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Terrestrial animals are
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s that live predominantly or entirely on land (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, spiders), as compared with aquatic animals, which live predominantly or entirely in the water (e.g. fish,
lobster Lobsters are a family (biology), family (Nephropidae, Synonym (taxonomy), 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 ...
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, cuttle ...
es), and amphibians, which rely on a combination of aquatic and terrestrial habitats (e.g.
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" '' Triadobatrachus'' is ...
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 ...
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 t ...
, earwigs, cockroaches, grasshoppers and many others, while other groups are partially aquatic, such as mosquitoes and dragonflies, 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. Th ...
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 species, which live primarily in trees. There are other less common terms that apply to specific groups of terrestrial animals: * Saxicolous creatures are rock dwelling. "Saxicolous" is derived from the Latin word ''saxum'', meaning a rock. * Arenicolous creatures live in the sand. * Troglofauna predominantly live in caves.


Taxonomy

Terrestrial invasion is one of the most important events in the history of life. 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 Phyl ...
, among which arthropods, vertebrates and mollusks are representatives of more successful groups of terrestrial animals. Terrestrial animals do not form a unified clade; 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 during the Paleozoic 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 Creta ...
, whereas few animals became fully terrestrial during the Cenozoic. If internal
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
s are excluded, free living species in terrestrial environments are represented by the following eleven phyla: * Gastrotrichs (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 (roundworms) by going dormant during desiccation * Eutardigrades (water bears) live in transient terrestrial water and go dormant during desiccation * Flatworms ( 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, a class of worms characterized by the presence of a peculiar armature of spines or plates in their proboscis. Families * Acteonemertidae * Amphiporidae * Carcinoneme ...
) require moist habitats and have restricted range * Onychophora (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 Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s (fully terrestrial members: hexapods, arachnids, myriapods, woodlice, sandhoppers, and terrestrial crabs; 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, and Seed Shrimp) *
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:
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 as ...
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 sm ...
s) * Chordates ( Tetrapods; semiterrestrial members: Amphibious fish) 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, seals, and walruses sleep on land and feed in the ocean, yet they are all considered terrestrial. Many insects, e.g. mosquitos, 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 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. Th ...
. There are crab species that are completely aquatic, crab species that are amphibious, and crab species that are terrestrial. Fiddler crabs 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 est ...
. Many hundreds of gastropod 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 ag ...
. 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. By the late Ordovician, 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, sandhoppers, and terrestrial crabs. Additionally, the sister panarthropodan groups Onychophora (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 and amphipods and seed shrimp 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, 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 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 as ...
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 sm ...
s. Terrestrial invasion of gastropod mollusks has occurred in Neritopsina, Cyclophoroidea, Littorinoidea,
Rissooidea Rissooidea, originally named Rissoacea by Gray, 1847, is a taxonomic superfamily of small and minute marine snails, belonging to the clade Littorinimorpha.Gofas, S. (2013). Rissooidea. Accessed through: World Register of Marine Species at http ...
, Ellobioidea, Onchidioidea,
Veronicelloidea Veronicelloidea is a superfamily of air-breathing land slugs. They are terrestrial pulmonate gastropod molluscs in the clade Systellommatophora. Taxonomy The following two families were recognized in the taxonomy of Bouchet & Rocroi (20 ...
,
Succineoidea Succineoidea is a taxonomic superfamily of air-breathing land snails, terrestrial pulmonate gastropod mollusks in the suborder Helicina. MolluscaBase eds. (2021). MolluscaBase. Succineoidea Beck, 1837. Accessed through: World Register of Marine ...
, 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 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 Creta ...
. Gastropods are especially unique due to several fully terrestrial and epifaunal lineages that evolved during the Cenozoic. 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 ...
,
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 are considered to have colonized to land during the Cenozoic. Most truncatellid and assimineid snails amphibiously live in intertidal and supratidal zones from brackish water to pelagic areas. Terrestrial lineages likely evolved from such ancestors. The
rissooidea Rissooidea, originally named Rissoacea by Gray, 1847, is a taxonomic superfamily of small and minute marine snails, belonging to the clade Littorinimorpha.Gofas, S. (2013). Rissooidea. Accessed through: World Register of Marine Species at http ...
n gastropod family Pomatiopsidae is one of the few groups that have evolved fully terrestrial taxa during the late Cenozoic in the Japanese Archipelago 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 (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
. 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 ''Xerocrassa seetzeni'' is a species of air-breathing land snail, a terrestrial animal, terrestrial pulmonate gastropod mollusk in the family Geomitridae. MolluscaBase eds. (2022). MolluscaBase. Xerocrassa seetzeni (L. Pfeiffer, 1847). Accessed t ...
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 and amphipods and seed shrimps are known to go dormant when dry and live in transient bodies of water too.


See also

* Aquatic animal * Aquatic ecosystem * Aquatic locomotion * Aquatic mammal * Aquatic plant * Marine invertebrates * Marine mammal * Terrestrial * Terrestrial ecosystem * Terrestrial locomotion * Terrestrial plant * * Wetland indicator status * Zoology


Further reading

* Clack J. A. (2002). ''Gaining ground: the origin and evolution of tetrapods''. Indiana University Press, 369 pp., . * Cloudsley-Thompson J. L. (1988). ''Evolution and adaptation of terrestrial arthropods''. Springer, 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 fi ...
, 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 Henry VIII of England, King Henry VIII in 1534, it is the oldest university press in the world. It is also the King's Printer. Cambr ...
, 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'' 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