of a gastropod is usually adapted to the food that a The upper pair of tentacles on the
species eats. The simplest gastropods are the limpets and head of Helix pomatia have eye
abalones, herbivores that use their hard radula to rasp at seaweeds spots, but the main sensory organs
of the snail are sensory receptors
on rocks.
for olfaction, situated in the
epithelium of the tentacles.
Many marine gastropods are burrowers, and have a siphon that
extends out from the mantle edge. Sometimes the shell has a
siphonal canal to accommodate this structure. A siphon enables the animal to draw water into their
mantle cavity and over the gill. They use the siphon primarily to "taste" the water to detect prey from a
distance. Gastropods with siphons tend to be either predators or scavengers.

Respiratory system
Almost all marine gastropods breathe with a gill, but many freshwater species, and the majority of
terrestrial species, have a pallial lung. The respiratory protein in almost all gastropods is hemocyanin,
but one freshwater pulmonate family, the Planorbidae, have hemoglobin as the respiratory protein.

In one large group of sea slugs, the gills are arranged as a rosette of feathery plumes on their backs,
which gives rise to their other name, nudibranchs. Some nudibranchs have smooth or warty backs
with no visible gill mechanism, such that respiration may likely take place directly through the skin.

Circulatory system
Gastropods have open circulatory system and the transport fluid is hemolymph. Hemocyanin is
present in the hemolymph as the respiratory pigment.

Excretory system
The primary organs of excretion in gastropods are nephridia, which produce either ammonia or uric
acid as a waste product. The nephridium also plays an important role in maintaining water balance in
freshwater and terrestrial species. Additional organs of excretion, at least in some species, include
pericardial glands in the body cavity, and digestive glands opening into the stomach.

Reproductive system
Courtship is a part of mating behavior in some gastropods, including some of the Helicidae. Again, in
some land snails, an unusual feature of the reproductive system of gastropods is the presence and
utilization of love darts.
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Gastropods exhibit an important degree of variation in mitochondrial gene organization when

compared to other animals.[26] Main events of gene rearrangement occurred at the origin of
Patellogastropoda and Heterobranchia, whereas fewer changes occurred between the ancestors of
Vetigastropoda (only tRNAs D, C and N) and Caenogastropoda (a large single inversion, and
translocations of the tRNAs D and N).[26] Within Heterobranchia, gene order seems relatively
conserved, and gene rearrangements are mostly related with transposition of tRNA genes.[26]

Geological history and evolution

The first gastropods were exclusively marine, with the earliest
representatives of the group appearing in the Late Cambrian
(Chippewaella, Strepsodiscus),[27] though their only gastropod
character is a coiled shell, so they could lie in the stem lineage, if
they are gastropods at all.[28] Earliest Cambrian organisms like
Helcionella, Barskovia and Scenella are no longer considered
gastropods, and the tiny coiled Aldanella of earliest Cambrian
time is probably not even a mollusk. Trochonema sp., an early gastropod
from the Middle Ordovician of the
As such, it's not until the Ordovician that the first crown-group Galena Group of Minnesota.
members arise.[29] By the Ordovician period the gastropods were a
varied group present in a range of aquatic habitats. Commonly,
fossil gastropods from the rocks of the early Palaeozoic era are too
poorly preserved for accurate identification. Still, the Silurian
genus Poleumita contains fifteen identified species. Fossil
gastropods were less common during the Palaeozoic era than
bivalves.[29]

Most of the gastropods of the Palaeozoic era belong to primitive

groups, a few of which still survive. By the Carboniferous period
many of the shapes seen in living gastropods can be matched in Fossil gastropod and attached
mytilid bivalves on a Jurassic
the fossil record, but despite these similarities in appearance the
limestone bedding plane of the
majority of these older forms are not directly related to living
Matmor Formation in southern
forms. It was during the Mesozoic era that the ancestors of many Israel.
of the living gastropods evolved.[29]

One of the earliest known terrestrial (land-dwelling) gastropods is Anthracopupa (=Maturipupa),[30]

which is found in the Coal Measures of the Carboniferous period in Europe, but relatives of the
modern land snails are rare before the Cretaceous period.[29]

In rocks of the Mesozoic era, gastropods are slightly more common as fossils; their shells are often
well preserved. Their fossils occur in ancient beds deposited in both freshwater and marine
environments. The "Purbeck Marble" of the Jurassic period and the "Sussex Marble" of the early
Cretaceous period, which both occur in southern England, are limestones containing the tightly
packed remains of the pond snail Viviparus.[29]

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⁠

Neritimorpha

⁠Vetigastropoda
⁠

⁠Patellogastropoda ►
⁠

Cocculiniformia, Neomphalina and Lower Heterobranchia are not included in the above cladogram.

Taxonomy

Current classification
The present backbone classification of gastropods relies on the results of phylogenomic analyses.
Consensus has not been reached yet considering the relationships at the very base of the gastropod
tree of life, but otherwise the major groups are known with confidence.[32][33][34]

Gastropoda

Adenogonogastropoda[33] (Angiogastropoda[32])
Apogastropoda
Caenogastropoda
Heterobranchia
Neritimorpha
Patellogastropoda
Vetigastropoda (including Neomphaliones)

History
Since Darwin, biological taxonomy has attempted to reflect the
phylogeny of organisms, i.e., the tree of life. The classifications
used in taxonomy attempt to represent the precise
interrelatedness of the various taxa. However, the taxonomy of the
Gastropoda is constantly being revised and so the versions shown
in various texts can differ in major ways.

In the older classification of the gastropods, there were four

A group of fossil shells of Turritella
subclasses:[35]
cingulifera from the Pliocene of
Cyprus
Opisthobranchia (gills to the right and behind the heart).

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Many gastropod species face threats from habitat destruction, pollution, and climate change. Some
species are endangered or have become extinct due to these factors. Conservation efforts often focus
on protecting their habitats, especially in freshwater and terrestrial ecosystems.

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External links
Gastropod reproductive behavior (http://www.scholarpedia.org/article/Gastropod_reproductive_be
havior)
2004 Linnean taxonomy of gastropods (http://www.manandmollusc.net/advanced_introduction/gas
tropod_taxonomy_1.html)
Webster, S.; Fiorito, G. (2001). "Socially guided behaviour in non-insect invertebrates". Animal
Cognition. 4 (2): 69. doi:10.1007/s100710100108 (https://doi.org/10.1007%2Fs100710100108).
S2CID 25373798 (https://api.semanticscholar.org/CorpusID:25373798). – Article about social
learning also in gastropods.
Gastropod photo gallery (https://www.flickr.com/photos/jsjgeology/sets/72157647577987145),
mostly fossils, a few modern shells
A video of a crawling Garden Snail (https://www.youtube.com/watch?v=LKD5DZ4f6z0) (Cornu
aspersum), YouTube
Grove, S.J. (2018). A Guide to the Seashells and other Marine Molluscs of Tasmania: Molluscs of
Tasmania with images (https://molluscsoftasmania.org.au/)

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