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Terrestrial animals are
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s that live predominantly or entirely on
land Land, also known as dry land, ground, or earth, is the solid terrestrial surface of the planet Earth that is not submerged by the ocean or other bodies of water. It makes up 29% of Earth's surface and includes the continents and various isla ...
(e.g.
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s, dogs, ants,
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s), as compared with
aquatic animal An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic ...
s, which live predominantly or entirely in the water (e.g.
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
,
lobster Lobsters are a family (Nephropidae, synonym Homaridae) of marine crustaceans. They have long bodies with muscular tails and live in crevices or burrows on the sea floor. Three of their five pairs of legs have claws, including the first pair, ...
s,
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, ...
es), and
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s, which rely on a combination of aquatic and terrestrial
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s (e.g.
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s and
newt A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aqua ...
s). Some groups of
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
are terrestrial, such as ants,
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
, earwigs,
cockroaches Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known ...
,
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
s and many others, while other groups are partially aquatic, such as
mosquitoes Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small Diptera, flies within the family Culicidae (from the Latin ''culex'' meaning "gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish ...
and
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
, which pass their
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
l stages in water. Terrestrial animals tend to be more developed and intelligent than aquatic animals.


Terrestrial classes

The term "terrestrial" is typically applied to species that live primarily on the ground, in contrast to
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...
species, which live primarily in trees. There are other less common terms that apply to specific groups of terrestrial animals: *
Saxicolous A saxicolous lichen is a lichen that grows on rock. The prefix "sax" from the Latin means "rock" or "stone". Characteristics Saxicolous lichens exhibit very slow growth rates. They may develop on rock substrates for long periods of time, give ...
creatures are rock dwelling. "Saxicolous" is derived from the Latin word ''saxum'', meaning a rock. * Arenicolous creatures live in the sand. *
Troglofauna Troglofauna are small cave-dwelling animals that have adapted to their dark surroundings. Troglofauna and stygofauna are the two types of subterranean fauna (based on life-history). Both are associated with subterranean environments – troglofa ...
predominantly live in caves.


Taxonomy

Terrestrial invasion is one of the most important events in the
history of life The history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', for ''gigaannum'') and evide ...
. Terrestrial lineages evolved in several animal
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
, among which arthropods, vertebrates and mollusks are representatives of more successful groups of terrestrial animals. Terrestrial animals do not form a unified
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
; rather, they share only the fact that they live on land. The transition from an aquatic to terrestrial life by various groups of animals has occurred independently and successfully many times. Most terrestrial lineages originated under a mild or
tropical climate Tropical climate is the first of the five major climate groups in the Köppen climate classification identified with the letter A. Tropical climates are defined by a monthly average temperature of 18 °C (64.4 °F) or higher in the cool ...
during the
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...
and
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
, whereas few animals became fully terrestrial during the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
. If internal
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...
s are excluded, free living species in terrestrial environments are represented by the following eleven phyla: *
Gastrotrich The gastrotrichs (phylum Gastrotricha), commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, acoelomate animals, and are widely distributed and abundant in freshwater and marine environmen ...
s (hairy-backs) live in transient terrestrial water and go dormant during desiccation *
Rotifer The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John H ...
s (wheel animals) live in transient terrestrial water and go dormant during desiccation *
Nematodes The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
(roundworms) by going dormant during desiccation *
Eutardigrades Eutardigrada are a class of tardigrades (Tardigrada) without lateral appendices. Primarily freshwater bound, some species have secondarily gained the ability to live in marine environments ('' Halobiotus''). By cryptobiosis many species are able ...
(water bears) live in transient terrestrial water and go dormant during desiccation *
Flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegmen ...
s ( land planarians) require moist habitats and have restricted range * Nemerteans (ribbon worms in
Monostilifera Monostilifera is a suborder of nemertean worms belonging to the class Enopla Enopla is one of the classes of the worm phylum Nemertea, characterized by the presence of a peculiar armature of spines or plates in the proboscis. Evolution a ...
) require moist habitats and have restricted range *
Onychophora Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...
(velvet worms) require moist habitats and have restricted range, the only solely terrestrial phylum *
Annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s (
Clitellates The Clitellata are a class (biology), class of annelid worms, characterized by having a clitellum - the 'collar' that forms a reproductive cocoon during part of their life cycles. The clitellates comprise around 8,000 species. Unlike the class o ...
) require moist habitats, highly diverse and derived from their marine relatives *
Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s (fully terrestrial members: hexapods,
arachnids Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
, myriapods,
woodlice A woodlouse (plural woodlice) is an isopod crustacean from the polyphyleticThe current consensus is that Oniscidea is actually triphyletic suborder Oniscidea within the order Isopoda. They get their name from often being found in old wood ...
, sandhoppers, and
terrestrial crabs A number of Lineage (evolution), lineages of crabs have evolved to live predominantly on land. Examples of terrestrial crabs are found in the families Gecarcinidae and Gecarcinucidae, as well as in selected genera from other families, such as ' ...
; semi-terrestrial members include
Water Fleas The Diplostraca or Cladocera, commonly known as water fleas, are a superorder of small crustaceans that feed on microscopic chunks of organic matter (excluding some predatory forms). Over 1000 species have been recognised so far, with many more ...
,
Copepods Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (inhabiting sea waters), some are benthic (living on the ocean floor), a number of species have p ...
, and
Seed Shrimp Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typica ...
) *
Mollusks Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
(
Gastropods The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. The ...
:
land snail A land snail is any of the numerous species of snail that live on land, as opposed to the sea snails and freshwater snails. ''Land snail'' is the common name for terrestrial gastropod mollusks that have shells (those without shells are known ...
s and
slug Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a ...
s) *
Chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
s (
Tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...
; semiterrestrial members:
Amphibious fish Amphibious fish are fish that are able to leave water for extended periods of time. About 11 distantly related genera of fish are considered amphibious. This suggests that many fish genera independently evolved amphibious traits, a process known ...
) Roundworms, gastrotrichs, tardigrades, rotifers and some smaller species of arthropods and annelids are microscopic animals that require a film of water to live in, and are therefore considered semi-terrestrial.The Terrestrial Plankton , NZETC
/ref> Flatworms, ribbon worms, velvet worms and annelids all depend on more or less moist habitats. The three remaining phyla, arthropods, mollusks, and chordates, all contain species that have adapted totally to dry terrestrial environments, and which have no aquatic phase in their life cycles.


Difficulties

Labeling an animal species "terrestrial" or "aquatic" is often obscure and becomes a matter of judgment. Many animals considered terrestrial have a life-cycle that is partly dependent on being in water.
Penguins Penguins (order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adapt ...
,
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
, and
walruses The walrus (''Odobenus rosmarus'') is a large flippered marine mammal with a discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. The walrus is the only living species in the fami ...
sleep on land and feed in the ocean, yet they are all considered terrestrial. Many insects, e.g.
mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...
s, and all terrestrial crabs, as well as other clades, have an aquatic life cycle stage: their eggs need to be laid in and to hatch in water; after hatching, there is an early aquatic form, either a
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...
or
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
. There are crab species that are completely aquatic, crab species that are amphibious, and crab species that are terrestrial.
Fiddler crabs The fiddler crab or calling crab may be any of more than one hundred species of semiterrestrial marine crabs in the family Ocypodidae, well known for their sexually dimorphic claws; the males' major claw is much larger than the minor claw, while ...
are called "semi-terrestrial" since they make burrows in the muddy substrate, to which they retreat during high tides. When the tide is out, fiddler crabs search the beach for food. The same is true in the
mollusca Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is es ...
. Many hundreds of
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
genera and species live in intermediate situations, such as for example, ''Truncatella''. Some gastropods with gills live on land, and others with a lung live in the water. As well as the purely terrestrial and the purely aquatic animals, there are many borderline species. There are no universally accepted criteria for deciding how to label these species, thus some assignments are disputed.


Terrestrial panarthropods

Fossil evidence has shown that sea creatures, likely arthropods, first began to make forays onto land around 530 million years ago, in the Early
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
. There is little reason to believe, however, that animals first began living reliably on land around that time. A more likely hypothesis is that these early arthropods' motivation for venturing onto dry land was to mate (as modern horseshoe crabs do) or to lay eggs out of the reach of predators. Three groups of arthropods had independently adapted to land by the end of the Cambrian: myriapods, hexapods and
arachnids Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
. By the late
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
, they may have fully terrestrialized. There are other groups of arthropods, all from
malacostraca Malacostraca (from New Latin; ) is the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobs ...
n crustaceans, which independently became terrestrial at a later date:
woodlice A woodlouse (plural woodlice) is an isopod crustacean from the polyphyleticThe current consensus is that Oniscidea is actually triphyletic suborder Oniscidea within the order Isopoda. They get their name from often being found in old wood ...
, sandhoppers, and
terrestrial crabs A number of Lineage (evolution), lineages of crabs have evolved to live predominantly on land. Examples of terrestrial crabs are found in the families Gecarcinidae and Gecarcinucidae, as well as in selected genera from other families, such as ' ...
. Additionally, the sister
panarthropoda Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphyletic ...
n groups
Onychophora Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...
(velvet worms) are also terrestrial, while the Eutardigrada are also adapted for land to some degree; both groups probably becoming so during the Early Devonian. Among arthropods, many microscopic crustacean groups like
copepods Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (inhabiting sea waters), some are benthic (living on the ocean floor), a number of species have p ...
and
amphipods Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far describ ...
and
seed shrimp Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typica ...
can go dormant when dry and live in transient bodies of water.


Vertebrate terrestrialization

By approximately 375 million years ago the bony fish best adapted to life in shallow coastal/swampy waters (such as ''Tiktaalik roseae''). Thanks to relatively strong, muscular limbs (which were likely weight-bearing, thus making them a preferable alternative to traditional fins in extremely shallow water),Hohn-Schulte, Bianca, Holger Preuschoft, Ulrich Witzel, and Claudia Distler-Hoffman. "Biomechanics and Functional Preconditions for Terrestrial Lifestyle in Basal Tetrapods, with Special Consideration of ''Tiktaalik Roseae''." Historical Biology 25.2 (2013): 167-81. Web. and lungs which existed in conjunction with gills, ''Tiktaalik'' and animals like it were able to establish a strong foothold on land by the end of the Devonian period. In the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
, tetrapods (losing their gills) became fully terrestrialized, allowing their expansion into most terrestrial niches, though later on some will return back to being aquatic and conquer the air also.


Terrestrial gastropods

Gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
mollusks are one of the most successful animals that have diversified in the fully terrestrial habitat. They have evolved terrestrial taxa in more than nine lineages. They are commonly referred to as
land snail A land snail is any of the numerous species of snail that live on land, as opposed to the sea snails and freshwater snails. ''Land snail'' is the common name for terrestrial gastropod mollusks that have shells (those without shells are known ...
s and
slug Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a ...
s. Terrestrial invasion of gastropod mollusks has occurred in Neritopsina, Cyclophoroidea,
Littorinoidea Littorinoidea are a superfamily of both sea snails and land snails which have a gill and an operculum, terrestrial and marine gastropod mollusks in the clade Littorinimorpha. The terrestrial family within this group, the Pomatiidae, are some ...
, Rissooidea,
Ellobioidea Ellobiidae, common name the hollow-shelled snails, is a family of small air-breathing land snails, terrestrial pulmonate gastropod mollusks in the clade Eupulmonata.Bouchet, P. (2012). Ellobiidae. Accessed through: World Register of Marine Spec ...
, Onchidioidea,
Veronicelloidea Veronicelloidea is a superfamily of air-breathing land slugs. They are terrestrial pulmonate gastropod molluscs in the clade Systellommatophora The Systellommatophora (synonym Gymnomorpha) is a clade of primitive, air-breathing slugs, accord ...
,
Succineoidea Succineoidea is a taxonomic superfamily of air-breathing land snails, terrestrial pulmonate gastropod mollusks in the suborder Helicina ''Helicina'' is a genus of tropical and subtropical land snails with an operculum, terrestrial gastropo ...
, and
Stylommatophora Stylommatophora is an orderPhilippe Bouchet, Jean-Pierre Rocroi, Bernhard Hausdorf, Andrzej Kaim, Yasunori Kano, Alexander Nützel, Pavel Parkhaev, Michael Schrödl and Ellen E. Strong. 2017. Revised Classification, Nomenclator and Typification ...
, and in particular, each of Neritopsina, Rissooidea and Ellobioidea has likely achieved land invasion more than once. Most terrestrialization events have occurred during the
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...
or
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
. Gastropods are especially unique due to several fully terrestrial and epifaunal lineages that evolved during the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
. Some members of rissooidean families
Truncatellidae Truncatellidae, common name the "looping snails", is a family of small amphibious snails, with gills and an operculum, semi-marine gastropod mollusks or micromollusks. Shell description This family of snails have small shells which lose thei ...
,
Assimineidae Assimineidae is a family of minute snails, also known as palmleaf snails, with an operculum, gastropod mollusks or micromollusks in the superfamily Rissoidae. Many of these very small snails live in intermediate habitats, being amphibious betwee ...
, and
Pomatiopsidae Pomatiopsidae is a family of small, mainly freshwater snails, (some also occur in other habitats) that have gills and an operculum, aquatic gastropod mollusks in the superfamily Truncatelloidea (according to the taxonomy of the Gastropoda b ...
are considered to have colonized to land during the Cenozoic. Most truncatellid and assimineid snails amphibiously live in
intertidal The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...
and supratidal zones from
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
to pelagic areas. Terrestrial lineages likely evolved from such ancestors. The rissooidean gastropod family
Pomatiopsidae Pomatiopsidae is a family of small, mainly freshwater snails, (some also occur in other habitats) that have gills and an operculum, aquatic gastropod mollusks in the superfamily Truncatelloidea (according to the taxonomy of the Gastropoda b ...
is one of the few groups that have evolved fully terrestrial taxa during the late Cenozoic in the
Japanese Archipelago The Japanese archipelago (Japanese: 日本列島, ''Nihon rettō'') is a group of 6,852 islands that form the country of Japan, as well as the Russian island of Sakhalin. It extends over from the Sea of Okhotsk in the northeast to the East Chin ...
only. Shifts from aquatic to terrestrial life occurred at least twice within two Japanese endemic lineages in Japanese Pomatiopsidae and it started in the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The ...
. About one-third of gastropod species are terrestrial. In terrestrial habitats they are subjected to daily and seasonal variation in temperature and water availability. Their success in colonizing different habitats is due to physiological, behavioral, and morphological adaptations to water availability, as well as ionic and thermal balance. They are adapted to most of the habitats on Earth. The shell of a snail is constructed of calcium carbonate, but even in acidic soils one can find various species of shell-less slugs. Land-snails, such as Xerocrassa seetzeni and Sphincterochila boissieri, also live in deserts, where they must contend with heat and aridity. Terrestrial gastropods are primarily herbivores and only a few groups are carnivorous. Carnivorous gastropods usually feed on other gastropod species or on weak individuals of the same species; some feed on insect larvae or earthworms.


Semi-terrestrial animals

Semi-terrestrial animals are macroscopic animals that rely on very moist environments to thrive, they may be considered a transitional point between true terrestrial animals and aquatic animals. Among vertebrates, amphibians have this characteristic relying on a moist environment and breathing through their moist skin while reproducing in water. Many other animal groups solely have terrestrial animals that live like this: land planarians, land ribbon worms, roundworms (nematodes), and land annelids (clitellates) who are very primitive and breathe through skin.
Clitellates The Clitellata are a class (biology), class of annelid worms, characterized by having a clitellum - the 'collar' that forms a reproductive cocoon during part of their life cycles. The clitellates comprise around 8,000 species. Unlike the class o ...
or terrestrial annelids demonstrate many unique terrestrial adaptations especially in their methods of reproduction, they tend towards being simpler than their marine relatives, the bristleworms, lacking many of the complex appendages the latter have. Velvet worms are prone to desiccation not due to breathing through their skin but due to their spiracles being inefficient at protecting from desiccation, like clitellates they demonstrate extensive terrestrial adaptations and differences from their marine relatives including live birth.


Geoplankton

Many animals live in terrestrial environments by thriving in transient often microscopic bodies of water and moisture, these include rotifers and gastrotrichs which lay resilient eggs capable of surviving years in dry environments, and some of which can go dormant themselves. Nematodes are usually microscopic with this lifestyle. Although eutardigrades only have lifespans of a few months, they famously can enter suspended animation during dry or hostile conditions and survive for decades, which allows them to be ubiquitous in terrestrial environments despite needing water to grow and reproduce. Many microscopic crustacean groups like
copepods Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (inhabiting sea waters), some are benthic (living on the ocean floor), a number of species have p ...
and
amphipods Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far describ ...
and seed shrimps are known to go dormant when dry and live in transient bodies of water too.


See also

*
Aquatic animal An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic ...
*
Aquatic ecosystem An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms that are dependent on each other and on their environment. The t ...
*
Aquatic locomotion Aquatic locomotion or swimming is biologically propelled motion through a liquid medium. The simplest propulsive systems are composed of cilia and flagella. Swimming has evolved a number of times in a range of organisms including arthropods, ...
*
Aquatic mammal Aquatic and semiaquatic mammals are a diverse group of mammals that dwell partly or entirely in bodies of water. They include the various marine mammals who dwell in oceans, as well as various freshwater species, such as the European otter. They ...
*
Aquatic plant Aquatic plants are plants that have adapted to living in aquatic environments ( saltwater or freshwater). They are also referred to as hydrophytes or macrophytes to distinguish them from algae and other microphytes. A macrophyte is a plant that ...
* Marine invertebrates *
Marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their ...
* Terrestrial *
Terrestrial ecosystem Terrestrial ecosystems are ecosystems which are found on land. Examples include tundra, taiga, temperate deciduous forest, tropical rain forest, grassland, deserts. Terrestrial ecosystems differ from aquatic ecosystems by the predominant pres ...
*
Terrestrial locomotion Terrestrial locomotion has evolved as animals adapted from aquatic to terrestrial environments. Locomotion on land raises different problems than that in water, with reduced friction being replaced by the increased effects of gravity. As viewe ...
*
Terrestrial plant A terrestrial plant is a plant that grows on, in, or from land. Other types of plants are aquatic (living in water), epiphytic (living on trees) and lithophytic (living in or on rocks). The distinction between aquatic and terrestrial plants i ...
* *
Wetland indicator status Wetland indicator status denotes the probability of individual species of vascular plants occurring in freshwater, brackish and saltwater wetlands in the United States. The wetland status of 7,000 plants is determined upon information contained i ...
*
Zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, an ...


Further reading

* Clack J. A. (2002). ''Gaining ground: the origin and evolution of tetrapods''.
Indiana University Press Indiana University Press, also known as IU Press, is an academic publisher founded in 1950 at Indiana University that specializes in the humanities and social sciences. Its headquarters are located in Bloomington, Indiana. IU Press publishes 140 ...
, 369 pp., . * Cloudsley-Thompson J. L. (1988). ''Evolution and adaptation of terrestrial arthropods''.
Springer Springer or springers may refer to: Publishers * Springer Science+Business Media, aka Springer International Publishing, a worldwide publishing group founded in 1842 in Germany formerly known as Springer-Verlag. ** Springer Nature, a multinationa ...
, 141 pp., . * Dejours P. et al. (1987). ''Comparative physiology: life in water and on land''. Liviana Editrice, Italy, 556 pp., . * Gordon M. S. & Olson E. C. (1995). ''Invasions of the land: the transitions of organisms from aquatic to terrestrial life''.
Columbia University Press Columbia University Press is a university press based in New York City, and affiliated with Columbia University. It is currently directed by Jennifer Crewe (2014–present) and publishes titles in the humanities and sciences, including the fie ...
, 312 pp., . * Little C. (1983). ''The colonisation of land: Origins and adaptations of terrestrial animals''.
Cambridge University Press Cambridge University Press is the university press of the University of Cambridge. Granted letters patent by King Henry VIII in 1534, it is the oldest university press in the world. It is also the King's Printer. Cambridge University Pr ...
, Cambridge. 290 pp., . * Little C. (1990). ''The terrestrial invasion. An ecophysiological approach to the origin of land animals''. Cambridge University Press, Cambridge. 304 pp. . *


References

This article incorporates CC-BY-2.0 text from the referenceKameda Y. & Kato M. (2011). "Terrestrial invasion of pomatiopsid gastropods in the heavy-snow region of the Japanese Archipelago". ''
BMC Evolutionary Biology ''BMC Ecology and Evolution'' (since January 2021), previously ''BMC Evolutionary Biology'' (2001–2020), is a peer-reviewed open access scientific journal covering all fields of evolutionary biology, including phylogenetics and palaeontology ...
'' 11: 118. .
and CC-BY-2.5 text from the referenceRaz S., Schwartz N. P., Mienis H. K., Nevo E. & Graham J. H. (2012). "Fluctuating Helical Asymmetry and Morphology of Snails (Gastropoda) in Divergent Microhabitats at ‘Evolution Canyons I and II,’ Israel". '' PLoS ONE'' 7(7): e41840. . and CC-BY-3.0 text from the reference {{DEFAULTSORT:Terrestrial Animal Zoology Terrestrial